Seasonal distribution and dive behaviour of harp seals (<Emphasis Type="Italic">Pagophilus groenlandicus</Emphasis>) of the White Sea–Barents Sea stock

Eight adult female harp seals (Pagophilus groenlandicus) of the White Sea–Barents Sea stock were tagged with satellite-linked dive recorders during the nursing period and followed from breeding in late February 1995 until moulting in late April 1995. Another ten adult harp seals of both sexes were tagged and followed from moult in early May 1996 until breeding in late February the following year. Between breeding and moult the seals were distributed along the coasts of Kola of Russia and eastern Finnmark of Norway, coinciding in time and space with the spawning capelin (Mallotus villosus). Between moulting and breeding they encircled the entire Barents Sea, mostly in open water, using the water column from 20 to 300 m, and in so doing by and large reflecting the annual migrations of the capelin. Capelin is therefore assumed to be the main source of prey for the White Sea–Barents Sea stock of harp seals, to be substituted, in part, by amphipods (e.g. Themisto libellula) in mid-summer and polar cod (Boreogadus saida) and herring (Clupea pallasii) in late autumn and winter. These data provide a baseline for the evaluation of the effects of future climatic change in the rich Barents Sea ecosystem.     

Remarkable development of diving performance and migrations of hooded seals (Cystophora cristata) during their first year of life

Newborn hooded seals (Cystophora cristata) have smaller weight-specific oxygen stores than adults, but nothing is known about how this affects their diving behaviour. Here, we present data on the diving behaviour and migrations of seven weaned hooded seal pups of the Greenland Sea stock during their first year of life, as collected by use of satellite telemetry. The pups started diving 1–2 days after tagging, and during a tracking period of 25–398 days they dispersed over vast areas of the Greenland and Norwegian Seas in a manner similar to adults. The initial development of diving depths and durations in April–May was rapid, and pups reached depths of >100 m and dived for >15 min within 3 weeks of age. During early summer (May–June) this development was temporarily discontinued, to be resumed throughout autumn and winter, during which time maximum depths and durations of >700 m and >30 min, respectively, were reached. Depths and durations were significantly related to age/season, location and time of day. The dive behaviour in early summer, with relatively shallow and short dives without diurnal variations, resembles that of adults and probably reflects the vertical distribution of prey rather than physiological constraints. Dives of pups were nevertheless shallower and shorter than those of adults, but relative to body mass both hooded seal pups and adults display a remarkable diving capacity which makes the species particularly suited for studies of defence mechanisms against hypoxia insult in mammals.     

Harp seal foraging behaviour during summer around Svalbard in the northern Barents Sea: diet composition and the selection of prey

The harp seal Pagophilus groenlandicus is a major high trophic level predator in the Barents Sea, and to better understand their function in the Barents Sea ecosystem, we need to understand their foraging behaviour during their most intensive feeding period. We analysed the diet composition and prey preference of 184 harp seals and 94 faeces samples, sampled in the northern Barents Sea (around Svalbard) during the period May–August in 1996, 1997, and 2004–2006. Concurrent with the sampling of seals, prey availability was assessed in one area in 1996 and 1997 and in two areas in 2006 using standard acoustic methods. Our study showed that harp seal diet composition varied significantly both in time (year) and space, and that their diets appeared to be size dependent. Both subadult (<150 cm) and adult seals were associated with pelagic crustaceans (particularly krill), whereas primarily adult seals were associated with fish (capelin, gadoids and flatfish). Krill was the most important prey group (63 %) followed by polar cod (16 %) and other fish species (10 %). The prey preference of harp seals varied in time and space; polar cod was often preferred by the seals whereas krill was commonly consumed in lower proportion than observed in the survey area. Gadoids and capelin had either been exploited in the same or less proportion as observed in the survey sea. This study emphasises the ecological significance of krill as prime food for harp seals during their intensive feeding period in summer.     

