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

Vocal Repertoire, Social Structure and Feeding Preferences of Australian and Antarctic Killer Whales (Orcinus orca)

This thesis aimed to improve the understanding of killer whale populations in the Australian and Antarctic regions. Visual surveys, photo-identification and passive acoustic recordings were combined to study these populations. This study describes the call repertoire of killer whales found in Australian and Antarctic waters, presents an acoustic comparison between sympatric ecotypes in Antarctic waters and investigates the sociality and feeding preferences of killer whales in the Bremer Sub-Basin

Physical constraints of cultural evolution of dialects in killer whales

Data collection was supported by a variety of organizations, including the Russian Fund for the Fundamental Research (Grant No. 15-04-05540), the Rufford Small Grants Fund, Whale and Dolphin Conservation, the Fundação para a Ciência e a Tecnologia (Grant No. SFRH/BD/30303/2006), Russell Trust Award of the University of St. Andrews, the Office of Naval Research, the Icelandic Research Fund (i. Rannsóknasjóður), the National Geographic Society Science and Exploration Europe (Grant No. GEFNE65-12), Vancouver Aquarium Marine Science Centre, the Canadian Ministry of Fisheries and Oceans, and the North Gulf Oceanic Society.Odontocete sounds are produced by two pairs of phonic lips situated in soft nares below the blowhole; the right pair is larger and is more likely to produce clicks, while the left pair is more likely to produce whistles. This has important implications for the cultural evolution of delphinid sounds: the greater the physical constraints, the greater the probability of random convergence. In this paper the authors examine the call structure of eight killer whale populations to identify structural constraints and to determine if they are consistent among all populations. Constraints were especially pronounced in two-voiced calls. In the calls of all eight populations, the lower component of two-voiced (biphonic) calls was typically centered below 4 kHz, while the upper component was typically above that value. The lower component of two-voiced calls had a narrower frequency range than single-voiced calls in all populations. This may be because some single-voiced calls are homologous to the lower component, while others are homologous to the higher component of two-voiced calls. Physical constraints on the call structure reduce the possible variation and increase the probability of random convergence, producing similar calls in different populations.PostprintPeer reviewe

Correlates of variability in killer whale stereotyped call repertoires.

Killer whales vocalisations include repertoires of stereotyped call types (Ford 1984). There is strong evidence that these vocalisations are learnt (Hoelzel and Osborne 1986; Bain 1989; Deecke et al. 2000; Yurk et al. 2002). Call types can be group specific or shared amongst a number of groups, depending upon the social structure of the population or the call type (Ford 1991; Deecke 2003). It is thought that these call types function in the group cohesion and coordination (Hoelzel and Osborne 1986; Ford 1989 1991; Miller 2000, 2002). Some call types contain two overlapping, independently modulated, components each having different transmission properties (Miller 2002), these call types have a higher estimated active space than single- component call types (Miller 2006).This thesis investigates the evolution of these call type repertoires, focusing on call type usage and structure of the Southern Resident population over a period of 27 years, but including comparisons with other populations. I present evidence of hetero-specific mimicry and further evidence for vocal production and usage learning in killer whales. I compared the relative frequency of use of call types between two time periods (1977-81 & 2001-2003) and between contexts, such as direction changes with directional travel and multi-pod aggregations with single pods. I found a strong correlation of relative call type usage for each pod between the two time periods and each pod was easily acoustically distinguishable from the other two pods in both periods. The implications of these results for a role of call type repertoires in kin recognition are discussed. The least cohesive pod produced a significantly higher proportion of two-component call types than the other two more cohesive pods. Lone whales separated from their pod also used a rare subset of two-component call types rather than their pod's main call types. In recordings of multi-pod aggregations I recorded a high proportion of the same subset of two-component call types not commonly produced by any of the three pods individually, these call types were used in significantly higher proportions when all three pods were converging or socialising rather than travelling. These contextual correlates suggest that call types are selectively used and shared between groups based on their transmission properties. Each of the Southern Resident pods, J к and L, were found to increase the duration of their primary call type 10-15% in the presence of vessel noise in recordings made between 2001 & 2003. This response was not detected in recording from two earlier time periods, (1977 or 1989-1992). This change in behaviour correlated with an increase over the past decade in vessel presence around this population and may be an anti-masking strategy. I also compared the range and mean minimum and maximum fundamental frequency of the call types within the repertoires of six North Pacific killer whale populations. There was a degree of homogeneity in the range of call type fundamental frequencies within the repertoires of populations of the same ecotype, but differences between ecotypes. Offshore call types generally had a higher pitch fundamental frequency than transient or resident call types. All three resident populations had call types in their repertoires that had a maximum fundamental frequency 3 kHz higher than found in any transient call type

