North Atlantic killer whale Orcinus orca populations: a review of current knowledge and threats to conservation

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

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.     

Dietary variation within and between populations of northeast Atlantic killer whales,Orcinus orca,inferred from δ13C and δ15N analyses

Epidermal skin samples from eastern North Atlantic killer whales, Orcinus orca, were analyzed for carbon and nitrogen stable isotope ratios. From those, comparisons within a data set of 17 samples collected from Tysfjord, Norway, in November suggested that diet is relatively specialized during this time period at this location. There were significant differences between a small set of samples from Iceland and those collected from Norway, which had all been assigned to the same population by a previous population genetics study. The results would be consistent with matrilines feeding on either the Norwegian or Icelandic stocks of Atlantic herring (Clupea harengus). There was no significant difference within Icelandic samples between those assigned to the population known to feed upon herring and those assigned to a population hypothesized to follow Atlantic mackerel (Scomber scombrus). The greatest differences were between the epidermal samples analyzed in this study and tooth and bone collagen samples from the North Sea that were analyzed previously, which also showed significantly more variation in isotopic ratios than found for skin samples. These differences could reflect differences in turnover rate, differences in diet‐tissue fractionation and discrimination due to the amino acid composition of the different tissues, and/or greater competition promoting dietary variation between groups in the North Sea.     

Mitochondrial sequence divergence among Antarctic killer whale ecotypes is consistent with multiple species

Recently, three visually distinct forms of killer whales (Orcinus orca) were described from Antarctic waters and designated as types A, B and C. Based on consistent differences in prey selection and habitat preferences, morphological divergence and apparent lack of interbreeding among these broadly sympatric forms, it was suggested that they may represent separate species. To evaluate this hypothesis, we compared complete sequences of the mitochondrial control region from 81 Antarctic killer whale samples, including 9 type A, 18 type B, 47 type C and 7 type-undetermined individuals. We found three fixed differences that separated type A from B and C, and a single fixed difference that separated type C from A and B. These results are consistent with reproductive isolation among the different forms, although caution is needed in drawing further conclusions. Despite dramatic differences in morphology and ecology, the relatively low levels of sequence divergence in Antarctic killer whales indicate that these evolutionary changes occurred relatively rapidly and recently.     

Estimated field metabolic rates and prey requirements of resident killer whales

Killer whales are large animals that often feed in groups and thus have the potential to deplete prey populations. Determining predator energy requirements is essential to assessing whether prey availability is sufficient. This is important because one risk factor facing the endangered Southern Resident killer whale distinct population segment is limited prey availability. Body mass, field metabolic rate (FMR), and daily prey energy requirements (DPERs) were estimated for each individual in the population. FMRs were calculated from body mass, assuming they range from five to six times Kleiber‐predicted basal metabolic rates. FMRs of adults were also calculated from resident killer whale activity budgets and the metabolic cost of swimming at speeds associated with daily activities. These two methods yielded similar results. Total FMRs varied by age and sex, which is partly due to the long developmental period and sexual dimorphism in killer whales. FMRs for males (465–4,434 kg) ranged from 35,048 to 228,216 kcal/d while FMRs for females (465–3,338 kg) ranged from 35,048 to 184,444 kcal/d. DPERs were calculated from FMRs assuming a standard digestive efficiency. Corresponding DPERs ranged from 41,376 to 269,458 kcal/d and 41,376 to 217,775 kcal/d, respectively.     

Ecotype and geographical variation in carbon and nitrogen stable isotope values in western North Pacific killer whales (Orcinus orca)

Killer whales are top predators in marine trophic chains, and therefore their feeding preferences can substantially affect the abundance of species on the lower trophic levels. Killer whales are known to feed on many different types of prey from small fish to large whales, but a given killer whale population usually focuses on a specific type of prey. Stable isotope analysis is widely used to study whale diets, because direct observations are often impossible. Killer whale feeding habits in the western North Pacific are poorly studied, and the large-scale stable isotope analysis provides a unique opportunity to gain insights into the trophic links of this top predator. In this study, we compare the δ¹³C and δ¹⁵N stable isotope values from killer whale skin samples obtained in different areas of the western North Pacific from fish-eating (R-type) and mammal-eating (T-type) killer whale ecotypes. The effect of ecotype was highly significant: both carbon and nitrogen stable isotope values were lower in R-type whales than in T-type whales. The geographical variation also affected killer whale stable isotope values due to both the differences in killer whale diet and the variation in baseline stable isotope values across the study areas.     

