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.     

Winter distribution and migrations of beluga whales (<Emphasis Type="Italic">Delphinapterus leucas</Emphasis>) in the White Sea based on satellite tracking data

Based on satellite tracking of eight beluga whale males in the White Sea, their habitats in the autumn, winter, and spring periods have been identified. A correlation between the distribution of beluga whales, ice dynamics, and migration of Atlantic salmon has been revealed. It has been found than beluga whale males do not leave the White Sea during the entire ice period. The results obtained confirm the hypothesis that the White Sea population of beluga whales is isolated.     

Satellite telemetry reveals population specific winter ranges of beluga whales in the Bering Sea

At least five populations (stocks) of beluga whales (Delphinapterus leucas) are thought to winter in the Being Sea, including the Bristol Bay, Eastern Bering Sea (Norton Sound), Anadyr, Eastern Chukchi Sea, and Eastern Beaufort Sea (Mackenzie) populations. Belugas from each population have been tagged with satellite‐linked transmitters, allowing us to describe their winter (January–March) distribution. The objectives of this paper were to determine: (1) If each population winters in the Bering Sea, and if so, where? (2) Do populations return to the same area each year (i.e., are wintering areas traditional)? (3) To what extent do the winter ranges of different populations overlap? Tagged belugas from all five populations either remained in, or moved into, the Bering Sea and spent the winter there. Each population wintered in a different part of the Bering Sea and populations with multiple years of data (four of five) returned to the same regions in multiple years. When data were available from two populations that overlapped in the same year, they did not occupy the shared area at the same time. Although our sample sizes were small, the evidence suggests belugas from different populations have traditional winter ranges that are mostly exclusive to each population.     

Distribution of beluga whales (Delphinapterus leucas) in the Russian Arctic seas according to the results of expedition aboard RV Mikhail Somov, September–November 2010

Data on the distribution of marine mammals, including beluga whales (Delphinapterus leucas Pallas, 1766), in the Arctic are scarce because of various causes and conditions, including the vast expanses of the region, its poor accessibility, severe climate, long polar night, and high cost of research. Nevertheless, the results of aerial observations during ice reconnaissance and onboard observations during sea voyages (Kleinenberg et al., 1964; Geptner et al., 1976; Belikov, Boltunov, and Gorbunov, 2002; Belikov and Boltunov, 2002; Ezhov, 2005; Matishov and Ognetov, 2006; Biologiya i okeanografiya??, 2007; Lukin and Ognetov, 2009) have provided a general idea of the distribution pattern of beluga whales in the Russian Arctic seas. More detailed data concern the distribution of these whales in the White Sea, where aerial surveys of the water area were performed previously and have been resumed in recent years (Nazarenko et al., 2008; Glazov et al., 2010, 2011). The relevant data on the Barents, Kara, Laptev, and East Siberian seas are much poorer. In the summer (ice-free) period, beluga whales concentrate in coastal waters. They have been recorded most frequently off Franz Josef Land, Novaya Zemlya, Vaygach Island, and in Czech Bay in the Barents Sea; in Baydaratskaya Bay, Gulf of Ob, and Yenisei Gulf in the Kara Sea; off the northeastern coast of Taimyr and in estuaries of the Anabar, Olenyok, and Lena rivers in the Laptev Sea; and in the estuaries of the Indigirka (where the whales come from the west) and the Kolyma and Ked??ma rivers (where they come from the east) in the East Siberian Sea. The amount of information obtained in other seasons is very limited. In autumn, mass migration of beluga whales from the Kara Sea to the Barents Sea have been recorded in the Karskie Vorota Strait and off Cape Zhelaniya in the north of Novaya Zemlya. In winter, almost no records of these whales have been made in the Kara, Laptev, and East Siberian seas. These data are based on previous observations and have practically not been complemented in recent years.     

