Marilyn E. Dahlheim

Low worldwide genetic diversity in the killer whale (Orcinus orca): implications for demographic history.

A low level of genetic variation in mammalian populations where the census population size is relatively large has been attributed to various factors, such as a naturally small effective population size, historical bottlenecks and social behaviour. The killer whale (Orcinus orca) is an abundant, highly social species with reduced genetic variation. We find no consistent geographical pattern of global diversity and no mtDNA variation within some regional populations. The regional lack of variation is likely to be due to the strict matrilineal expansion of local populations. The worldwide pattern and paucity of diversity may indicate a historical bottleneck as an additional factor.

Social cohesion among kin, gene flow without dispersal and the evolution of population genetic structure in the killer whale (Orcinus orca)

In social species, breeding system and gregarious behavior are key factors influencing the evolution of large‐scale population genetic structure. The killer whale is a highly social apex predator showing genetic differentiation in sympatry between populations of foraging specialists (ecotypes), and low levels of genetic diversity overall. Our comparative assessments of kinship, parentage and dispersal reveal high levels of kinship within local populations and ongoing male‐mediated gene flow among them, including among ecotypes that are maximally divergent within the mtDNA phylogeny. Dispersal from natal populations was rare, implying that gene flow occurs without dispersal, as a result of reproduction during temporary interactions. Discordance between nuclear and mitochondrial phylogenies was consistent with earlier studies suggesting a stochastic basis for the magnitude of mtDNA differentiation between matrilines. Taken together our results show how the killer whale breeding system, coupled with social, dispersal and foraging behaviour, contributes to the evolution of population genetic structure.

VARIATION IN FEEDING VOCALIZATIONS OF HUMPBACK WHALES MEGAPTERA NOVAEANGLIAE FROM SOUTHEAST ALASKA

ABSTRACT Vocalizations of feeding humpback whales from southeast Alaska were analyzed to characterize quantitatively the predominant vocalization associated with feeding and assess variation among vocalizations. Whales uttered series of cries similar in acoustic structure to those described previously as stereotyped, rhythmic ‘feeding calls’. Individual cries ranged in duration from 0.4 to 8.2 sec. (median = 2.6 sec). Cries typically had a short, strongly frequency modulated (FM) introductory and ending component (labeled Section A and C, respectively). Cries had relatively little FM over the main body of the call (labeled Section B) which ranged in fundamental frequency from 360 to 988 Hz (median = 553 Hz) and sometimes exhibited a frequency oscillation over a bandwidth of approximately 16 to 65 Hz. Principle components analysis indicated that most variation in the data-set (over 35%) could be attributed to measures of absolute frequency, however a substantial amount of variation was also due to other acoustic parameters such as duration, frequency oscillation and average slope of cry sections A and C. Within series, cries were stereotyped and varied little, whereas there was statistically significant variation in cries between series. Furthermore, overlapping cries, which are considered to represent the vocalizations of different individuals, varied significantly. These results suggest that whales may produce individually specific cries, and we propose two alternative hypotheses to account for our observations: 1) cries may carry individual signature information, and 2) simultaneously vocalizing animals may actively mismatch cries to maximize a herding effect on prey.

Killer Whale Nuclear Genome and mtDNA Reveal Widespread Population Bottleneck during the Last Glacial Maximum

Ecosystem function and resilience is determined by the interactions and independent contributions of individual species. Apex predators play a disproportionately determinant role through their influence and dependence on the dynamics of prey species. Their demographic fluctuations are thus likely to reflect changes in their respective ecological communities and habitat. Here, we investigate the historical population dynamics of the killer whale based on draft nuclear genome data for the Northern Hemisphere and mtDNA data worldwide. We infer a relatively stable population size throughout most of the Pleistocene, followed by an order of magnitude decline and bottleneck during the Weichselian glacial period. Global mtDNA data indicate that while most populations declined, at least one population retained diversity in a stable, productive ecosystem off southern Africa. We conclude that environmental changes during the last glacial period promoted the decline of a top ocean predator, that these events contributed to the pattern of diversity among extant populations, and that the relatively high diversity of a population currently in productive, stable habitat off South Africa suggests a role for ocean productivity in the widespread decline.

Effects of Masking Noise on Detection Thresholds of Killer Whales

(2006). Does intense ship noise disrupt foraging in deep-diving Cuviers beaked whales (Ziphius cavirostris)? sounds induce acoustic trauma in cephalopods. (2006). Long-range effects of airgun noise on marine mammals: responses as a function of received sound level and distance. Cambridge, UK.

Population genomics of the killer whale indicates ecotype evolution in sympatry involving both selection and drift

The evolution of diversity in the marine ecosystem is poorly understood, given the relatively high potential for connectivity, especially for highly mobile species such as whales and dolphins. The killer whale (Orcinus orca) has a worldwide distribution, and individual social groups travel over a wide geographic range. Even so, regional populations have been shown to be genetically differentiated, including among different foraging specialists (ecotypes) in sympatry. Given the strong matrifocal social structure of this species together with strong resource specializations, understanding the process of differentiation will require an understanding of the relative importance of both genetic drift and local adaptation. Here we provide a high‐resolution analysis based on nuclear single‐nucleotide polymorphic markers and inference about differentiation at both neutral loci and those potentially under selection. We find that all population comparisons, within or among foraging ecotypes, show significant differentiation, including populations in parapatry and sympatry. Loci putatively under selection show a different pattern of structure compared to neutral loci and are associated with gene ontology terms reflecting physiologically relevant functions (e.g. related to digestion). The pattern of differentiation for one ecotype in the North Pacific suggests local adaptation and shows some fixed differences among sympatric ecotypes. We suggest that differential habitat use and resource specializations have promoted sufficient isolation to allow differential evolution at neutral and functional loci, but that the process is recent and dependent on both selection and drift.

