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

Spatial and temporal occurrence of killer whale ecotypes off the outer coast of Washington State, USA

Three killer whale Orcinus orca ecotypes inhabit the northeastern Pacific: residents, transients, and offshores. To investigate intraspecific differences in spatial and temporal occur-rence off the outer coast of Washington State, USA, 2 long-term acoustic recorders were deployed from July 2004 to August 2013: one off the continental shelf in Quinault Canyon (QC) and the other on the shelf, off Cape Elizabeth (CE). Acoustic encounters containing pulsed calls were analyzed for call types attributable to specific ecotypes, as no calls are shared between ecotypes. Both sites showed killer whale presence year-round, although site CE had a higher number of days with en- counters overall. Transients were the most common ecotype at both sites and were encountered mainly during the spring and early summer. Residents were encountered primarily at site CE and showed potential seasonal segregation between the 2 resident communities, with northern resi- dents present mainly during summer and early fall when southern residents were not encountered. Offshore encounters were higher at site QC, with little evidence for seasonality. Spatial and temporal variability of residents and transients matches the distribution of their prey and can potentially be used for further inferences about prey preferences for different transient groups

Genome-wide association and Mendelian randomisation analysis provide insights into the pathogenesis of heart failure

Heart failure (HF) is a leading cause of morbidity and mortality worldwide. A small proportion of HF cases are attributable to monogenic cardiomyopathies and existing genome-wide association studies (GWAS) have yielded only limited insights, leaving the observed heritability of HF largely unexplained. We report results from a GWAS meta-analysis of HF comprising 47,309 cases and 930,014 controls. Twelve independent variants at 11 genomic loci are associated with HF, all of which demonstrate one or more associations with coronary artery disease (CAD), atrial fibrillation, or reduced left ventricular function, suggesting shared genetic aetiology. Functional analysis of non-CAD-associated loci implicate genes involved in cardiac development (MYOZ1, SYNPO2L), protein homoeostasis (BAG3), and cellular senescence (CDKN1A). Mendelian randomisation analysis supports causal roles for several HF risk factors, and demonstrates CAD-independent effects for atrial fibrillation, body mass index, and hypertension. These findings extend our knowledge of the pathways underlying HF and may inform new therapeutic strategies

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

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