The whaling issue: Conservation, confusion, and casuistry

Morishita’s “multiple analysis”of the whaling issue [Morishita J. Multiple analysis of the whaling issue: Understanding the dispute by a matrix. Marine Policy 2006;30:802–8] is essentially a restatement of the Government of Japan’s whaling policy, which confuses the issue through selective use of data, unsubstantiated facts, and the vilification of opposing perspectives. Here, we deconstruct the major problems with Morishita’s article and provide an alternative view of the whaling dispute. For many people in this debate, the issue is not that some whales are not abundant, but that the whaling industry cannot be trusted to regulate itself or to honestly assess the status of potentially exploitable populations. This suspicion has its origin in Japan’s poor use of science, its often implausible stock assessments, its insistence that culling is an appropriate way to manage marine mammal populations, and its relatively recent falsification of whaling and fisheries catch data combined with a refusal to accept true transparency in catch and market monitoring. Japanese policy on whaling cannot be viewed in isolation, but is part of a larger framework involving a perceived right to secure unlimited access to global marine resources. Whaling is inextricably tied to the international fisheries agreements on which Japan is strongly dependent; thus, concessions made at the IWC would have potentially serious ramifications in other fora.

The Phylogenetic Relationships and Biogeography of True Porpoises (Mammalia: Phocoenidae) Based On Morphological Data

Prior studies of phylogenetic relationships among phocoenids based on morphology and molecular sequence data conflict and yield unresolved relationships among species. This study evaluates a comprehensive set of cranial, postcranial, and soft anatomical characters to infer interrelationships among extant species and several well-known fossil phocoenids, using two different methods to analyze polymorphic data: polymorphic coding and frequency step matrix. Our phylogenetic results confirmed phocoenid monophyly. The division of Phocoenidae into two subfamilies previously proposed was rejected, as well as the alliance of the two extinct genera Salumiphocaena and Piscolithax with Phocoena dioptrica and Phocoenoides dalli. Extinct phocoenids are basal to all extant species. We also examined the origin and distribution of porpoises within the context of this phylogenetic framework. Phocoenid phylogeny together with available geologic evidence suggests that the early history of phocoenids was centered in the North Pacific during the middle Miocene, with subsequent dispersal into the southern hemisphere in the middle Pliocene. A cooling period in the Pleistocene allowed dispersal of the southern ancestor of Phocoena sinusinto the North Pacific (Gulf of California).

Diets of Baird’s Beaked Whales, Berardius bairdii, in the Southern Sea Of Okhotsk and Off the Pacific Coast Of Honshu, Japan

Stomach contents were analyzed from 127 Baird’s beaked whales, Berardizls bairdii, taken in coastal waters of Japan. During late July-August of 1985- 1987, 1989, and 1991, 107 samples were collected from off the Pacific coast of Honshu. An additional 20 samples were collected from whales taken in the southern Sea of Okhotsk during late August-September of 1988 and 1989. Prey identification using fish otoliths and cephalopod beaks revealed the whales fed primarily on deep-water gadiform fishes and cephalopods in both regions. Prey species diversity and the percentage of cephalopods and fish differed between the two regions. Off the Pacific coast of Honshu the whales fed primarily on benthopelagic fishes (81.8%) and only 18.0% on cephalopods. Eight species of fish representing two families, the codlings (Moridae) and the grenadiers (Macrouridde), collectively made up 81.3% of the total. Thirty species of cephalopods representing 14 families made up 12.7%. In the southern Sea of Okhotsk, cephalopods accounted for 87.1% of stomach contents. The families Gonatidae and Cranchiidae were the predominant cephalopod prey, accounting for 86.7% of the diet. Gadiform fish accounted for only 12.9% of the diet. Longfin codling, Laernonma longipes, was the dominant fish prey in both regions. Depth distribution of the two commonly consumed fish off the Pacific coast of Honshu indicate the whales in this region fed primarily at depths ranging from 800 to 1,200 m.

Book Review of Cetacean Societies: Field Studies Of Whales And Dolphins Edited by I. Mann, R. C. Connor, P. L. Tyack & H. Whitehead

Studying the sociobiology and behavioral ecology of cetaceans is particularly challenging due in large part to the aquatic environment in which they live. Nevertheless, many of the obstacles traditionally associated with data gathering on tree-ranging whales, dolphins and porpoises are rapidly being overcome, and are now far less formidable. During the past several decades, marine mammal scientists equipped with innovative research methods and new technologies have taken field-based behavioral studies to a new level of sophistication. In some cases, as is true for bottlenose dolphins, killer whales, sperm whales and humpback whales, modern research paradigms in the marine environment are comparable to present-day studies of terrestrial mammal social systems. Cetacean Society stands testament to the relatively recent advances in marine mammal science, and to those scientists, past and present, whose diligence has been instrumental in shaping the discipline.

