Horizontal transmission of vocal traditions in killer whale (<Emphasis Type="Italic">Orcinus orca</Emphasis>) dialects

Unlike most other mammals, killer whales are capable of vocal learning and learn the dialect of their natal pod from their mothers. The classical model of killer whale dialect development suggests that the repertoire of calls is learned only “vertically” from mother to offspring, and calls evolve gradually with time by random drift caused by the accumulation of copying errors. However, some observations suggest that not only “vertical” (from mother to offspring) vocal learning can occur in killer whales, but also “horizontal” (between adult animals). In this study we analyzed the distribution of different call types and similarity of calls from the same type in different pods of killer whales from Kamchatka waters to estimate the probability of existence of interpod horizontal transmission of vocal traditions in killer whales. We found that the degree of similarity of K1 calls and K5 calls in different pods can differ. This situation contradicts the classical hypothesis and is possible in two cases: if different call types change with various speed in different pods, or if horizontal transmission of call features takes place. The distribution of K4 and K10 call types across pods also suggests the existence of horizontal transmission: K4 calls occur in the dialects of five of ten pods, and K10 calls, in six of ten pods, but only one pod has both K4 and K10 calls. Our results suggest that the real picture of the distribution of call features and call types in killer whale dialects contradicts the classical hypothesis of killer whale dialect evolution through the accumulation of copying errors.     

Temporal and Contextual Patterns of Killer Whale (Orcinus orca) Call Type Production

Fish-eating killer whales Orcinus orca in the northeastern Pacific live in highly stable matrifocal social groups called pods. Each pod produces a repertoire of seven or more stereotyped call types. We compared the relative production of call types of free-ranging killer whale pods over time and between social contexts. The relative production of call types by each pod during directional travel was distinct over a 27-yr period; however, both temporal stability and pod distinctiveness were strongly influenced by a subset of dominant call types within the repertoire of each pod. Some call types within the repertoires contain biphonation (two overlapping independently modulated tones) and have a higher estimated active space than call types containing just one tone. In multi-pod aggregations the relative production of the dominant call types of each pod decreased and the relative production of a subset of call types that are rarely recorded from single-pod groupings increased. The majority of these contained biphonation. The data suggest a distinction between a subset of dominant call types that may function to identify the pod and a subset of less common call types including several call types containing biphonation that are more commonly produced during inter-pod affiliations.     

Cultural transmission within maternal lineages: vocal clans in resident killer whales in southern Alaska

Cultural lineages are based on learned social traditions that are stable for several generations. When cultural lineages also reflect common ancestry and/or are shared by individuals that live together they are called clans. The existence of clans among killer whales has been previously proposed but has not been confirmed. Here, we show that clans exist among resident type killer whales, Orcinus orca, in southern Alaska. Resident killer whales live in stable matrilines from which emigration of either sex has not been observed. Matrilines that associate regularly (≥50% observation time) are called pods. Pods are believed to consist of closely related matrilines and share a unique repertoire of discrete call types. Pods that share parts of their repertoire form what Ford (1991, Canadian Journal of Zoology,69, 1454-1483) called an acoustic clan. Here, we identified discrete call types of seven pods from southern Alaska, using a method based on human discrimination of distinct aural and visual (spectrogram) differences. Mitochondrial DNA of members of each pod was also analysed. The repertoires of the seven pods were compared and two acoustically distinct groups of pods were identified. Each group was monomorphic for a different mitochondrial D-loop haplotype. Nevertheless, pods from different clans associated frequently. It thus appears that the acoustic similarities within groups, which we presume to be cultural, reflect common ancestry, and that these groups therefore meet the above definition of clans. We also argue that a combination of cultural drift and selection are the main mechanisms for the maintenance of clans. Copyright 2002 The Association for the Study of Animal Behaviour. Published by Elsevier Science Ltd. All rights reserved.     

Usage of monophonic and biphonic calls by free-ranging resident killer whales (Orcinus orca) in Kamchatka, Russian Far East

Killer whale discrete calls include types containing an overlapping high-frequency component (biphonic calls) and types without an overlapping high-frequency component (monophonic calls). In the resident killer whales of the Northeast Pacific, biphonic discrete calls exhibit higher source levels than monophonic calls, which suggests different active space and consequently different functions for monophonic and biphonic call types. In this study we investigate the potential communicative functions of monophonic and biphonic discrete calls produced by killer whales from Kamchatka (Northwest Pacific). We analyze how the usage of these calls depends on the number of pods present in the area and type of activity. Our results show that the usage of monophonic and biphonic calls in Kamchatkan killer whales depends on the number of pods in the area and is less dependent on the type of activity. Biphonic calls are more common when more than one pod is present in the area and could therefore function as markers of pod and matriline affiliation, serving mainly as cohesion signals. Monophonic calls dominated the vocalizations when a single pod was present, while in the presence of more than one pod both categories were used in equal proportions.     

