Olga A. Filatova

Call diversity in the North Pacific killer whale populations: implications for dialect evolution and population history

Although killer whale, Orcinus orca, dialects have been studied in detail in several populations, little attempt has been made to compare dialect characteristics between populations. In this study we investigated geographical variation in monophonic and biphonic calls among four resident populations from the North Pacific Ocean: Northern and Southern residents from British Columbia and Washington State, southern Alaska residents, and eastern Kamchatka residents. We tested predictions generated by the hypothesis that call variation across populations is the result of an accumulation of random errors and innovation by vertical cultural transmission. Call frequency contours were extracted and compared using a dynamic time-warping algorithm. We found that the diversity of monophonic calls was substantially higher than that of biphonic calls for all populations. Repertoire diversity appeared to be related to population size: in larger populations, monophonic calls were more diverse and biphonic calls less diverse. We suggest that the evolution of both monophonic and biphonic calls is caused by an interaction between stochastic processes and directional selection, but the relative effect of directional selection is greater for biphonic calls. Our analysis revealed no direct correlation between call repertoire similarity and geographical distance. Call diversity within predefined call categories, types and subtypes, showed a high degree of correspondence between populations. Our results indicate that dialect evolution is a complex process influenced by an interaction among directional selection, horizontal transmission and founder effects. We suggest several scenarios for how this might have arisen and the implications of these scenarios for call evolution and population history.

THE STRUCTURE OF THE DISCRETE CALL REPERTOIRE OF KILLER WHALES ORCINUS ORCA FROM SOUTHEAST KAMCHATKA

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.

Cross-cultural and cross-ecotype production of a killer whale 'excitement' call suggests universality

Facial and vocal expressions of emotion have been found in a number of social mammal species and are thought to have evolved to aid social communication. There has been much debate about whether such signals are culturally inherited or are truly biologically innate. Evidence for the innateness of such signals can come from cross-cultural studies. Previous studies have identified a vocalisation (the V4 or ‘excitement’ call) associated with high arousal behaviours in a population of killer whales in British Columbia, Canada. In this study, we compared recordings from three different socially and reproductively isolated ecotypes of killer whales, including five vocal clans of one ecotype, each clan having discrete culturally transmitted vocal traditions. The V4 call was found in recordings of each ecotype and each vocal clan. Nine independent observers reproduced our classification of the V4 call from each population with high inter-observer agreement. Our results suggest the V4 call may be universal in Pacific killer whale populations and that transmission of this call is independent of cultural tradition or ecotype. We argue that such universality is more consistent with an innate vocalisation than one acquired through social learning and may be linked to its apparent function of motivational expression.

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.

Is killer whale dialect evolution random

The killer whale is among the few species in which cultural change accumulates over many generations, leading to cumulative cultural evolution. Killer whales have group-specific vocal repertoires which are thought to be learned rather than being genetically coded. It is supposed that divergence between vocal repertoires of sister groups increases gradually over time due to random learning mistakes and innovations. In this case, the similarity of calls across groups must be correlated with pod relatedness and, consequently, with each other. In this study we tested this prediction by comparing the patterns of call similarity between matrilines of resident killer whales from Eastern Kamchatka. We calculated the similarity of seven components from three call types across 14 matrilines. In contrast to the theoretical predictions, matrilines formed different clusters on the dendrograms made by different calls and even by different components of the same call. We suggest three possible explanations for this phenomenon. First, the lack of agreement between similarity patterns of different components may be the result of constraints in the call structure. Second, it is possible that call components change in time with different speed and/or in different directions. Third, horizontal cultural transmission of call features may occur between matrilines.

Individual and Sexual Differences in the Calls of the Monomorphic White-Faced Whistling Duck Dendrocygna viduata

Abstract. The White-faced Whistling Duck is a waterfowl species lacking any differences in ornamentation, coloration, size or behavior between the sexes. For distant communication, this species uses loud whistles. We analyzed 12 spectral parameters of 344 whistles from 23 captive adult ducks (14 males and 9 females). Discriminant analysis showed 94% correct assignment to an individual (N = 279 calls from 14 birds; 15–22 calls per bird). Separately for 8 males (162 calls) and for 6 females (117 calls), discriminant analysis showed 99% and 93% correct assignment to individuals respectively. Discriminant analysis for sex (N = 86; 3 calls from each of 14 males and 5 calls from each of 9 females) showed 100% correct assignment. Intersexual differences were governed by frequency parameters, the values of which were significantly higher in females than in males. Cluster analysis showed that differences between sexes were expressed significantly more strongly than the individual differences. The fact that the “acoustical keys” differed as regards the identification of individual birds or their sex may significantly enhance the reliability of acoustical recognition systems in the White-faced Whistling Duck. The data are discussed in the context of the biology of the White-faced Whistling Duck and significant intersexual differences in syringial and tracheal anatomy, which may be responsible for the sharp distinctions between the sexes in the calls of this species.

Ultrasonic whistles of killer whales (Orcinus orca) recorded in the North Pacific (L)

Ultrasonic whistles were previously found in North Atlantic killer whales and were suggested to occur in eastern North Pacific killer whales based on the data from autonomous recorders. In this study ultrasonic whistles were found in the recordings from two encounters with the eastern North Pacific offshore ecotype killer whales and one encounter with the western North Pacific killer whales of unknown ecotype. All ultrasonic whistles were highly stereotyped and all but two had downsweep contours. These results demonstrate that specific sound categories can be shared by killer whales from different ocean basins.

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.

Cultural evolution of killer whale calls: background, mechanisms and consequences

Cultural evolution is a powerful process shaping behavioural phenotypes of many species including our own. Killer whales are one of the species with relatively well-studied vocal culture. Pods have distinct dialects comprising a mix of unique and shared call types; calves adopt the call repertoire of their matriline through social learning. We review different aspects of killer whale acoustic communication to provide insights into the cultural transmission and gene-culture co- evolution processes that produce the extreme diversity of group and population repertoires. We argue that the cultural evolution of killer whale calls is not a random process driven by steady error accumulation alone: temporal change occurs at different speeds in different components of killer whale repertoires, and constraints in call structure and horizontal transmission often degrade the phylogenetic signal. We discuss the implications from bird song and human linguistic studies, and propose several hypotheses of killer whale dialect evolution.

Horizontal transmission of vocal traditions in killer whale (Orcinus orca) 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.