Phylogenomics of the killer whale indicates ecotype divergence in sympatry

For many highly mobile species, the marine environment presents few obvious barriers to gene flow. Even so, there is considerable diversity within and among species, referred to by some as the ‘marine speciation paradox’. The recent and diverse radiation of delphinid cetaceans (dolphins) represents a good example of this. Delphinids are capable of extensive dispersion and yet many show fine-scale genetic differentiation among populations. Proposed mechanisms include the division and isolation of populations based on habitat dependence and resource specializations, and habitat release or changing dispersal corridors during glacial cycles. Here we use a phylogenomic approach to investigate the origin of differentiated sympatric populations of killer whales (Orcinus orca). Killer whales show strong specialization on prey choice in populations of stable matrifocal social groups (ecotypes), associated with genetic and phenotypic differentiation. Our data suggest evolution in sympatry among populations of resource specialists.

Ecological aspects of transient killer whales Orcinus orca as predators in southeastern Alaska

Abstract In this study we present empirical data on predator numbers, movements and area usage, and predation obtained from tracking transient killer whales Orcinus orca throughout the inland waters of southeastern Alaska, USA. During 1991-2007, we documented 155 transient killer whales via photo-identification methodology within the large study area (27,808 km2). Transient killer whales were distributed throughout southeastern Alaska and were present during all seasons, although not all individuals were seen every year. Resighting data suggested that within southeastern Alaska, maternal groups may partition area usage of their environment. By following whales for 1,467 km, we calculated a mean travel speed of 7.2 km/hour with mean daily movements of 134 km ± 88 km/24 hours and ranging within 59-240 km/24 hours. Photographic matches demonstrated that most of the transient killer whales (86%) identified in southeastern Alaska also utilized British Columbia and Washington State waters. In contrast, photographic matches between whales in southeastern Alaska and whales seen off of California, USA, were rare, suggesting that different transient killer whale stocks occupy these two regions. Transient killer whales preyed upon Dalls porpoise Phocoenoides dalli, Pacific white-sided dolphins Lagenoryhncus obliquidens, harbor porpoise Phocoena phocoena, minke whales Balaenoptera acutorostrata, Steller sea lions Eumetopias jubatus, harbor seals Phoca vitulina and seabirds. Potential prey species that were available, but not targeted, included humpback whales Megaptera novaeangliae, elephant seals Mirounga angustirostris and sea otters Enhydra lutris. Prey-handling techniques varied depending on the prey being targeted with no evidence of prey specialization. During 114 encounters totaling 332.5 hours of direct observations of transient killer whales, we documented 36 predation events for a calculated kill rate of 0.62 prey items/24-hour period/whale. The data we present in this article provide a foundation of transient killer whale ecology aimed at improving our ability to understand the impact of transient killer whale predation on southeastern Alaska prey populations.

Balancing and Directional Selection at Exon-2 of the MHC DQB1 Locus among Populations of Odontocete Cetaceans

The diversity of exon-2 (peptide-binding region) of the DQB1 locus (Class II, major histocompatibility complex, MHC) was investigated on an extended sample of populations of three focal cetacean species (two sibling delphinid species and another in the same family). We tested the hypothesis that dolphin populations with a worldwide distribution across different habitats and geographic regions will be under differential selective pressure by comparing DQB1 variation with variation at neutral markers and by investigating putative functional residues within the exon-2 sequence at the population level. Variation at the DQB1 locus was not correlated to neutral differentiation (assessed by comparison with microsatellite DNA markers), and overall F(ST) values were significantly lower for the MHC locus, consistent with expectations for balancing selection. Measures of heterozygosity and d(n)/d(s) ratios were also consistent with balancing selection. However, outliers in the F(ST) comparisons and the analysis of putative functional residues suggested incidences of directional selection in local populations.

The Sounds of Sperm Whale Calves

The development of the click sounds of sperm whales (Physeter catodon) has been investigated through comparisons of these vocalizations from calves of different sizes. The observations include sounds from four small stranded calves held for short periods in aquaria at Miami, Florida, and Seattle, Washington, and in a bay on Long Island, New York. These vocalizations were compared with those of larger calves encountered at sea. All the calves produced typical sperm whale sounds -- clicks with broadband spectra, often produced in short series. Vocalizations from the smaller calves sometimes included slightly noisy, tonal components -- similar noisy click sounds were also occasionally heard in the presence of calves at sea, and they were interpreted as the result of improperly formed clicks. The smallest calves produced few temporally repetitive click patterns, but the larger (older) ones produced sequences with stereotyped “coda”-like temporal patterns. The use of such patterned click sequences increased with the apparent age of the calves. In the larger calves, the sounds appeared more organized in sequences similar to the communicative signals of adults. None of the calves appeared to use their sounds in ways that were related to echolocation.