DISSOLVING STOCK DISCRETENESS WITH SATELLITE TRACKING: BOWHEAD WHALES IN BAFFIN BAY

Nine bowhead whales (Balaena mysticetus) were instrumented with satellite transmitters in West Greenland in May 2002 and 2003. Transmitters were either encased in steel cans or imbedded in floats attached to wires. Transmitters mounted in steel cans had a high initial failure rate, yet those that were successful provided tracking durations up to seven months. Float tags had a low initial failure rate and initially provided large numbers of positions; however, they had deployment durations of only 2–33 d. All tracked whales departed from West Greenland and headed northwest towards Lancaster Sound in the end of May. Three tags with long tracking durations (197–217 d) recorded movements of whales (1 ♂, 2 ♀) into December in 2002 and 2003. All of these individuals remained within the Canadian High Arctic or along the east coast of Baffin Island in summer and early fall. By the end of October, all three whales moved rapidly south along the east coast of Baffin Island and entered Hudson Strait, an apparent wintering ground for the population. One of the whales did not visit Isabella Bay on east Baffin Island, the locality used for abundance estimation from photographic reidentification of individuals. The movements of whales tagged in this study raise critical questions about the assumed stock discreteness of bowhead whales in Foxe Basin, Hudson Strait, and Davis Strait and indicate current estimates of abundance are negatively biased.     

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.     

Seasonal migrations of Sea of Okhotsk beluga whales (<Emphasis Type="Italic">Delphinapterus leucas</Emphasis>) of the Sakhalin-Amur summer aggregation

Seasonal migrations of beluga whales from the Sakhalin-Amur aggregation in the Sea of Okhotsk were investigated with the use of satellite telemetry. Satellite tags were attached to four females captured near Chkalova Island, Sakhalin Bay, in August 2007. At 5 weeks after tagging, the belugas left the Chkalova Island area and moved to the Nikolaya and Ul’banskii Bays in the Shantar Sea. The animals stayed in these bays (mainly in Nikolaya Bay) until the end of autumn and then traveled northward to deeper waters. In winter and spring, they preferred to stay in the regions with dense ice cover or close to the ice edge. During the winter migration, the tagged whales, as a rule, did not keep together, although they followed the same course with a little time lag. The female whose tag transmitted the longest (9.5 months) returned to Chkalova Island in late May and thus completed the seasonal migration cycle. Based on the data on the migration routes of the tagged belugas, we suggest that the Sakhalin-Amur and Shantar aggregations interact in autumn. It is also possible that they have similar winter migratory paths     

Autumn migration and wintering of northern fulmars (<Emphasis Type="Italic">Fulmarus glacialis</Emphasis>) from the Canadian high Arctic

Five northern fulmars (Fulmarus glacialis) were tracked by satellite transmitters from their breeding colony in the Canadian high Arctic (Cape Vera, Devon Island, NT) to their wintering grounds in the northwest Atlantic Ocean. In both 2004 and 2005, fulmars left northern Baffin Bay in mid- to late September, and migrated south to Davis Strait in less than 1 week, after which movements were erratic. In October and November, the birds were widely distributed, but by December through March, they tended to remain in the Labrador Sea between 50 and 55°N. Average flight speed was 35 km/h with a maximum of 64 km/h, and over their entire transmission periods, the five traveled on average 84 km/day. Our work suggests that the North Atlantic northern fulmar population may be panmictic in winter, with the Labrador Sea as a key wintering site for fulmars from high Arctic Canada.     

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.     

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.     