Multi-decadal trends in biomarkers in harp seal teeth from the North Atlantic reveal the influence of prey availability on seal trophic position

Arctic food webs are being impacted by borealisation and environmental change. To quantify the impact of these multiple forcings, it is crucial to accurately determine the temporal change in key ecosystem metrics, such as trophic position of top predators. Here, we measured stable nitrogen isotopes (δ¹⁵N) in amino acids in harp seal teeth from across the North Atlantic spanning a period of 60 years to robustly assess multi-decadal trends in harp seal trophic position, accounting for changes in δ¹⁵N at the base of the food web. We reveal long-term variations in trophic position of harp seals which are likely to reflect fluctuations in prey availability, specifically fish- or invertebrate-dominated diets. We show that the temporal trends in harp seal trophic position differ between the Northwest Atlantic, Greenland Sea and Barents Sea, suggesting divergent changes in each local ecosystem. Our results provide invaluable data for population dynamic and ecotoxicology studies.     

DIET OF WEANED PUPS AND SEASONAL VARIATIONS IN BODY CONDITION OF JUVENILE BARENTS SEA HARP SEALS PHOCA GROENLANDICA

Data on blubber depth, body condition, and diet were collected from young of year and juvenile harp seals ( Phoca groenlandica ) in the Barents Sea in the periods February-April and June-October 1990–1997. Harp seal pups feed independently shortly after weaning. In the southern parts of the Barents Sea the diet of harp seal pups consisted mainly of euphausiids ("krill") Thysanoessa sp. and amphipods of the genus Parathemisto . The change in body condition of harp seal pups between weaning (mid-March) and June indicated that considerable amounts of blubber energy, deposited during suckling, were mobilized. This suggests that the early food intake of weaned pups was insufficient to meet the energy requirements. The pattern of low spring and earlysummer body condition, followed by a subsequent rapid improvement in condition during late summer and autumn appeared to occur not only among pups-of-the-year, but also in one- and two-year old harp seals.     

Movements and dive behaviour of two leopard seals (<Emphasis Type="Italic">Hydrurga leptonyx</Emphasis>) off Queen Maud Land,Antarctica

Two adult female leopard seals (Hydrurga leptonyx) were tagged with satellite-linked dive recorders off Queen Maud Land, Antarctica, just after moulting in mid-February. The transmitters transmitted for 80 and 220 days, respectively. Both seals remained within the pack ice relatively close to the Antarctic Continent until early May, when contact was lost with one seal. The one remaining seal then migrated north, to the east side of the South Sandwich Islands in 3 weeks, whereafter it headed east, until contact was lost at 55°S in early September. From mid-May to late September this animal always stayed close to the edge of the pack ice. Both seals made mostly short (<5 min) dives to depths of 10–50 m and only occasionally dove deeper than 200 m, the deepest dive recorded being 304 m. A nocturnal diving pattern was evident in autumn and early winter, while day-time diving prevailed in mid-winter. Haul out probability was highest at mid-day (about 40% in late February and more than 80% in March and April). From May till September the remaining animal mainly stayed at sea, in the vicinity of the pack ice, with only occasional haul outs. These data suggest that a portion of the adult leopard seals may spend the winter mainly in open water, off the edge of the pack ice, where they primarily hunt near the surface. In that case, it is likely that krill (Euphausia superba), as well as penguins, young crabeater seals (Lobodon carcinophaga) and a variety of fish are important prey items.     

Trophic level and fatty acids in harp seals compared with common minke whales in the Barents Sea

The objectives of this study were to explore trophic levels and possible diet overlap between harp seals (Pagophilus groenlandicus) and common minke whales (Balaenoptera acutoroostrata) in the Barents Sea using stable isotopes of nitrogen (δ¹⁵N) and carbon (δ¹³C) and fatty acid analyses, and to explore the energy pathways from the plankton to the top predators. Blubber and muscle samples from 93 harp seals and 20 minke whales were collected in the southern Barents Sea in May 2011. The study showed that harp seals were at a higher trophic level than minke whales during spring. This supported previous diet studies suggesting a more fish-dominant diet for seals, as compared with the whales, at this time of the year. The stable isotopes and fatty acids indicated niche separation between the seals and the whales, and between different age groups of the harp seals. Older seals had fatty acid profiles more equal to minke whales as compared with younger seals. Furthermore, while the fatty acid profiles suggested that krill were of particular importance for the young seals, the profiles from older seals and whales suggested that fish dominated their diets.     

