FEEDING, SOCIAL AND MIGRATION BEHAVIOR OF BOWHEAD WHALES, BALAENA MYSTICETUS, IN BAFFIN BAY VS. THE BEAUFORT SEA—REGIONS WITH DIFFERENT AMOUNTS OF HUMAN ACTIVITY

This paper compares the behavior of bowhead whales of the Davis Strait/Baffin Bay stock, as observed along the east coast of Baffin Island in 1979–1986, with behavior of the Bering/Chukchi/Beaufort Sea stock observed in the Beaufort Sea in 1980–1986. All data used here were collected during late summer and early autumn in the absence of acute human disturbance. The behavioral repertoires of the two populations were similar. However, quantitative differences were found for whales engaged in all three activities studied: (1) Bowheads feeding in deep water off Isabella Bay, Baffin Island, had longer dives and surfacings, on average, than noted for bowheads feeding in the Beaufort Sea. (2) Among whales socializing in shallow water, we saw sexual interactions more often at Isabella Bay than in the Beaufort Sea. Calls emitted by socializing whales off Baffin Island were similar to those heard in the Chukchi and Beaufort Seas. However, pulsed tonal calls were longer off Baffin Island, and previously undescribed mechanical "crunch" sounds were recorded there near socializing bowheads. (3) During autumn migration, "fluke-out" dives were less common, and dive durations were longer, in the Beaufort Sea than off Baffin Island (P<0.001). Multivariate and other analyses indicated that some but not all differences can be ascribed to regional differences in the natural environment or in whale activities, However, during 1974–1986, Bering/Chukchi/Beaufort bowheads were exposed to more industrial, hunting and other human activity than Davis Strait/Baffin Bay bowheads. The "inconspicuous" behavior during autumn migration in the Beaufort may have been attributable to human activities, but causative links cannot be isolated.     

Increasing abundance of bowhead whales in West Greenland

In April 2006, a dedicated survey of bowhead whales (Balaena mysticetus) was conducted on the former whaling ground in West Greenland to determine the current wintering population abundance. This effort included a double platform aerial survey design, satellite tracking of the movements of nine whales, and estimation of high-resolution surface time from 14 whales instrumented with time-depth recorders. Bowhead whales were estimated to spend an average of 24% (cv=0.03) of the time at or above 2m depth, the maximum depth at which they can be seen on the trackline. This resulted in a fully corrected abundance estimate of 1229 (95% CI: 495-2939) bowhead whales when the availability factor was applied and sightings missed by observers were corrected. This surprisingly large population estimate is puzzling given that the change in abundance cannot be explained by a recent or rapid growth in population size. One possible explanation is that the population, which demonstrates high age and sex segregation, has recently attained a certain threshold size elsewhere, and a higher abundance of mature females appears on the winter and spring feeding ground in West Greenland. This in combination with the latest severe reduction in sea ice facilitating access to coastal areas might explain the surprising increase in bowhead whale abundance in West Greenland.     

An estimate of the fraction of belugas (<Emphasis Type="Italic">Delphinapterus leucas</Emphasis>) in the Canadian high Arctic that winter in West Greenland

Five belugas, or white whales (Delphinapterus leucas), were tracked by satellite from Creswell Bay, Somerset Island, in the Canadian high Arctic towards West Greenland in autumn 2001. After 1 October, three of the whales stayed in the North Water polynya and the other two whales moved to West Greenland. One of the whales that moved to Greenland migrated south along the west coast, following a route and timing similar to another beluga tracked in 1996. The belugas that moved towards West Greenland from Canada did so before or near 1 October. The movements of both these whales followed a similar timing and assumed migratory route of belugas hunted in autumn in West Greenland. In Greenland, the hunt begins in September, where the first whales are taken in the northernmost community of Qaanaaq. Hunting takes place farther south in Upernavik in October, and finally in November and December, belugas are taken even farther south in Uummannaq and Disko Bay. The whales that remain in the North Water after 1 October most likely do not contribute to the harvest in West Greenland. Based on the total number of belugas satellite-tracked in Canada between 1995 and 2001 with tags that lasted beyond 1 October, approximately 0.15 (95% CI 0.06-0.35; n=26) of the summering stock of belugas in the Canadian high Arctic move to West Greenland for the winter. Genetic studies have indicated that belugas moving east through Lancaster Sound are significantly differentiated from belugas taken in the autumn hunt in West Greenland. These conflicting results suggest molecular genetics cannot be solely relied on to reveal the stock identity of these belugas.     

