Vocalizations of bearded seals (Erignathus barbatus) and their influence on the soundscape of the western Canadian Arctic

The soundscape is a crucial habitat feature for marine mammals. This study investigates the contribution of bearded seal vocalizations to the soundscape in the western Canadian Arctic, and also the vocal characteristics of bearded seals relative to sea ice conditions. Passive acoustic data were recorded near Sachs Harbour between August 2015 and July 2016. Sound pressure levels (SPL) in the 50–1,000 Hz and 1–10 kHz bands increased as the total duration of all bearded seal vocalizations increased, and this relationship was moderated by sea ice concentration. Bearded seals in this region had an overlapping vocal repertoire with bearded seals in other areas of the Arctic, and had seven additional vocalizations that have not been previously documented for this region. This study is the first detailed assessment of the influence of bearded seal calls on SPL, which shows the high potential of bearded seals to influence underwater sound levels during the mating season. Bearded seals live in a changing Arctic seascape, and their influence on the soundscape may shift as sea ice continues to diminish. It is imperative that acoustic monitoring continues within the Arctic, and this study provides a baseline for future monitoring as the Arctic continues to change.     

Observational Evidences of Vocal Learning in Southern Elephant Seals: a Longitudinal Study

Vocal learning in mammals is sparsely documented, and there are few reports of vocal learning by wild mammals. In particular, no information based on longitudinal data for identified individuals exists, even for well‐studied highly social species in which vocal communication is an important aspect of social life. We present such information for the southern elephant seal (Mirounga leonina). We studied a small breeding population in the Falkland Islands over 8 yr (1995–2002). We recorded approx. 2400 agonistic vocalizations from approx. 285 different males, including many recorded over more than one breeding season (55 males were recorded for >2 yr and 29 were recorded throughout vocal development). Vocalizations are a key element of male agonistic behaviour in this species, the world's most polygynous vertebrate. For male agonistic vocalizations, we show that: (1) a limited number of discrete vocal types exists; (2) proportions of vocal types change over time; (3) observed trends of increase or decrease of vocal types are well explained by a process of vocal learning, in which younger peripheral males imitate vocal types of older more successful breeders.     

CLASSIFICATION OF DIVERSE CALL TYPES USING CLUSTER ANALYSIS TECHNIQUES

ABSTRACT

We investigated the problem of categorizing the repertoire of a group of highly varied vocalizations. A set of Weddell seal Leptonychotes weddellim-air calls recorded near Davis, Antarctica, were examined. The repertoire size was estimated by first subjectively assigning each call (based on auditory and spectrographic patterns) to one of a large number of provisional call types. A set of frequency, duration and waveform measurements were made on every call. For each provisional call type, the mean value of each of these measures was calculated. These mean values were used to perform Cluster Analyses of the provisional call types. Beginning with calls clustered closest together, the most similar provisional call types were successively amalgamated until further joining would link two clearly dissimilar vocalizations (rising versus falling frequency sweeps in this case). The Weddell seal repertoire contained 12 call types ranging from long sinusoidal upsweeps to growls. This procedure provides a method of estimating the minimum repertoire size of a sample of calls. Dichotomous Sorting using Principal Components Analysis can also be used to categorize calls but will probably be of greater value when applied to finding subdivisions within a single call type.     

Examining harp seal behavioural patterns via their underwater vocalizations

Recordings of harp seal (Pagophilus groenlandicus) underwater vocalizations, obtained at 3-h intervals over 7 days during the breeding season, were examined with regard to the relative occurrence of high frequency calls. An increase of high frequency calls at mid-March is possibly associated with the onset of courtship and mating. A daily pattern indicated that the seals are vocally active all night and least active at daybreak and in the early afternoon. The inactive afternoon period coincided with the time that the greatest number of seals were resting on the ice. Remote and/or automated monitoring of vocalizations may provide a means of examining various underwater activities of marine mammals.     

Polar bear responses to the underwater vocalizations of ringed seals

Summary Two captive polar bears were exposed to six different marine mammal vocalizations, in an attempt to determine if polar bears would selectively respond to the underwater vocalization of their primary prey, ringed seals. Vocalizations were played to each bear for a total of eight minutes while their behavioral responses were observed. An analysis of activity indicated that the bears displayed a significantly stronger response to ringed seals' vocalizations. It is theorized that the ability to recognize the underwater vocalizations of their primary prey would increase a bear's probability of a successful capture when a ringed seal rises to breath at an ice bound breathing hole.     

