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.     

Ecosystem regime shifts have not affected growth and survivorship of eastern Beaufort Sea belugas

Large-scale ocean-atmosphere physical dynamics can have profound impacts on the structure and organization of marine ecosystems. These changes have been termed “regime shifts”, and five different episodes have been detected in the North Pacific Ocean, with concurrent changes also occurring in the Bering and Beaufort Seas. Belugas from the Eastern Beaufort Sea (EBS) use the Bering Sea during winter and the Beaufort Sea during summer, yet the potential effects of regime shifts on belugas have not been assessed. We investigated whether body size and survivorship of EBS belugas harvested in the Mackenzie River delta region between 1993 and 2003 have been affected by previous purported regime shifts in the North Pacific. Residuals from the relationship between body length and age were calculated and compared among belugas born between 1932 and 1989. Residual body size was not significantly related to birth year for any regime, nor to the age group individuals belonged to during any regime. The percentage deviation in number of belugas born in any given year that survived to be included in the hunt (survivorship) did not show any significant trend within or between regimes. Accounting for lags of 1–5 years did not reveal any evidence of delayed effects. Furthermore, neither population index was significantly related to changes in major climatic variables that precede regime shifts. Our results suggest that EBS beluga body size and survivorship have not been affected by the major regime shifts of the North Pacific and the adjacent Bering and Beaufort Seas. EBS belugas may have been able to modify their diet without compromising their growth and survivorship. Diet and reproductive analyses over large and small time scales can help understand the mechanisms enabling belugas to avoid significant growth and reproductive effects of past regime shifts. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Intraspecific variability in the “vowel”‐like sounds of beluga whales (Delphinapterus leucas): Intra‐ and interpopulation comparisons

The beluga whale (Delphinapterus leucas) has a rich and complicated vocal repertoire. However, different populations use similar and common types of signals. We studied physical features of one of these types, “vowels,” in three Russian populations: the White Sea population (European North), the Chukotka population (the Bering Sea, Chukotka), and the Okhotsk Sea population (Russian Far East) as well as in four summer aggregations of the White Sea belugas over several years in duration. The pulse repetition rate (PRR) at half of the duration of the signal was measured. We found that the PRR of “vowels” collected in the same summer aggregation during different years is stable in time but varies between locations. The degree of variation corresponds with the geographic distance between different locations. Significant differences were discovered between populations separated by thousands of kilometers, and to a lesser extent, between summer aggregations inhabiting different bays of the White Sea. The variation in PRR between the locations can be caused by the divergence of signals owing to the accumulation of random errors during transmission of these signals from generation to generation, which progressed independently in different summer aggregations and populations.     

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.     

Autotomy and regeneration of limbs in the male blue king crabParalithodes platypus from the Bering Sea and the Sea of Okhotsk

Limb loss and the frequency of regeneration was investigated in the male blue king crabParalithodes platypus collected in the Sea of Okhotsk and the Bering Sea. TheP. platypus population in the Bering Sea is characterized by an anomalously high proportion of crabs with missing limbs, which is largely due to the intensive fishing. The present approach to the exploitation of the Bering Sea population, in which large intact crabs are removed and injured crabs are left to recover to the normal state, may result in a decline in the reproductive potential of this population as early as this coming year.     

Migratory routes of red‐necked phalaropes Phalaropus lobatus breeding in southern Chukotka revealed by geolocators

The migration routes of red‐necked phalaropes breeding around the Bering Sea are poorly known, despite the fact that the Bering Sea could mark the boundary between the East Palearctic populations that winter in the Pacific Ocean around the East Indies and the West Nearctic populations that winter in the Pacific Ocean off the coast of South America. Geolocator data retrieved from two male phalaropes tagged in southern Chukotka, Far Eastern Russia, confirm that birds breeding in this region belong to the East Palearctic population and winter in the East Indies, suggesting that the division line with the West Nearctic population is farther to the east. The routes taken by the two phalaropes were almost entirely pelagic, totaling around 18 000–20 000 km round‐trip, with the birds continuously on the move during migration, rather than resident in any particular stopover site, contrary to most other migratory shorebirds.     