Autumn movements, home ranges, and winter density of narwhals (Monodon monoceros) tagged in Tremblay Sound, Baffin Island

Seven narwhals (Monodon monoceros) were instrumented with satellite transmitters in Tremblay Sound, northeast Canada in August 1999. The whales were tracked for 5-218 days with positions received until 17 March 2000. All whales stayed in the fjord system where they were tagged until the end of August. Three whales went northwest visiting adjacent fjords before moving south, together with the three other whales, along the east coast of Baffin Island. The narwhals arrived on the wintering ground in northern Davis Strait in late October. Speed and range of movements declined once the wintering ground was reached. Dive depths increased from summer to autumn, and reached at least 1,500 m. Late summer and winter kernel home ranges were approximately 3,400 km² and 12,000 km², respectively. The relative abundance of whales on the wintering ground was 936 narwhals. Assuming that the home range defines the winter distribution of the stock, an estimated 5,348 narwhals (corrected for perception and availability bias) were present in this area.     

The Northwest Passage opens for bowhead whales

The loss of Arctic sea ice is predicted to open up the Northwest Passage, shortening shipping routes and facilitating the exchange of marine organisms between the Atlantic and the Pacific oceans. Here, we present the first observations of distribution overlap of bowhead whales (Balaena mysticetus) from the two oceans in the Northwest Passage, demonstrating this route is already connecting whales from two populations that have been assumed to be separated by sea ice. Previous satellite tracking has demonstrated that bowhead whales from West Greenland and Alaska enter the ice-infested channels of the Canadian High Arctic during summer. In August 2010, two bowhead whales from West Greenland and Alaska entered the Northwest Passage from opposite directions and spent approximately 10 days in the same area, documenting overlap between the two populations.     

Population structure of North American beluga whales (Delphinapterus leucas) based on nuclear DNA microsatellite variation and contrasted with the population structure revealed by mitochondrial DNA variation

Beluga whales ( Delphinapterus leucas ) in North American waters migrate seasonally between wintering areas in broken pack ice and summering locations in estuaries and other open water areas in the Arctic and sub-Arctic. Results from our previous investigation of beluga whale mitochondrial DNA (mtDNA) revealed genetic heterogeneity among beluga from different summering locations that was interpreted as representing a high degree of summering site philopatry. However, mtDNA is maternally inherited and does not reflect mating that may occur among beluga from different summering locations in wintering areas or during annual migrations. To test the possibility that breeding occurs among beluga from different summering locations, genetic variability at five nuclear DNA (nDNA) microsatellite loci was examined in the same animals tested in the mtDNA study. Beluga samples ( n = 640) were collected between 1984 and 1994 from 24 sites across North America, mostly during the summer. Whales from the various sites were categorized into eight summering locations as identified by mtDNA analysis, as well as four hypothesized wintering areas: Bering Sea, Hudson Strait (Hudson Strait, Labrador Sea, southwest Davis Strait), Baffin Bay (North Water, east Davis Strait), and St Lawrence River. Microsatellite allele frequencies indicated genetic homogeneity among animals from summering sites believed to winter together but differentiation among whales from some of the wintering areas. In particular, beluga from western North America (Bering Sea) were clearly distinguished from beluga from eastern North America (Hudson Strait, Baffin Bay, and St Lawrence River). Based upon the combined data set, the population of North American beluga whales was divided into two evolutionarily significant units. However, the population may be further subdivided into management units to reflect distinct groups of beluga at summering locations.     

