Behavioral context of echolocation and prey-handling sounds produced by killer whales (Orcinus orca) during pursuit and capture of Pacific salmon (Oncorhynchus spp.).

Availability of preferred salmonid prey and a sufficiently quiet acoustic environment in which to forage are critical to the survival of resident killer whales (Orcinus orca) in the northeastern Pacific. Although piscivorous killer whales rely on echolocation to locate and track prey, the relationship between echolocation, movement, and prey capture during foraging by wild individuals is poorly understood. We used acoustic biologging tags to relate echolocation behavior to prey pursuit and capture during successful feeding dives by fish-eating killer whales in coastal British Columbia, Canada. The significantly higher incidence and rate of echolocation prior to fish captures compared to afterward confirms its importance in prey detection and tracking. Extremely rapid click sequences (buzzes) were produced before or concurrent with captures of salmon at depths typically exceeding 50 m, and were likely used by killer whales for close-range prey targeting, as in other odontocetes. Distinctive crunching and tearing sounds indicative of prey-handling behavior occurred at relatively shallow depths following fish captures, matching concurrent observations that whales surfaced with fish prior to consumption and often shared prey. Buzzes and prey-handling sounds are potentially useful acoustic signals for estimating foraging efficiency and determining if resident killer whales are meeting their energetic requirements.

Low-frequency signals produced by Northeast Atlantic killer whales (Orcinus orca).

Killer whale acoustic behavior has been extensively investigated; however, most studies have focused on pulsed calls and whistles. This study reports the production of low-frequency signals by killer whales at frequencies below 300 Hz. Recordings were made in Iceland and Norway when killer whales were observed feeding on herring and no other marine mammal species were nearby. Low-frequency sounds were identified in Iceland and ranged in duration between 0.14 and 2.77 s and in frequency between 50 and 270 Hz, well below the previously reported lower limit for killer whale tonal sounds of 500 Hz. Low-frequency sounds appeared to be produced close in time to tail slaps, which are indicative of feeding attempts, suggesting that these sounds may be related to a feeding context. However, their precise function is unknown, and they could be the by-product of a non-vocal behavior rather than a vocal signal deliberately produced by the whales. Although killer whales in Norway exhibit similar feeding behavior, this sound has not been detected in recordings from Norway to date. This study suggests that, like other delphinids, killer whales produce low-frequency sounds, but further studies will be required to understand whether similar sounds exist in other killer whale populations.

Grey seals use anthropogenic signals from acoustic tags to locate fish: evidence from a simulated foraging task.

Anthropogenic noise can have negative effects on animal behaviour and physiology. However, noise is often introduced systematically and potentially provides information for navigation or prey detection. Here, we show that grey seals (Halichoerus grypus) learn to use sounds from acoustic fish tags as an indicator of food location. In 20 randomized trials each, 10 grey seals individually explored 20 foraging boxes, with one box containing a tagged fish, one containing an untagged fish and all other boxes being empty. The tagged box was found after significantly fewer non-tag box visits across trials, and seals revisited boxes containing the tag more often than any other box. The time and number of boxes needed to find both fish decreased significantly throughout consecutive trials. Two additional controls were conducted to investigate the role of the acoustic signal: (i) tags were placed in one box, with no fish present in any boxes and (ii) additional pieces of fish, inaccessible to the seal, were placed in the previously empty 18 boxes, making possible alternative chemosensory cues less reliable. During these controls, the acoustically tagged box was generally found significantly faster than the control box. Our results show that animals learn to use information provided by anthropogenic signals to enhance foraging success.

Divergent foraging strategies between populations of sympatric matrilineal killer whales.

In cooperative species, human-induced rapid environmental change may threaten cost-benefit tradeoffs of group behavioral strategies that evolved in past environments. Capacity for behavioral flexibility can increase population viability in novel environments. Whether the partitioning of individual responsibilities within social groups is fixed or flexible across populations is poorly understood, despite its relevance for predicting responses to global change at the population and species levels and designing successful conservation programs. We leveraged bio-logging data from two populations of fish-eating killer whales (Orcinus orca) to quantify patterns of fine-scale foraging movements and their relationships with demography. We reveal striking interpopulation differences in patterns of individual foraging behavior. Females from the endangered Southern Resident (SRKW) population captured less prey and spent less time pursuing prey than SRKW males or Northern Resident (NRKW) females, whereas NRKW females captured more prey than NRKW males. The presence of a calf (≤3 years) reduced the number of prey captured by adult females from both populations, but disproportionately so for SRKW. SRKW adult males with a living mother captured more prey than those whose mother had died, whereas the opposite was true for NRKW adult males. Across populations, males foraged in deeper areas than females, and SRKW captured prey deeper than NRKW. These population-level differences in patterns of individual foraging behavior challenge the existing paradigm that females are the disproportionate foragers in gregarious resident killer whales, and demonstrate considerable variation in the foraging strategies across populations of an apex marine predator experiencing different environmental stressors.