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     

A genetic analysis of the beluga whale Delphinapterus leucas (Cetacea: Monodontidae) from summer aggregations in the Russian Far East

The genetic structure of four summer aggregations of the Beluga Whale, Delphinapterus leucas, in Sakhalin Bay and Udskaya Bay, off the western coast of Kamchatka in the Sea of Okhotsk and in the Anadyr Estuary of the Bering Sea was analyzed through nucleotide sequencing of the mtDNA control region and detection of the allelic composition of nine microsatellite loci in nuclear DNA. It has been shown that each of the aggregations features a unique set of maternal lines, which indicates a high degree of philopatry in this species. Beluga whales of the Anadyr Estuary are genetically isolated from those of the Sea of Okhotsk. Beluga whales of the summer aggregations of Sakhalin Bay and those from Udskaya Bay share a common gene pool and belong to a single population, while the whales that summer off western Kamchatka with great consistency may be attributed to a different population. Comparison of nucleotide sequences of the mtDNA in beluga whales from various waters of the Russian Far East and North America allowed us to propose a hypothesis about how the structure of beluga whale populations formed in the North Pacific during the postglacial period.     

Fall movements of Red‐headed Woodpeckers in South Carolina

Fall migration of Red‐headed Woodpeckers (Melanerpes erythrocephalus) can be erratic, with departure rates, directions, and distances varying among populations and individuals. We report fall migration departure dates, rates, and routes, and the size of fall home ranges of 62 radio‐tagged Red‐headed Woodpeckers in western South Carolina. Rates of fall migration differed among years; all radio‐tagged woodpeckers migrated in 2005 (15 of 15), none (0 of 23) migrated in 2006, and 54.2% (13 of 24) migrated in 2007. Of 28 woodpeckers that left their breeding territories, we relocated eight either en route or on their fall home ranges. These woodpeckers migrated short distances (4.3–22.2 km) south along the floodplain forest of a large creek. The variable migration patterns we observed indicate that Red‐headed Woodpeckers may best be described as facultative migrants. We determined the home range sizes of 13 woodpeckers in both seasons, regardless of whether they migrated, and fall home ranges were smaller (mean = 1.12 ha) than summer home ranges (mean = 3.23 ha). Fall‐winter movements of Red‐headed Woodpeckers were concentrated on mast‐producing oak (Quercus spp.) trees, which may have restricted home range sizes. The partial migration we observed in 2007 suggests that factors other than mast crop variability may also influence migration patterns because woodpeckers that year responded to the same annual mast crop in different ways, with some migrating and some remaining on breeding season home ranges during the fall.     

Home range of the raccoon dog (Nyctereutes procyonoides) in southern Finland

Home ranges, relationships between individuals and dispersal among raccoon dogs ( Nyctereutes procyonoides ) were studied in southern Finland in 1989-91. The average maximum home range, calculated by the harmonic mean method. was 9·5 km² and the core area (85% utilization) 3·4 km². There were no statistically significant annual. seasonal or sexual differences in the size of the average core area of adults, but the home ranges of juveniles in autumn were larger than those of adults. However. the maximum home ranges of adults were larger in autumn than in summer, especially those of males, which were conspicuously small in summer and large in autumn. The core areas of adjacent pairs did not usually overlap in the pup-rearing season. but in autumn some home ranges shifted and there was much more overlap. The home ranges of the male and female of a pair overlapped almost totally, and a male and a female sharing the same home range also travelled together or close to each other, thus supporting the idea that the raccoon dog is monogamous in Finland. None of the adults left the study area, but 38% of the juveniles were recovered further than 10 km from the marking place during the first autumn.     

长江口及其毗邻海域鱼类群落优势种的生态位宽度与重叠

根据“北斗号”调查船2006年6月、8月和10月在长江口及其毗邻海域进行渔业资源底拖网调查的资料,通过相对重要性指数确定鱼类群落优势种,对优势种的生态位测度指标及季节变化进行分析.结果表明： 3航次的优势种共有10种,它们的生态位宽度和生态位重叠指标呈现出明显的季节性变化,通过Bray-Curtis相似性聚类与非度量多维标度(MDS)分析可分为两组.第一组为鳀鱼、鳄齿鱼、发光鲷,其生态位宽度和生态位重叠指标夏季大于秋季,呈现出由夏季到秋季从长江口及其毗邻海域向外部深水区移动的趋势;第二组为带鱼、矛尾虾虎鱼、细条天竺鲷、小黄鱼、刺鲳、白姑鱼、七星底灯鱼,其生态位宽度和生态位重叠指标秋季大于夏季,呈现出由夏季到秋季从黄海南部和东海北部水域向长江口及其毗邻海域移动的趋势.这两组鱼类不同的移动趋势与其生态习性和环境因素有关.     