Population structure of North American beluga whales (Delphinapterus leucas) based on nuclear DNA microsatellite variation and contrasted with the population structure revealed by mitochondrial DNA variation

Beluga whales ( Delphinapterus leucas ) in North American waters migrate seasonally between wintering areas in broken pack ice and summering locations in estuaries and other open water areas in the Arctic and sub-Arctic. Results from our previous investigation of beluga whale mitochondrial DNA (mtDNA) revealed genetic heterogeneity among beluga from different summering locations that was interpreted as representing a high degree of summering site philopatry. However, mtDNA is maternally inherited and does not reflect mating that may occur among beluga from different summering locations in wintering areas or during annual migrations. To test the possibility that breeding occurs among beluga from different summering locations, genetic variability at five nuclear DNA (nDNA) microsatellite loci was examined in the same animals tested in the mtDNA study. Beluga samples ( n = 640) were collected between 1984 and 1994 from 24 sites across North America, mostly during the summer. Whales from the various sites were categorized into eight summering locations as identified by mtDNA analysis, as well as four hypothesized wintering areas: Bering Sea, Hudson Strait (Hudson Strait, Labrador Sea, southwest Davis Strait), Baffin Bay (North Water, east Davis Strait), and St Lawrence River. Microsatellite allele frequencies indicated genetic homogeneity among animals from summering sites believed to winter together but differentiation among whales from some of the wintering areas. In particular, beluga from western North America (Bering Sea) were clearly distinguished from beluga from eastern North America (Hudson Strait, Baffin Bay, and St Lawrence River). Based upon the combined data set, the population of North American beluga whales was divided into two evolutionarily significant units. However, the population may be further subdivided into management units to reflect distinct groups of beluga at summering locations.     

New insights into the northward migration route of gray whales between Vancouver Island,British Columbia,and southeastern Alaska

The route taken by northward migrating gray whales during spring between Vancouver Island and southeastern Alaska, a distance of about 575 km, has long been uncertain. It is generally believed that the whales closely follow the western, outer coastline of Haida Gwaii (formerly the Queen Charlotte Islands), an archipelago lying between Vancouver Island and southeastern Alaska, consistent with their pattern of migrating close to shore over the majority of their northward migratory corridor. By tracking satellite‐tagged individuals and surveying whales from shore bases, we provide evidence that this is not the primary migratory corridor, but instead that most whales migrate through Hecate Strait and Dixon Entrance, broad waterways that lie to the east and north of Haida Gwaii. By using this route, northbound gray whales potentially face a wider range of industrial activities and developments than they would by migrating along the outer coast.     

Assessing polar bear (Ursus maritimus) population structure in the Hudson Bay region using SNPs

Defining subpopulations using genetics has traditionally used data from microsatellite markers to investigate population structure; however, single‐nucleotide polymorphisms (SNPs) have emerged as a tool for detection of fine‐scale structure. In Hudson Bay, Canada, three polar bear (Ursus maritimus) subpopulations (Foxe Basin (FB), Southern Hudson Bay (SH), and Western Hudson Bay (WH)) have been delineated based on mark–recapture studies, radiotelemetry and satellite telemetry, return of marked animals in the subsistence harvest, and population genetics using microsatellites. We used SNPs to detect fine‐scale population structure in polar bears from the Hudson Bay region and compared our results to the current designations using 414 individuals genotyped at 2,603 SNPs. Analyses based on discriminant analysis of principal components (DAPC) and STRUCTURE support the presence of four genetic clusters: (i) Western—including individuals sampled in WH, SH (excluding Akimiski Island in James Bay), and southern FB (south of Southampton Island); (ii) Northern—individuals sampled in northern FB (Baffin Island) and Davis Strait (DS) (Labrador coast); (iii) Southeast—individuals from SH (Akimiski Island in James Bay); and (iv) Northeast—individuals from DS (Baffin Island). Population structure differed from microsatellite studies and current management designations demonstrating the value of using SNPs for fine‐scale population delineation in polar bears.     