Key Factors Determining the Distribution of Beluga Whales, <Emphasis Type="Italic">Delphinapterus leucas</Emphasis> (Pallas, 1776), in River Estuaries of Western Kamchatka

The results of observations on the distribution of beluga whales, Delphinapterus leucas (Pallas, 1776), in three large rivers of western Kamchatka in the summer and autumn seasons are discussed. In the summer, the number of beluga whales in the Khairyuzova, Belogolovaya, and Moroshechnaya rivers reaches 111–250 individuals. Most of the belugas enter the rivers during the flood tidal phase: the number of animals in the estuaries increases along with the rising water level to the maximum value at spring tide. The belugas do not move upstream out of the estuaries and tend to remain in the zone of mixing riverine and marine waters, where 20 species of fish and three species of invertebrates have been identified. At ebb tide, the belugas leave for the sea; however, during a large run of salmon some individuals remain in the estuaries and continue hunting in deep-water areas. The main issue that causes beluga whales to form summer aggregations in Kamchatkan rivers is the hunt for salmon. The distribution of beluga whales in river estuaries is defined by the dynamics and intensity of salmon spawning runs. The preference of beluga whales for these rivers can be explained by the channel type of their estuaries.     

Movements of beluga whales (Delphinapterus leucas) in Bristol Bay,Alaska

We describe the annual distribution of beluga whales (Delphinapterus leucas) in Bristol Bay, Alaska, using data from 31 satellite‐linked transmitters during 2002–2011. Bristol Bay has one of the largest and best studied Pacific salmon (Oncorhynchus spp.) fisheries in the world, allowing us to link the seasonal distribution of belugas to that of salmon. During salmon migrations, beluga movements were restricted to river entrances. Belugas generally did not relocate to different river entrances or change bays during peak salmon periods. However, the location of belugas was not related to the number of salmon passing counting towers, suggesting that belugas were either selecting locations that were good for catching salmon or there were simply more salmon than belugas needed to supply their nutritional needs. The distribution of belugas expanded after salmon runs ended, and was greatest in winter when belugas ranged beyond the inner bays, traveling as far west as Cape Constantine. Belugas continued to frequent the inner bays in winter whenever sea ice conditions allowed, e.g., when winds moved sea ice offshore; however, they were never located south of the southern ice edge in open water or outside of Bristol Bay.     

Assessing the abundance of Bristol Bay belugas with genetic mark‐recapture methods

The Bristol Bay stock of beluga whales (Delphinapterus leucas) is genetically distinct and resides in Bristol Bay year‐round. We estimated the abundance of this population using genetic mark‐recapture, whereby genetic markers from skin biopsies, collected between 2002 and 2011, were used to identify individuals. We identified 516 individual belugas in two inner bays, 468 from Kvichak Bay and 48 from Nushagak Bay, and recaptured 75 belugas in separate years. Using a POPAN Jolly‐Seber model, abundance was estimated at 1,928 belugas (95% CI = 1,611–2,337), not including calves, which were not sampled. Most belugas were sampled in Kvichak Bay at a time when belugas are also known to occur in Nushagak Bay. The pattern of genetic recaptures and data from belugas with satellite transmitters suggested that belugas in the two bays regularly mix. Hence, the estimate of abundance likely applies to all belugas within Bristol Bay. Simulations suggested that POPAN estimates of abundance are robust to most forms of emigration, but that emigration causes negative bias in both capture and survival probabilities. Because it is likely that some belugas do not enter the sampling area during sampling, our estimate of abundance is best considered a minimum population size.     

Population-specific home ranges and migration timing of Pacific Arctic beluga whales (Delphinapterus leucas)

Two populations of beluga whales (Delphinapterus leucas), the Eastern Beaufort Sea (BS) and Eastern Chukchi Sea (ECS), make extensive seasonal migrations into the Pacific Arctic. However, the extent to which these populations overlap in time and space is not known. We quantified distribution and migration patterns for BS and ECS belugas using daily locations from whales tracked with satellite-linked transmitters. Home ranges and core areas in summer (July and August) and in each month (July–November), daily displacement, dispersal from core areas, and autumn migration timing were estimated. Distinct summer and fall distribution patterns and staggered autumn migration timing were identified for BS and ECS whales. Summer home ranges for each population had less than 10 % overlap. Monthly home ranges were also relatively distinct between populations except in September (up to 88 % home range overlap). A distinct east–west shift in focal area use occurred in September that persisted into October, with the two populations essentially switching longitudinal positions. Highest daily displacements occurred during the migratory period in September for BS whales and October for ECS whales, further indicating westward fall migration was offset between populations. Sexual segregation of males and females within a population also varied monthly. Autumn migration timing as well as differences in spatial and temporal segregation between BS and ECS beluga populations may be a result of maternally driven philopatry and population-specific adaptations to dynamically available resources. Our results contribute to the management of these populations by identifying seasonal area use and differences in migration patterns.     

