Memory for own behaviour in pinnipeds.

Pinnipeds are aquatic predators feeding on a vast range of prey, and their social behaviour differs greatly between species (from extreme polygyny in some sea lions to monogamy in some true seals). It has been hypothesised that the foraging and social complexity of their lifestyle should drive the evolution of their cognitive abilities. To investigate how aware pinnipeds are of their own behaviour, a grey seal (Halichoerus grypus), two harbour seals (Phoca vitulina) and four South American sea lions (Otaria flavescens) were trained to repeat their own behaviour on command. Three already trained behaviours were used, and the animal was asked to repeat the behaviour twice to ensure that the animal recalled its own behaviour and not the command given for the previous behaviour. All three species could recall their own behaviour significantly better than by chance. The duration for which the animals could recall their behaviour was tested using a staircase paradigm. A delay was implemented between the completion of the behaviour and the command to repeat it. The delay was increased after correct responses and decreased after incorrect responses. The performance of all species fell towards chance level after 12-18 s, with no significant difference between species. These results indicate that sea lions and true seals are aware of their own behaviour and that true seals have similar short-term memory abilities. It also shows that pinnipeds have less developed short-term memory abilities compared to other aquatic predators, such as the bottlenose dolphin. The complexity of pinniped foraging and social behaviour does not seem to have driven the evolution of short-term memory abilities in these animals but might have contributed to their ability to recall their own behaviour.

Does Enrichment Improve Well Being in Animals under Human Care? A Case Study of Two Harbor Seals (Phoca Vitulina).

Harbor seals in the wild live in a stimulating environment; therefore, nonhuman-animal caretakers have increasingly been using environmental enrichment to improve the well being of seals under human care. The purpose of this study was to evaluate an object-based environmental enrichment program during a four-month period on stimulating exploration and play and improving conspecific social interactions and human-animal relationships (HAR). Zoo staff conducted the environmental enrichment program as part of the animal care program. Seals were given objects haphazardly and were observed for 20 minutes, and seals' responsiveness during training sessions before and after enrichment was assessed. Seals showed interest in objects throughout the study and interacted more times per session with objects during the later months. Seals showed preferences for objects that were suspended in the water column (e.g., rope). Seals did not show more affiliative behavior but did show some aggressive behavior during enrichment sessions in comparison with free-swimming sessions. One seal showed better responsiveness to trainers in training sessions that followed an enrichment session than in other trainings sessions. Overall, the enrichment program was successful in increasing intrinsically motivated behaviors and showed that object-based enrichment has the potential to improve HAR between seals and their trainers.

Effect of pile-driving sounds on harbor seal (Phoca vitulina) hearing.

Seals exposed to intense sounds may suffer hearing loss. After exposure to playbacks of broadband pile-driving sounds, the temporary hearing threshold shift (TTS) of two harbor seals was quantified at 4 and 8 kHz (frequencies of the highest TTS) with a psychoacoustic technique. The pile-driving sounds had: a 127 ms pulse duration, 2760 strikes per h, a 1.3 s inter-pulse interval, a ∼9.5% duty cycle, and an average received single-strike unweighted sound exposure level (SELss) of 151 dB re 1 µPa2s. Exposure durations were 180 and 360 min [cumulative sound exposure level (SELcum): 190 and 193 dB re 1 µPa2s]. Control sessions were conducted under low ambient noise. TTS only occurred after 360 min exposures (mean TTS: seal 02, 1-4 min after sound stopped: 3.9 dB at 4 kHz and 2.4 dB at 8 kHz; seal 01, 12-16 min after sound stopped: 2.8 dB at 4 kHz and 2.6 dB at 8 kHz). Hearing recovered within 60 min post-exposure. The TTSs were small, due to the small amount of sound energy to which the seals were exposed. Biological TTS onset SELcum for the pile-driving sounds used in this study is around 192 dB re 1 µPa2s (for mean received SELss of 151 dB re 1 µPa and a duty cycle of ∼9.5%).

Classification of underwater vocalizations of wild spotted seals (Phoca largha) in Liaodong Bay, China.