Jellyfish of the Far Eastern Seas of Russia. 3. Biomass and abundance

The biomass and abundance of large jellyfish (Cnidaria: Scyphozoa, Hydrozoa) was estimated and their seasonal and interannual dynamics was studied based on the data of trawl surveys conducted by the Pacific Research Fisheries Center (TINRO Center) in the Sea of Okhotsk, Bering Sea, Sea of Japan, and the Northwestern Pacific Ocean (NWPO) in 1991–2009. Most of the jellyfish biomass (over 95%) in the Sea of Okhotsk, Bering Sea, and NWPO was formed by Chrysaora spp., Cyanea capillata, Aequorea spp., Phacellophora camtschatica, and Aurelia limbata. The same species along with Calycopsis nematophora predominated in abundance in the Bering Sea and NWPO, while Ptychogena lactea, C. capillata, and Chrysaora spp. were most abundant in the Sea of Okhotsk. In the northwestern Sea of Japan, Aurelia aurita, C. capillata, and Aequorea spp. predominated both in abundance and biomass. Generally, the jellyfish abundance reached the highest values in the summer and fall and decreased abruptly in the winter. Meanwhile, the seasonal dynamics proved to be specific for each species and were manifested in some of them by reaching maximum values at various periods of the warm season, whereas the other (Tima sachalinensis and P. lactea) showed the reverse pattern of seasonal variations, with the highest abundance in cold months. Jellyfish biomass and abundance varied greatly from year to year, which was related to the short lifecycle and alternation between sexual and asexual generations, in which reproductive success was predetermined by various environmental factors. In the fall, year-to-year fluctuations of the relative biomass could increase by ten times. In 1991–2009, it varied from 200 to 2000 kg/km² in the northern Sea of Okhotsk, from 500 to 4200 kg/km² in the northwestern Bering Sea, and from 300 to 3700 kg/km² in the southwestern Bering Sea. Taking the jellyfish abundance estimates into account, along with the vertical distribution and the seasonal dynamics, the overall biomass of large species that occurred in trawl catches in Far Eastern seas and adjacent Pacific waters during the warm season could reach 13.0–15.0 million tons, of which up to about 6.0 million tons would be concentrated in the western Bering Sea and 5.5–6.0 million tons in the Sea of Okhotsk.     

Sex-related variability of rostrum morphometry of Aristeus antennatus (Decapoda: Aristeidae) from the Ionian Sea (Eastern Mediterranean, Greece)

Sex-related rostral variability was studied in the aristeid shrimp Aristeus antennatusfrom the Eastern Ionian Sea (Mediterranean). Shrimps were collected on a monthly basis from December 1996 to November 1997 using a commercial bottom trawl in a depth range of 446–728 m. Female relative growth of rostrum proved to be negative allometric both seasonally and in the pooled annual data set. Males on the other hand, showed no or negative correlation of rostrum length with size. Mature males with short rostra dominated in the male population all year around. The appearance of males with long and intermediate rostra during winter, which disappear thereafter in favour of those with short rostra, indicates that rostrum shortening takes place during the end of winter. The increase of mated females during spring supports the hypothesis already addressed by other authors on the function of the male short rostrum in this species mating behaviour. Nevertheless, the paucity, in comparison to other Mediterranean populations, of males with long or intermediate rostra could indicate that for the bulk of the male population, the process of rostrum shortening in the Eastern Ionian Sea occurs outside the geographical locality or depth range sampled.     