Responses of Kamchatkan fish‐eating killer whales to playbacks of conspecific calls

Killer whales produce repertoires of stereotyped call types that are primarily transmitted vertically through social learning, leading to dialects between sympatric pods. The potential function of these call repertoires remains untested. In this study, we compared the reaction of Kamchatkan fish‐eating killer whales to the playbacks of calls from the same and different pods. After the playback of recordings from a different pod, in three cases whales changed the direction of their movement toward the boat, and in three cases no changes in direction were observed. After the playback of recordings from the same pod (either from the same or a different unit within the pod), in seven cases whales changed the direction of their movement toward the boat, and in only one case no change in direction was observed. Whales remained silent after all six playbacks of recordings from a different pod, even when they changed direction toward the boat. After the playback of recordings from the same pod, however, in all eight cases whales started calling in response. Our playback study shows that killer whales may react to playbacks of conspecific sounds and that reactions are dependent on the type of playback stimuli.     

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.     

Preliminary analysis of the social structure of killer whales,Orcinus orca,at subantarctic Marion Island

Studies of social differentiation between populations of killer whales (Orcinus orca) are important due to the cosmopolitan nature of the species, both in terms of distribution and feeding habits. The following research provides preliminary findings describing the social structure of the killer whale, Orcinus orca, population at subantarctic Marion Island. We provide evidence for consistent, observable patterns of social interactions with animals associating and disassociating in nonrandom patterns. We show that the social structure of this population may follow a new pattern of association, displaying a blend of the traditional resident/transient model displayed in the Northern Hemisphere. However, we emphasize the critical need for further studies related to the sociality, biology, and life history of Southern Ocean killer whales.     

Depredation of Patagonian toothfish (Dissostichus eleginoides) by two sympatrically occurring killer whale (Orcinus orca) ecotypes: Insights on the behavior of the rarely observed type D killer whales

Sympatric forms of ecologically distinctive killer whales (Orcinus orca) have been documented worldwide. This study focused on a new case of such sympatric occurrence of the “Crozet” type and the recently described “type D” killer whales off the Crozet Islands. The two ecotypes are morphologically and genetically distinct, but they both depredate the same local longline fishery. We used observational, photo‐identification, and fishing data, collected between 2003 and 2015, to examine differences in their patterns of depredation. Of the 828 sets where ecotype could be confirmed, type D killer whales interacted with 82 (11%) of the sets, including 9 (1%) sets that were simultaneously depredated by both ecotypes. Associations between the two types were never observed. Type D killer whales typically occurred in larger groups and both ecotypes preferentially depredated Patagonian toothfish (Dissostichus eleginoides). GLMM modeling revealed that the probability of type D depredation significantly increased throughout the study period, especially in deep waters, and photo‐identification data suggested that a subset of all individuals were habituating to depredation. This study documents the partitioning of resources between two distinct ecotypes of killer whales and provides preliminary insight into the feeding ecology of the rare type D killer whale.     

ISOLATION AND EXPRESSION OF THE INTERLEUKIN-2 GENE FROM THE KILLER WHALE, ORCINUS ORCA

Complementary DNA encoding interleukin-2 (IL-2) was isolated, cloned, and sequenced from the killer whale, Orcinus orca . The sequence of the killer whale IL-2 coding region consists of 455 nucleotides which translate into a polypeptide containing 151 amino acids. Killer whale IL-2 displays 88%, 88%, 87%, 87%, 85%, 84%, 80%, and 71% nucleotide sequence homology and 76%, 76%, 76%, 73%, 68%, 73%, 64%, and 56% amino acid homology with the cow, sheep, pig, red deer, horse, human, manatee, and mouse, respectively. High levels of killer whale recombinant IL-2 were generated by transiently transfecting killer whale IL-2/pCMV Blue plasmid DNA into cultured monkey kidney cells (Cos-1). Generation of this recombinant IL-2 will allow the development of assays useful for assessing IL-2 levels in the serum and from isolated lymphocytes of killer whales and possibly other species of cetaceans. A major contribution and significance of the in vitro expression of killer whale IL-2 exists in its potential to be used as a therapeutic agent.     