Underwater vocalizations were recorded and classified from wild spotted seals (Phoca largha) in Liaodong Bay, China. The spotted seals exhibited an extensive underwater vocal repertoire but with limited complexity. Four major call types, representing 77.8% of all calls recorded, were identified using multivariate analyses of ten acoustic parameters; knock, growl, drum, and sweep. The calls were relatively brief (12-270 ms, mean of -10 dB duration) pulsating sounds of low-frequency (peak frequency <600 Hz) and narrow bandwidth (169-232 Hz, mean of -3 dB bandwidth; 237-435 Hz, mean of -6 dB bandwidth). Frequency variables (-3/-6 dB frequency bandwidth, center frequency, and top three peak frequencies) were the primary descriptors used to differentiate the call types. Comparing the spotted seal underwater vocalizations with those of the closely related Pacific harbor seal (Phoca vitulina richardii) indicated that the two species use similar bandwidths and peak frequencies but spotted seal calls were generally shorter. Knowledge of underwater vocalizations of wild spotted seals is important for understanding the species behavior and for planning future acoustic surveys of its distribution and occurrence.

Source levels and call parameters of harbor seal breeding vocalizations near a terrestrial haulout site in Glacier Bay National Park and Preserve.

Source levels of harbor seal breeding vocalizations were estimated using a three-element planar hydrophone array near the Beardslee Islands in Glacier Bay National Park and Preserve, Alaska. The average source level for these calls was 144 dBRMS re 1 µPa at 1 m in the 40-500 Hz frequency band. Source level estimates ranged from 129 to 149 dBRMS re 1 µPa. Four call parameters, including minimum frequency, peak frequency, total duration, and pulse duration, were also measured. These measurements indicated that breeding vocalizations of harbor seals near the Beardslee Islands of Glacier Bay National Park are similar in duration (average total duration: 4.8 s, average pulse duration: 3.0 s) to previously reported values from other populations, but are 170-220 Hz lower in average minimum frequency (78 Hz).

A harbor seal can transfer the same/different concept to new stimulus dimensions.

We investigated the formation of an abstract concept of same/different in a harbor seal by means of a two-item same/different task. Stimuli were presented on a TFT monitor. The subject was trained to respond according to whether two horizontally aligned white shapes presented on a black background were the same, or different from each other, by giving a no-go or go response. Training comprised of four stages. First, the same/different task was trained with two shapes forming two same problems (A-A and B-B) and two different problems (A-B and B-A). After the learning criterion was reached, training proceeded with new pairs of shapes. In the second experimental stage, every problem was presented just five times before new problems were introduced. We showed that training to criterion with just two shapes resulted in item-specific learning, whereas reducing the number of presentations to five per problem led to the formation of a same/different learning set as well as some restricted relational learning. Training with trial-unique problems in the third stage of this study resulted in the formation of an abstract concept of same/different which was indicated by a highly significant performance in transfer tests with 120 novel problems. Finally, extra-dimensional transfer of the concept was tested. The harbor seal showed a significantly correct performance on transfer tests with 30 unfamiliar pattern and 60 unfamiliar brightness same/different problems, thus demonstrating that the concept is not restricted to the shape dimension originally learned, but can be generalized across stimulus dimensions.

Vibrissal touch sensing in the harbor seal (Phoca vitulina): how do seals judge size?

"Whisker specialists" such as rats, shrews, and seals actively employ their whiskers to explore their environments and extract object properties such as size, shape, and texture. It has been suggested that whiskers could be used to discriminate between different sized objects in one of two ways: (i) to use whisker positions, such as angular position, spread or amplitude to approximate size; or (ii) to calculate the number of whiskers that contact an object. This study describes in detail how two adult harbor seals use their whiskers to differentiate between three sizes of disk. The seals judged size very fast, taking <400 ms. In addition, they oriented their smaller, most rostral, ventral whiskers to the disks, so that more whiskers contacted the surface, complying to a maximal contact sensing strategy. Data from this study supports the suggestion that it is the number of whisker contacts that predict disk size, rather than how the whiskers are positioned (angular position), the degree to which they are moved (amplitude) or how spread out they are (angular spread).

Are harbour seals (Phoca vitulina) able to perceive and use polarised light?

Harbour seals are active at night and during the day and see well in both air and water. Polarised light, which is a well-known visual cue for orientation, navigation and foraging, is richly available in harbour seal habitats, both above and below the water surface. We hypothesised that an ability to detect and use polarised light could be valuable for seals, and thus tested if they are able to see this property of light. We performed two behavioural experiments, one involving object discrimination and the other involving object detection. These objects were presented to the seals as two-dimensional stimuli on a specially modified liquid crystal display that generated objects whose contrast was purely defined in terms of polarisation (i.e. objects lacked luminance contrast). In both experiments, the seals' performance did not deviate significantly from chance. In contrast, the seals showed a high baseline performance when presented with objects on a non-modified display (whose contrast was purely defined in terms of luminance). We conclude that harbour seals are unable to use polarised light in our experimental context. It remains for future work to elucidate if they are polarisation insensitive per se.

