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.     

Lycodes nishimurai, a new deep-sea eelpout (Teleostei: Zoarcidae) from the Sea of Japan

A new zoarcid fish, Lycodes nishimuari, is described on the basis of four specimens from the Yamato Bank, the Sea of Japan, at depths of 642–669 m. Although this species resembles Lycodes diapterus, L. hubbsi, L. pectoralis, L. macrochir, and L. nakamurae in having a deep notch in the pectoral fin and a single lateral line on the body, it is distinguished from them by the following combination of characters: no scales on pectoral fin base; scales on dorsum before dorsal fin origin; suborbital, preopercular, and last postorbital pores with risen rim; whitish lips; no whitish transverse bars on body; pectoral fin membrane and anterior part of dorsal fin membrane without black spots; basal color of vertical fins black. Supplementary material to this paper is available in electronic format at .     

A new zoarcid, Lycodes mcallisteri, from eastern Arctic Canada (Teleostei: Perciformes)

A new zoarcid fish, Lycodes mcallisteri, is described on the basis of nine specimens (230–375 mm SL) from eastern Arctic Canada, Nunavut (Hudson Strait, 298 m; Cumberland Sound and Baffin Bay, 579 m). It resembles about 15 other Lycodes spp. in having a long tail and a "ventromediolateral" type of lateral line. It is distinguished from Arctic members of this group (Lycodes marisalbi, L. palearis, and L. sagittarius) by the high number of pectoral fin rays (22–23), short pelvic fins, low submental crests, and coloration (6–10 light patches dorsally). It resembles two Pacific species (L. macrochir and L. soldatovi) by the high number of pectoral fin rays, but differs by coloration, meristics, and shape of the pectoral fin. Received: April 13, 2000 / Revised: September 7, 2000 / Accepted: January 11, 2001     

THE DISTRIBUTION OF BOWHEAD WHALES, BALAENA MYSTICETUS, IN THE CHUKCHI SEA

Under a U.S.-U.S.S.R. cooperative Marine Mammal Project, shipboard cruises were made in 1979, 1980 and 1982, primarily to determine whether there is a substock of western Arctic bowhead whales ( Balaena mysticetus ) that summers in the western Chukchi Sea instead of migrating to the Beaufort Sea. More than 100 bowheads were sighted along the north Chukotka coast of Siberia in October 1979, and more than 200 bowheads were sighted there in September 1980. None were seen anywhere in the Chukchi Sea in late July and August 1982. We conclude that the September and October sightings were of animals returning early from the Beaufort Sea and that, other than occurrences peripheral to the main migration, there are no large concentrations in the Chukchi in the summer and there is apparently no western Chukchi substock.     

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.     

Phylogeny of the eelpout genus Lycodes (Pisces,Zoarcidae) as inferred from mitochondrial cytochrome b and 12S rDNA

The bottom-dwelling and species-rich eelpout genus Lycodes Reinhardt has a great potential for the study of Arctic marine speciation. Subdivision of the genus has been based on single or few morphological characters (e.g., lateral line configuration) with contradicting results and phylogenetic approaches have not been attended. Here we present the first phylogenetic analysis of the genus employing DNA sequences of the mitochondrial genes cytochrome b and 12S rDNA (714 bp). The analysis with the two genes combined resulted in two equally parsimonious trees. In both cladograms most of the previously suggested subgroups are para- or polyphyletic, except for the so-called short-tailed Lycodes spp., with a short tail, a single mediolateral lateral line configuration and a shallow or filled otolith sulcus. The group of long-tailed Lycodes spp., with ventral or ventro-medio-lateral types of lateral line configuration and a deep otolith sulcus, appears to be paraphyletic, since Pacific and Atlantic species in this group are not each other's closest relatives. Thus, the short-tailed species are placed in a derived clade, indicating a secondary shortening of the tail, and a "slope to shore" type of evolution. This is not in accordance with earlier assumptions of the more elongate, deeper living species being the more derived. The basal position of long-tailed Pacific species supports earlier theories of Pacific origin of the genus/family. Small genetic differences between Arctic/Atlantic species indicate a rather recent radiation in these areas after the opening of the Bering Strait 3.0-3.5 million years ago.     