Stomach contents of bowhead whales (Balaena mysticetus) from four locations in the Canadian Arctic

The stomach contents of four bowhead whales (Balaena mysticetus) harvested between 1994 and 2008 from the Canadian Arctic were examined to assess diet composition. Three samples were collected from bowhead whales of the Eastern Canada–West Greenland (EC–WG) population and represent, according to our knowledge, the first diet analysis from this bowhead whale stock. We also examined the stomach content of one bowhead whale from the Bering-Chukchi-Beaufort (BCB) population hunted in 1996. All four whales had food in their stomachs and their diet varied from exclusively pelagic (BCB whale), with Limnocalanus macrurus being the main prey, to epibenthic and benthic (EC–WG) with Mysis oculata playing an important role. These results indicate broad foraging spectrum of the bowhead whales and add to a basic knowledge of their diet.     

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

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Ovary development in Greenland halibut Reinhardtius hippoglossoides in west Greenland waters

Maturity in adult female Greenland halibut Reinhardtius hippoglossoides was studied in three areas in west Greenland waters: the inshore area in Disko Bay and two offshore areas, Baffin Bay and Davis Strait. The aim was to monitor maturity changes in the inshore fjords of Disko Bay over an extended period from winter to autumn and compare these findings with specimens from Baffin Bay and the presumed spawning area in Davis Strait. A significant difference in maturity level was observed in and between the three areas. In Disko Bay maturity indices increased significantly in August and September both with respect to the gonado‐somatic index ( I _G) and the size in the leading oocyte cohort. In the period February to May no significant changes were observed. Mature ovaries were only observed among fish >80 cm total length and only among a fraction of these large fish. Offshore areas of Baffin Bay, even though poorly sampled, showed similar signs in the maturity indices as in Disko Bay. Relative to Disko Bay and Baffin Bay, female fish in Davis Strait had more progressed maturity indices. Furthermore, almost all fish in Davis Strait showed signs of progressed maturity contrary to Disko and Baffin Bay. A large proportion of the Greenland halibut in Disko and Baffin Bay apparently did not begin the maturation cycle until very late in their life history or were repeat spawners with a multi‐year maturation cycle. These observations could thus support the hypothesis that Greenland halibut have a prolonged adolescent phase. Atresia was highest in the early phases of maturation in Greenland halibut but relatively high levels of atresia were also observed in fish in more advanced maturity phase. The first was ascribed to fecundity regulation while the latter could be linked to the fish's fitness condition but it was not possible to show this with the available condition index.     

Trends in bowhead whales in West Greenland: Aerial surveys vs. genetic capture‐recapture analyses

We contrast two methods for estimating the trends of bowhead whales (Balaena mysticetus) in West Greenland: (1) double platform visual aerial survey, corrected for missed sightings and the time the whales are available at the surface; and (2) a genetic capture‐recapture approach based on a 14‐yr‐long biopsy sampling program in Disko Bay. The aerial survey covered 39,000 km² and resulted in 58 sightings, yielding an abundance estimate of 744 whales (CV = 0.34, 95% CI: 357–1,461). The genetic method relied on determining sex, mitochondrial haplotypes and genotypes of nine microsatellite markers. Based on samples from a total of 427 individuals, with 11 recaptures from previous years in 2013, this resulted in an estimate of 1,538 whales (CV = 0.24, 95% CI: 827–2,249). While the aerial survey is considered a snapshot of the local spring aggregation in Disko Bay, the genetic approach estimates the abundance of the source of this aggregation. As the whales in Disko Bay primarily are adult females that do not visit the bay annually, the genetic method would presumably yield higher estimates. The studies indicate that an increase in abundance observed between 1998 and 2006 has leveled off.     

