Seasonal change in foraging areas and dive depths of breeding king penguins at Heard Island

The seasonal variation in the foraging behaviour of king penguins (Aptenodytes patagonicus) was studied at Heard Island (53°05′S, 73°30′E) during 1992/1993. On seven occasions throughout the breeding cycle, time-depth-light recorders were deployed on breeding adults to record the dive activities and foraging. Foraging locations changed with season: in autumn and spring 1992, adults foraged between 48–52°S and 74–78°E, about 370 km NNE of Heard Island close to the Polar Front. Two penguins tracked in winter travelled 2220 km east of Heard Island (95°E) along the northern ice limit, and 1220 km south of Heard Island to approximately 65°S, respectively. In spring (October), the penguins again foraged further north than during winter. The foraging area utilised in October overlapped the area where the penguins foraged in March/April. The penguins' diving behaviour also varied seasonally: the modal depth of deep dives (>50 m) increased from about 100 m in February to 220 m in October. Mean dive depths increased from 70 ± 52 m in March 1992 to 160 ± 68 m in August 1992. Penguins dived deep (>50 m) only during daylight hours (16 h in February, 9 h in July). Mean dive durations ranged from 2.9 ± 1.1 min in March 1992 to 5.1 ± 1.2 min in August 1992. Associated with changes in foraging location and dive behaviour was a change in diet composition: during summer the penguins ingested mainly myctophid fish (>90%) while in winter the most important diet item was squid. Accepted: 19 October 1998     

Movements and behavior of satellite-tagged spotted seals (Phoca largha) in the Bering and Chukchi Seas

Satellite-linked tags were attached to 12 spotted seals (Phoca largha) captured at a coastal lagoon in the eastern Chukchi Sea during August 1991–1993. Movements of seals were tracked for 32–298 days using the Argos system. Of 9,651 total location records obtained, 7,268 were usable. Individual seals were located on 41–96% of the days that tags were operational. During August–November, tagged seals alternated haul-outs at coastal sites lasting 1–304 h with trips to sea of 14–901 h. Coastal haul-outs occurred at 14 sites in western Alaska and eastern Russia. On several trips to sea, seals covered distances of more than 1,000 km. Movement southward from the Chukchi Sea generally began in October, with most of the seals passing through the Bering Strait during November. Seals first hauled out on sea ice in October (Chukchi Sea) or November (Bering Sea), and generally moved southward during October–December as sea-ice coverage increased. Seven seals, whose transmitters were still operating, spent December to June in the Bering Sea region between Kuskokwim Bay and Anadyr Gulf, which corresponded to the location of the ice front. The seals made active east-west movements within the ice front. Spotted seals are unlike other ice-breeding seals in that they regularly use coastal haul-outs during summer and autumn. Compared to the closely related Pacific harbor seal (Phoca vitulina richardsi), spotted seals make much longer trips to sea and spend longer continuous periods at their haul-outs during summer and autumn. Received: 9 April 1997 / Accepted: 30 September 1997     

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     

Distribution and diving behaviour of harp seals (<Emphasis Type="Italic">Pagophilus groenlandicus</Emphasis>) from the Greenland Sea stock

The distribution and diving behaviour of 16 adult harp seals (Pagophilus groenlandicus) from the Greenland Sea stock were studied in 1993 and 1999, using satellite-linked dive recorders (SDRs). The seals remained near the pack-ice edge in the Greenland Sea between breeding and moulting (April/May 1993; 6F) and during the first 7 weeks after moulting (June/July, 1999; 4F, 6M), there diving to depths of <100 m. In mid-July 1999, seven out of eight seals with active SDRs migrated into the Barents Sea, there diving to <400 m and sharing feeding grounds with the Barents Sea harp seal stock. Between September and December, six of these seals joined the eighth seal in the Denmark Strait until March 2000, there diving to depths of 100–400 m. Overall, dives were significantly deeper in the day and in winter than at night and in summer, with some regional differences. Harp seals are considered pack-ice-associated seals, but our tagged seals spent a considerable proportion of their time in open water, their distribution largely overlapping with that of capelin (Mallotus villosus).     

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.     

