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.

Killer Whale Predation on Belugas in Cook Inlet, Alaska: Implications for a Depleted Population

Killer whale predation on belugas in Cook Inlet, Alaska, has become a concern since the decline of these belugas was documented during the 1990s. Accordingly, killer whale sightings were compiled from systematic surveys, observer databases, and anecdotal accounts. Killer whales have been relatively common in lower Cook Inlet (at least 100 sightings from 1975 to 2002), but in the upper Inlet, north of Kalgin Island, sightings were infrequent (18 in 27 yr), especially prior to the 1990s. Beach cast beluga carcasses with teeth marks and missing flesh also provided evidence of killer whale predation. Most observed killer whale/beluga interactions were in the upper Inlet. During 11 of 15 observed interactions, belugas were obviously injured or killed, either through direct attacks or indirectly as a result of stranding. Assuming at least one beluga mortality occurred during the other four encounters, we can account for 21 belugas killed between 1985 and 2002. This would suggest a minimum estimate of roughly l/yr and does not include at least three instances where beluga calves accompanied an adult that was attacked.

Estimating abundance of killer whales in the nearshore waters of the Gulf of Alaska and Aleutian Islands using line-transect sampling

Killer whale (Orcinus orca Linnaeus, 1758) abundance in the North Pacific is known only for a few populations for which extensive longitudinal data are available, with little quantitative data from more remote regions. Line-transect ship surveys were conducted in July and August of 2001–2003 in coastal waters of the western Gulf of Alaska and the Aleutian Islands. Conventional and Multiple Covariate Distance Sampling methods were used to estimate the abundance of different killer whale ecotypes, which were distinguished based upon morphological and genetic data. Abundance was calculated separately for two data sets that differed in the method by which killer whale group size data were obtained. Initial group size (IGS) data corresponded to estimates of group size at the time of first sighting, and post-encounter group size (PEGS) corresponded to estimates made after closely approaching sighted groups.

Eastern temperate North Pacific offshore killer whales (Orcinus orca): Occurrence, movements, and insights into feeding ecology

Beginning in the late 1980s, large groups of previously unidentified killer whales (Orcinus orca) were sighted off the west coast of Vancouver Island and in the Queen Charlotte Islands, British Columbia. Scientists working in this region produced two killer whale photo-identification catalogues that included both transient (mammal-eating) whales and 65 individual whales that investigators believed represented a distinct killer whale community (Ford et al. 1992, Heise et al. 1993). It was thought that these killer whales maintained a generally offshore distribution and were provisionally termed “offshores”; a term that has since been used as a population identifier for the eastern temperate North Pacific offshore killer whale population. Then in September 1992, 75 unidentified whales entered the Strait of Juan de Fuca just south and east of Victoria, British Columbia (Walters et al. 1992). Although most of these whales had not been seen before, two were matched to killer whales in the Queen Charlotte photo-identification catalogue (Ford et al. 1992, Heise et al. 1993) and were thus listed as “offshore” killer whales. During a similar time period, other large groups of killer whales, previously unidentified, were also being sighted off Alaska and California (Dahlheim et al. 1997; Nancy Black and Alisa Schulman- Janiger, unpublished data, respectively).

Killer Whales and Marine Mammal Trends in The North Pacific—A Re-Examination of Evidence for Sequential Megafauna Collapse and the Prey-Switching Hypothesis

Springer et al. (2003) contend that sequential declines occurred in North Pacific populations of harbor and fur seals, Steller sea lions, and sea otters. They hypothesize that these were due to increased predation by killer whales, when industrial whaling’s removal of large whales as a supposed primary food source precipitated a prey switch. Using a regional approach, we reexamined whale catch data, killer whale predation observations, and the current biomass and trends of potential prey, and found little support for the prey-switching hypothesis. Large whale biomass in the Bering Sea did not decline as much as suggested by Springer et al., and much of the reduction occurred 50–100 yr ago, well before the declines of pinnipeds and sea otters began; thus, the need to switch prey starting in the 1970s is doubtful. With the sole exception that the sea otter decline followed the decline of pinnipeds, the reported declines were not in fact sequential. Given this, it is unlikely that a sequential megafaunal collapse from whales to sea otters occurred. The spatial and temporal patterns of pinniped and sea otter population trends are more complex than Springer et al. suggest, and are often inconsistent with their hypothesis. Populations remained stable or increased in many areas, despite extensive historical whaling and high killer whale abundance. Furthermore, observed killer whale predation has largely involved pinnipeds and small cetaceans; there is little evidence that large whales were ever a major prey item in high latitudes. Small cetaceans (ignored by Springer et al.) were likely abundant throughout the period. Overall, we suggest that the Springer et al. hypothesis represents a misleading and simplistic view of events and trophic relationships within this complex marine ecosystem.