Underwater calling rates of harp and Weddell seals as a function of hydrophone location

Stereo recordings of harp seal (Pagophilus groenlandicus) and Weddell seal (Leptonychotes weddellii) underwater vocalizations were made near breeding groups using separations between two hydrophones of 159–339 m. Within a large herd, harp seal call numbers varied slightly between channels. Counts of Weddell seal calls were higher near a small herd on the ice than 159–180 m away. Repeat counts of harp seal calls by a single observer differed significantly. Source levels of Weddell seal calls varied and higher amplitude calls would be detected up to 2 orders of magnitude farther away than quieter calls. Hydrophone location (especially near small groups), observer variability and call source level differences will bias the use of monitoring underwater seal vocalizations to index locations, population size or underwater behaviours. Using hydrophone arrays and multiple observers may mitigate these problems. Accepted: 15 September 2000     

Embryonic vocalizations mediate parental care in Masked Lapwings (Vanellus miles) but not Red-capped Plovers (Charadrius ruficapillus)

Embryonic vocalizations in birds may enable communication between embryos and incubating parents but responses of parents to embryonic vocalizations have not been measured experimentally. In two ground-nesting shorebirds, we compare parental care of eggs without embryonic vocalizations, those with natural embryonic vocalizations, and those in which we experimentally introduced embryonic vocalizations, while accounting for prevailing temperatures. We continuously recorded nests (one block of up to 24 h per nest) of two ground-nesting shorebirds, the Red-capped Plover Charadrius ruficapillus and Masked Lapwing Vanellus miles and quantified parental behaviours. We predict an increase in egg thermoregulation behaviours (time spent incubating or shading) and ‘fussing’ (a combination of rising and re-settling, shuffling, orientation change and egg turning) when vocalizations were naturally present or when they were experimentally introduced in comparison to when no vocalizations were present (control). In Lapwings, the rate of fussing increased at the nest only when natural vocalizations were present; however, preening increased when natural or introduced vocalizations were present. Additionally, the frequency of bill to egg touches (inspections) increased only in the presence of introduced vocalizations. For Plovers, treatment had no influence on parental behavioural; however, the frequency of fussing, bill to egg touches and shell tossing increased with air temperature. For both species, there was no influence of the presence of the embryo’s vocalizations (natural or introduced) on the parent’s egg thermoregulation behaviours. Our results highlight that acoustic communication across the eggshell may occur and can influence parental care in shorebirds, but this appears to be species-specific.     

Killing of Cape fur seal (Arctocephalus pusillus pusillus) pups by brown hyenas (Parahyaena brunnea) at mainland breeding colonies along the coastal Namib Desert

Brown hyenas (Parahyaena brunnea) scavenge and kill seal pups at mainland Cape fur seal (Arctocephalus pusillus pusillus) colonies. The prey encounter interval and interval between kills depended on seal density, and increased density resulted in an increase of the capture rate and increase in hunting efficiency from 14% in November to 47% in January. The time brown hyenas spent at the seal colony decreased with increasing seal density and increasing air temperatures. Nevertheless, they were regularly active during the day when less adult seals were present at the colony, which indicates that the attendance of adult seals might play a role in the choice of foraging time. Brown hyenas killed seal pups throughout the study period. The predation rate was independent of the availability of non-violent mortalities, but the absolute number of kills was positively density-dependent. Mass kill events were recorded throughout the study period and are therefore not unusual occurrences. The overabundance of easy and vulnerable prey may lead to an over stimulus situation that triggers killing independent of the consumption of the prey or the hunger state.     