Limited use of sea ice by the Ross seal (<Emphasis Type="Italic">Ommatophoca rossii</Emphasis>), in Amundsen Sea,Antarctica, using telemetry and remote sensing data

To understand the use and importance of the Antarctic sea ice to the Ross seal (Ommatophoca rossii), four adult females were tagged with Argos satellite transmitters in the Amundsen Sea, Antarctica. The Ross seal is the least studied of the Antarctic seal species and nothing was previously known about their behaviour in the Amundsen Sea. During almost 1 year, their movements, haul out behaviour and time spent at different temperatures were logged. By comparing their movements with daily ice maps, distances to the ice edge were calculated, and seals dependence on sea ice for resting, breeding and moulting was analysed. The tagged seals spent on average 70.8 % (range 66.8–77.8 %) of their time in the water and hauled out mainly during the moult in December–January, and in late October–mid-November during breeding. During the pelagic period, they were on average 837.5 km (range 587–1,282 km) from the ice edge indicating a fully pelagic life during several months. Their pelagic behaviour suggests that Ross seals, although being an ice obligate species, may adapt comparatively easy to climate change involving ice melting and recession and thereby potentially being less sensitive to the reduction of sea ice than other Antarctic seal species. Although nothing is known about their mating behaviour, they appear to be relatively stationary during moulting and breeding, hence requiring a small ice surface. Although previous studies in other parts of Antarctica have found similar results, still many questions remain about this peculiar species.     

Movements and diving of adult ringed seals (Phoca hispida) in Svalbard

Seven post-moulting adult ringed seals (Phoca hispida) were equipped with Satellite Linked Dive Recorders in Svalbard in July 1996 to determine if ringed seals conduct long-distance post-moulting feeding excursions, and to obtain details of their diving behaviour. The mean duration of tags was 206 days (range 103–325). Two seals swam 400 km north to the drifting pack ice (82°N). The rest undertook more local movements. Forty-eight percent of all dives were shallower than 20 m and 90% were shallower than 100 m. Ninety-five percent of all dive durations were shorter than 10 min, and 99.5% were shorter than 15 min. This study has shown that adult ringed seals undertake varying patterns of post-moulting excursions. Accepted: 1 April 2000     

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.     

Postrelease dive ability in rehabilitated harbor seals

The efficacy of seal rehabilitation is examined in a postrelease study of dive ability in harbor seal pups (Phoca vitulina) in the Wash, United Kingdom. Six rehabilitated seals were fitted with Sea Mammal Research Unit (SMRU) Argos Satellite Relay Data Logger tags and their individual dive behavior was monitored for an average of 122 d. The upper 90 percentile edge of dive behavior (dive duration [DD₉₀] and percentage of time at‐sea spent in a dive [PD₉₀]), in 7 d bins, was used as a proxy for physiological dive ability. The results are compared with data from five wild adult harbor seals. There was no statistically significant difference between (1) the mean track duration of rehabilitated seals (126.20 ± 27.48 [SD] d) and adult seals (150.2 ± 24.62 d) (P= 0.108), indicating no evidence that short‐term survival was less in the rehabilitated group; (2) the mean mass‐scaled DD₉₀ of rehabilitated seals (3.95 ± 0.37 min) and adult seals (4.09 ± 0.55 min) (P= 0.632); and (3) the mean PD₉₀ of rehabilitated seals (81.62 ± 1.21%) and adult seals (81.48 ± 3.93%) (P= 0.943). These three results all suggest the success of the rehabilitation program in terms of short‐term survival and dive ability.     

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 and haulout behavior of crabeater seals in the Weddell Sea,Antarctica, during March 1986

Summary Time-depth recorders were used to study the diving and haulout behavior of six crabeater seals in the marginal. ice edge zone of the Weddell Sea during March 1986. Haulout patterns revealed the seals' clear preference for diving during darkness and hauling out onto sea ice during daylight. Seals did not necessarily haul out every day; individual seals hauled out on 80–100% of days during the study period. Four general dive types were identified: 1) traveling dives, 2) foraging dives, 3) crepuscular foraging dives, and 4) exploratory dives. Nearly continual diving occurred for extended periods (about 16 h) nightly, with one individual diving up to 44 h without interruption. Foraging dives occurring during crepuscular periods were deeper than those made during the darkest hours. The authors suggest that the distinct diel pattern of dive timing and depth may be related to possible predator avoidance behavior by the seals' principal prey, Antarctic Krill.     

