Graeme M. Ellis

Linking killer whale survival and prey abundance: food limitation in the oceans' apex predator?

Killer whales (Orcinus orca) are large predators that occupy the top trophic position in the worlds oceans and as such may have important roles in marine ecosystem dynamics. Although the possible top-down effects of killer whale predation on populations of their prey have received much recent attention, little is known of how the abundance of these predators may be limited by bottom-up processes. Here we show, using 25 years of demographic data from two populations of fish-eating killer whales in the northeastern Pacific Ocean, that population trends are driven largely by changes in survival, and that survival rates are strongly correlated with the availability of their principal prey species, Chinook salmon (Oncorhynchus tshawytscha). Our results suggest that, although these killer whales may consume a variety of fish species, they are highly specialized and dependent on this single salmonid species to an extent that it is a limiting factor in their population dynamics. Other ecologically specialized killer whale populations may be similarly constrained to a narrow range of prey species by culturally inherited foraging strategies, and thus are limited in their ability to adapt rapidly to changing prey availability.

PCB-associated changes in mRNA expression in killer whales (Orcinus orca) from the NE Pacific Ocean.

Killer whales in the NE Pacific Ocean are among the worlds most PCB-contaminated marine mammals, raising concerns about implications for their health. Sixteen health-related killer whale mRNA transcripts were analyzed in blubber biopsies collected from 35 free-ranging killer whales in British Columbia using real-time quantitative polymerase chain reaction. We observed PCB-related increases in the expression of five gene targets, including the aryl hydrocarbon receptor (AhR; r(2) = 0.83; p < 0.001), thyroid hormone α receptor (TRα; r(2) = 0.64; p < 0.001), estrogen α receptor (ERα; r(2) = 0.70; p < 0.001), interleukin 10 (IL-10; r(2) = 0.74 and 0.68, males and females, respectively; p < 0.001), and metallothionein 1 (MT1; r(2) = 0.58; p < 0.001). Best-fit models indicated that population (dietary preference), age, and sex were not confounding factors, except for IL-10, where males differed from females. While the population-level consequences are unclear, the PCB-associated alterations in mRNA abundance of such pivotal end points provide compelling evidence of adverse physiological effects of persistent environmental contaminants in these endangered killer whales.

Geographic and temporal dynamics of a global radiation and diversification in the killer whale

Global climate change during the Late Pleistocene periodically encroached and then released habitat during the glacial cycles, causing range expansions and contractions in some species. These dynamics have played a major role in geographic radiations, diversification and speciation. We investigate these dynamics in the most widely distributed of marine mammals, the killer whale (Orcinus orca), using a global data set of over 450 samples. This marine top predator inhabits coastal and pelagic ecosystems ranging from the ice edge to the tropics, often exhibiting ecological, behavioural and morphological variation suggestive of local adaptation accompanied by reproductive isolation. Results suggest a rapid global radiation occurred over the last 350 000 years. Based on habitat models, we estimated there was only a 15% global contraction of core suitable habitat during the last glacial maximum, and the resources appeared to sustain a constant global effective female population size throughout the Late Pleistocene. Reconstruction of the ancestral phylogeography highlighted the high mobility of this species, identifying 22 strongly supported long‐range dispersal events including interoceanic and interhemispheric movement. Despite this propensity for geographic dispersal, the increased sampling of this study uncovered very few potential examples of ancestral dispersal among ecotypes. Concordance of nuclear and mitochondrial data further confirms genetic cohesiveness, with little or no current gene flow among sympatric ecotypes. Taken as a whole, our data suggest that the glacial cycles influenced local populations in different ways, with no clear global pattern, but with secondary contact among lineages following long‐range dispersal as a potential mechanism driving ecological diversification.

Estimating Carrying Capacity for Sea Otters in British Columbia

Abstract We estimated carrying capacity for sea otters (Enhydra lutris) in the coastal waters of British Columbia, Canada, by characterizing habitat according to the complexity of nearshore intertidal and sub-tidal contours. We modeled the total area of complex habitat on the west coast of Vancouver Island by first calculating the complexity of the Checleset Bay–Kyuquot Sound (CB–KS) region, where sea otters have been at equilibrium since the mid-1990s. We then identified similarly complex areas on the west coast of Vancouver Island (WCVI model), and adapted the model to identify areas of similar complexity along the entire British Columbia coast (BC model). Using survey data from the CB–KS region, we calculated otter densities for the habitat predicted by the 2 models. The density estimates for CB–KS were 3.93 otters/km2 and 2.53 otters/km2 for the WCVI and BC models, respectively, and the resulting 2 estimates of west coast of Vancouver Island complex habitat carrying capacity were not significantly different (WCVI model: 5,123, 95% CI = 3,337–7,104; BC model: 4,883, 95% CI = 3,223–6,832). The BC model identified the region presently occupied by otters on the central British Columbia coast, but the amount of coast-wide habitat it predicted (5,862 km2) was relatively small, and the associated carrying capacity estimate (14,831, 95% CI = 9,790–20,751) was low compared to historical accounts. We suggest that our model captured a type of high-quality or optimum habitat prevalent on the west coast of Vancouver Island, typified by the CB–KS region, and that suitable sea otter habitat elsewhere on the coast must include other habitat characteristics. We therefore calculated a linear, coast-wide carrying capacity of 52,459 sea otters (95% CI = 34,264–73,489)—a more realistic upper limit to sea otters in British Columbia. Our carrying capacity estimates are helping set population recovery targets for sea otters in Canada, and our habitat predictions represent a first step in Critical Habitat identification. This habitat-based approach to estimating carrying capacity is likely suitable for other nonmigratory, density-dependent species.

