Individual differences in the behavioral characteristics of beluga whales (Dephinapterus leucas).

The topic of individual differences in animal behavior has garnered a great deal of recent attention across many species, but questions remain concerning the degree to which behavioral differences vary over time or by age and sex. The present study focused on white whales (Delphinapterus leucas), a species in which a high degree of behavioral variability may be expected due to the fact that belugas are large-brained, long-lived, and highly social in nature. A suite of 23 behavioral measures related to boldness, playfulness, sociability, and other traits were assessed in 41 seaquarium-based belugas that were housed in mixed age/sex grouping. The goals were to assess consistency within individuals over time as well as variations by age and sex. Nineteen of the 23 measures showed significant within-subject consistency over time, suggesting that stable behavioral differences appear to exist in this species. However, very few measures showed significant correlations with each other, indicating that they could not be grouped into identifiable factors that comprised recognizable clusters. It is suggested, therefore, that individual differences are best viewed as a complex array of characteristics that depend on an individual's age and contextual variables that influence the degree to which specific sets of behaviors are expressed and co-vary.

A beluga whale socialized with bottlenose dolphins imitates their whistles.

The research on imitation in the animal kingdom has more than a century-long history. A specific kind of imitation, auditory-vocal imitation, is well known in birds, especially among songbirds and parrots, but data for mammals are limited to elephants, marine mammals, and humans. Cetaceans are reported to imitate various signals, including species-specific calls, artificial sounds, and even vocalizations from other species if they share the same habitat. Here we describe the changes in the vocal repertoire of a beluga whale that was housed with a group of bottlenose dolphins. Two months after the beluga's introduction into a new facility, we found that it began to imitate whistles of the dolphins, whereas one type of its own calls seemed to disappear. The case reported here may be considered as an interesting phenomenon of vocal accommodation to new social companions and cross-species socialization in cetaceans.

Behavioral responses of beluga whales (Delphinapterus leucas) to environmental variation in an Arctic estuary.

Some Arctic estuaries serve as substrate rubbing sites for beluga whales (Delphinapterus leucas) in the summer, representing a specialized resource for the species. Understanding how environmental variation affects the species' behavior is essential to management of these habitats in coming years as the climate changes. Spatiotemporal and environmental variables were recorded for behavioral observations, during which focal groups of whales in an estuary were video-recorded for enumeration and behavioral analysis. Multiple polynomial linear regression models were optimized to identify the effects of spatiotemporal and environmental conditions on group size, composition, and the frequency of behaviors being performed. Results suggest that belugas take advantage of environmental variation to express behaviors that 1) protect young, e.g., bringing calves close to shore during cloudier days, obscuring visualization from terrestrial predators; 2) avoid predation, e.g., rubbing against substrates at higher Beaufort sea states to obscure visualization, and resting during low tides while swimming on outgoing tides to avoid stranding; and 3) optimize bioenergetic resources, e.g., swimming during lower Beaufort sea states and clearer days. Predictive models like the ones presented in this study can inform conservation management strategies as environmental conditions change in future years.

Contextual imitation of intransitive body actions in a Beluga whale (Delphinapterus leucas): A "do as other does" study.

Cetaceans are remarkable for exhibiting group-specific behavioral traditions or cultures in several behavioral domains (e.g., calls, behavioral tactics), and the question of whether they can be acquired socially, for example through imitative processes, remains open. Here we used a "Do as other does" paradigm to experimentally study the ability of a beluga to imitate familiar intransitive (body-oriented) actions demonstrated by a conspecific. The participant was first trained to copy three familiar behaviors on command (training phase) and then was tested for her ability to generalize the learned "Do as the other does" command to a different set of three familiar behaviors (testing phase). We found that the beluga (1) was capable of learning the copy command signal "Do what-the-other-does"; (2) exhibited high matching accuracy for trained behaviors (mean = 84% of correct performance) after making the first successful copy on command; (3) copied successfully the new set of three familiar generalization behaviors that were untrained to the copy command (range of first copy = 12 to 35 trials); and (4) deployed a high level of matching accuracy (mean = 83%) after making the first copy of an untrained behavior on command. This is the first evidence of contextual imitation of intransitive (body-oriented) movements in the beluga and adds to the reported findings on production imitation of sounds in this species and production imitation of sounds and motor actions in several cetaceans, especially dolphins and killer whales. Collectively these findings highlight the notion that cetaceans have a natural propensity at skillfully and proficiently matching the sounds and body movements demonstrated by conspecifics, a fitness-enhancing propensity in the context of cooperative hunting and anti-predatory defense tactics, and of alliance formation strategies that have been documented in these species' natural habitats. Future work should determine if the beluga can also imitate novel motor actions.