Seasonal diving behaviour in lactating subantarctic fur seals on Amsterdam Island

Diving behaviour was investigated in female subantarctic fur seals (Arctocephalus tropicalis) breeding on Amsterdam Island, Indian Ocean. Data were collected using electronic Time Depth Recorders on 19 seals during their first foraging trip after parturition in December, foraging trips later in summer, and during winter. Subantarctic fur seals at Amsterdam Island are nocturnal, shallow divers. Ninety-nine percent of recorded dives occurred at night. The diel dive pattern and changes in dive parameters throughout the night suggest that fur seals follow the nycthemeral migrations of their main prey. Seasonal changes in diving behaviour amounted to the fur seals performing progressively deeper and longer dives from their first foraging trip through winter. Dive depth and dive duration increased from the first trip after parturition (16.6 ± 0.5 m and 62.1 ± 1.6 s respectively, n=1000) to summer (19.0 ± 0.4 m and 65 ± 1 s, respectively, n=2000) through winter (29.0 ± 1.0 m and 91.2 ± 2.2 s, respectively, n=800). In summer, subantarctic fur seals increased the proportion of time spent at the bottom during dives of between 10 and 20 m, apparently searching for prey when descending to these depths, which corresponded to the oceanic mixed layer. In winter, fur seals behaved similarly when diving between 20 and 50 m, suggesting that the most profitable depths for feeding moved down during the study period. Most of the dives did not exceed the physiological limits of individuals. Although dive frequency did not vary (10 dives/h of night), the vertical travel distance and the time spent diving increased throughout the study period, while the post-dive interval decreased, indicating that subantarctic fur seals showed a greater diving effort in winter, compared to earlier seasons. Accepted: 1 August 1999     

Different habitat use strategies by subadult and adult ringed seals (<Emphasis Type="Italic">Phoca hispida</Emphasis>) in the Bering and Chukchi seas

Habitat partitioning by adult and subadult ringed seals (Phoca hispida) is poorly understood. Conclusions about displacement of subadult seals to suboptimal offshore habitat are largely based on nearshore observations as few satellite tagging studies include data from winter months. In this study, movement patterns of 14 subadult and 11 adult ringed seals were monitored in the Bering and Chukchi seas using satellite-linked telemetry. Seals were captured in Kotzebue Sound, Alaska, during October 2007 and 2008 and tracked for 17–297 days. Subadult ringed seals traveled south from the Chukchi Sea into the Bering Sea ([`(x)] \bar{x}  = 36 km/day) as sea ice coverage increased during November and December, remained ~1,000 km south near the ice edge during winter and returned north in the spring with the receding ice edge. Adults remained in the Chukchi and northern Bering seas, where their movements were more localized ([`(x)] \bar{x}  = 22 km/day). Adults were on average 322 km farther from the ice edge and 48 km closer to land and shorefast ice than were subadults. During winter, adult ringed seals construct and maintain breathing holes through the ice, and in spring, females give birth in subnivean lairs, mostly in shorefast ice; adult males defend breeding territories around those lairs. Our results show that subadult ringed seals, unconstrained by the need to maintain territories that contain stable breeding/pupping habitat, moved south to the Bering Sea ice edge, where there are better feeding opportunities, lower energetic costs (no breathing hole maintenance), and less exposure to predation.     