Kin-directed food sharing promotes lifetime natal philopatry of both sexes in a population of fish-eating killer whales, Orcinus orca

The vast majority of social animals exhibit sex-biased dispersal as a strategy to reduce kin competition and avoid inbreeding. Piscivorous ‘resident’ killer whales, Orcinus orca, of the eastern North Pacific, however, are unusual in that both sexes remain philopatric throughout life, forming highly stable, multigeneration matrilines that are closed to immigration. We conducted a 12-year study documenting extensive cooperative prey sharing within these matrilines, and hypothesized that extreme natal philopatry in resident killer whales arose due to inclusive fitness benefits gained by provisioning maternal kin. We found that prey sharing was nonreciprocal, and even though whales routinely foraged in mixed associations containing multiple matrilines, prey sharing among individuals belonging to different matrilines was very infrequent. Furthermore, maternal relatedness was a significant predictor of the frequency of prey sharing between individuals, with close maternal kin sharing more often than distant relatives or nonkin. Adult females were much more likely to share prey than adult males or subadults, probably because they mainly provisioned their offspring. However, food sharing was not limited solely to maternal care; all age–sex classes engaged in this behaviour by sharing with close maternal relatives, such as siblings and mothers. We also investigated the frequency of prey sharing between mothers and their offspring as a function of offspring sex and age, and found that maternal food sharing with daughters declined after daughters reached reproductive maturity, which could help to explain matriline fission events. The evolution of kin-directed food sharing requires the ability to reliably discriminate kin, which resident killer whales likely achieve through social familiarity and vocal dialect recognition. We propose that lifetime philopatry of both sexes has been selectively favoured in this population due to the inclusive fitness benefits of kin-directed food sharing, a cooperative behaviour that may also inhibit dispersal by reducing resource competition among kin.

You Are What You Eat: Foraging Specializations and Their Influence on the Social Organization and Behavior of Killer Whales

The feeding ecology of predators can have a profound effect on their life history and behaviour. The killer whale—the apex marine predator—has a cosmopolitan distribution throughout the world’s oceans. Globally, it is a generalist predator with a diverse diet, but regionally, different socially and genetically isolated killer whale populations can have highly specialized foraging strategies involving only a few types of prey. In the eastern North Pacific, the three sympatric killer whale lineages have distinct dietary specializations: one feeds primarily on marine mammals, another on salmon, and the third appears to specialize on sharks. These ecological specializations are associated with distinct patterns of seasonal distribution, group size, social organization, foraging behavior, and acoustic activity. Divergent foraging strategies may have played a major role in the social isolation and genetic divergence of killer whale populations.

Fine-scale foraging movements by fish-eating killer whales (Orcinus orca) relate to the vertical distributions and escape responses of salmonid prey (Oncorhynchus spp.)

BackgroundWe sought to quantitatively describe the fine-scale foraging behavior of northern resident killer whales (Orcinus orca), a population of fish-eating killer whales that feeds almost exclusively on Pacific salmon (Oncorhynchus spp.). To reconstruct the underwater movements of these specialist predators, we deployed 34 biologging Dtags on 32 individuals and collected high-resolution, three-dimensional accelerometry and acoustic data. We used the resulting dive paths to compare killer whale foraging behavior to the distributions of different salmonid prey species. Understanding the foraging movements of these threatened predators is important from a conservation standpoint, since prey availability has been identified as a limiting factor in their population dynamics and recovery.ResultsThree-dimensional dive tracks indicated that foraging (N = 701) and non-foraging dives (N = 10,618) were kinematically distinct (Wilks’ lambda: λ16 = 0.321, P < 0.001). While foraging, killer whales dove deeper, remained submerged longer, swam faster, increased their dive path tortuosity, and rolled their bodies to a greater extent than during other activities. Maximum foraging dive depths reflected the deeper vertical distribution of Chinook (compared to other salmonids) and the tendency of Pacific salmon to evade predators by diving steeply. Kinematic characteristics of prey pursuit by resident killer whales also revealed several other escape strategies employed by salmon attempting to avoid predation, including increased swimming speeds and evasive maneuvering.ConclusionsHigh-resolution dive tracks reconstructed using data collected by multi-sensor accelerometer tags found that movements by resident killer whales relate significantly to the vertical distributions and escape responses of their primary prey, Pacific salmon.