Movements and diving of bearded seal (Erignathus barbatus) mothers and pups during lactation and post-weaning

Eleven bearded seals (Erignathus barbatus) were tagged with satellite-linked dive recorders in Kongsfjorden, Svalbard, Norway, in May 1994. These animals included four mother-pup pairs and three single pups. The seals were tracked for 21–258 days. A total of ˜207,000 dives were recorded. Bearded seal mothers showed limited movements during the nursing and moulting periods. After weaning, the pups moved out of the tagging area and dispersed coastally. One pup left Svalbard and moved far offshore to Greenland and Jan Mayen. Bearded seal adults displayed a bi-modal dive behaviour, with peaks of activity that were shallower than 10 m or from 50 to 70 m. Most dives for adult seals (97%) were shorter than 10 min. Young pups performed dives that were shallower and shorter in duration than their accompanying mothers, but diving skills improved rapidly with age. Six of the seven pups dived deeper than 448 m by the time they were 2 months old. Analyses of movement data with respect to separation of mother-pup pairs suggest a lactation period of about 24 days. Accepted: 31 January 2000     

Distribution and diving behaviour of crabeater seals (Lobodon carcinophagus) off Queen Maud Land

Eight crabeater seals (Lobodon carcinophagus) (three females, five males), ranging in body mass between 125 and 220 kg, were captured off Queen Maud Land (70–72°S, 7–16°W) during the last week of February, just after moulting, and tagged with Argos satellite-linked dive recorders to provide data on location and diving depth and duration. During the first few weeks of March the seals were moving in the pack ice along the continental shelf edge, close to the coast of Queen Maud Land. In April and May, when the pack ice extended northwards, most of the seals moved north, one reaching 63°S in late May. In the first half of June the two remaining seals turned south and moved back deep into the pack ice. The seals made about 150 dives per day each throughout the study period. Ninety percent of these were made to depths of less than 52 m. Individual maximum diving depths varied between 288 and 528 m. In March the seals were most active at night, when the dive depth was shallower than during the day. In April and May the seals were more active during day-time, with an absence of any diurnal change in divng depth. These results support the notion that crabeater seals predominantly feed on krill in Antarctic pack ice, even when winter returns to the waters off Queen Maud Land.Publication no. 134 of the Norwegian Antarctic Research Expedtion 1992/1993     

Gut length, food transit time and diving habit in phocid seals

The southern elephant seal (Mirounga leonina) has the ability to dive for 2 h and reach depths of 1200 m. This creature is also exceptional in having a small intestine that is 25 times body length. Krockenberger and Bryden advanced the hypothesis that the long small intestine has developed to compensate for the extended periods with reduced or even abolished intestinal blood perfusion during diving. To test this hypothesis we have measured small-intestinal lengths in crabeater (Lobodon carcinophagus), Weddell (Leptonychotes weddellii), Ross (Ommatophoca rossi), leopard (Hydrurga leptonyx), harp (Phoca groenlandica), ringed (Phoca hispida) and hooded (Cystophora cristata) seals and related them to available data on their maximal dive duration. We found no significant correlation (P > 0.05) between intestinal length relative to body length and diving ability, but we found that small-intestinal internal area was significantly (P < 0.05) related to body length. A crude scanning electron microscopical examination of the small intestines of Weddell, crabeater, hooded and harp seals failed to reveal any gross anatomical differences between small-intestinal surfaces. This suggests that gut dimension in this variety of phocid species with widely differing diving ability is not related to diving habit, but is instead related to body size. The transit time of digesta was determined in two 1-year-old harp seals by use of radiopaque polyethylene rings of 4-mm diameter followed by X-ray examination, as markers for the solid phase passage, and chromium ethylene-diaminetetra acetic acid (Cr-EDTA) as a marker for the liquid phase. The transit time for the Cr-EDTA marker was 6.9 h ± 0.5 SE (range 4.5–8 h, n= 7), while 80% of the polyethylene markers appeared in the colon after 17.6 h ± 1.0 SE (range 14–21.5 h, n= 6) and were sometimes retained in the colon for several hours before defecation. These transit times did not change significantly (P > 0.05) in response to repetitive diving over a period of 8 h. This indicates that the often-used Cr-EDTA is not a good measure for digesta passage time when used alone in seals, and that the hypothesis of Krockenberger and Bryden is most likely wrong. Received: 17 December 1997 / Accepted: 4 May 1998     