The effects of El Niño–La Niña on reproductive parameters of elephant seals feeding in the Bellingshausen Sea

Aim To assess the impacts of El Niño–La Niña events on the pup weaning mass and diet of female southern elephant seals (Mirounga leonina) feeding in the Bellingshausen Sea, Antarctica, and to understand the ecological processes that drive these impacts. Location Atlantic southern elephant seal weaning mass and diet were measured at King George Island (62º14′ S, 58º30′ W). Feeding areas for pregnant female seals from King George Island are located west of Alexander Island in the Bellingshausen Sea. Methods Data on weaning mass were collected between 1985 and 1994 during the breeding season (September–November). Moulting females were anaesthetized and cephalopod beaks were isolated and identified from stomach contents obtained from stomach lavages. Sea‐surface temperature anomaly (SSTA) data for the ‘El Niño 3.4’ geographical region (5º N–5º S, 120º W–170º W) were used to define El Niño–Southern Oscillation (ENSO) event years (grouped as El Niño, La Niña and Neutral) as well as the strength of each ENSO event year. Using data from the US National Center for Environmental Prediction, temperature, sea ice concentration and atmospheric pressure anomalies in the Bellingshausen Sea were calculated from March to August, corresponding to the feeding period of pregnant female seals. Results Positive temperature anomalies and negative pressure anomalies in the Bellingshausen Sea were observed during La Niña years and negative temperature anomalies and positive pressure anomalies during El Niño years. These data correlate with sea ice concentration anomalies, which are highly negative during La Niña years and highly positive during El Niño years. Warm temperature conditions in the Bellingshausen Sea during La Niña years are strongly related to both higher weaning mass in elephant seals and to an increase in squid beaks in the stomach contents of females. Main conclusions It is possible that higher elephant seal weaning masses in La Niña years correlate with warmer waters in the Bellingshausen Sea leading to the rapid growth of squid and their more frequent descents to depths frequented by elephant seals. This results in increased predation by pregnant females, leading to a greater mass among weaned pups. This hypothesis may guide future research about interactions between climate and the marine biosphere.     

Diving behaviour of hooded seals (Cystophora cristata) in the Greenland and Norwegian Seas

Satellite-linked dive recorders were used to collect data on depths and durations of ∼120,000 dives by 16 hooded seals (Cystophora cristata). Following tagging after moult (four males, eight females) and breeding (four females) off east Greenland, seals dispersed widely in the northeast Atlantic during 172 ± 97 days (mean satellite-linked dive recorder lifetime ± SD). Meso/bathypelagic dives of 5- to 25-min duration to 100–600 m dominated (75%), but some very deep (≥1016 m) and long (>52 min) dives occurred. Diving in open ocean was continuous, with an estimated 90.7±0.8% (mean±SE) of time spent submerged. The proportion of time spent submerged was similar during night and day, but dives during the day were generally deeper and longer (P < 0.05) than during the night. Also, dives in winter were deeper and longer than in summer. Published data on the distribution of likely prey suggest that Greenland halibut (Reinhardtius hippoglossoides), redfish (Sebastes spp.), polar cod (Boreogadus saida), herring (Clupea harengus), squid (Gonatus fabricii) and blue whiting (Micromesistius poutassou) are important prey of hooded seals. Accepted: 22 January 1999     

Record of a new northern range of Sowerby's beaked whale (Mesoplodon bidens )

Two beaked whales identified as Sowerby's beaked whale (Mesoplodon bidens) were observed in sea state 0–1 on 16 July 1995 at 71°30′N 04°00′E in the Norwegian Sea. A number of morphological features, such as dentition, were clearly seen during the encounter. The Sowerby's beaked whale's core distribution is in the North Sea and the previous northernmost sighting was made at 63°06′N 00°41′E. According to the literature, the species has never been recorded in the polar zone. The sighting in the Norwegian Sea suggests that the current data on the species distribution is uncertain, and that its range may include the polar waters of the Norwegian Sea. Received: 24 April 1996 / Accepted: 25 August 1996     

Distribution and diving of harbour seals (Phoca vitulina) in Svalbard

The distribution, movements and diving of high-arctic harbour seals (Phoca vitulina) were studied in Svalbard, Norway, from 1992 to 1995. A total of 14 seals were equipped with satellite transmitters at Prins Karls Forland (ca. 78°30′N 12°E). These gave data on position, but ten also gave information on dive depths (N ∼ 160,000) and dive durations (N ∼ 162,000). Dive-depth frequencies show that ∼50% of the diving is shallower than 40 m, and that 95% of the diving is shallower than 250 m. Based on dive-duration frequencies, ∼50% of the dives lasted 2–4 min, 90% of the dives lasted less than 7 min, and 97% were shorter than 10 min. All but three seals stayed in the tagging area. Accepted: 6 October 2000     