Genetic differentiation among North Atlantic killer whale populations

Population genetic structure of North Atlantic killer whale samples was resolved from differences in allele frequencies of 17 microsatellite loci, mtDNA control region haplotype frequencies and for a subset of samples, using complete mitogenome sequences. Three significantly differentiated populations were identified. Differentiation based on microsatellite allele frequencies was greater between the two allopatric populations than between the two pairs of partially sympatric populations. Spatial clustering of individuals within each of these populations overlaps with the distribution of particular prey resources: herring, mackerel and tuna, which each population has been seen predating. Phylogenetic analyses using complete mitogenomes suggested two populations could have resulted from single founding events and subsequent matrilineal expansion. The third population, which was sampled at lower latitudes and lower density, consisted of maternal lineages from three highly divergent clades. Pairwise population differentiation was greater for estimates based on mtDNA control region haplotype frequencies than for estimates based on microsatellite allele frequencies, and there were no mitogenome haplotypes shared among populations. This suggests low or no female migration and that gene flow was primarily male mediated when populations spatially and temporally overlap. These results demonstrate that genetic differentiation can arise through resource specialization in the absence of physical barriers to gene flow.     

GENETIC ANALYSIS OF KILLER WHALE (ORCINUS ORCA) HISTORICAL BONE AND TOOTH SAMPLES TO IDENTIFY WESTERN U.S. ECOTYPES

Little is known about the historical range of killer whale ecotypes in the eastern North Pacific (ENP). It is possible that ranges have shifted in the last few decades because of changes in availability of food. In particular, the southern resident ecotype, currently found primarily in the inland waters of Washington State, is known to prey extensively on salmon, which have declined in recent decades along the outer coasts of Washington, Oregon, and California. To investigate historical distributions of this and the other ENP ecotypes, samples of teeth and bones were obtained from NMFS and museum collections. We amplified a short section of the mitochondrial DNA control region that contains four diagnostic sites that differentiate between haplotypes found in ecotypes of ENP killer whales. Results did not show any southern resident haplotypes in samples from south of the Washington State inland waterways. One whale genetically identified as a northern resident extends the known southernmost distribution of the population from Oregon to California. Items of diet identified from stomach contents of six of the whales genetically identified to ecotype conformed with what is known of the feeding habits of the various ecotypes.     

Social behavior increases in multipod aggregations of southern Alaska resident killer whales (Orcinus orca)

Killer whales (Orcinus orca) are highly social and occasionally gather in large aggregations that reach 150 individuals. During 338 encounters with Southern Alaska resident killer whales, we collected 1,352 hr of behavioral data to assess the probability of various behaviors based on season, number of pods present, presence of rarely sighted pods, and number of mitochondrial DNA haplotypes present. A binomial generalized linear model was used to estimate the role of these factors in the probability of four behaviors, foraging, resting, socializing, and traveling. The presence of “rarely sighted” pods (sighted in <5% of encounters) significantly increased probability of social behavior, and significantly decreased probability of resting. The number of pods present also significantly increased probability of increased social behavior. The presence of rarely sighted pods and the number of pods present did not have a significant interaction. Ordinal day and number of mitochondrial DNA haplotypes appears to not have changed the probability of any behavior. Foraging remained the predominant behavior throughout all factors. The concurrent increase in social behavior and decrease in resting behavior with rarely sighted pods present implies an unusually high importance of social behavior in the lives of resident killer whales.     

A CETACEAN BIOPSY SYSTEM USING LIGHTWEIGHT PNEUMATIC DARTS, AND ITS EFFECT ON THE BEHAVIOR OF KILLER WHALES

Lightweight untethered pneumatic darts were used to biopsy killer whales, Orcinus orca , for genetic and toxicological analysis. Samples of epidermal, dermal, and hypodermal tissue weighing approximately 0.5 g were obtained by 65% of the 91 darts fired during the study. Sufficient DNA for multiple analyses was extracted from the biopsies, which were also used for fatty acid and toxic contaminant analyses. Reactions such as momentary shakes or accelerations were observed after 81% of the dart hits and 53% of the misses. Aversion to the research vessel was assessed by reapproaching target whales after the sampling attempts. In 6% of the hits and 8% of the misses aversion to the research boat increased immediately following the attempt. No similar increases in aversion were seen when killer whales were reapproached one day to one year after being hit. The darts were also tested successfully on humpback whales, Megaptera novaeangliae . In view of the simplicity of the system, its effectiveness in acquiring multipurpose samples, and the apparently short-term disturbance it caused, it is recommended for future cetacean biopsy studies.     