Increased blubber cortisol in ice-entrapped beluga whales (<Emphasis Type="Italic">Delphinapterus</Emphasis><Emphasis Type="Italic">leucas</Emphasis>)

Entrapments of whales in sea ice occur occasionally in the Arctic and often last several weeks or months, resulting in emaciation or death of whales. These events provide a unique opportunity for investigating the physiological response to a prolonged or chronic stress in an otherwise healthy population of marine mammals. By measuring cortisol in blubber, a peripheral tissue, we expect to see a reflection of long-term or chronic stress rather than short-term or acute stress. Adipose tissue should be less subject to rapid changes compared to blood cortisol, reflecting stressors experienced over a longer period of time, and should not be affected by potential stress associated with sampling. We measured blubber cortisol of 29 beluga whales (Delphinapterus leucas) entrapped in November 2006 in Husky Lakes basin and 26 whales from the same population (Eastern Beaufort Sea) during regular seasonal harvests in July of 2006 and 2007. Mean cortisol concentrations (±SEM) were seven times higher in blubber from entrapped whales (1.76 ± 0.32 ng/g wet weight) compared to whales from regular seasonal harvests (0.26 ± 0.042 ng/g wet weight) and appeared to increase with whale age. Our results provide a measure of blubber cortisol from a prolonged stress and demonstrate blubber cortisol as a useful indicator of longer-term exposure to stress in beluga whales.     

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.     

Parental behavior of the beluga whale (Delphinapterus leucas) in natural environment

The parental behavior of beluga whales observed in a summer reproductive gathering (near Solovetsky Island, Onega Bay, White Sea) is described, including the parturition process. A tight association between the female and its newborn infant has been traced by the case study of a stillborn calf. Individual behavioral patterns of a mother and calves observable only in natural habitats are detailed.     

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.     

Home range and activity patterns of male red deer (Cervus elaphus L.) in the alps

Summary From 1978 to 1981 in the Bavarian Alps (Southern West Germany) the home range and activity patterns of nine male red deer have been studied using radio-telemetry. The home range patterns definetely change with age. Younger stags first follow the patterns of their mothers, then often emigrate from these home ranges and establish new ones elsewhere. Except for the change in range at about 2^1/2 years of age, these patterns seem to be very constant in both spatial as well as seasonal position and the size of the home ranges. Winter and rutting ranges are relatively small, averaging 113 ha and 134 ha, respectively, whereas the mean size of the home ranges used from spring to autumn amounts to some 386 ha. Just as do the home range patterns, so also do activity patterns exhibit a marked annual cycle. The daily sum total of activity varies from about 9 h in winter to some 15 h in summer. The daily distribution of activity reveals a typical bimodal 24-h rhythm which in the course of the year also shows modifications according to the seasonally varying LD-ratio. In the discussion, earlier results on female red deer are compared to those of this study. Notable differences between sexes occur in the home range patterns and the annual cycle of daily activity.     

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.     

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.     

Identifying gray whale (Eschrichtius robustus) foraging grounds along the Chukotka Peninsula, Russia, using satellite telemetry

The purpose of this study was to evaluate summer and fall residency and habitat selection by gray whales, Eschrichtius robustus, together with the biomass of benthic amphipod prey on the coastal feeding grounds along the Chukotka Peninsula. Thirteen gray whales were instrumented with satellite transmitters in September 2006 near the Chukotka Peninsula, Russia. Nine transmitters provided positions from whales for up to 81?days. The whales travelled within 5?km of the Chukotka coast for most of the period they were tracked with only occasional movements offshore. The average daily travel speeds were 23?km?day^?1 (range 9–53?km?day^?1). Four of the whales had daily average travel speeds <1?km?day^?1 suggesting strong fidelity to the study area. The area containing 95% of the locations for individual whales during biweekly periods was on average 13,027?km² (range 7,097–15,896?km²). More than 65% of all locations were in water <30?m, and between 45 and 70% of biweekly kernel home ranges were located in depths between 31 and 50?m. Benthic density of amphipods within the Bering Strait at depths <50?m was on average ~54?g wet wt m^?2 in 2006. It is likely that the abundant benthic biomass is more than sufficient forage to support the current gray whale population. The use of satellite telemetry in this study quantifies space use and movement patterns of gray whales along the Chukotka coast and identifies key feeding areas.     

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.     

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.     

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.