Summer at-sea distribution of little auks Alle alle and harp seals Pagophilus (Phoca) groenlandica in the Fram Strait and the Greenland Sea: impact of small-scale hydrological events

Among the most numerous seabird and pinniped species of the Fram Strait and the Greenland Sea, little auks Alle alle and harp seals Pagophila (Phoca) groenlandica are very abundant in the mixed Polar/Arctic Waters at the front between the two water masses. This must reflect the presence of very high concentrations of their food, Arctic zooplankton and nekton, massively attracting their predators. Such a high biological production seems to be depending on new primary production based on upwelling and high nutrient concentration. This usually takes place at the ice edge (e.g. July 2005), but hydrological conditions such as eddies can modify its position, east of the front in open water as caused by a subsurface eddy (August 2005), or in ice-covered areas if westerly winds push the pack ice to the east, eventually covering an eddy and causing very high concentrations of little auks and harp seals (July 2008). On the other hand, a dramatic decrease of pack ice coverage can move this water mass farther north and west, making it inaccessible to little auks during their breeding season, and apparently causing breeding failure in Jan Mayen in July 2005. In future years, if a much stronger diminution in sea ice coverage will take place, similar to the retreat in 2005 and 2007, the failure might affect the whole Spitsbergen population, as well as other seabird species feeding mainly at the ice edge.     

Combined Genetic and Telemetry Data Reveal High Rates of Gene Flow,Migration, and Long-Distance Dispersal Potential in Arctic Ringed Seals (Pusa hispida)

Ringed seals (Pusa hispida) are broadly distributed in seasonally ice covered seas, and their survival and reproductive success is intricately linked to sea ice and snow. Climatic warming is diminishing Arctic snow and sea ice and threatens to endanger ringed seals in the foreseeable future. We investigated the population structure and connectedness within and among three subspecies: Arctic (P. hispida hispida), Baltic (P. hispida botnica), and Lake Saimaa (P. hispida saimensis) ringed seals to assess their capacity to respond to rapid environmental changes. We consider (a) the geographical scale of migration, (b) use of sea ice, and (c) the amount of gene flow between subspecies. Seasonal movements and use of sea ice were determined for 27 seals tracked via satellite telemetry. Additionally, population genetic analyses were conducted using 354 seals representative of each subspecies and 11 breeding sites. Genetic analyses included sequences from two mitochondrial regions and genotypes of 9 microsatellite loci. We found that ringed seals disperse on a pan-Arctic scale and both males and females may migrate long distances during the summer months when sea ice extent is minimal. Gene flow among Arctic breeding sites and between the Arctic and the Baltic Sea subspecies was high; these two subspecies are interconnected as are breeding sites within the Arctic subspecies.     

A lifetime at depth: vertical distribution of southern elephant seals in the water column

Although numerous studies have addressed the migration and dive behaviour of southern elephant seals (Mirounga leonina), questions remain about their habitat use in the marine environment. We report on the vertical use of the water column in the species and the potential lifetime implications for southern elephant seals from Marion Island. Long-term mark-resight data were used to complement vertical habitat use for 35 known individuals tagged with satellite-relay data loggers, resulting in cumulative depth use extrapolated for each individual over its estimated lifespan. Seals spent on average 77.59% of their lives diving at sea, 7.06% at the sea surface, and 15.35% hauled out on land. Some segregation was observed in maximum dive depths and depth use between male and female animals—males evidently being physiologically more capable of exploiting increased depths. Females and males spent 86.98 and 80.89% of their lives at sea, respectively. While at sea, all animals spent more time between 300 and 400 m depth, than any other depth category. Males and females spent comparable percentages of their lifetimes below 100 m depth (males: 65.54%; females: 68.92%), though males spent 8.98% of their lives at depths in excess of 700 m, compared to females’ 1.84% at such depths. Adult males often performed benthic dives in excess of 2,000 m, including the deepest known recorded dive of any air-breathing vertebrate (>2,133 m). Our results provide a close approximation of vertical habitat use by southern elephant seals, extrapolated over their lifespans, and we discuss some physiological and developmental implications of their variable depth use.