A comparison of Northeast Atlantic killer whale (Orcinus orca) stereotyped call repertoires

Killer whale call repertoires can provide information on social connections among groups and populations. Killer whales in Iceland and Norway exhibit similar ecology and behavior, are genetically related, and are presumed to have been in contact before the collapse of the Atlanto-Scandian herring stock in the 1960s. However, photo-identification suggests no recent movements between Iceland and Norway but regular movement between Iceland and Shetland. Acoustic recordings collected between 2005 and 2016 in Iceland, Norway, and Shetland were used to undertake a comprehensive comparison of call repertoires of Northeast Atlantic killer whales. Measurements of time and frequency parameters of calls from Iceland (n = 4,037) and Norway (n = 1,715) largely overlapped in distribution, and a discriminant function analysis had low correct classification rate. No call type matches were confirmed between Iceland and Norway or Shetland and Norway. Three call types matched between Iceland and Shetland. Therefore, this study suggests overall similarities in time and frequency parameters but some divergence in call type repertoires. This argues against presumed past contact between Icelandic and Norwegian killer whales and suggests that they may not have been one completely mixed population.     

Within-pod variation in the sound production of a pod of killer whales, Orcinus orca

Pod-specific calling behaviour of resident killer whales has been shown to include: discrete call types not shared among pods, different production rates of shared call types, and differences in the detailed structure of shared call types. To investigate the mechanisms leading to pod-specific calling, we compared the repertoire and structure of calls produced by three different matrilineal units within the same pod, and described call features encoding matrilineal-unit distinctiveness. The three matrilineal units had different production rates of shared calls, including one call type used almost exclusively by one matrilineal unit. Cross-validated discriminant function analyses revealed matrilineal-unit distinctive structure in five of the six shared call types examined, with duration of the terminal component being the most distinctive feature for all call types containing a terminal component. Calls generally consist of low- and high-frequency components that may follow different time-frequency contours. In our sample, a particular high-frequency contour was consistently paired with a particular low-frequency contour, both contours had roughly equal overall variability, and each contained independent matrilineal-unit distinctive information. The only call type that did not differ structurally between matrilineal units is reportedly used more in interpod meetings than in intrapod contexts. The differences in calling behaviour between matrilineal units were similar in form to previously described differences between pods, although more subtle. These results suggest that pod-specific calling behaviour in resident killer whales arises primarily as a consequence of accumulated drift or divergence between highly cohesive matrilineal units as they gradually separate into different pods. Copyright 2000 The Association for the Study of Animal Behaviour.     

Pilot Whales Attracted to Killer Whale Sounds: Acoustically-Mediated Interspecific Interactions in Cetaceans

In cetaceans’ communities, interactions between individuals of different species are often observed in the wild. Yet, due to methodological and technical challenges very little is known about the mediation of these interactions and their effect on cetaceans’ behavior. Killer whales (Orcinus orca) are a highly vocal species and can be both food competitors and potential predators of many other cetaceans. Thus, the interception of their vocalizations by unintended cetacean receivers may be particularly important in mediating interspecific interactions. To address this hypothesis, we conducted playbacks of killer whale vocalizations recorded during herring-feeding activity to free-ranging long-finned pilot whales (Globicephala melas). Using a multi-sensor tag, we were able to track the whales and to monitor changes of their movements and social behavior in response to the playbacks. We demonstrated that the playback of killer whale sounds to pilot whales induced a clear increase in group size and a strong attraction of the animals towards the sound source. These findings provide the first experimental evidence that the interception of heterospecific vocalizations can mediate interactions between different cetacean species in previously unrecognized ways.     