Satellite tracking of a killer whale (Orcinus orca) in the eastern Canadian Arctic documents ice avoidance and rapid, long-distance movement into the North Atlantic

Killer whales (Orcinus orca) occur in the eastern Canadian Arctic during the open-water season, but their seasonal movements in Arctic waters and overall distribution are poorly understood. During August 2009, satellite transmitters were deployed onto two killer whales in Admiralty Inlet, Baffin Island, Canada. A whale tracked for 90 days remained in Admiralty and Prince Regent Inlets from mid-August until early October, when locations overlapped aggregations of marine mammal prey species. While in Admiralty and Prince Regent Inlets, the whale traveled 96.1 ± 45.3 km day⁻¹ (max 162.6 km day⁻¹) and 120.1 ± 44.5 km day⁻¹ (max 192.7 km day⁻¹), respectively. Increasing ice cover in Prince Regent Inlet in late September and early October was avoided, and the whale left the region prior to heavy ice formation. The whale traveled an average of 159.4 ± 44.8 km day⁻¹ (max 252.0 km day⁻¹) along the east coast of Baffin Island and into the open North Atlantic by mid-November, covering over 5,400 km in approximately one month. This research marks the first time satellite telemetry has been used to study killer whale movements in the eastern Canadian Arctic and documents long-distance movement rarely observed in this species.     

Comparing marine mammal acoustic habitats in Atlantic and Pacific sectors of the High Arctic: year-long records from Fram Strait and the Chukchi Plateau

During the International Polar Year (IPY), acoustic recorders were deployed on oceanographic moorings in Fram Strait and on the Chukchi Plateau, representing the first coordinated year-round sampling of underwater acoustic habitats at two sites in the High Arctic. Examination of species-specific marine mammal calls recorded from autumn 2008–2009 revealed distinctly different acoustic habitats at each site. Overall, the Fram Strait site was acoustically complex compared with the Chukchi Plateau site. In Fram Strait, calls from bowhead whales (Balaena mysticetus) and a variety of toothed whales (odontocetes) were recorded year-round, as were airgun pulses from seismic surveys. In addition, calls from blue whales (Balaenoptera musculus) and fin whales (B. physalus) were recorded from June to October and August to March, respectively. Conversely, at the Chukchi Plateau site, beluga (Delphinapterus leucas) and bowhead whale calls were recorded primarily from May to August, with airgun signals detected only in September–October. Ribbon seal (Phoca fasciata) calls were detected in October–November, with no marine mammals calls at all recorded from December to February. Of note, ice-adapted bearded seals (Erignathus barbatus) were recorded at both sites, primarily in spring and summer, corresponding with the mating season for that species. Differences in acoustic habitats between the two sites were related to contrasts in sea ice cover, temperature, patterns of ocean circulation and contributions from anthropogenic noise sources. These data provide a provisional baseline for the comparison of underwater acoustic habitats between Pacific and Atlantic sectors of the High Arctic.     

Fractal analysis of narwhal space use patterns

Quantifying animal movement in response to a spatially and temporally heterogeneous environment is critical to understanding the structural and functional landscape influences on population viability. Generalities of landscape structure can easily be extended to the marine environment, as marine predators inhabit a patchy, dynamic system, which influences animal choice and behavior. An innovative use of the fractal measure of complexity, indexing the linearity of movement paths over replicate temporal scales, was applied to satellite tracking data collected from narwhals (Monodon monoceros) (n = 20) in West Greenland and the eastern Canadian high Arctic. Daily movements of individuals were obtained using polar orbiting satellites via the ARGOS data location and collection system. Geographic positions were filtered to obtain a daily good quality position for each whale. The length of total pathway was measured over seven different temporal length scales (step lengths), ranging from one day to one week, and a seasonal mean was calculated. Fractal dimension (D) was significantly different between seasons, highest during summer (D = 1.61, SE 0.04) and winter (D = 1.69, SE 0.06) when whales made convoluted movements in focal areas. Fractal dimension was lowest during fall (D = 1.34, SE 0.03) when whales were migrating south ahead of the forming sea ice. There were no significant effects of size category or sex on fractal dimension by season. The greater linearity of movement during the migration period suggests individuals do not intensively forage on patchy resources until they arrive at summer or winter sites. The highly convoluted movements observed during summer and winter suggest foraging or searching efforts in localized areas. Significant differences between the fractal dimensions on two separate wintering grounds in Baffin Bay suggest differential movement patterns in response to the dynamics of sea ice.     