Assessment of primary production in the White Sea

The literature and original data on the primary production of phytoplankton in the White Sea are analyzed. By this parameter, the White Sea is significantly inferior only to the Chukchi Sea; it is similar to the Barents Sea, and exceeds other Russian Arctic seas by two to three times (the Kara Sea, Laptev Sea, and East Siberian Sea).     

Summer diet of beluga whales inferred by fatty acid analysis of the eastern Beaufort Sea food web

Beluga whales (Delphinapterus leucas) are the most abundant odontocetes in Arctic waters and are thus thought to influence food web structure and function. The diet of the Beaufort Sea beluga population is not well known, partly due to the inherent difficulty of observing feeding behaviour in Arctic marine cetaceans. To determine which prey items are critical to the Beaufort Sea beluga diet we first examine and describe the Mackenzie Delta and Beaufort Sea food web using fatty acid analyses. Fatty acid profiles effectively partitioned prey items into groups associated with their habitat and feeding ecology. Next, the relative contribution of various prey items to beluga diet was investigated using fatty acids. Finally, beluga diet variability was examined as a function of body size, a known correlate of habitat use. Beluga appeared to feed predominantly on Arctic cod (Boreogadus saida) collected from near shore and offshore regions. Size related dietary differences suggested larger sized beluga preferred offshore Arctic cod given the shared high levels of long chain monounsaturates, whereas smaller sized beluga appeared to feed on prey in their near shore habitats that included near shore Arctic cod. The presence of Arctic cod groups in shallow near shore and deep offshore habitats may facilitate the behavioural segregation of beluga habitat use as it relates to their size and resource requirements. Given Arctic cod are a sea ice associated fish combined with the accelerated sea ice loss in this region, beluga whales may need to adapt to new dietary regimes.     

Composition of the summer photosynthetic pico and nanoplankton communities in the Beaufort Sea assessed by T-RFLP and sequences of the 18S rRNA gene from flow cytometry sorted samples

The composition of photosynthetic pico and nanoeukaryotes was investigated in the North East Pacific and the Arctic Ocean with special emphasis on the Beaufort Sea during the MALINA cruise in summer 2009. Photosynthetic populations were sorted using flow cytometry based on their size and pigment fluorescence. Diversity of the sorted photosynthetic eukaryotes was determined using terminal-restriction fragment length polymorphism analysis and cloning/sequencing of the 18S ribosomal RNA gene. Picoplankton was dominated by Mamiellophyceae, a class of small green algae previously included in the prasinophytes: in the North East Pacific, the contribution of an Arctic Micromonas ecotype increased steadily northward becoming the only taxon occurring at most stations throughout the Beaufort Sea. In contrast, nanoplankton was more diverse: North Pacific stations were dominated by Pseudo-nitzschia sp. whereas those in the Beaufort Sea were dominated by two distinct Chaetoceros species as well as by Chrysophyceae, Pelagophyceae and Chrysochromulina spp.. This study confirms the importance of Arctic Micromonas within picoplankton throughout the Beaufort Sea and demonstrates that the photosynthetic picoeukaryote community in the Arctic is much less diverse than at lower latitudes. Moreover, in contrast to what occurs in warmer waters, most of the key pico- and nanoplankton species found in the Beaufort Sea could be successfully established in culture.     

Arctic waters: Needs and options for CANADIAN‐AMERICAN cooperation

Abstract

Four major factors bid the United States and Canada to move toward more formalized arrangements for cooperative ocean management in the Arctic. Ocean currents in the Beaufort Sea region have the potential of transporting marine pollutants from one country to the other. Living resources, such as bowhead and beluga whales, undertake extensive transboundary migrations. Alaskan and Canadian Inuit depend heavily on renewable marine resources and raise the need for ocean management on an ecological basis. Cost savings could occur by coordinating development of offshore cold‐water technologies and shipping safety systems. This paper suggests six options for moving toward a more regionalized approach to the management of Arctic waters: a Beaufort Sea Boundary Agreement, a Beaufort Marine Cooperation Agreement, a Northwest Passage Agreement, an Equal Access Agreement, a Marine Mammal Cooperation Agreement, and an Arctic Regional Action Plan.     