BEHAVIORAL SIGNIFICANCE OF UNDERWATER VOCALIZATIONS OF CAPTIVE LEOPARD SEALS, HYDURGA LEPTONYX

Two groups of underwater vocalizations were identified in a three-year study of two captive leopard seals, Hydurga leptonyx (one female and one male at Taronga Zoo, Sydney). This was supplemented by recordings over three months from a male at Marineland, New Zealand. The sexual state of the seals at Taronga was deduced from serum hormonal concentrations: the female was considered to be in estrus at specific times during the breeding season. The seal at Marineland, New Zealand was assumed to be sexually mature on the basis of size and age. Of 12 different underwater sound types recorded, six were produced by the seals at Taronga Zoo during agonistic interactions (local calls) and were heard through most of the year. The other six sound types were produced by lone seals. These broadcast calls were produced by the female only when sexually receptive, and by the mature male during December and January, months believed to be the breeding season of wild leopard seals. We propose that underwater acoustic behavior is important in the mating system of this species, and that broadcast calls are used by mature females to advertise their sexual receptivity, and possibly by mature males in search of mates.     

Geographical variation in temporal and spatial vocalization patterns of male harbour seals in the mating season

In the aquatically mating harbour seal, Phoca vitulina, oestrous females show marked differences in spatial and temporal distribution between geographical areas. This suggests that the males' display behaviour may also vary between areas. We recorded male vocalizations in two areas, the Moray Firth and Orkney, U.K. In the Moray Firth, females haul out on a few intertidal sandbars and travel along predictable routes to forage at sea. In Orkney, female haul out sites are much less influenced by tidal availability and females are much more dispersed. In the Moray Firth, males vocalized only during a short mating season, from 1 July to 12 August. Vocalizations varied significantly with the tide, the peak at high tide clearly coinciding with the period when most females were in the water. In contrast, vocalizations in Orkney were significantly related to both tidal and diel patterns. We suggest that the timing of male vocalizations reflects differences in female availability between sites. In the inner Moray Firth, vocalizations were heard throughout the females' range, whereas vocalizations in Orkney were heard only in two discrete areas. However, at both sites the density of vocalizing males was highest in narrow channels and/or along predictable female travel routes. Therefore, males clearly adapt their temporal and spatial behaviour patterns to variations in female distribution and density. These results suggest that male mating strategies in aquatically mating pinnipeds are more variable than was previously envisaged. Copyright 1999 The Association for the Study of Animal Behaviour.     

Vocalizations and Behaviour of Pacific Humpback Dolphins Sousa chinensis

Very little is known about the acoustic repertoire of the Pacific humpback dolphin Sousa chinensis . This study, off eastern Australia, used concurrent observations of surface behaviour and acoustic recordings to gain an insight into the behavioural significance of humpback dolphin vocalizations. Humpback dolphins exhibit five different vocalization categories: broad band clicks; barks; quacks; grunts; and whistles. Broad band clicks were high in frequency (8 kHz to > 22 kHz), were directly related to foraging behaviour and may play a role in social behaviour. Barks and quacks were burst pulse sounds (frequency: 0.6 kHz to > 22 kHz, duration: 0.1–8 s) and were associated with both foraging and social behaviour. The grunt vocalization is a low frequency narrow band sound (frequency 0.5–2.6 kHz, duration 0.06–2 s) and was only heard during socializing. There were 17 different types of whistles, ranging widely in frequency (0.9–22 kHz) and vocal structure (n=329). The predominant whistle types used by the groups were type 1 (46%) and type 2 (17%). Most whistles were heard during both socializing and foraging. The number of whistles recorded in a group increased significantly as the number of mother–calf pairs increased, suggesting that whistles may be used as contact calls. Few vocalizations were heard during either travelling or milling behaviours. Broad band clicks, barks and whistle type 1 were the only vocalizations recorded during either travelling or milling.     

A demonstration of the ability of the harbour seal PhocaVitulina (L.) to discriminate among pup vocalizations

A captive adult harbour seal demonstrated that it could distinguish between vocalizations recorded from different pups. This supports the implication of an earlier study which showed that the pup's call is individually characteristic and might therefore provide one means by which the mother could recognize her offspring and monitor its location. This remote monitoring would be especially important when the pair are in the water where the risk of separation is high.     