Responses to familiar and unfamiliar objects by belugas (Delphinapterus leucas), bottlenose dolphins (Tursiops truncatus), and Pacific white-sided dolphins (Lagenorhynchus obliquidens).

Previous research with bottlenose dolphins (Tursiops truncatus) demonstrated their ability to discriminate between familiar and unfamiliar stimuli. Dolphins gazed longer at unfamiliar stimuli. The current study attempted to extend this original research by examining the responses of three species of cetaceans to objects that differed in familiarity. Eleven belugas from two facilities, five bottlenose dolphins and five Pacific white-sided dolphins housed at one facility were presented different objects in a free-swim scenario. The results indicated that the animals gazed the longest at unfamiliar objects, but these gaze durations did not significantly differ from gaze durations when viewing familiar objects. Rather, the animals gazed longer at unfamiliar objects when compared to the apparatus alone. Species differences emerged with longer gaze durations exhibited by belugas and bottlenose dolphins and significantly shorter gaze durations for Pacific white-sided dolphins. It is likely that the animals categorized objects into familiar and unfamiliar categories, but the free-swim paradigm in naturalistic social groupings did not elicit clear responses. Rather this procedure emphasized the importance of attention and individual preferences when investigating familiar and unfamiliar objects, which has implications for cognitive research and enrichment use.

Auditory evoked potentials in the auditory system of a beluga whale Delphinapterus leucas to prolonged sound stimuli.

The effects of prolonged (up to 1500 s) sound stimuli (tone pip trains) on evoked potentials (the rate following response, RFR) were investigated in a beluga whale. The stimuli (rhythmic tone pips) were of frequencies of 45, 64, and 90 kHz at levels from 20 to 60 dB above threshold. Two experimental protocols were used: short- and long-duration. For the short-duration protocol, the stimuli were 500-ms-long pip trains that repeated at a rate of 0.4 trains/s. For the long-duration protocol, the stimuli were continuous pip successions lasting up to 1500 s. The RFR amplitude gradually decreased by three to seven times from 10 ms to 1500 s of stimulation. Decrease of response amplitude during stimulation was approximately proportional to initial (at the start of stimulation) response amplitude. Therefore, even for low stimulus level (down to 20 dB above the baseline threshold) the response was never suppressed completely. The RFR amplitude decay that occurred during stimulation could be satisfactorily approximated by a combination of two exponents with time constants of 30-80 ms and 3.1-17.6 s. The role of adaptation in the described effects and the impact of noise on the acoustic orientation of odontocetes are discussed.

Evoked-potential recovery during double click stimulation in a beluga whale: implications for biosonar gain control.

Auditory evoked potentials (AEPs) were recorded in a beluga whale Delphinapterus leucas using a double-pulse stimulation paradigm, specifically measuring the recovery (release from masking) of the second (test) response as a function of delay after the first (conditioning) pulse at various levels of the conditioning and test stimuli. The conditioning/test stimulus level ratio influenced the recovery time (the higher the ratio, the longer the recovery). This interrelation was used to evaluate the intensity/time trade in release from forward masking. Trade was evaluated as 32.2 dB per time decade. Data were considered as simulating interactions between the transmitted pulse and echo during echolocation, assuming that a transmitted sonar pulse produces forward masking of the echo response. With increased target distance, the attenuation of the echo may be compensated by the release from masking. According to the model, the compensation results in substantial stabilization of the echo response even if the intensity/time trade of release from masking is not precisely equal to the rate of echo attenuation with distance.

Frequency tuning of hearing in the beluga whale: discrimination of rippled spectra.