New Data on the Distribution of Lamprey <Emphasis Type="Italic">Petromyzon marinus</Emphasis> and <Emphasis Type="Italic">Lethenteron camtschaticum</Emphasis> (Petromyzontidae) in Barents and White Seas

Information about the sites of catches of the sea lamprey Petromyzon marinus in the western Barents Sea and Arctic lamprey Lethenteron camtschaticum in the Barents and White seas is presented based on the data of trawl surveys performed in 2004?2016. It is demonstrated that sea lamprey is occasionally encountered in the western Barents Sea; nine specimens have been recorded during the entire period of surveys. The northernmost point of a capture of sea lamprey is located near 76° N and the easternmost point is at 31°15′ E. Arctic lamprey is not numerous in the Barents and White seas; a total of 66 and 17 specimens have been caught, respectively. Its local aggregations are found in the southeastern part of the Barents Sea and in Dvina Bay in the White Sea. Arctic lamprey penetrates to the north to 76° N and into the central part of the Barents Sea.     

Presence and behavior of bowhead whales (Balaena mysticetus) in the Alaskan Beaufort Sea in July 2011

The Western Arctic bowhead whale (Balaena mysticetus) is highly adapted to sea ice and annually migrates through the Bering, Chukchi, and Beaufort seas. While the overall distribution and seasonal movements of bowhead whales are mostly understood, information about their distribution in the Alaskan Beaufort Sea in early to mid-summer has not been well documented. In July 2011, we conducted an exploratory flight in the Alaskan Beaufort Sea, north of Camden Bay (71°N 144°W), near the location of a single satellite-tagged bowhead whale. Eighteen bowhead whales were observed, and behavior consistent with feeding was documented. To our knowledge, this is the first documentation of behavior consistent with feeding north of Camden Bay in mid-July. Few studies have focused on bowhead whale distribution in the Alaskan Beaufort Sea in early to mid-summer, and no long-term, region-wide surveys have been conducted during summer. Bowhead whales are already exposed to anthropogenic disturbance in the Canadian Beaufort Sea in summer, the Alaskan Beaufort Sea in fall, and the Chukchi and Bering seas from fall through spring. The presence of bowhead whale aggregations in the Alaskan Beaufort Sea in summer should be considered when assessing the cumulative effects of human-related activities.     

Ringed seal (Pusa hispida) tooth annuli as an index of reproduction in the Beaufort Sea

Multi-decadal time-series of biological indices that reflect the state of a population are rare in ecological studies, but invaluable for assessing environmental regulation of population dynamics. We utilized canine teeth extracted from ringed seals (Pusa hispida) killed by polar bears (Ursus maritimus) in the Beaufort Sea, Canada, in 1985–2011, to obtain widths of annual growth layers in the cementum. Canine teeth for 75 individuals were measured and compared across years using a proportional width index (PWI) spanning 1965–2007. PWI was positively correlated with ringed seal ovulation rate obtained independently from other studies and was significantly lower than normal during ringed seal reproductive declines in 1974–75, 1984–87, 1991–93, and 2004–05, suggesting that PWI reflects ringed seal reproductive capacity. The PWI was also examined against climatic and sea ice factors to assess environmental regulation of ringed seal reproduction. Results suggest that ringed seals benefit from cyclonic circulation regimes in the Beaufort Sea, and an earlier breakup of sea ice in summer that may positively influence the quality and quantity of food during the open water season. Results highlight how cementum annuli in the canine teeth of ringed seals can provide an index of body state and linkages to sea ice conditions. Canine teeth from ringed seals can function as a means to monitor the effects of past Arctic marine variability on area-specific populations for which there are few independent empirical data.     