Use of chemical tracers in assessing the diet and foraging regions of eastern North Pacific killer whales

Top predators in the marine environment integrate chemical signals acquired from their prey that reflect both the species consumed and the regions from which the prey were taken. These chemical tracers—stable isotope ratios of carbon and nitrogen; persistent organic pollutant (POP) concentrations, patterns and ratios; and fatty acid profiles—were measured in blubber biopsy samples from North Pacific killer whales (Orcinus orca) (n = 84) and were used to provide further insight into their diet, particularly for the offshore group, about which little dietary information is available. The offshore killer whales were shown to consume prey species that were distinctly different from those of sympatric resident and transient killer whales. In addition, it was confirmed that the offshores forage as far south as California. Thus, these results provide evidence that the offshores belong to a third killer whale ecotype. Resident killer whale populations showed a gradient in stable isotope profiles from west (central Aleutians) to east (Gulf of Alaska) that, in part, can be attributed to a shift from off-shelf to continental shelf-based prey. Finally, stable isotope ratio results, supported by field observations, showed that the diet in spring and summer of eastern Aleutian Island transient killer whales is apparently not composed exclusively of Steller sea lions.

Use of chemical tracers in assessing the diet and foraging regions of eastern North Pacific killer whales

Top predators in the marine environment integrate chemical signals acquired from their prey that reflect both the species consumed and the regions from which the prey were taken. These chemical tracers—stable isotope ratios of carbon and nitrogen; persistent organic pollutant (POP) concentrations, patterns and ratios; and fatty acid profiles—were measured in blubber biopsy samples from North Pacific killer whales (Orcinus orca) (n = 84) and were used to provide further insight into their diet, particularly for the offshore group, about which little dietary information is available. The offshore killer whales were shown to consume prey species that were distinctly different from those of sympatric resident and transient killer whales. In addition, it was confirmed that the offshores forage as far south as California. Thus, these results provide evidence that the offshores belong to a third killer whale ecotype. Resident killer whale populations showed a gradient in stable isotope profiles from west (central Aleutians) to east (Gulf of Alaska) that, in part, can be attributed to a shift from off-shelf to continental shelf-based prey. Finally, stable isotope ratio results, supported by field observations, showed that the diet in spring and summer of eastern Aleutian Island transient killer whales is apparently not composed exclusively of Steller sea lions.

Draft U.S. Pacific Marine Mammal Stock Assessments: 2009

Under the 1994 amendments to the Marine Mammal Protection Act (MMPA), the National Marine Fisheries Service (NMFS) and the U.S. Fish and Wildlife Service (USFWS) are required to publish Stock Assessment Reports for all stocks of marine mammals within U.S. waters, to review new information every year for strategic stocks and every three years for non-strategic stocks, and to update the stock assessment reports when significant new information becomes available. This report presents stock assessments for 13 Pacific marine mammal stocks under NMFS jurisdiction, including 8 “strategic” stocks and 5 “non-strategic” stocks (see summary table). A new stock assessment for humpback whales in American Samoa waters is included in the Pacific reports for the first time. New or revised abundance estimates are available for 9 stocks, including Eastern North Pacific blue whales, American Samoa humpback whales, five U.S. west coast harbor porpoise stocks, the Hawaiian monk seal, and southern resident killer whales. A change in the abundance estimate of Eastern North Pacific blue whales reflects a recommendation from the Pacific Scientific Review Group to utilize mark-recapture estimates for this population, which provide a better estimate of total population size than the average of recent line-transect and mark-recapture estimates. The ‘Northern Oregon/Washington Coast Stock’ harbor porpoise stock assessment includes a name change (‘Oregon’ is appended to ‘Northern Oregon’) to reflect recent stock boundary changes. Changes in abundance estimates for the two stocks of harbor porpoise that occur in Oregon waters are the result of these boundary changes, and do not reflect biological changes in the populations. Updated information on the three stocks of false killer whales in Hawaiian waters is also included in these reports. Information on the remaining 50 Pacific region stocks will be reprinted without revision in the final 2009 reports and currently appears in the 2008 reports (Carretta et al. 2009). Stock Assessments for Alaskan marine mammals are published by the National Marine Mammal Laboratory (NMML) in a separate report. Pacific region stock assessments include those studied by the Southwest Fisheries Science Center (SWFSC, La Jolla, California), the Pacific Islands Fisheries Science Center (PIFSC, Honolulu, Hawaii), the National Marine Mammal Laboratory (NMML, Seattle, Washington), and the Northwest Fisheries Science Center (NWFSC, Seattle, WA). Northwest Fisheries Science Center staff prepared the report on the Eastern North Pacific Southern Resident killer whale. National Marine Mammal Laboratory staff prepared the Northern Oregon/Washington coast harbor porpoise stock assessment. Pacific Islands Fisheries Science Center staff prepared the report on the Hawaiian monk seal. Southwest Fisheries Science Center staff prepared stock assessments for 9 stocks. The stock assessment for the American Samoa humpback whale was prepared by staff from the Center for Coastal Studies, Hawaiian Islands Humpback National Marine Sanctuary, the Smithsonian Institution, and the Southwest Fisheries Science Center. Draft versions of the stock assessment reports were reviewed by the Pacific Scientific Review Group at the November 2008, Maui meeting. The authors also wish to thank those who provided unpublished data, especially Robin Baird and Joseph Mobley, who provided valuable information on Hawaiian cetaceans. Any omissions or errors are the sole responsibility of the authors. This is a working document and individual stock assessment reports will be updated as new information on marine mammal stocks and fisheries becomes available. Background information and guidelines for preparing stock assessment reports are reviewed in Wade and Angliss (1997). The authors solicit any new information or comments which would improve future stock assessment reports. These Stock Assessment Reports summarize information from a wide range of sources and an extensive bibliography of all sources is given in each report. We strongly urge users of this document to refer to and cite original literature sources rather than citing this report or previous Stock Assessment Reports. If the original sources are not accessible, the citation should follow the format: [Original source], as cited in [this Stock Assessment Report citation].