Temperature- and size-dependent growth and movement of the North Pacific giant octopus (Enteroctopus dofleini) in the Bering Sea

Octopus growth and movement occurs during all life stages and have implications for survival, food web dynamics and reproduction. From 2009 to 2011, 1714 North Pacific giant octopus (Enteroctopus dofleini) were tagged and recaptured in the eastern Bering Sea with visible implant elastomers to better understand the ecology of this data-poor species. Over this period, 246 of the tagged individuals were recaptured. In autumn, when temperatures were warmest, E. dofleini had higher growth rates and moved more than in the colder winter months. For both short- and long-term recaptures, small octopus grew faster than large octopus. Movement of octopus over short- and long-term periods was low for very small and very large octopus and high for median-sized octopus, which is likely to be a function of maturity status and reproductive activities. Approximately 80% of recaptures moved less than 2?km from the initial tagging location for time periods up to a year, suggesting long-term site fidelity. As temperature and size may be tightly linked to growth and movement rates for E. dofleini in the eastern Bering Sea, predicted climate warming will likely alter ecological processes for the species and impact their distribution.     

Population and size-specific distribution of Atlantic salmon Salmo salar in the Baltic Sea over five decades

Population-specific assessment and management of anadromous fish at sea requires detailed information about the distribution at sea over ontogeny for each population. However, despite a long history of mixed-stock sea fisheries on Atlantic salmon, Salmo salar, migration studies showing that some salmon populations feed in different regions of the Baltic Sea and variation in dynamics occurs among populations feeding in the Baltic Sea, such information is often lacking. Also, current assessment of Baltic salmon assumes equal distribution at sea and therefore equal responses to changes in off-shore sea fisheries. Here, we test for differences in distribution at sea among and within ten Atlantic salmon Salmo salar populations originating from ten river-specific hatcheries along the Swedish Baltic Sea coast, using individual data from >125,000 tagged salmon, recaptured over five decades. We show strong population and size-specific differences in distribution at sea, varying between year classes and between individuals within year classes. This suggests that Atlantic salmon in the Baltic Sea experience great variation in environmental conditions and exploitation rates over ontogeny depending on origin and that current assessment assumptions about equal exploitation rates in the offshore fisheries and a shared environment at sea are not valid. Thus, our results provide additional arguments and necessary information for implementing population-specific management of salmon, also when targeting life stages at sea.     

Hematology and serum chemistry values in the beluga (Delphinapterus leucas)

Normal values and ranges for 31 clinical hematology and serum chemistry tests are reported for the beluga or white whale (Delphinapterus leucas). The values were collected over a 6-yr period from eight belugas maintained for display at Sea World (San Diego, California, USA) facilities and represent long-term evaluations for each animal in a controlled environment. They represent the first report for a number of serum chemistry values for the beluga. Normal values such as these provide an important data base from which to detect diagnostically important changes in health status for belugas in a zoological setting. They also establish a baseline from which to evaluate differences in normal values in free-ranging belugas and from which to diagnose disease problems in wild populations.     

Post‐breeding movements of northeast Atlantic ivory gull Pagophila eburnea populations

The post‐breeding movements of three northeast Atlantic populations (north Greenland, Svalbard and Franz Josef Land) of the ivory gull Pagophila eburnea, a threatened high‐Arctic sea‐ice specialist, were studied between July and December 2007 using 31 satellite transmitters. After leaving their breeding grounds, all birds first dispersed eastward in August–September, to an area extending from the Fram Strait to the northwestern Laptev Sea (off Severnaya Zemlya). Most returned along the same flyway in October–November, hence describing a loop migration before moving south, off east Greenland. Wintering grounds were reached in December, in southeast Greenland and along the Labrador Sea ice‐edge, where Canadian birds also overwinter. One to two birds from each population however continued eastwards towards a third wintering area in the Bering Strait region, hence demonstrating a bi‐directional migration pattern for the populations and elucidating the origin of the birds found in the north Pacific during winter time. Overall, all birds breeding in the northeast Atlantic region used the same flyways, had similar rates of travel, and showed a peak in migratory activity in November. Though the total length of the main flyway, to the Labrador Sea, is only and at most 7500 km on a straight line, the mean total distance travelled by Greenland birds between July and December was 50 000 km when estimated from hourly rates of travel. Our study presents the first comprehensive and complete picture for the post‐breeding movements of the different ivory gull populations breeding in the northeast Atlantic.     