Long-term assessment of relationships between changing environmental conditions and the physiology of southern Beaufort Sea polar bears (Ursus maritimus)

Climate change is influencing polar bear (Ursus maritimus) habitat, diet, and behavior but the effects of these changes on their physiology is not well understood. Blood-based biomarkers are used to assess the physiologic health of individuals but their usefulness for evaluating population health, especially as it relates to changing environmental conditions, has rarely been explored. We describe links between environmental conditions and physiologic functions of southern Beaufort Sea polar bears using data from blood samples collected from 1984 to 2018, a period marked by extensive environmental change. We evaluated associations between 13 physiologic biomarkers and circumpolar (Arctic oscillation index) and regional (wind patterns and ice-free days) environmental metrics and seasonal and demographic co-variates (age, sex, season, and year) known to affect polar bear ecology. We observed signs of dysregulation of water balance in polar bears following years with a lower annual Arctic oscillation index. In addition, liver enzyme values increased over time, which is suggestive of potential hepatocyte damage as the Arctic has warmed. Biomarkers of immune function increased with regional-scale wind patterns and the number of ice-free days over the Beaufort Sea continental shelf and were lower in years with a lower winter Arctic oscillation index, suggesting an increased allocation of energetic resources for immune processes under these conditions. We propose that the variation in polar bear immune and metabolic function is likely indicative of physiologic plasticity, a response that allows polar bears to remain in homeostasis even as they experience changes in nutrition and habitat in response to changing environments.     

Spring fasting behavior in a marine apex predator provides an index of ecosystem productivity

The effects of declining Arctic sea ice on local ecosystem productivity are not well understood but have been shown to vary inter‐specifically, spatially, and temporally. Because marine mammals occupy upper trophic levels in Arctic food webs, they may be useful indicators for understanding variation in ecosystem productivity. Polar bears (Ursus maritimus) are apex predators that primarily consume benthic and pelagic‐feeding ice‐associated seals. As such, their productivity integrates sea ice conditions and the ecosystem supporting them. Declining sea ice availability has been linked to negative population effects for polar bears but does not fully explain observed population changes. We examined relationships between spring foraging success of polar bears and sea ice conditions, prey productivity, and general patterns of ecosystem productivity in the Beaufort and Chukchi Seas (CSs). Fasting status (≥7 days) was estimated using serum urea and creatinine levels of 1,448 samples collected from 1,177 adult and subadult bears across three subpopulations. Fasting increased in the Beaufort Sea between 1983–1999 and 2000–2016 and was related to an index of ringed seal body condition. This change was concurrent with declines in body condition of polar bears and observed changes in the diet, condition and/or reproduction of four other vertebrate consumers within the food chain. In contrast, fasting declined in CS polar bears between periods and was less common than in the two Beaufort Sea subpopulations consistent with studies demonstrating higher primary productivity and maintenance or improved body condition in polar bears, ringed seals, and bearded seals despite recent sea ice loss in this region. Consistency between regional and temporal variation in spring polar bear fasting and food web productivity suggests that polar bears may be a useful indicator species. Furthermore, our results suggest that spatial and temporal ecological variation is important in affecting upper trophic‐level productivity in these marine ecosystems.     

Population structure and variability of Pacific herring (<Emphasis Type="Italic">Clupea pallasii</Emphasis>) in the White Sea,Barents and Kara Seas revealed by microsatellite DNA analyses

Pacific herring, Clupea pallasii, have recently colonised the northeast Atlantic and Arctic Oceans in the early Holocene. In a relatively short evolutionary time, the herring formed a community with a complex population structure. Previous genetic studies based on morphological, allozyme and mitochondrial DNA data have supported the existence of two herring subspecies from the White Sea and eastern Barents and Kara Seas (C. p. marisalbi and C. p. suworowi, respectively). However, the population structure of the White Sea herring has long been debated and remains controversial. The analyses of morphological and allozyme data have previously identified local spawning groups of herring in the White Sea, whereas mtDNA markers have not revealed any differentiation. We conducted one of the first studies of microsatellite variation for the purpose of investigating the genetic structure and relationship of Pacific herring among ten localities in the White Sea, the Barents Sea and the Kara Sea. Using classical genetic variance-based methods (hierarchical AMOVA, overall and pairwise F _ST comparisons), as well as the Bayesian clustering, we infer considerable genetic diversity and population structure in herring at ten microsatellite loci. Genetic differentiation was the most pronounced between the White Sea (C. p. marisalbi) versus the Barents and Kara seas (Chesha–Pechora herring, C. p. suworowi). While microsatellite variation in all C. pallasii was considerable, genetic diversity was significantly lower in C. p. suworowi, than in C. p. marisalbi. Also, tests of genetic differentiation were indicating significant differentiation within the White Sea herring between sympatric summer- and spring-spawning groups, in comparison with genetic homogeneity of the Chesha–Pechora herring.     