Frequency tuning was measured in the beluga whale (Delphinapterus leucas) using rippled-noise test stimuli in conjunction with an auditory evoked potential (AEP) technique. The test stimulus was a 2-octave-wide rippled noise with frequency-proportional ripple spacing. The rippled-noise signal contained either a single reversal or rhythmic (1-kHz rate) reversals of the ripple phase. Single or rhythmic phase reversals evoked, respectively, a single auditory brainstem response (ABR) or a rhythmic AEP sequence-the envelope following response (EFR). The response was considered as an indication of resolvability of the ripple pattern. The rhythmic phase-reversal test with EFR recording revealed higher resolution than the single phase-reversal test with single ABR recording. The limit of ripple-pattern resolution with the single phase-reversal test ranged from 17 ripples per octave (rpo) at 32 kHz to 24 rpo at 45 to 64 kHz; for the rhythmic phase-reversal test, the limit ranged from 20 to 32 rpo. An interaction model of a ripple spectrum with frequency-tuned filters suggests that the ripple-pattern resolution limit of 20 to 32 rpo requires a filter quality Q of 29 to 46. Possible causes of disagreement of these estimates with several previously published data are discussed.

Visual laterality in belugas (Delphinapterus leucas) and Pacific white-sided dolphins (Lagenorhynchus obliquidens) when viewing familiar and unfamiliar humans.

Lateralization of cognitive processes and motor functions has been demonstrated in a number of species, including humans, elephants, and cetaceans. For example, bottlenose dolphins (Tursiops truncatus) have exhibited preferential eye use during a variety of cognitive tasks. The present study investigated the possibility of visual lateralization in 12 belugas (Delphinapterus leucas) and six Pacific white-sided dolphins (Lagenorhynchus obliquidens) located at two separate marine mammal facilities. During free swim periods, the belugas and Pacific white-sided dolphins were presented a familiar human, an unfamiliar human, or no human during 10-15 min sessions. Session videos were coded for gaze duration, eye presentation at approach, and eye preference while viewing each stimulus. Although we did not find any clear group level lateralization, we found individual left eye lateralized preferences related to social stimuli for most belugas and some Pacific white-sided dolphins. Differences in gaze durations were also observed. The majority of individual belugas had longer gaze durations for unfamiliar rather than familiar stimuli. These results suggest that lateralization occurs during visual processing of human stimuli in belugas and Pacific white-sided dolphins and that these species can distinguish between familiar and unfamiliar humans.

Passive acoustic monitoring of Cook Inlet beluga whales (Delphinapterus leucas).

The endangered beluga whale (Delphinapterus leucas) population in Cook Inlet, AK faces threats from a variety of anthropogenic factors, including coastal development, oil and gas exploration, vessel traffic, and military activities. To address existing gaps in understanding about the occurrence of belugas in Cook Inlet, a project was developed to use passive acoustic monitoring to document the year-round distribution of belugas, as well as killer whales (Orcinus orca), which prey on belugas. Beginning in June 2009, ten moorings were deployed throughout the Inlet and refurbished every two to eight months. Despite challenging conditions consisting of strong tidal currents carrying debris and seasonal ice cover, 83% of mooring deployments were successfully recovered. Noise from water flow, vessel traffic, and/or industrial activities was present at several sites, potentially masking some signals. However, belugas were successfully detected at multiple locations. Detections were relatively common in the upper inlet and less common or absent at middle and lower inlet locations. Killer whale signals were also recorded. Some seasonal variability in the occurrence of both belugas and killer whales was evident.

[Multipurpose probe for underwater assessment of marine basin ecology].

[The multipurpose probe for real-time assessment of behavior reactions of marine mammals and concurrent temperature and noise pollution measurements was subjected to field testing]

Allonursing in captive belugas (Delphinapterus leucas).

This study describes allonursing (females nursing offspring that are not their own) in captive belugas (Delphinapterus leucas). In addition to the calf's mother, two females that were not pregnant or nursing at the time of the calf's birth spontaneously lactated and nursed the male calf intermittently throughout 34 months of his life at the Vancouver Aquarium Marine Science Center. These observations suggest that allonursing may also take place in their wild counterparts and add to existing evidence of alloparental care in this species. Alloparental care, the care of nonoffspring, has been observed in every major mammalian taxon; the protection of calves through alloparental care may be a strong evolutionary benefit.