A bowhead whale in the eastern North Pacific

Bowhead whales occur in the Arctic year‐round. Their movements are largely correlated with seasonal expansions and reductions of sea ice, but a few recent extralimital sightings have occurred in the eastern and western North Atlantic and one was also documented in the western North Pacific over 50 years ago. Here we present details of a juvenile bowhead whale that was photographed and filmed from above and below the water while it was skim‐feeding in Caamaño Sound, BC, Canada on May 31, 2016. This sighting occurred over 2000 km southeast from the nearest known range for this species in the Bering Sea at a time that most bowhead whales in that region would have been migrating northeast. This sighting represents the first and only documentation of a bowhead whale in the eastern North Pacific to date.     

Dietary habits of polar bears in Foxe Basin,Canada: possible evidence of a trophic regime shift mediated by a new top predator

Polar bear (Ursus maritimus) subpopulations in several areas with seasonal sea ice regimes have shown declines in body condition, reproductive rates, or abundance as a result of declining sea ice habitat. In the Foxe Basin region of Nunavut, Canada, the size of the polar bear subpopulation has remained largely stable over the past 20 years, despite concurrent declines in sea ice habitat. We used fatty acid analysis to examine polar bear feeding habits in Foxe Basin and thus potentially identify ecological factors contributing to population stability. Adipose tissue samples were collected from 103 polar bears harvested during 2010–2012. Polar bear diet composition varied spatially within the region with ringed seal (Pusa hispida) comprising the primary prey in northern and southern Foxe Basin, whereas polar bears in Hudson Strait consumed equal proportions of ringed seal and harp seal (Pagophilus groenlandicus). Walrus (Odobenus rosmarus) consumption was highest in northern Foxe Basin, a trend driven by the ability of adult male bears to capture large‐bodied prey. Importantly, bowhead whale (Balaena mysticetus) contributed to polar bear diets in all areas and all age and sex classes. Bowhead carcasses resulting from killer whale (Orcinus orca) predation and subsistence harvest potentially provide an important supplementary food source for polar bears during the ice‐free period. Our results suggest that the increasing abundance of killer whales and bowhead whales in the region could be indirectly contributing to improved polar bear foraging success despite declining sea ice habitat. However, this indirect interaction between top predators may be temporary if continued sea ice declines eventually severely limit on‐ice feeding opportunities for polar bears.     

Spatial and seasonal distribution of American whaling and whales in the age of sail

American whalemen sailed out of ports on the east coast of the United States and in California from the 18(th) to early 20(th) centuries, searching for whales throughout the world's oceans. From an initial focus on sperm whales (Physeter macrocephalus) and right whales (Eubalaena spp.), the array of targeted whales expanded to include bowhead whales (Balaena mysticetus), humpback whales (Megaptera novaeangliae), and gray whales (Eschrichtius robustus). Extensive records of American whaling in the form of daily entries in whaling voyage logbooks contain a great deal of information about where and when the whalemen found whales. We plotted daily locations where the several species of whales were observed, both those caught and those sighted but not caught, on world maps to illustrate the spatial and temporal distribution of both American whaling activity and the whales. The patterns shown on the maps provide the basis for various inferences concerning the historical distribution of the target whales prior to and during this episode of global whaling.     

PATTERNS OF BOWHEAD WHALE DISTRIBUTION AND ABUNDANCE NEAR BARROW, ALASKA, IN FALL 1982-1989

Although the distribution and relative abundance of bowhead whales varied annually within the fall whaling area near Barrow, Alaska, the distance of whales from shore was not significantly different among years 1982-1989 (ANOVA, F = 0.5, P > 0.5). The minimum detectable distance for the ANOVA was 12 km (α= 0.05, β= 0.1). Annual median distance of random bowhead sightings from shore ranged from 23 to 39 km, with an eight-year median of 32 km. Highest annual bowhead sighting rates were positively associated with the proportion of feeding whales, indicating that whale feeding opportunities may affect the availability of whales within hunting range each fall.