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.     

Does winter region affect spring arrival time and body mass of king eiders in northern Alaska?

Events during the non-breeding season may affect the body condition of migratory birds and influence performance during the following breeding season. Migratory birds nesting in the Arctic often rely on endogenous nutrients for reproductive efforts, and are thus potentially subject to such carry-over effects. We tested whether king eider (Somateria spectabilis) arrival time and body mass upon arrival at breeding grounds in northern Alaska were affected by their choice of a winter region in the Bering Sea. We captured birds shortly after arrival on breeding grounds in early June 2002–2006 at two sites in northern Alaska and determined the region in which individuals wintered using satellite telemetry or stable isotope ratios of head feathers. We used generalized linear models to assess whether winter region explained variation in arrival body mass among individuals by accounting for sex, site, annual variation, and the date a bird was captured. We found no support for our hypothesis that either arrival time or arrival body mass of king eiders differed among winter regions. We conclude that wintering in different regions in the Bering Sea is unlikely to have reproductive consequences for king eiders in our study areas.     

Molecular identification of Ammodytes (PISCES, Ammodytidae) larvae, with ontogenetic evidence on separating populations

A total of ten larval Ammodytes were collected from the Yellow (5 specimens) and Bering Seas (5), and a 638 base pair sequence from their mitochondrial DNA COI genes was analyzed for species identification. Sequences were compared with those of adult Ammodytes personatus from the East (4), the Yellow (2) and the East China Seas (2), and Japan (3). Average genetic distance was 0.064 between the Yellow Sea larvae and the Bering Sea larvae, but was 0.0043 between the Yellow Sea larvae and all the adult A. personatus except two individuals from the East Sea. NJ-tree showed that five Yellow Sea larvae were closely clustered with adult A. personatus, except for two individuals. The five Bering Sea larvae were located distantly from adult A. personatus, and are thought to be A. hexapterus. We found morphological differences among two populations of A. personatus larvae and A. hexapterus larvae in morphometric characters and pigmentation. Our results indicated that the East Sea population of A. personatus is closer to A. hexapterus than to the Yellow Sea population of A. personatus, suggesting the East Sea population may be part of separate species.     

Scales and otoliths as identity cards of the Indian oil sardine Sardinella longiceps (Teleostei: Clupeiformes) populations: Ultrastructure and ornamentation characteristics using light and scanning electron microscopy

Scale and otolith morphology and morphometry of Indian oil sardine Sardinella longiceps (Clupeidae) were investigated and described using light and scanning electron microscopy from eight different body regions for scales and the right and left otoliths. Scales of the Indian oil sardine show general characteristics of the other studied clupeids sand that are easily distinguishable from other fish groups, by having striae in the posterior field. The studied cycloid scales of S. longiceps were classified into three types based on the overall shape including circular (e.g. true circular and cordate), pentagonal and quadrilateral in the different body regions. The circular shape was the most common shape (87.5%), while the quadrilateral and pentagonal forms constituted 6.25% each. The results also showed that the relative scale size (J-index) plays a desirable contribution in separating the examined populations. The results showed that the mean (or relative) scale size for all the eight regions in the Oman Sea population is larger than the Arabian Sea population. Also, another scale variable, the scale shape index (Si index), demonstrated variation (a mean of 0.86 to 1.1) in different regions of both populations from the Oman and Arabian Seas. Interestingly, here, we found that scale characters of S. longiceps not only differ from its other congeneric species, but also differ in the populations from both sides of the Oman Sea (Iran and Oman) and the Arabian Sea. It shows a positive signal for the presence of different taxonomic and management unit in the Oman and Arabian Seas. The idea should be approved by using integrated molecular and morphological traits. The otolith morphology of S. longiceps from the Oman and Arabian Seas was more conservative than the scales, which can be due to its function actin primarily as a balance organ and also enhancing hearing. The overall shape of S. longiceps otolith was lanceolate, with an elongated morphology and a well-developed rostrum, an ostial sulcus acusticus that opens to the anterior/ dorsal margin. These morphological characters are also found in the Iranian population of S. longiceps. However, otolith displayed variation in biometric parameters among two populations and left and right otoliths and the RRL parameter were important characters to discriminate the Oman and Arabian Sea populations. Thus, the structural/biometrical variability of the otoliths may be used for population distinctness, especially in water bodies with various environmental factors, and the otolith has turned out to be a useful tool to track the life history of teleostean fishes in environments with physicochemical gradients.     