Cetaceans of Southeast Alaska: Distribution and Seasonal Occurrence

Aim To assess the distribution, group size, seasonal occurrence and annual trends of cetaceans. Location The study area included all major inland waters of Southeast Alaska. Methods Between 1991 and 2007, cetacean surveys were conducted by observers who kept a constant watch when the vessel was underway and recorded all cetaceans encountered. For each species, we examined distributional patterns, group size, seasonal occurrence and annual trends. Analysis of variance (anova F) was used to test for differences in group sizes between multiple means, and Student’s t-test was used to detect differences between pairwise means. Cetacean seasonal occurrence and annual trends were investigated using a generalized linear model framework. Results Humpback whales (Megaptera novaeangliae) were seen throughout the region, with numbers lowest in spring and highest in the fall. Fin whale (Balaenoptera physalus) and minke whale (Balaenoptera acutorostrata) distributions were more restricted than that reported for humpback whales, and the low number of sightings precluded evaluating seasonal trends. Three killer whale (Orcinus orca) eco-types were documented with distributions occurring throughout inland waters. Seasonal patterns were not detected or could not be evaluated for resident and offshore killer whales, respectively; however, the transient eco-type was more abundant in the summer. Dall’s porpoise (Phocoenoides dalli) were distributed throughout the region, with more sightings in spring and summer than in fall. Harbour porpoise (Phocoena phocoena) distribution was clumped, with concentrations occurring in the Icy Strait/Glacier Bay and Wrangell areas and with no evidence of seasonality. Pacific white-sided dolphins (Lagenorhynchus obliquidens) were observed only occasionally, with more sightings in the spring. For most species, group size varied on both an annual and seasonal basis. Main conclusions Seven cetacean species occupy the inland waters of Southeast Alaska, with distribution, group size, seasonal occurrence and annual trends varying by species. Future studies that compare spatial and temporal patterns with other features (e.g. oceanography, prey resources) may help in identifying the key factors that support the high density and biodiversity of cetaceans found in this region. An increased understanding of the region’s marine ecology is an essential step towards ensuring the long-term conservation of cetaceans in Southeast Alaska.

The traceability/trackability of illegal trade in whale products: a proposal to evaluate the technical functionality of DNA registers

The Government of Japan, through the Institute for Cetacean Research (Tokyo), has established a DNA register for whales taken under special permit or otherwise destined for commercial markets (IWC 2005; IWC 2010a). The functionality of this DNA register, for the purposes of traceability/trackability, is critical to the current negotiations on the future of the IWC (IWC 2010b). Here we request access to the DNA register for 3 species of whales (fin, sei and Antarctic minke) for the purposes of tracking the origins of whale products purchased at commercial outlets in Seoul, South Korea and Santa Monica, US, as described in the Baker et al. (2010). The attached proposal was included as Supplementary Material to this published article and submitted for consideration to the IWC Data Availability Group (DAG) on 12 April 2010. However, the DAG declined to forward the proposal to the data holders, recommending that we “wait until the Scientific Committee has reviewed the proposed DNA register/market sampling text in the draft Consensus Decision in accordance with the Commission's instructions and then reported to the Commission itself” (email 16 May 2010). We assume that this will take place at SC/62 in Agadir and request that this proposal be considered for endorsement by the DNA subcommittee.