Ringed and bearded seal densities in the eastern Chukchi Sea, 1999–2000

Aerial surveys were conducted in 1999 and 2000 to estimate the densities of ringed (Phoca hispida) and bearded (Erignathus barbatus) seals in the eastern Chukchi Sea. Survey lines were focused mainly on the coastal zone within 37 km of the shoreline, with additional lines flown 148–185 km offshore to assess how densities of seals changed as a function of distance from shore. Satellite-linked time-depth recorders were attached to ringed seals in both years to evaluate the time spent basking on the ice surface. Haulout patterns indicated that ringed seals transitioned to basking behavior in late May and early June, and that the largest proportion of seals (60–68%) was hauled out between 0830 and 1530 local solar time. Ringed seals were relatively common in nearshore fast ice and pack ice, with lower densities in offshore pack ice. The average density of ringed seals was 1.91 seals km-2 in 1999 (range 0.37– 16.32) and 1.62 seals km-2 in 2000 (range 0.42–19.4), with the highest densities of ringed seals found in coastal waters south of Kivalina and near Kotzebue Sound. The estimated abundance of ringed seals for the entire study area was similar in 1999 (252,488 seals, SE=47,204) and 2000 (208,857 seals, SE=25,502). Bearded seals were generally more common in offshore pack ice, with the exception of high bearded seal numbers observed near the shore south of Kivalina. Bearded seal densities were not adjusted for haulout behavior, and therefore, abundance was not estimated. Unadjusted average bearded seal density was 0.07 seals km-2 in 1999 (range 0.011–0.393) and 0.14 seals km-2 in 2000 (range 0.009– 0.652). Levels of primary productivity, benthic biomass, and fast ice distribution may influence the distributions of ringed and bearded seals in the Chukchi Sea. Information on movement and haulout behavior of ringed and bearded seals would be very useful for designing future surveys.

Provisioning Strategies of Antarctic Fur Seals and Chinstrap Penguins Produce Different Responses to Distribution of Common Prey and Habitat

Central-place foragers that must return to a breeding site to deliver food to offspring are faced with trade-offs between prey patch quality and distance from the colony. Among colonial animals, pinnipeds and seabirds may have different provisioning strategies, due to differences in their ability to travel and store energy. We compared the foraging areas of lactating Antarctic fur seals and chinstrap penguins breeding at Seal Island, Antarctica, to investigate whether they responded differently to the distribution of their prey (Antarctic krill and myctophid fish) and spatial heterogeneity in their habitat. Dense krill concentrations occurred in the shelf region near the colony. However, only brooding penguins, which are expected to be time-minimizers because they must return frequently with whole food for their chicks, foraged mainly in this proximal shelf region. Lactating fur seals and incubating penguins, which can make longer trips to increase energy gain per trip, and so are expected to be energy-maximizers, foraged in the more distant (>20 km from the island) slope and oceanic regions. The shelf region was characterized by more abundant, but lower-energy-content immature krill, whereas the slope and oceanic regions had less abundant but higher-energy-content gravid krill, as well as high-energy-content myctophids. Furthermore, krill in the shelf region undertook diurnal vertical migration, whereas those in the slope and oceanic regions stayed near the surface throughout the day, which may enhance the capture rate for visual predators. Therefore, we sug- gest that the energy-maximizers foraged in distant, but potentially more profitable feeding regions, while the time-minimizers foraged in closer, but potentially less profitable regions. Thus, time and energy constraints derived from different provisioning strategies may result in sympatric colonial predator species using different foraging areas, and as a result, some central-place foragers use sub- optimal foraging habitats, in terms of the quality or quantity of available prey.

Hunting and social behaviour of leopard seals (Hydrurga leptonyx) at Seal Island, South Shetland Islands, Antarctica