Gene flow on ice: the role of sea ice and whaling in shaping Holarctic genetic diversity and population differentiation in bowhead whales (Balaena mysticetus)

Sea ice is believed to be a major factor shaping gene flow for polar marine organisms, but it remains unclear to what extent it represents a true barrier to dispersal for arctic cetaceans. Bowhead whales are highly adapted to polar sea ice and were targeted by commercial whalers throughout Arctic and subarctic seas for at least four centuries, resulting in severe reductions in most areas. Both changing ice conditions and reductions due to whaling may have affected geographic distribution and genetic diversity throughout their range, but little is known about range‐wide genetic structure or whether it differed in the past. This study represents the first examination of genetic diversity and differentiation across all five putative stocks, including Baffin Bay‐Davis Strait, Hudson Bay‐Foxe Basin, Bering‐Beaufort‐Chukchi, Okhotsk, and Spitsbergen. We also utilized ancient specimens from Prince Regent Inlet (PRI) in the Canadian Arctic and compared them with modern stocks. Results from analysis of molecular variance and demographic simulations are consistent with recent and high gene flow between Atlantic and Pacific stocks in the recent past. Significant genetic differences between ancient and modern populations suggest PRI harbored unique maternal lineages in the past that have been recently lost, possibly due to loss of habitat during the Little Ice Age and/or whaling. Unexpectedly, samples from this location show a closer genetic relationship with modern Pacific stocks than Atlantic, supporting high gene flow between the central Canadian Arctic and Beaufort Sea over the past millennium despite extremely heavy ice cover over much of this period.     

Redescription of large-scale eelpout <Emphasis Type="Italic">Lycodes macrolepis</Emphasis> Taranetz et Andriashev, 1935 (Perciformes: Zoarcidae)

Morphology of a rare species—large-scale eelpout Lycodes macrolepis Taranetz et Andriashev, 1935—has been studied. It was shown that the main lateral line of this species has a ventrolateral configuration rather than medial, as was considered previously. Large-scale eelpout is an endemic species of the Sea of Okhotsk; evidence of its finding in the Sea of Japan is based on the erroneous classification of another eelpout species.     

Long-term increases in young-of-the-year growth of Arctic cisco Coregonus autumnalis and environmental influences

Arctic cisco Coregonus autumnalis young-of-year (YOY) growth was used as a proxy to examine the long-term response of a high-latitude fish population to changing climate from 1978 to 2004. YOY growth increased over time (r2 = 0·29) and was correlated with monthly averages of the Arctic oscillation index, air temperature, east wind speed, sea-ice concentration and river discharge with and without time lags. Overall, the most prevalent correlates to YOY growth were sea-ice concentration lagged 1 year (significant correlations in 7 months; r2 = 0·14-0·31) and Mackenzie River discharge lagged 2 years (significant correlations in 8 months; r2 = 0·13-0·50). The results suggest that decreased sea-ice concentrations and increased river discharge fuel primary production and that life cycles of prey species linking increased primary production to fish growth are responsible for the time lag. Oceanographic studies also suggest that sea ice concentration and fluvial inputs from the Mackenzie River are key factors influencing productivity in the Beaufort Sea. Future research should assess the possible mechanism relating sea ice concentration and river discharge to productivity at upper trophic levels.     

Mitochondrial Lineages of the Beluga Whale <Emphasis Type="Italic">Delphinapterus leucas</Emphasis> in the Russian Arctic

The data on mtDNA sequences of the beluga whale (Delphinapterus leucas) from the central Eastern Arctic as well as coastal waters of the Chukchi Peninsula and different parts of the White Sea are presented and analyzed for the first time. Certain sequences found in the region form a separate phylogenetic clade. The distinctness of composition of maternal lineages found for the White Sea is noted. However, the set of sequences found in the Russian Arctic in whole cannot be characterized as region-specific.