Population histories of right whales (Cetacea: Eubalaena) Inferred from Mitochondrial Sequence Diversities and Divergences of Their Whale Lice (Amphipoda: Cyamus)

Right whales carry large populations of three ‘whale lice’ (Cyamus ovalis, Cyamus gracilis, Cyamus erraticus) that have no other hosts. We used sequence variation in the mitochondrial COI gene to ask (i) whether cyamid population structures might reveal associations among right whale individuals and subpopulations, (ii) whether the divergences of the three nominally conspecific cyamid species on North Atlantic, North Pacific, and southern right whales (Eubalaena glacialis, Eubalaena japonica, Eubalaena australis) might indicate their times of separation, and (iii) whether the shapes of cyamid gene trees might contain information about changes in the population sizes of right whales. We found high levels of nucleotide diversity but almost no population structure within oceans, indicating large effective population sizes and high rates of transfer between whales and subpopulations. North Atlantic and Southern Ocean populations of all three species are reciprocally monophyletic, and North Pacific C. erraticus is well separated from North Atlantic and southern C. erraticus. Mitochondrial clock calibrations suggest that these divergences occurred around 6 million years ago (Ma), and that the Eubalaena mitochondrial clock is very slow. North Pacific C. ovalis forms a clade inside the southern C. ovalis gene tree, implying that at least one right whale has crossed the equator in the Pacific Ocean within the last 1–2 million years (Myr). Low-frequency polymorphisms are more common than expected under neutrality for populations of constant size, but there is no obvious signal of rapid, interspecifically congruent expansion of the kind that would be expected if North Atlantic or southern right whales had experienced a prolonged population bottleneck within the last 0.5 Myr.

The western gray whale: a review of past exploitation, current status and potential threats

Gray whales (Eschrichtius robustus) occur along the eastern and western coastlines of the North Pacific as two geographically isolated populations and have traditionally been divided into the eastern (California-Chukchi) and western (Korean-Okhotsk) populations. Recent molecular comparisons confirm, based on differences in haplotypic frequencies, that these populations are genetically separated at the population-level. Both populations were commercially hunted, but only the eastern gray whale has returned to near pre-exploitation numbers. In contrast, the western population remains highly depleted, shows no apparent signs of recovery and its future survival remains uncertain. Research off Sakhalin Island, Russia between 1995 and 1999 has produced important new information on the present day conservation status of western gray whales and provided the basis for the World Conservation Union (IUCN) to list the population as 'Critically Endangered in 2000. The information presented here, in combination with potential impacts from anthropogenic threats throughout the range of this population, raises strong concerns about the recovery and continued survival of the western gray whale.

Recovery Plan for The Blue Whale (Balaenoptera musculus)

Recovery plans identify reasonable actions which are believed to be required to recover and/or protect endangered species. Plans are prepared by the National Marine Fisheries Service (NMFS) and the U.S. Fish and Wildlife Service (FWS) and sometimes with the assistance of recovery teams, contractors, State agencies, and others. This plan was prepared by Randall R. Reeves, Phillip J. Clapham, Robert L. Brownell, Jr., and Gregory K. Silber for NMFS. Recovery plans do not necessarily represent the views nor the official positions or approvals of any individuals or agencies, other than those of NMFS, and they represent the views of NMFS only after they have been approved by the Assistant Administrator for Fisheries. Objectives will only be attained and funds expended contingent upon appropriations, priorities, and other budgetary constraints. Approved recovery plans are subject to modification as dictated by new findings, changes in species status, and the completion of recovery tasks described in the plan.

A whale of a new species

Although four new species of whale have been identified since 1937, the absence of new ones for the past 28 years might be taken to mean that they had finally all been dis- covered. Not so - a new species of beaked whale, Mesoplodonperuvianus, is described by J. C. Reyes and colleagues in the latest issue of Marine Mammal Science.

Whale Meat in the Japanese Diet

Junghans (Letters, 4 Apr., p. 6) points out several errors in Beary’s letter (14 December 1979, p. 1260) regarding the high concentrations of mercury in whale meat eaten by the Japanese. We would like to call attention to some additional errors in Beary’s letter.