Decadal shifts in autumn migration timing by Pacific Arctic beluga whales are related to delayed annual sea ice formation

Migrations are often influenced by seasonal environmental gradients that are increasingly being altered by climate change. The consequences of rapid changes in Arctic sea ice have the potential to affect migrations of a number of marine species whose timing is temporally matched to seasonal sea ice cover. This topic has not been investigated for Pacific Arctic beluga whales (Delphinapterus leucas) that follow matrilineally maintained autumn migrations in the waters around Alaska and Russia. For the sympatric Eastern Chukchi Sea (‘Chukchi’) and Eastern Beaufort Sea (‘Beaufort’) beluga populations, we examined changes in autumn migration timing as related to delayed regional sea ice freeze‐up since the 1990s, using two independent data sources (satellite telemetry data and passive acoustics) for both populations. We compared dates of migration between ‘early’ (1993–2002) and ‘late’ (2004–2012) tagging periods. During the late tagging period, Chukchi belugas had significantly delayed migrations (by 2 to >4 weeks, depending on location) from the Beaufort and Chukchi seas. Spatial analyses also revealed that departure from Beaufort Sea foraging regions by Chukchi whales was postponed in the late period. Chukchi beluga autumn migration timing occurred significantly later as regional sea ice freeze‐up timing became later in the Beaufort, Chukchi, and Bering seas. In contrast, Beaufort belugas did not shift migration timing between periods, nor was migration timing related to freeze‐up timing, other than for southward migration at the Bering Strait. Passive acoustic data from 2008 to 2014 provided independent and supplementary support for delayed migration from the Beaufort Sea (4 day yr^?1) by Chukchi belugas. Here, we report the first phenological study examining beluga whale migrations within the context of their rapidly transforming Pacific Arctic ecosystem, suggesting flexible responses that may enable their persistence yet also complicate predictions of how belugas may fare in the future.     

ICEBREAKER NOISE IN LANCASTER SOUND, N.W.T., CANADA: IMPLICATIONS FOR MARINE MAMMAL BEHAVIOR

Abstract: In 1986, we recorded the MV Arctic , CCGS des Groseilliers and MV Lady Franklin during routine icebreaking operations and travel to and from the mine at Nanisivik, Baffin Island, Northwest Territories, Canada. We found that the Arctic generated more high frequency noise than did the other vessels we recorded. Monitoring of vessel noise levels indicated that belugas and, probably, narwhals should be able to detect the high frequency components of Arctic noise at least as far as 25 to 30 km from the source. The ability of whales to detect the MV Arctic at long distances may explain why belugas and narwhals in Lancaster Sound seem to react to ships at longer distances than do other stocks of arctic whales.     

Occurrence of killer whale Orcinus orca rake marks on Eastern Canada-West Greenland bowhead whales Balaena mysticetus

Killer whales (Orcinus orca) are increasing in occurrence and residence time in the eastern Canadian Arctic (ECA) in part due to a decrease in sea ice associated with global climate change. Killer whales prey on bowhead whales (Balaena mysticetus) of the Eastern Canada-West Greenland (EC-WG) population, but their patterns of predation pressure and effect on the EC-WG population’s ability to recover from historical whaling remain unknown. We analyzed photographs of individual bowhead whale flukes from five regions within the EC-WG population’s geographic range (Cumberland Sound, Foxe Basin, Isabella Bay, Repulse Bay and Disko Bay), taken during 1986 and from 2007 to 2012, to estimate the occurrence of rake marks (parallel scars caused by killer whale teeth). Of 598 identified whales, 10.2 % bore rake marks from killer whales. A higher occurrence of rake marks was found in Repulse and Disko Bays, where primarily adult bowhead whales occur seasonally, than in Foxe Basin, where juveniles and females with calves occur. Older bowheads, which have had greater exposure time to killer whales due to their age, had higher occurrences of rake marks than juveniles and calves, which may indicate that younger whales do not survive killer whale attacks. A high proportion of adult females also had rake marks, perhaps due to protecting their calves from killer whale predation. In order to quantify the effect of killer whales on EC-WG population recovery, further research is needed on the relationship between the occurrence of rake marks and bowhead adult, calf, and juvenile mortality in the ECA, as well as more information about Arctic killer whale ecology.     