Dependence of killer whale (Orcinus orca) acoustic signals on the type of activity and social context

We investigated the influence of the type of activity and the social context on the proportion of four different structural categories of stereotyped calls in the acoustic communication of Kamchatkan killer whales. Using generalized linear models, we described the dependence of each sound category on the type of activity, the number of killer whale pods and the presence of mixed-pod groups. We found that the proportion of different sound categories depended on the number of pods and the presence of mixed-pod groups, while the type of activity did not affect the proportion of sounds of different categories. Based on the observed differences we suggest that biphonic and high-frequency monophonic calls are mainly used as family and pod markers, and help to track the position of family members at long ranges, and low-frequency monophonic calls are used as close-range intra-group signals to maintain contact between pod members in the conditions of limited underwater visibility.     

Killer whales are capable of vocal learning

The production learning of vocalizations by manipulation of the sound production organs to alter the physical structure of sound has been demonstrated in only a few mammals. In this natural experiment, we document the vocal behaviour of two juvenile killer whales, Orcinus orca, separated from their natal pods, which are the only cases of dispersal seen during the three decades of observation of their populations. We find mimicry of California sea lion (Zalophus californianus) barks, demonstrating the vocal production learning ability for one of the calves. We also find differences in call usage (compared to the natal pod) that may reflect the absence of a repertoire model from tutors or some unknown effect related to isolation or context.     

Call types of Bigg’s killer whales (Orcinus orca) in western Alaska: using vocal dialects to assess population structure

Bigg’s killer whales (Orcinus orca; i.e. ‘transient’ ecotype), as apex predators, are important to the dynamics of marine ecosystems, but little is known about their population structure in western Alaska. Currently, all Bigg’s killer whales in western Alaska are ascribed to a single broad stock for management under the US Marine Mammal Protection Act. However, recent nuclear microsatellite and mitochondrial DNA analyses indicate that this stock is likely comprised of genetically distinct sub-populations. In accordance with what is known about killer whale vocal dialects in other locations, we used the spatial distribution of group-specific call types to investigate the population structure of Bigg’s killer whales in this part of Alaska. Digital audio recordings were collected from 33 Bigg’s killer whale encounters throughout the Aleutian and Pribilof Islands in the summers of 2001–2007 and 2009–2010. Recorded calls were qualitatively classified into discrete types and then quantitatively described using 12 structural and time-frequency measures. Resulting call categories were validated using a random forest approach. A total of 36 call types and subtypes were identified across the entire study area, and regional patterns of call type use revealed three distinct dialects which correspond to proposed genetic delineations. Our results suggest that there are at least three acoustically and genetically distinct sub-populations in western Alaska, and we present an initial catalogue for this area describing the regional vocal repertoires of Bigg’s killer whale call types.     

Assessment of meaning and the detection of unreliable signals by vervet monkeys

Free-ranging vervet monkeys, Cercopithecus aethiops, who had learned to ignore playbacks of one type of call by an unreliable signaller subsequently also ignored playback of an acoustically different call by the same individual if the calls had similar referents. Such transfer did not occur if either the identity of the signaller changed or if the two calls had different referents. After repeatedly being played an intergroup call in the absence of other groups, vervets also ignored an acoustically different intergroup call given by the same individual. The monkeys did not transfer habituation, however, if the new call was given by a different individual. In tests where the referents of two calls were different, the monkeys also failed to transfer habituation across call types. Vervet monkeys who had learned to ignore an unreliable leopard alarm call did not later ignore an eagle alarm call, even when the signaller remained the same. Results suggest that vervets, like humans, process information at a semantic, and not just an acoustic, level.     

Using a multi-disciplinary approach to identify a critically endangered killer whale management unit

A key goal for wildlife managers is identifying discrete, demographically independent conservation units. Previous genetic work assigned killer whales that occur seasonally in the Strait of Gibraltar (SoG) and killer whales sampled off the Canary Islands (CI) to the same population. Here we present new analyses of photo-identification and individual genotypes to assess the level of contemporary gene flow and migration between study areas, and analyses of biomarkers to assess ecological differences. We identified 47 different individuals from 5 pods in the SoG and 16 individuals in the CI, with no matches found between the areas. Mitochondrial DNA control region haplotype was shared by all individuals sampled within each pod, suggesting that pods have a matrifocal social structure typical of this species, whilst the lack of shared mitogenome haplotypes between the CI and SoG individuals suggests that there was little or no female migration between groups. Kinship analysis detected no close kin between CI and SoG individuals, and low to zero contemporary gene flow. Isotopic values and organochlorine pollutant loads also suggest ecological differences between study areas. We further found that one individual from a pod within the SoG not seen in association with the other four pods and identified as belonging to a potential migrant lineage by genetic analyses, had intermediate isotopic values and contaminant between the two study areas. Overall our results suggest a complex pattern of social and genetic structuring correlated with ecological variation. Consequently at least CI and SoG should be considered as two different management units. Understanding this complexity appears to be an important consideration when monitoring and understanding the viability of these management units. Understand the viability will help the conservation of these threatened management units.     