Feeding habits of harp seals (Phoca groenlandica) during early summer and autumn in the northern Barents Sea

The feeding habits of harp seals (Phoca groenlandica) in the Barents Sea were examined in studies conducted during June 1991, September 1990 and 1991, and October 1992. Analyses of stomach and intestinal contents were carried out and concurrent estimates of prey abundance were made using trawl gear. Harp seals appeared to feed at low intensity in the pack ice belt during the first half of June. There was little potential prey in the water column, but prawns (Pandalus borealis), capelin (Mallotus villosus) and polar cod (Boreogadus saida) were abundant close to the bottom. In September, the seals sampled in the northern pack ice areas of the Barents Sea fed on the pelagic amphipod Parathemisto libellula, krill (Thysanoessa spp.), prawns and, to a lesser extent, on fish species such as polar cod, sculpins (Cottidae) and snailfish (Liparidae). Trawling revealed that large quantities of Parathemisto libellala were present in the upper layers of the water column. Fish, mainly capelin and polar cod, were less abundant and occurred in deeper waters. In mid-October, the diet of seals in the northern Barents Sea consisted mainly of amphipods (Parathemisto sp.). Later in October, when increasing pack ice cover forced the harp seals to move south, the diet seemed to change from amphipods to fish prey, predominantly capelin and polar cod.     

Global change effects on the long‐term feeding ecology and contaminant exposures of East Greenland polar bears

Rapid climate changes are occurring in the Arctic, with substantial repercussions for arctic ecosystems. It is challenging to assess ecosystem changes in remote polar environments, but one successful approach has entailed monitoring the diets of upper trophic level consumers. Quantitative fatty acid signature analysis (QFASA) and fatty acid carbon isotope (δ¹³C‐FA) patterns were used to assess diets of East Greenland (EG) polar bears (Ursus maritimus) (n = 310) over the past three decades. QFASA‐generated diet estimates indicated that, on average, EG bears mainly consumed arctic ringed seals (47.5 ± 2.1%), migratory subarctic harp (30.6 ± 1.5%) and hooded (16.7 ± 1.3%) seals and rarely, if ever, consumed bearded seals, narwhals or walruses. Ringed seal consumption declined by 14%/decade over 28 years (90.1 ± 2.5% in 1984 to 33.9 ± 11.1% in 2011). Hooded seal consumption increased by 9.5%/decade (0.0 ± 0.0% in 1984 to 25.9 ± 9.1% in 2011). This increase may include harp seal, since hooded and harp seal FA signatures were not as well differentiated relative to other prey species. Declining δ¹³C‐FA ratios supported shifts from more nearshore/benthic/ice‐associated prey to more offshore/pelagic/open‐water‐associated prey, consistent with diet estimates. Increased hooded seal and decreased ringed seal consumption occurred during years when the North Atlantic Oscillation (NAO) was lower. Thus, periods with warmer temperatures and less sea ice were associated with more subarctic and less arctic seal species consumption. These changes in the relative abundance, accessibility, or distribution of arctic and subarctic marine mammals may have health consequences for EG polar bears. For example, the diet change resulted in consistently slower temporal declines in adipose levels of legacy persistent organic pollutants, as the subarctic seals have higher contaminant burdens than arctic seals. Overall, considerable changes are occurring in the EG marine ecosystem, with consequences for contaminant dynamics.     