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

Funding for data collection was provided by the BBC Natural History Unit, Fundação para a Ciência e a Tecnologia (grant number SFRH/BD/30303/2006), the Icelandic Research Fund (i. Rannsóknasjóður) through a START Postdoctoral Fellowship (grant number 120248042) and a Project Grant (grant number 163060‐051), the National Geographic Global Exploration Fund (grant number GEFNE65‐12), a Marie Curie International Incoming Fellowship (project number 297116), the Office of Naval Research (grant number N00014‐08‐1‐0984), and a Russell Trust Award from the University of St. Andrews.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.PostprintPeer reviewe

Comparing call-based versus subunit-based methods for categorizing Norwegian killer whale, Orcinus orca, vocalizations

Author Posting. © The Author(s), 2010. This is the author's version of the work. It is posted here by permission of Elsevier B.V. for personal use, not for redistribution. The definitive version was published in Animal Behaviour 81 (2011): 377-386, doi:10.1016/j.anbehav.2010.09.020.Students of animal communication face significant challenges when deciding how to categorise calls into subunits, calls, and call series. Here, we use algorithms designed to parse human speech to test different approaches for categorising calls of killer whales. Killer whale vocalisations have traditionally been categorised by humans into discrete call types. These calls often contain internal spectral shifts, periods of silence, and synchronously produced low and high frequency components, suggesting that they may be composed of subunits. We describe and compare three different approaches for modelling Norwegian killer whale calls. The first method considered the whole call as the basic unit of analysis. Inspired by human speech processing techniques, the second and third methods represented the calls in terms of subunits. Subunits may provide a more parsimonious approach to modelling the vocal stream since (1) there were fewer subunits than call types; (2) nearly 75% of all call types shared at least one subunit. We show that contour traces from stereotyped Norwegian killer whale calls yielded similar automatic classification performance using either whole calls or subunits. We also demonstrate that subunits derived from Norwegian stereotyped calls were detected in some Norwegian variable (non-stereotyped) calls as well as the stereotyped calls of other killer whale populations. Further work is required to test whether killer whales use subunits to generate and categorize their vocal repertoire.The undergraduate students were supported by the Massachusetts Institute of Technology Undergraduate Research Opportunities Program office and the Ocean Life Institute (OLI) at the Woods Hole Oceanographic Institution (WHOI). Field work was financed by the OLI, National Geographic Society and WWF Sweden. A. D. Shapiro was funded by a National Defense Science and Engineering Graduate Fellowship and the WHOI Academic Programs Office

The social organization of resident-type killer whales (Orcinus orca) in Avacha Gulf, Northwest Pacific, as revealed through association patterns and acoustic similarity

Abstract Northeast Pacific resident-type killer whales (Orcinus orca) are known to form stable associations based on kinship between maternal relatives (matrilines) with a system of vocal dialects thought to reflect kinship relationships. We analyzed association patterns and acoustic similarity to study the social organization of killer whales in Avacha Gulf (Kamchatka, Russia), in the Northwest Pacific. The resident-type killer whales of Avacha Gulf formed temporally stable units that included maternal relatives with no dispersal observed. Acoustically, the killer whale community of Avacha Gulf was characterized by a system of dialects comparable to the communities of Northeast Pacific residenttype killer whales. Different units rarely associated with each other and these associations were nonrandom. Associations at different spatial levels did not always coincide with each other and with the patterns of acoustic similarity. Associations between units could change quickly irrespective of kinship relationships. The vocal dialect of a unit, which is more stable than the association patterns between units, might better reflect the overall kinship relationships. The stability and frequency of associations between units depended on the number of mature males in a unit, which could contribute to differences in the speed of change in vocal dialects and association patterns

How Selective is Social Learning in Dolphins?

Social learning is an important aspect of dolphin social life and dolphin behavioral development. In addition to vocal social learning, dolphins discover behaviors for foraging, play, and social interactions by observing other members of their social group. But dolphins neither indiscriminately observe nor mindlessly mimic other dolphins. To the contrary, dolphin calves are quite selective in their choices of who to observe and/or imitate. Calves are most likely to learn foraging behaviors from their mothers, but they are more likely to watch and reproduce the play behaviors of other calves than the play behaviors of adult dolphins (including their mothers). But not all calves are equally likely to be good models. Instead, calves are more likely to observe and mimic the behaviors of other calves that are producing either novel behaviors or more complex forms of behaviors that the observing calf already knows. As a result, there is a general tendency for calves to watch and learn from calves that are older than they are. But differences in age are only part of the story. In fact, dolphin personality may be more important than dolphin age in determining the efficacy of a model