Long-term variability in the abundance of Antarctic fur seals Arctocephalus gazella at Signy Island, South Orkneys

The number of Antarctic fur seals Arctocephalus gazella hauled out at Signy Island in the South Orkneys was monitored annually between 1977 and 2008. Over the study period seal abundance showed a tenfold increase, from a minimum of 1,643 seals in 1978 to a maximum of 21,303 in 1994. The majority of individuals observed were young adult males, likely to be migrants from South Georgia, with small numbers of female seals and only 65 pups recorded during the survey period. Variability in counts showed a similar pattern to Laurie Island, also in the South Orkneys archipelago, suggesting a similar annual immigration of seals to these two islands. The date of first seal arrival was correlated with the date of fast-ice breakout at Factory Cove, Signy Island, and years in which break out was exceptionally late (>21 December) corresponded with years of reduced seal abundance. While the presence of fast-ice during the early breeding season may currently inhibit the establishment of a major breeding population of fur seals at Signy Island, it is important that routine monitoring should continue, particularly in the light of current patterns of climate warming in the Antarctic.     

Effects of reproductive state on vocalizations in squirrel monkeys (Saimiri sciureus)

Vocalizations and behavior of a group of 6 squirrel monkeys, 2 males and 4 females, were recorded during the nonbreeding and breeding seasons. Behavioral and physical criteria were used to determine the presence of estrus. During the breeding season the types of vocalizations uttered by estrous females changed, and the adult male increased his rate of vocalizing. Err vocalizations by estrous females were associated with increased following and initiation of affiliative behavior with the adult male, and may have functioned to facilitate these interactions. Errs appeared to be related to changes in female reproductive state rather than to the behavior of others. The adult male increased vocalizations associated with sexual and aggressive behavior (squeals and cackles), primarily in response to the estrous females' persistent initiation of interactions with him. We concluded that certain vocalizations in Saimiri reflected changes in the reproductive state of males and females, and functioned to mediate changes in social bonding during the breeding season.     

The effect of a large Danish offshore wind farm on harbor and gray seal haul-out behavior

This study investigates the effects of the construction and operation of a large Danish offshore wind farm on harbor and gray seal haul-out behavior within a nearby (4 km) seal sanctuary. Time-lapse photography, visual monitoring, and aerial surveys were used to monitor the number of seals on land in daylight hours. Seals were monitored during two preconstruction periods (19 June–31 August 2001 and April–August 2002), a construction period of the wind farm (August 2002–December 2003), and a period of operation of the wind farm (December 2003–December 2004). Monthly aerial surveys were conducted to estimate the proportion of seals in the sanctuary relative to neighboring haul-out sites. From preconstruction to construction and through the first year of operation the number of harbor seals in the sanctuary increased at the same rate as the number of seals at the neighboring haul-out sites. No long-term effects on haul-out behavior were found due to construction and operation of the wind farm. However, a significant short-term decrease was seen in the number of seals present on land during sheet pile driving in or near the wind farm. Acoustic deterrents were utilized simultaneously to avoid hearing damage.     

Variations in underwater vocalizations of Weddell seals (Leptonychotes weddelli) at the Vestfold Hills as a measure of breeding population discreteness

Weddell seal vocalizations from Davis Station showed similarities to those from McMurdo Sound and Palmer Peninsula, but none were identical. One vocalization, DD1, was unique to Davis Station. At all sites trills, or territorial defense calls, were the most common and had more types than other calls. Chugs, an aggressive sound, were common at all sites. Weddell seal calls from Davis Station showed similarities to sounds from McMurdo by sharing the use of prefixes and suffixes. Sounds from Davis Station shared the use of both ascending and descending trills and whistles with calls from Palmer Peninsula.Commonalities in underwater vocalizations may indicate the degree of mixing between breeding populations, in which case, Weddell seals in the Davis area probably are from a breeding population distinct from those at either McMurdo Sound or Palmer Peninsula.     

Individual variation in pup vocalizations and absence of behavioral signs of maternal vocal recognition in Weddell seals (Leptonychotes weddellii)

Individually stereotyped vocalizations often play an important role in relocation of offspring in gregarious breeders. In phocids, mothers often alternate between foraging at sea and attending their pup. Pup calls are individually distinctive in various phocid species. However, experimental evidence for maternal recognition is rare. In this study, we recorded Weddell seal (Leptonychotes weddellii) pup vocalizations at two whelping patches in Atka Bay, Antarctica, and explored individual vocal variation based on eight vocal parameters. Overall, 58% of calls were correctly classified according to individual. For males (n= 12) and females (n= 9), respectively, nine and seven individuals were correctly identified based on vocal parameters. To investigate whether mothers respond differently to calls of familiar vs. unfamiliar pups, we conducted playback experiments with 21 mothers. Maternal responses did not differ between playbacks of own, familiar, and unfamiliar pup calls. We suggest that Weddell seal pup calls may need to contain only a critical amount of individually distinct information because mothers and pups use a combination of sensory modalities for identification. However, it cannot be excluded that pup developmental factors and differing environmental factors between colonies affect pup acoustic behavior and the role of acoustic cues in the relocation process.     