ESTIMATING STRUCK AND LOSS RATES FOR HARP SEALS (PAGOPHILUS GROENLANDICUS) IN THE NORTHWEST ATLANTIC

One of the major sources of unreported mortality during the commercial harp seal ( Pagophilus groenlandicus ) hunt in the Northwest Atlantic is the number of animals that are killed but not recovered or reported, commonly referred to as struck and lost. With the significant expansion of the hunt in Canada and in Greenland during the mid-1990s, there is a concern that the number of seals lost may have reached a level that is not sustainable relative to current harvest levels. To address this problem a study examining struck and loss rates was initiated in 1998 and 1999. Loss rates for young seals taken on the ice varied from 0% to 1.9% and from 0% to 10.0% when taken in the water. Seals greater than one year of age had loss rates of 0%–4.9% when taken on the ice and 13.8%-50.0% when taken in the water. These estimates are lower than those recently reported for Northwest Atlantic harp seals.     

Movements and diving behavior of weaned Weddell seal (Leptonychotes weddellii ) pups

Between 1993 and 1995, the diving behavior and movement patterns of 23 weaned Weddell seal pups (Leptonychotes weddellii) were tracked in the Ross Sea. Antarctica, using satellite-linked time-depth recorders. Regression analyses revealed that for seals of between 8 and 27 weeks old, age was poorly correlated with the dive depth, duration, or frequency. However, changes in dive parameters suggested that Weddell seal pups were attempting to maximize dive time, but the manner in which this was done depended on age and time of day. Movement patterns indicated that most Weddell seal pups left their natal area by the end of February, and traveled north along the Antarctic continent coastline. Several individuals returned to McMurdo Sound, but others were last located more than 400 km from McMurdo. Routes followed suggest that pups can use the pack ice habitat, but prefer to remain closer to the coastline than do adults. Accepted: 21 July 1998     

Diving physiology and winter foraging behavior of a juvenile leopard seal (Hydrurga leptonyx)

Diving physiology and at-sea behavior of a juvenile leopard seal (Hydrurga leptonyx) were opportunistically measured in the Antarctic Peninsula during winter 2002. Total body oxygen stores were estimated from measures of hematocrit, hemoglobin, myoglobin, and total blood volume and were used to calculate an aerobic dive limit (ADL). Movement patterns and diving behavior were measured by equipping the seal with a Satellite Relay Data Logger that transmitted data from 8–31 August 2002. The seal remained in a focal area, in contrast to crabeater seals tracked simultaneously. The seal displayed short, shallow dives (mean 2.0±1.4 min, 44±48 m) and spent 99.9% of its time within the estimated ADL of 7.4 min. The shallow diving behavior contradicts previous diet research suggesting Antarctic krill (Euphausia superba) is the primary prey of leopard seals during the winter months as krill were found at deeper depths during this period. These measurements of diving and movement of a leopard seal provide valuable preliminary data necessary to develop future research on the at-sea behavior of an apex predator in the Antarctic ecosystem.     

Diet composition of polar bears in Svalbard and the western Barents Sea

We estimated both the numerical and biomass composition of the prey of polar bears (Ursus maritimus) from 135 opportunistic observations of kills in Svalbard and the western Barents Sea collected from March to October 1984-2001. By number, the prey composition was dominated by ringed seals (Phoca hispida) (63%), followed by bearded seals (Erignathus barbatus) (13%), harp seals (P. groenlandica) (8%) and unknown species (16%). However, when known prey were converted to biomass, the composition was dominated by bearded seals (55%), followed by ringed seals (30%) and harp seals (15%). Results indicated that bearded seals are an important dietary item for polar bears in the western Barents Sea. We believe that different patterns of space use by different bears may result in geographic variation of diet within the same population.     

A PREHISTORIC BREEDING POPULATION OF HARP SEALS (PHOCA GROENLANDICA) IN THE BALTIC SEA

The pelagic and gregarious, low Arctic harp seal ( Phoca groenlandica ) is the most common seal species in most refuse faunas from coastal hunter-gatherer sites dating from the late Atlantic to the early Subboreal period ( ca. 4000-2000 cal B. C.) in the Baltic Sea. Our main objective was to examine the migration contra breeding population hypotheses regarding the Baltic harp seals. Analyses of epiphyseal fusion data and osteometry of archeological harp seal remains from 25 dwelling-sites suggest that a local breeding population established itself in the early Subboreal period. In the Middle Neolithic the rookery possibly was situated in the Baltic proper, south of Aland and west of Gotland. The mean adult size of the Baltic harp seals decreased, suggesting minimal genetic exchange with the north Atlantic Ocean population. Genetic drift, interspecific competition, and over-hunting by humans are all factors likely to have contributed to the eventual extinction of harp seals in the Baltic Sea.     