Serologic evidence of influenza A infection in marine mammals of arctic Canada.

A serologic survey of influenza A antibodies was undertaken on 1,611 blood samples from five species of marine mammals collected from Arctic Canada from 1984-98. Sampling was done in 24 locations throughout the Canadian Arctic encompassing Sachs Harbor (72 degrees N, 125 degrees W), Northwest Territories in the west to Loks Land (63 degrees N, 64 degrees W), Nunavut in the east, to Eureka (80 degrees N, 86 degrees W), Nunavut in the north to Sanikiluaq (56 degrees N, 79 degrees W), Nunavut in the south. A competitive ELISA using a monoclonal antibody (Mab) against influenza A nucleoprotein (NP) was used. Five of 418 (1.2%) belugas (Delphinapterus leucas) and 23 of 903 (2.5%) ringed seals (Phoca hispida) were serologically positive. None of the 210 walruses (Odobenus rosmarus rosmarus), 76 narwhals (Monodon monoceros) and four bowhead whales (Balaena mysticetus) had detectable antibodies to influenza A. Positive belugas were identified from communities on southeast Baffin Island while positive ringed seals came from communities in the eastern, western and high Arctic. Virus isolation attempts on lung tissue from a seropositive beluga were unsuccessful. We believe that influenza A infection in marine mammals is sporadic, the infection is probably self-limiting, and it may not be able to be maintained in these animals. Although the predominant hemagglutinin (H) type was not determined and therefore the pathogenicity of the strains to humans is unknown, the hunting and consumption of marine mammals by the Inuit, may put them at risk for influenza A infection.     

The origin of Arctic terrestrial and freshwater tardigrades

The tardigrade faunas of six Arctic sites (Canadian Axel Heiberg I., east and west coasts of Greenland, Iceland, Svalbard, Novaya Zemlya and the Russian Taimyr Peninsula) form, with those of northern North America, a coherent “Nearctic-Arctic” biogeographic cluster. This cluster is distinct from that of “Northern and Alpine Europe”. Few, if any, Arctic tardigrades survived Pleistocene glaciation in situ amongst ice-free refugia. Similarly, few/none moved south ahead of the advancing ice-cap into the deglaciated Palaearctic and Nearctic and subsequently returned north during the Holocene deglaciation. It is more probable that most Arctic tardigrades are derived from wind-blown Nearctic propagules that colonized the region during the Holocene. Received: 2 July 1997 / Accepted: 3 October 1997     

Geographic variation in cranial morphology of narwhals (Monodon monoceros) from Greenland and the eastern Canadian Arctic

We analyzed variation in nine non-metric and eight metric variables in the skulls of 132 narwhals (Monodon monoceros) from five localities in Greenland (Inglefield Bredning, Melville Bay, Uummannaq, Disko Bay, and Scoresby Sound) and one in the eastern Canadian Arctic (Eclipse Sound). Metric variables were used to compare the combined Disko Bay and Uummannaq samples with the samples from Inglefield Bredning and Scoresby Sound using three different multivariate techniques for each sex. None of the results were significant. Seven of the non-metric variables were independent of age and sex and were used in comparing samples from the six localities. No differences were found among the four localities in West Greenland, but differences were found in two of the non-metric variables between the combined West Greenland sample and the one from Scoresby Sound. A major shortcoming of the analysis based on metric data was the small sample size from several of the areas, which resulted in low statistical power. Genetic as well as environmental factors could explain the differences detected here between narwhals living along the west and the east coasts of Greenland.     