Population genetic structure of walleye pollock <Emphasis Type="Italic">Theragra chalcogramma</Emphasis> (Gadidae,Pisces) from the Bering Sea and Sea of Okhotsk

Walleye pollock Theragra chalcogramma Pallas occupies a central place in ecosystems of the North Pacific and is an important target species of fisheries. The species is characterized by daily vertical, spawning, feeding, and wintering migrations and spawning occurring under the sea ice. Since population structure estimation by the tagging with recapture is inefficient in walleye pollock, the pollock resources are difficult to estimate by conventional methods, requiring population genetic studies with molecular markers. The population genetic structure of five spawning aggregations from the Bering Sea was for the first time studied with ten microsatellite loci: Tch5, Tch10, Tch11, Tch12, Tch14, Tch16, Tch17, Tch19, Tch20, and Tch22. A spatially distant sample from the Sea of Okhotsk was used as a reference group. Polymorphism for the markers reached 100%, and heterozygosity of individual loci ranged from 41 to 95% in different populations. It was shown the aggregations of interest are in goodness-to-fit the Hardy-Weinberg equilibrium (HWE) at hole, while the Sea of Okhotsk sample demonstrated a sex bias: the heterozygosity at Tch16 in males was significantly lower than in females. The highest discriminative power was observed for Tch10, Tch20, and Tch22. F _ST genetic distances between populations were typical for marine fishes. A mixed composition was supposed for the sample from the region of the underwater Shirshov Ridge, which serves as a natural partial geographic barrier between the Olyutor-Karagin and Koryak walleye pollock stocks. With the Shirshov sample excluded, F _ST scatter plots and the spatial autocorrelation approach supported isolation by distance for the aggregations. An influence of abiotic factors on the population structure was assumed for walleye pollock of the Bering Sea.     

Confirmation of a wintering ground of Ross's Gull Rhodostethia rosea in the northern Labrador Sea

Ross's Gull Rhodostethia rosea is one of the world's least known seabirds; < 1% of the estimated global population can be accounted for at known breeding sites, and its wintering range has never been determined. Anecdotal reports over the last two centuries have prompted extensive speculation as to possible wintering areas used by this species in the north Pacific/Bering Sea region, but none has ever been confirmed. Using satellite and geolocator telemetry, we show that some Ross's Gulls from a colony in the Canadian Arctic winter in a restricted area of the northern Labrador Sea. Our discovery of a wintering area in the northwest Atlantic indicates that Ross's Gulls breeding in the Nearctic may be part of a disjunct population, or that birds breeding in the Palaearctic may winter off the east coast of North America.     

Genetic structuring among Alaskan Pacific herring populations identified using microsatellite variation

Five highly variable microsatellite loci were used to investigate population structuring in Pacific herring Clupea pallasi collected from Kodiak Island, two sites in the Bering Sea and four sites within Prince William Sound, Alaska. All loci revealed high levels of variability with heterozygosity estimates ranging from 86 to 97% (mean heterozygosity: 89%). The variation was structured significantly among sites suggesting that the samples investigated were genetically distinct from each other. Genetic divergence was greatest between populations from the Bering Sea and those from Prince William Sound. The Kodiak Island and Point Chalmers samples appeared to be distinct from the Prince William Sound and Bering Sea populations. The observed genetic distance relationships among samples could be explained largely in terms of geographical separation.