The hunting behavior of leopard seals Hydrurga leptonyx was monitored opportunistically at Seal Island, South Shetland Islands, during the austral summers from 1986/87 to 1994/95. Leopard seals used several methods to catch Antarctic fur seal pups Arctocephalus gazella and chinstrap penguins Pygoscelis antarctica, and individuals showed different hunting styles and hunting success. One to two leopard seals per year were responsible for an average of 60% of observed captures of fur seal pups. Leopard seals preyed on penguins throughout the summer, but preyed on fur seal pups only between late December and mid-February. Hunting behavior differed significantly between different locations on the island; fur seals were hunted only at one colony, and penguins were hunted in several areas. The relative abundance of prey types, size of prey in relation to predator, and specialization of individual leopard seals to hunt fur seal prey probably influence individual prey preferences among leopard seals. On five occasions, two leopard seals were seen together on Seal Island. Possible interpretations of the relationship between the interacting leopard seals included a mother-offspring relationship, a consorting male-female pair, and an adult leopard seal followed by an unrelated juvenile. In two incidents at Seal Island, two leopard seals were observed interacting while hunting: one seal captured fur seal pups and appeared to release them to the other seal. Observations of leopard seals interacting during hunting sessions were difficult to confirm as co-operative hunting, but they strongly implied that the two seals were not agonistic toward one another. The hunting success of individual leopard seals pursuing penguins or fur seals is probably high enough for co-operative hunting not to become a common hunting strategy; however, it may occur infrequently when it increases the hunting productivity of the seals.

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.

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.

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.

Determining spatial and temporal scales for management: lessons from whaling

Selection of the appropriate management unit is critical to the conservation of animal populations. Defining such units depends upon knowledge of population structure and upon the timescale being considered. Here, we examine the trajectory of eleven subpopulations of five species of baleen whales to investigate temporal and spatial scales in management. These subpopulations were all extirpated by commercial whaling, and no recovery or repopulation has occurred since. In these cases, time elapsed since commercial extinction ranges from four decades to almost four centuries. We propose that these subpopulations did not recover either because cultural memory of the habitat has been lost, because widespread whaling among adjacent stocks eliminated these as sources for repopulation, and/or because segregation following exploitation produced the abandonment of certain areas. Spatial scales associated with the extirpated subpopulations are frequently smaller than those typically employed in management. Overall, the evidence indicates that: (1) the time frame for management should be at most decadal in scope (i.e., <100 yr) and based on both genetic and nongenetic evidence of population substructure, and (2) at least some stocks should be defined on a smaller spatial scale than they currently are.

Historical Observations of Humpback And Blue Whales in the North Atlantic Ocean: Clues to Migratory Routes and Possibly Additional Feeding Grounds

The seasonal distributions of humpback and blue whales (Megaptera novaeangliae and Balaenoptera musculus, respectively) in the North Atlantic Ocean are not fully understood. Although humpbacks have been studied intensively in nearshore or coastal feeding and breeding areas, their migratory movements between these areas have been largely inferred. Blue whales have only been studied intensively along the north shore of the Gulf of St. Lawrence, and their seasonal occurrence and movements elsewhere in the North Atlantic are poorly known. We investigated the historical seasonal distributions of these two species using sighting and catch data extracted from American 18th and 19th century whaling logbooks. These data suggest that humpback whales migrated seasonally from low-latitude calving/ breeding grounds over a protracted period, and that some of them traveled far offshore rather than following coastal routes. Also, at least some humpbacks apparently fed early in the summer west of the Mid-Atlantic Ridge, well south of their known present-day feeding grounds. In assessing the present status of the North Atlantic humpback population, it will be important to determine whether such offshore feeding does in fact occur. Blue whales were present across the southern half of the North Atlantic during the autumn and winter months, and farther north in spring and summer, but we had too few data points to support inferences about these whales’ migratory timing and routes.

Seasonal home ranges and fidelity to breeding sites among ringed seals

Population structure and patterns of habitat use among ringed seals (Phoca hispida) are poorly known, in part because seasonal movements have not been adequately documented. We monitored the movements of 98 ringed seals in the Beaufort and Chukchi seas between 1990 and 2006 using three forms of telemetry. In the winter—spring period (when the seals were occupying shorefast ice), we used radio and ultra-sonic tags to track movements above and below the ice, respectively. We used satellite-linked transmitters in summer and fall (when the seals ranged away from their winter sites) to track at-sea movements. In the shorefast ice habitat, the home ranges of 27 adult males ranged from\1 to 13.9 km2 (median = 0.628) while the home ranges of 28 adult females ranged from \1 to 27.9 km2 (median = 0.652). The 3-dimensional volumes used by 9 seals tracked acoustically under the ice averaged 0.07 (SD = 0.04) km3 for subadults and adult males and 0.13 (SD = 0.04) km3 for adult females. Three of the radio-tracked seals and 9 tracked by satellite ranged up to 1,800 km from their winter/spring home ranges in summer but returned to the same small (1–2 km2) sites during the ice-bound months in the following year. The restricted movements of ringed seals during the ice-bound season— including the breeding season—limits their foraging activities for most of the year and may minimize gene flow within the species.

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.