Tool-use in the brown bear (Ursus arctos).

This is the first report of tool-using behaviour in a wild brown bear (Ursus arctos). Whereas the use of tools is comparatively common among primates and has also been documented in several species of birds, fishes and invertebrates, tool-using behaviours have so far been observed in only four species of non-primate mammal. The observation was made and photographed while studying the behaviour of a subadult brown bear in south-eastern Alaska. The animal repeatedly picked up barnacle-encrusted rocks in shallow water, manipulated and re-oriented them in its forepaws, and used them to rub its neck and muzzle. The behaviour probably served to relieve irritated skin or to remove food-remains from the fur. Bears habitually rub against stationary objects and overturn rocks and boulders during foraging and such rubbing behaviour could have been transferred to a freely movable object to classify as tool-use. The bear exhibited considerable motor skills when manipulating the rocks, which clearly shows that these animals possess the advanced motor learning necessary for tool-use. Advanced spatial cognition and motor skills for object manipulation during feeding and tool-use provide a possible explanation for why bears have the largest brains relative to body size of all carnivores. Systematic research into the cognitive abilities of bears, both in captivity and in the wild, is clearly warranted to fully understand their motor-learning skills and physical intelligence related to tool-use and other object manipulation tasks.

The structure of stereotyped calls reflects kinship and social affiliation in resident killer whales (Orcinus orca).

A few species of mammals produce group-specific vocalisations that are passed on by learning, but the function of learned vocal variation remains poorly understood. Resident killer whales live in stable matrilineal groups with repertoires of seven to 17 stereotyped call types. Some types are shared among matrilines, but their structure typically shows matriline-specific differences. Our objective was to analyse calls of nine killer whale matrilines in British Columbia to test whether call similarity primarily reflects social or genetic relationships. Recordings were made in 1985-1995 in the presence of focal matrilines that were either alone or with groups with non-overlapping repertoires. We used neural network discrimination performance to measure the similarity of call types produced by different matrilines and determined matriline association rates from 757 encounters with one or more focal matrilines. Relatedness was measured by comparing variation at 11 microsatellite loci for the oldest female in each group. Call similarity was positively correlated with association rates for two of the three call types analysed. Similarity of the N4 call type was also correlated with matriarch relatedness. No relationship between relatedness and association frequency was detected. These results show that call structure reflects relatedness and social affiliation, but not because related groups spend more time together. Instead, call structure appears to play a role in kin recognition and shapes the association behaviour of killer whale groups. Our results therefore support the hypothesis that increasing social complexity plays a role in the evolution of learned vocalisations in some mammalian species.

Killer whales (Orcinus orca) produce ultrasonic whistles.

This study reports that killer whales, the largest dolphin, produce whistles with the highest fundamental frequencies ever reported in a delphinid. Using wide-band acoustic sampling from both animal-attached (Dtag) and remotely deployed hydrophone arrays, ultrasonic whistles were detected in three Northeast Atlantic populations but not in two Northeast Pacific populations. These results are inconsistent with analyses suggesting a correlation of maximum frequency of whistles with body size in delphinids, indicate substantial intraspecific variation in whistle production in killer whales, and highlight the importance of appropriate acoustic sampling techniques when conducting comparative analyses of sound repertoires.

Body density of humpback whales (Megaptera novaengliae) in feeding aggregations estimated from hydrodynamic gliding performance.