Assessing the cross platform performance of marine mammal indicators between two collocated acoustic recorders

Equipment and deployment strategies for remote passive acoustic sensing of marine environments must balance memory capacity, power requirements, sampling rate, duty-cycle, deployment duration, instrument size, and environmental concerns. The impact of different parameters on the data and applicability of the data to the specific questions being asked should be considered before deployment. Here we explore the effect of recording and detection parameters on marine mammal acoustic data across two platforms. Daily classifications of marine mammal vocalizations from two passive acoustic monitors with different subsampling parameters, an AURAL and a Passive Aquatic Listener (PAL), collocated in the Bering Sea were compared. The AURAL subsampled on a pre-set schedule, whereas the PAL sampled via an adaptive protocol. Detected signals of interest were manually classified in each dataset independently. The daily classification rates of vocalizations were similar. Detections from the higher duty-cycle but lower sample rate AURAL were limited to species and vocalizations with energy below 4 kHz precluding detection of echolocation signals. Temporal coverage from the PAL audio files was limited by the adaptive sub-sampling protocol. A method for classifying ribbon (Histriophoca fasciata) and bearded seal (Erignathus barbatus) vocalizations from the sparse spectral time histories of the PAL was developed. Although application of the acoustic entropy as a rapid assessment of biodiversity was not reflective of the number of species detected, acoustic entropy was robust to changes in sample rate and window length.     

Behaviour Associated with Ultrasonic Vocalizations in Six Gerbilline Rodent Species

Encounters were staged between male and oestrus-induced females of the same or different taxa of the species/subspecies Gerbillurus paeba paeba, G. p. exilis, G. tytonis, G. setzeri, G. vallinus and Tatera brantsii. Each encounter lasted 20 min; converted ultrasonic emissions were recorded on videotape simultaneously with visual images. Videotapes were transcribed and sequences of acts and vocalizations were recorded. The frequency of acts was compared separately with the number of ultrasonic vocalizations during and after acts by χ² contingency table analysis, followed by partitioned χ² to identify individual acts which contributed significantly to the overall χ² value. Vocalizations occurred significantly more frequently than expected during sexual and/or huddling behaviour in all species-groups; and significantly less frequently than expected during exploratory, watching and/or submissive behaviour. Vocalizations were uncommon during investigative behaviour in all taxa, and aggressive behaviour in Gerbillurus taxa, but occurred significantly more frequently than expected after these behaviour categories in all taxa. Analysis of vocalizations immediately following acts verified the interpretations of function of vocalizations during acts. Ultrasonic vocalizations clearly play an important role in the communication of six taxa of southern African gerbils.     

Serotonin modulates responses to species-specific vocalizations in the inferior colliculus

Neuromodulators such as serotonin are capable of altering the neural processing of stimuli across many sensory modalities. In the inferior colliculus, a major midbrain auditory gateway, serotonin alters the way that individual neurons respond to simple tone bursts and linear frequency modulated sweeps. The effects of serotonin are complex, and vary among neurons. How serotonin transforms the responses to spectrotemporally complex sounds of the type normally heard in natural settings has been poorly examined. To explore this issue further, the effects of iontophoretically applied serotonin on the responses of individual inferior colliculus neurons to a variety of recorded species-specific vocalizations were examined. These experiments were performed in the Mexican free-tailed bat, a species that uses a rich repertoire of vocalizations for the purposes of communication as well as echolocation. Serotonin frequently changed the number of recorded calls that were capable of evoking a response from individual neurons, sometimes increasing (15% of serotonin-responsive neurons), but usually decreasing (62% of serotonin-responsive neurons), this number. A functional consequence of these serotonin-evoked changes would be to change the population response to species-specific vocalizations.