Harassment and killing of porpoises (“phocoenacide”) by fish-eating Southern Resident killer whales (Orcinus orca)

Endangered Southern Resident killer whales (Orcinus orca) are fish-eaters that preferentially prey on adult Chinook salmon (Oncorhynchus tshawytscha). Despite being salmon specialists, individuals from all three killer whale pods (J, K, L) have been observed harassing and killing porpoises (family Phocoenidae) without consuming them. Retrospectively, we identified and analyzed 78 episodes of Southern Resident killer whales harassing porpoises between 1962 and 2020, of which 28 resulted in the porpoise's death (“phocoenacide”). Fifty-six episodes involved harbor porpoise (Phocoena phocoena), 13 involved Dall's porpoise (Phocoenoides dalli), and the porpoise species was unreported for nine episodes. Southern Resident killer whales often targeted young porpoises that were similar in size to adult Chinook salmon. Both sexes participated in porpoise harassment. Juveniles engaged in the behavior the most; however, their rates of engagement were not found to differ significantly from most other age classes. The behavior was passed through generations and social groupings, as it was first observed in L pod and spread to the other two pods. Killer whales are highly complex animals known to exhibit social learning and cultural transmission of learned behaviors, but the reason(s) for this behavior is unknown. Hypotheses include the social and developmental benefits of play, hunting practice, or displaced epimeletic behavior.     

Call-type matching in vocal exchanges of free-ranging resident killer whales, Orcinus orca

Previous sound recordings of resident (fish-eating) killer whale groups have revealed matrilineal group-specific call repertoires and a strong tendency for calls of the same type to be produced in series. Vocal interactions between individual free-ranging animals, however, have remained unexplored because it has not been possible to identify signallers reliably with a single hydrophone. Here we link acoustic arrivals of calls on a towed hydrophone array with visual tracking of photo-identified individuals to ascribe calls to a focal animal when it was separated from other members of its matrilineal group by more than 35 m, and thereby out of visual range. We confirm that individual members of a matrilineal group share a repertoire of stereotyped calls, and we statistically examine timing of stereotyped calls produced by one individual relative to calls produced by other members of its group. Analysis of the intervals between stereotyped calls indicated that calls were produced in group bouts with a criterion interval of 19.6 s separating bouts. We were therefore careful to develop randomization tests that preserved call interval structure. Focal whales produced 36% of their calls within 5 s of a call from a nonfocal animal, four times more calls than expected by chance based upon a rotation randomization test. Consecutive calls produced by different individuals during group-calling bouts matched call type more than expected by chance. Vocal exchanges of stereotyped calls with type matching appear to be an important aspect of intragroup calling in killer whales, although the function of this calling behaviour remains to be explored.     

The influence of social affiliation on individual vocal signatures of northern resident killer whales (Orcinus orca)

Northern resident killer whales (Orcinus orca) live in highly stable groups and use group-specific vocal signals, but individual variation in calls has not been described previously. A towed beam-forming array was used to ascribe stereotyped pulsed calls with two independently modulated frequency contours to visually identified individual killer whales in Johnstone Strait, British Columbia. Overall, call similarity determined using neural networks differed significantly between different affiliation levels for both frequency components of all the call types analysed. This method distinguished calls from individuals within the same matriline better than different calls produced by a single individual and better than by chance. The calls of individuals from different matrilines were more distinctive than those within the same matriline, confirming previous studies based on group recordings. These results show that frequency contours of stereotyped calls differ among the individuals that are constantly associated with each other and use group-specific vocalizations, though across-group differences were substantially more pronounced.     

Stereotyped acoustic signals of the killer whale <Emphasis Type="Italic">Orcinus orca</Emphasis> (Cetacea: Delphinidae) from the Northwestern Pacific

In this paper the classification of the discrete calls of killer whales recorded in the Northwest Pacific is presented. It was shown that most of the calls can be divided into 20 discrete types easily distinguished by ear. The frequency parameters of the sounds made the maximum contribution into discrimination between call types, while their temporal parameters turned out to be less significant. The repertoires of stereotyped calls of killer whales from Chukot, Kamchatka, and the Commander and Kurile Islands were similar, due to the presence of shared types of calls. The greater the distance between the above-mentioned regions the fewer calls were shared by the resident killer whales. High vocal activity and the phenotypic features of the surveyed killer whales implied their attribution to the resident ecological type.     