Captive killer whale (Orcinus orca) survival

Killer whales (Orcinus orca) were first placed into captivity in 1961 and are now found in theme parks around the world. Despite successful breeding of captive killer whales since 1985 there is growing concern for their welfare in captivity, which often includes claims of poor survival. We employed Kaplan‐Meier and Cox Proportional hazards models and annual survival rate analyses on 201 captive killer whales to discern how sex, facility (U.S. vs. foreign), captive‐born vs. wild‐captured, pre‐ vs. post‐1 January 1985, and animal age upon entering captivity affect survival. Overall median survival estimate was 6.1 yr, with no difference between male and female survival. Killer whales in U.S. facilities (12.0 yr) demonstrated a significantly higher median survival than those in foreign facilities (4.4 yr), as did whales entering captivity post‐1 January 1985 (11.8 yr) vs. those entering prior to 1 January 1985 (3.9 yr). Median survival for captive‐born (14.1 yr) was significantly higher than wild‐captured killer whales (5.5 yr), though the two failed to differ among the post‐1 January 1985 cohort. Facility location and pre‐ vs. post‐1 January 1985 were predictors of the hazard rate. Survival of captive killer whale cohorts has generally improved through time, although survival to age milestones are poor when compared to wild killer whales.     

Behavioral context of echolocation and prey-handling sounds produced by killer whales (Orcinus orca) during pursuit and capture of Pacific salmon (Oncorhynchus spp.)

Availability of preferred salmonid prey and a sufficiently quiet acoustic environment in which to forage are critical to the survival of resident killer whales (Orcinus orca) in the northeastern Pacific. Although piscivorous killer whales rely on echolocation to locate and track prey, the relationship between echolocation, movement, and prey capture during foraging by wild individuals is poorly understood. We used acoustic biologging tags to relate echolocation behavior to prey pursuit and capture during successful feeding dives by fish-eating killer whales in coastal British Columbia, Canada. The significantly higher incidence and rate of echolocation prior to fish captures compared to afterward confirms its importance in prey detection and tracking. Extremely rapid click sequences (buzzes) were produced before or concurrent with captures of salmon at depths typically exceeding 50 m, and were likely used by killer whales for close-range prey targeting, as in other odontocetes. Distinctive crunching and tearing sounds indicative of prey-handling behavior occurred at relatively shallow depths following fish captures, matching concurrent observations that whales surfaced with fish prior to consumption and often shared prey. Buzzes and prey-handling sounds are potentially useful acoustic signals for estimating foraging efficiency and determining if resident killer whales are meeting their energetic requirements.     

Cooperative hunting behavior,prey selectivity and prey handling by pack ice killer whales (Orcinus orca), type B,in Antarctic Peninsula waters

Currently, there are three recognized ecotypes (or species) of killer whales (Orcinus orca) in Antarctic waters, including type B, a putative prey specialist on seals, which we refer to as “pack ice killer whale” (PI killer whale). During January 2009, we spent a total of 75.4 h observing three different groups of PI killer whales hunting off the western Antarctic Peninsula. Observed prey taken included 16 seals and 1 Antarctic minke whale (Balaenoptera bonaerensis). Weddell seals (Leptonychotes weddellii) were taken almost exclusively (14/15 identified seal kills), despite the fact that they represented only 15% of 365 seals identified on ice floes; the whales entirely avoided taking crabeater seals (Lobodon carcinophaga; 82% relative abundance) and leopard seals (Hydrurga leptonyx; 3%). Of the seals killed, the whales took 12/14 (86%) off ice floes using a cooperative wave‐washing behavior; they produced 120 waves during 22 separate attacks and successfully took 12/16 (75%) of the Weddell seals attacked. The mean number of waves produced per successful attack was 4.1 (range 1–10) and the mean attack duration was 30.4 min (range 15–62). Seal remains that we examined from one of the kills provided evidence of meticulous postmortem prey processing perhaps best termed “butchering.”