Spatiotemporal segregation among summer stocks of beluga (Delphinapterus leucas) despite nuclear gene flow: implication for the endangered belugas in eastern Hudson Bay (Canada)

Migratory connectivity between areas frequented by wide-ranging animals provides crucial information for conservation and management. In and around Hudson Bay (Canada), three stocks of beluga whales (Delphinapterus leucas) are associated with distinct summering areas. We analyzed genetic variation at mtDNA and 13 microsatellite loci among individuals (N > 1400) harvested by 23 Inuit communities to identify mating units and assess temporal and spatial differences in the way stocks use common migratory pathways. Strong structure at mtDNA and a lack of convincing evidence for nuclear genetic differentiation indicate that both males and females adopt distinct migratory routes towards summering grounds while probably interbreeding on wintering grounds. Spatiotemporal variation in stock composition indicates that subsistence hunting targets all three stocks. While representing ca. 5% of belugas in Hudson Bay, the endangered Eastern Hudson Bay stock accounts for 17% of the overall subsistence harvest by Inuit communities of northern Nunavik (Quebec), and ca. 30% of the spring harvest along northeastern Hudson Bay. Despite interbreeding, cultural conservatism of maternally transmitted migration routes seems to prevent the re-establishment of stocks in previously frequented estuaries. This phenomenon supports the current use of demographic population models based on stock composition for developing behavior-based management strategies.     

Population genetic structure and evolutionary history of North Atlantic beluga whales (Delphinapterus leucas) from West Greenland, Svalbard and the White Sea

Population structure in many Arctic marine mammal species reflects a dynamic interplay between physical isolating mechanisms and the extent to which dispersal opportunities are met. We examined variation within mtDNA and eight microsatellite markers to investigate population structure and demographic history in beluga whales in the North Atlantic. Genetic heterogeneity was observed between Svalbard and West Greenland that reveals limited gene flow over ecological time scales. Differentiation was also recorded between Atlantic belugas and two previously studied populations in the North Pacific, the Beaufort Sea and Gulf of Alaska. However, Bayesian cluster analysis of the nDNA data identified two population clusters that did not correspond to the respective ocean basins, as predicted, but to: (1) Arctic (Svalbard–White Sea–Greenland–Beaufort Sea) and (2) Subarctic (Gulf of Alaska) regions. Similarly, the deepest phylogeographic signal was between the Arctic populations and the Gulf of Alaska. Fitting an isolation-with-migration model yielded genetic abundance estimates that match census estimates and revealed that Svalbard and the Beaufort Sea likely diverged 7,600–35,400 years ago but have experienced recurrent periods with gene flow since then, most likely via the Russian Arctic during subsequent warm periods. Considering current projections of continued sea ice losses in the Arctic, this study identified likely routes of future contact among extant beluga populations, and other mobile marine species, which have implications for genetic introgression, health, ecology and behavior.     

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.     

Mating ecology of beluga (Delphinapterus leucas) and narwhal (Monodon monoceros) as estimated by reproductive tract metrics

Narwhal and beluga whales are important species to Arctic ecosystems, including subsistence hunting by Inuit, and little is understood about their mating ecology. Reproductive tract metrics vary across species in relation to mating strategy, and have been used to infer mating ecology. Reproductive tracts from beluga and narwhal were collected between 1997 and 2008 from five beluga stocks and two narwhal stocks across the Canadian Arctic. Tract length for males and females, relative testes mass for males, and tusk length for male narwhal were measured. We assessed variation relative to species, body size, stock, maturity, and season. Significant variation was found in testes mass across month and stock for beluga, and no significant difference between stock or date of harvest for narwhal. Beluga had significantly larger testes relative to body size than narwhal, suggesting they were more promiscuous than narwhal. A significant relationship was found between narwhal tusk length and testes mass, indicating the tusk may be important in female mate choice. No significant differences were found between narwhal and beluga reproductive tract length for males or females. The mating systems suggested for narwhal and belugas by our results mean the two species may respond differently to climate change.