Aggressive Thresholds of Male Pacific Treefrogs for Advertisement Calls Vary with Amplitude of Neighbors' Calls

In breeding choruses, male Pacific treefrogs (H. regilla) produce advertisement and encounter calls. The role of each call type in establishing intermale spacing patterns was investigated by playing back, at various amplitudes, recordings of these vocalizations to resident males. The aggressive threshold of a male for each call type was defined as the lowest amplitude of playback that elicited encounter calls. Pacific treefrogs consistently had lower aggressive thresholds to playbacks of encounter calls than to playbacks of advertisement calls. Aggressive thresholds to playbacks of advertisement calls were positively correlated with the maximum amplitude of the advertisement calls of neighboring males, as measured at the position of the focal male.     

FRIENDS OF WILDLIFE SOUND AT NSA

ABSTRACT

The problem of categorization arises in any classification system because classes should be discrete while the characteristics of most natural objects and aspects of nature are more or less gradual. In systematics, this problem usually is solved by creating several levels of categories, such as class, order, family, genus and species. In the existing killer whale discrete call classification, only two levels occur—call type and call subtype. In this paper we describe structural categories at a broader level than call type in the discrete sounds of killer whales and compare these categories between and within vocal clans in a community of resident killer whales from Southeast Kamchatka, Russian Far East, and also with killer whales outside this community. We found four main classes of discrete calls in the repertoire of resident killer whales from Southeast Kamchatka. The calls of Southeast Kamchatka transient killer whales and Sakhalin killer whales do not fall into these classes. This suggests that the resident killer whale community from Southeast Kamchatka has some rules defining the structure of calls which are typical for this community. Consequently, all resident killer whales from Southeast Kamchatka can be said to share the same vocal tradition.     

Analysis of a Blainville's beaked whale's movement response to playback of killer whale vocalizations

Increasing evidence links exposure to Navy sonar with certain mass stranding events of deep diving beaked whales. Although the cause of these strandings is unknown, one theory suggests that the animals confuse the sonar signals with vocalizations of killer whales, a known predator. Here we analyze the movement patterns of a tagged female Blainville's beaked whale in reaction to playback of killer whale predation calls. During a deep foraging dive, the whale was exposed to a playback of killer whale vocalizations with the source level slowly increased until the whale prematurely ceased foraging. The heading data from the tag were analyzed using a rotation test with a likelihood ratio calculated for a nonparametric kernel density estimate. We found a significant difference (P < 0.005) in the distribution of Δheading (the change in heading averaged over 200 s) after the cessation of the killer whale playback. A test of the angular standard deviation (SD) of the Δheading showed that after the playback, the SD was significantly reduced (P = 0.0064), which indicates that the animal maintained a straighter than normal course for an extended period of time. The prolonged directed avoidance response observed here suggests a behavioral reaction that could pose a risk factor for stranding.     

Dual Function of Chip Calls Depending on Changing Call Rate Related to Risk Level in Territorial Pairs of White‐Eared Ground‐Sparrows

Different mechanisms have been proposed for encoding information into vocalizations: variation of frequency or temporal characteristics, variation in the rate of vocalization production, and use of different vocalization types. We analyze the effect of rate variation on the dual function of chip calls (contact and alarm) produced by White‐eared Ground‐sparrows (Melozone leucotis). We conducted an acoustic playback experiment where we played back 1 min of four chip call rates (12, 36, 60, 84 calls/min). We measured the response of territorial pairs using behavioral responses, and fine structural features of calls produced in response to those playbacks. White‐eared Ground‐sparrows showed more intense behavioral responses to higher than lower call rate playbacks. Both individuals of the pair approached the source of the playback stimulus faster, produced the first vocalization faster, produced more vocalizations, and spent more time close to the stimulus in higher call rate than in lower call rate playbacks. Frequency and duration characteristics of calls (chip and tseet) were similar in response to all call rate playbacks. Our playback experiment elicited different intensity of behavioral responses, suggesting that risk‐based information is encoded in call rate. Our results suggest that variation in the rate of chip call production serves a dual function in this species; calls are used at lower rates for pair contact and at higher rates for alarm/mobbing signals.