Temporal processing of low-frequency sounds by seals (L).

In a recent study, Kastelein et al. [(2010) J. Acoust. Soc. Am. 127, 1135-1145] reported auditory integration times for harbor seals (Phoca vitulina) exceeding 3000 ms for 200 Hz tonal signals. This finding is unexpected and potentially significant given that time constants measured in mammals for tones above 1 kHz are typically less than 500 ms. To further explore this result, the hearing of another harbor seal was measured in air and water for 200 Hz tones with durations of 500 and 2500 ms. Threshold comparisons, as well as reaction time measures, revealed no gain in audibility as signal duration increased above 500 ms.

Hearing threshold shifts and recovery in harbor seals (Phoca vitulina) after octave-band noise exposure at 4 kHz.

Safety criteria for underwater sounds from offshore pile driving are needed to protect marine mammals. As a first step toward understanding effects of impulsive sounds, two harbor seals were exposed to octave-band white noise centered at 4 kHz at three mean received sound pressure levels (SPLs; 124, 136, and 148 dB re 1 µPa) at up to six durations (7.5, 15, 30, 60, 120, and 240 min); mean received sound exposure level (SEL) range was 166-190 dB re 1 µPa(2) s. Hearing thresholds were determined before and after exposure. Temporary hearing threshold shifts (TTS) and subsequent recovery were quantified as changes in hearing thresholds at 1-4, 4-8, 8-12, 48, and 96 min after noise exposure in seal 01, and at 12-16, 16-20, 20-24, 60, and 108 min after exposure in seal 02. Maximum TTS (1-4 min after 120 min exposure to 148 dB re 1 µPa; 187 dB SEL) was 10 dB. Recovery occurred within ~60 min. Statistically significant TTSs (>2.5 dB) began to occur at SELs of ~170 (136 SPL, 60 min) and 178 dB re 1 µPa(2) s (148 SPL, 15 min). However, SEL is not an optimal predictor of TTS for long duration, low SPL continuous noise, as duration and SPL play unequal roles in determining induced TTS.

Aversiveness of sounds in phocid seals: psycho-physiological factors, learning processes and motivation.

Aversiveness of sounds and its underlying physiological mechanisms in mammals are poorly understood. In this study we tested the influence of psychophysical parameters, motivation and learning processes on the aversiveness of anthropogenic underwater noise in phocid seals (Halichoerus grypus and Phoca vitulina). We compared behavioural responses of seals to playbacks of sounds based on a model of sensory unpleasantness for humans, sounds from acoustic deterrent devices and sounds with assumed neutral properties in different contexts of food motivation. In a captive experiment with food presentation, seals habituated quickly to all sound types presented at normalised received levels of 146 dB re. 1 microPa (r.m.s., root mean square). However, the fast habituation of avoidance behaviour was also accompanied by a weak sensitisation process affecting dive times and place preference in the pool. Experiments in the wild testing animals without food presentation revealed differential responses of seals to different sound types. We observed avoidance behaviour at received levels of 135-144 dB re. 1 microPa (sensation levels of 59-79 dB). In this experiment, sounds maximised for 'roughness' perceived as unpleasant by humans also caused the strongest avoidance responses in seals, suggesting that sensory pleasantness may be the result of auditory processing that is not restricted to humans. Our results highlight the importance of considering the effects of acoustic parameters other than the received level as well as animal motivation and previous experience when assessing the impacts of anthropogenic noise on animals.

Ringed seal post-moulting movement tactics and habitat selection.

Intra-specific and intra-population variation in movement tactics have been observed in many species, sometimes in association with alternative foraging techniques or large-scale habitat selection. However, whether animals adjust their small-scale habitat selection according to their large-scale tactics has rarely been studied. This study identified two large-scale movement tactics in ringed seals (Phoca hispida) during their non-breeding, post-moulting period. First-passage times (FPT) were used to explore these large-scale patterns. Subsequently, habitat selection was quantified by modelling the FPTs as a function of habitat attributes using Cox proportional hazards models. Some seals moved far offshore into areas preferentially containing 40-80% ice coverage, while other individuals spread along the coasts of Svalbard concentrating their time near glacier fronts. Both tactics resulted in ringed seals being in highly productive areas where they had access to ice-platforms to rest. When offshore, habitat selection was influenced mainly by sea ice concentration and season. Late in the season (autumn), increased risk of leaving an area was identified, even when ice conditions were still favourable, reflecting their need to return to over-wintering/breeding areas before the fjords of the archipelago freeze. For ringed seals that remained inshore, habitat use intensities were influenced mainly by the distance to glacier fronts and season. These animals were already close to their over-wintering habitat and hence their risk of leaving an area decreased as winter approached. This study of ringed seals habitat selection reveals how they fulfil their biological requirements in this dynamic, heterogeneous habitat. Individuals within the same population employed two distinct large-scale movement tactics, adjusting their decisions for small-scale habitat selection accordingly. This flexibility in ringed seal spatial ecology during summer and fall is expected to result in increased population viability in this high Arctic environment.