Many baleen whales undertake annual fasting and feeding cycles, resulting in substantial changes in their body condition, an important factor affecting fitness. As a measure of lipid-store body condition, tissue density of a few deep diving marine mammals has been estimated using a hydrodynamic glide model of drag and buoyancy forces. Here, we applied the method to shallow-diving humpback whales (Megaptera novaeangliae) in North Atlantic and Antarctic feeding aggregations. High-resolution 3-axis acceleration, depth and speed data were collected from 24 whales. Measured values of acceleration during 5 s glides were fitted to a hydrodynamic glide model to estimate unknown parameters (tissue density, drag term and diving gas volume) in a Bayesian framework. Estimated species-average tissue density (1031.6 ± 2.1 kg m-3, ±95% credible interval) indicates that humpback whale tissue is typically negatively buoyant although there was a large inter-individual variation ranging from 1025.2 to 1043.1 kg m-3. The precision of the individual estimates was substantially finer than the variation across different individual whales, demonstrating a progressive decrease in tissue density throughout the feeding season and comparably high lipid-store in pregnant females. The drag term (CDAm-1) was estimated to be relatively high, indicating a large effect of lift-related induced drag for humpback whales. Our results show that tissue density of shallow diving baleen whales can be estimated using the hydrodynamic gliding model, although cross-validation with other techniques is an essential next step. This method for estimating body condition is likely to be broadly applicable across a range of aquatic animals and environments.

Fine-scale foraging movements by fish-eating killer whales (Orcinus orca) relate to the vertical distributions and escape responses of salmonid prey (Oncorhynchus spp.).

BACKGROUND: We sought to quantitatively describe the fine-scale foraging behavior of northern resident killer whales (Orcinus orca), a population of fish-eating killer whales that feeds almost exclusively on Pacific salmon (Oncorhynchus spp.). To reconstruct the underwater movements of these specialist predators, we deployed 34 biologging Dtags on 32 individuals and collected high-resolution, three-dimensional accelerometry and acoustic data. We used the resulting dive paths to compare killer whale foraging behavior to the distributions of different salmonid prey species. Understanding the foraging movements of these threatened predators is important from a conservation standpoint, since prey availability has been identified as a limiting factor in their population dynamics and recovery. RESULTS: Three-dimensional dive tracks indicated that foraging (N = 701) and non-foraging dives (N = 10,618) were kinematically distinct (Wilks' lambda: λ16 = 0.321, P < 0.001). While foraging, killer whales dove deeper, remained submerged longer, swam faster, increased their dive path tortuosity, and rolled their bodies to a greater extent than during other activities. Maximum foraging dive depths reflected the deeper vertical distribution of Chinook (compared to other salmonids) and the tendency of Pacific salmon to evade predators by diving steeply. Kinematic characteristics of prey pursuit by resident killer whales also revealed several other escape strategies employed by salmon attempting to avoid predation, including increased swimming speeds and evasive maneuvering. CONCLUSIONS: High-resolution dive tracks reconstructed using data collected by multi-sensor accelerometer tags found that movements by resident killer whales relate significantly to the vertical distributions and escape responses of their primary prey, Pacific salmon.

Automated categorization of bioacoustic signals: avoiding perceptual pitfalls.

Dividing the acoustic repertoires of animals into biologically relevant categories presents a widespread problem in the study of animal sound communication, essential to any comparison of repertoires between contexts, individuals, populations, or species. Automated procedures allow rapid, repeatable, and objective categorization, but often perform poorly at detecting biologically meaningful sound classes. Arguably this is because many automated methods fail to address the nonlinearities of animal sound perception. We present a new method of categorization that incorporates dynamic time-warping and an adaptive resonance theory (ART) neural network. This method was tested on 104 randomly chosen whistle contours from four captive bottlenose dolphins (Tursiops truncatus), as well as 50 frequency contours extracted from calls of transient killer whales (Orcinus orca). The dolphin data included known biologically meaningful categories in the form of 42 stereotyped whistles produced when each individual was isolated from its group. The automated procedure correctly grouped all but two stereotyped whistles into separate categories, thus performing as well as human observers. The categorization of killer whale calls largely corresponded to visual and aural categorizations by other researchers. These results suggest that this methodology provides a repeatable and objective means of dividing bioacoustic signals into biologically meaningful categories.

Selective habituation shapes acoustic predator recognition in harbour seals.

Predation is a major force in shaping the behaviour of animals, so that precise identification of predators will confer substantial selective advantages on animals that serve as food to others. Because experience with a predator can be lethal, early researchers studying birds suggested that predator recognition does not require learning. However, a predator image that can be modified by learning and experience will be advantageous in situations where cues associated with the predator are highly variable or change over time. In this study, we investigated the response of harbour seals (Phoca vitulina) to the underwater calls of different populations of killer whales (Orcinus orca). We found that the seals responded strongly to the calls of mammal-eating killer whales and unfamiliar fish-eating killer whales but not to the familiar calls of the local fish-eating population. This demonstrates that wild harbour seals are capable of complex acoustic discrimination and that they modify their predator image by selectively habituating to the calls of harmless killer whales. Fear in these animals is therefore focused on local threats by learning and experience.