Annual distribution of hooded seals (Cystophora cristata) in the Greenland and Norwegian seas

 Nineteen hooded seals (Cystophora cristata) were tagged with satellite-linked platform terminal transmitters (PTT) on the sea ice near Jan Mayen. Fifteen were instrumented after completion of the moult in July 1992 (five males, ten females, at 71°N, 12°W), and four during breeding in March 1993 (four females, at 69°N, 20°W). Sixteen of the seals were tagged with Satellite-Linked Time-Depth-Recorders (SLTDR), yielding location, dive depth and dive duration data. The average (±SD) longevity of all PTTs was 199±84 days (n=19; range: 43–340 days), and they yielded 12,834 location fixes. Between tagging in July 1992 and pupping in March 1993, two seals remained in or near the ice off the east coast of Greenland for most of the tracking period. However, most of the seals made one or several trips away from the ice edge, mostly to distant waters. These excursions had an average (±SD) duration of 47±22 days (n=46; range: 4–99 days). Eight seals travelled to waters off the Faeroe Islands, three to the continental shelf break south of Bear Island, and three to the Irminger Sea southwest of Iceland. Eleven seals were tracked in the period between breeding (March/April) and moulting (July). Several of these spent extended periods at sea west of the British Isles, or in the Norwegian Sea. Received: 3 August 1994/Accepted: 4 July 1995     

PHOCINE DISTEMPER VIRUS INFECTION IN HARP SEALS (PHOCA GROENLANDICA)

Antibodies against phocine distemper virus were found in liver and serum specimens of harp seals from the Barents Sea and north of Jan Mayen (West Ice) in 1987 and 1989. By immunoperoxidase testing, antibodies against phocine distemper virus could be demonstrated in 34 of 74 liver specimens (46%) from 1987 and in 44 of 126 liver specimens (35%) from 1989. The prevalence of neutralizing serum antibodies against phocine distemper virus was found to be 98% in 1989. Both findings clearly demonstrate the presence of a widespread infection with phocine distemper virus or a closely related virus in harp seals. The significance of these findings to the recent outbreak of phocine distemper virus infection in harbor seals in Northwestern Europe is discussed and it is concluded that there may be an epizootiological link between this outbreak and the large migration of harp seals in 1987.     

The physiological and behavioural development of diving in Australian fur seal (<Emphasis Type="Italic">Arctocephalus pusillus doriferus</Emphasis>) pups

The physiological and behavioural development of diving was examined in Australian fur seal (Arctocephalus pusillus doriferus) pups to assess whether animals at weaning are capable of exploiting the same resources as adult females. Haematocrit, haemoglobin and myoglobin contents all increased throughout pup development though total body oxygen stores reached only 71% of adult female levels just prior to weaning. Oxygen storage components, however, did not develop at the same pace. Whereas blood oxygen stores had reached adult female levels by 9 months of age, muscle oxygen stores were slower to develop, reaching only 23% of adult levels by this age. Increases in diving behaviour corresponded to the physiological changes observed. Pups spent little time (<8%) in the water prior to moulting (age 1–2 months) whereas following the moult, they spent >27% of time in the water and made mid-water dives (maximum depth 35.7 ± 2.9 m) with durations of 0.35 ± 0.03 min. By 9 months (just prior to weaning), 30.5 ± 9.3% of all dives performed were U-shaped benthic dives (maximum depth 65.0 ± 6.0 m) with mean durations of 0.87 ± 0.25 min, significantly shorter than those of adult females. These results suggest that while Australian fur seal pups approaching the age of weaning are able to reach similar depths as adult females, they do not have the physiological capacity to remain at these depths for sufficient durations to exploit them to the same efficiency.