Deterring effects of 8-45 kHz tone pulses on harbour seals (Phoca vitulina) in a large pool.

The marine aquaculture industry suffers losses due to pinniped attacks which damage net enclosures and fish stocks. Acoustic harassment devices (AHDs) emit loud sounds which are intended to deter pinnipeds from approaching aquaculture enclosures. At present, many AHDs emit sounds in the 8-20 kHz frequency range. It is not known whether sounds of higher frequencies have a deterrent effect on seals. Therefore five captive harbour seals (Phoca vitulina) were subjected to four series of tone pulses together spanning a broad frequency range (8, 16, 32 and 45 kHz). Pulse duration was 250 ms and pulse interval was 5s. Each of the four sounds was made deterrent by increasing the amplitude. The seals reacted by swimming away from the sounds. The displacement effect of each sound was judged by comparing the animals' surface positions, and number of surfacings, during ten 45 min baseline periods with ten 45 min test periods per frequency (one frequency per day in rotation, 40 sessions in total). The seals were displaced by all four frequencies throughout the 40 trial days. The seals came to the surface more often when the test tones were produced than in the baseline periods. The initial displacement distances did not change over the 40 test days. This suggests that operating AHDs for only short periods will be more effective and less likely to result in habituation by the seals than operating them continuously. The discomfort threshold sound pressure level (SPL) was established for each of the four pulse frequencies. The acoustic discomfort threshold SPL is defined as the boundary SPL between the area that the animals generally occupied during the transmission of the sounds and the area that they generally did not enter during sound transmission. The discomfort threshold SPL may depend on the context.

The influence of underwater data transmission sounds on the displacement behaviour of captive harbour seals (Phoca vitulina).

To prevent grounding of ships and collisions between ships in shallow coastal waters, an underwater data collection and communication network (ACME) using underwater sounds to encode and transmit data is currently under development. Marine mammals might be affected by ACME sounds since they may use sound of a similar frequency (around 12 kHz) for communication, orientation, and prey location. If marine mammals tend to avoid the vicinity of the acoustic transmitters, they may be kept away from ecologically important areas by ACME sounds. One marine mammal species that may be affected in the North Sea is the harbour seal (Phoca vitulina). No information is available on the effects of ACME-like sounds on harbour seals, so this study was carried out as part of an environmental impact assessment program. Nine captive harbour seals were subjected to four sound types, three of which may be used in the underwater acoustic data communication network. The effect of each sound was judged by comparing the animals' location in a pool during test periods to that during baseline periods, during which no sound was produced. Each of the four sounds could be made into a deterrent by increasing its amplitude. The seals reacted by swimming away from the sound source. The sound pressure level (SPL) at the acoustic discomfort threshold was established for each of the four sounds. The acoustic discomfort threshold is defined as the boundary between the areas that the animals generally occupied during the transmission of the sounds and the areas that they generally did not enter during transmission. The SPLs at the acoustic discomfort thresholds were similar for each of the sounds (107 dB re 1 microPa). Based on this discomfort threshold SPL, discomfort zones at sea for several source levels (130-180 dB re 1 microPa) of the sounds were calculated, using a guideline sound propagation model for shallow water. The discomfort zone is defined as the area around a sound source that harbour seals are expected to avoid. The definition of the discomfort zone is based on behavioural discomfort, and does not necessarily coincide with the physical discomfort zone. Based on these results, source levels can be selected that have an acceptable effect on harbour seals in particular areas. The discomfort zone of a communication sound depends on the sound, the source level, and the propagation characteristics of the area in which the sound system is operational. The source level of the communication system should be adapted to each area (taking into account the width of a sea arm, the local sound propagation, and the importance of an area to the affected species). The discomfort zone should not coincide with ecologically important areas (for instance resting, breeding, suckling, and feeding areas), or routes between these areas.