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     

Individual Foraging Strategies Reveal Niche Overlap between Endangered Galapagos Pinnipeds

Most competition studies between species are conducted from a population-level approach. Few studies have examined inter-specific competition in conjunction with intra-specific competition, with an individual-based approach. To our knowledge, none has been conducted on marine top predators. Sympatric Galapagos fur seals (Arctocephalus galapagoensis) and sea lions (Zalophus wollebaeki) share similar geographic habitats and potentially compete. We studied their foraging niche overlap at Cabo Douglas, Fernandina Island from simultaneously collected dive and movement data to examine spatial and temporal inter- and intra-specific competition. Sea lions exhibited 3 foraging strategies (shallow, intermediate and deep) indicating intra-specific competition. Fur seals exhibited one foraging strategy, diving predominantly at night, between 0–80 m depth and mostly at 19–22 h. Most sea lion dives also occurred at night (63%), between 0–40 m, within fur seals'' diving depth range. 34% of sea lions night dives occurred at 19–22 h, when fur seals dived the most, but most of them occurred at dawn and dusk, when fur seals exhibited the least amount of dives. Fur seals and sea lions foraging behavior overlapped at 19 and 21 h between 0–30 m depths. Sea lions from the deep diving strategy exhibited the greatest foraging overlap with fur seals, in time (19 h), depth during overlapping time (21–24 m), and foraging range (37.7%). Fur seals foraging range was larger. Cabo Douglas northwest coastal area, region of highest diving density, is a foraging “hot spot” for both species. Fur seals and sea lions foraging niche overlap occurred, but segregation also occurred; fur seals primarily dived at night, while sea lions exhibited night and day diving. Both species exploited depths and areas exclusive to their species. Niche breadth generally increases with environmental uncertainty and decreased productivity. Potential competition between these species could be greater during warmer periods when prey availability is reduced.     

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     

Diving patterns of female macaroni penguins breeding on Marion Island, South Africa

Despite the large biomass of macaroni penguins Eudyptes chrysolophus in the Southern Ocean, their feeding ecology is poorly known at some important breeding localities. We investigated the diving behaviour and diet of female macaroni penguins feeding small chicks on Marion Island (46o52′S, 37o5′E), South Africa, one of the species’ most northerly breeding sites, supporting 4% of their global population. We then compared our results with similar studies from other localities. In December 2008, we collected information on 12 foraging trips from 6 individuals using time-depth recorders, as well as diet from 42 individuals. Median trip duration was 22.8 h (5.6–80.8 h). Penguins performed 42.8 ± 15.9 dives per hour at sea, with dive depths averaging 24.6 ± 8.6 m and lasting 40.8 ± 12.1 s, although 74.3% of dives were <10 m. Euphasids dominated their diet (86% by mass), mainly Thysanoessa vicina. A second peak in dive depths at 55–80 m might reflect the 12% of fish in their diet. The substantial proportion of shallow night dives (30% of total dives) suggests some foraging occurs at night. Differences in diving patterns of individual macaroni penguins in this study confirmed the behavioural flexibility of these birds reported from other breeding localities. However, most other studies assumed that dives <3–5 m were commuting dives whereas our study suggests that at least some prey are caught during shallow dives. We highlight how different analytical methods can change the outcome of studies. Despite macaroni penguins’ apparent flexibility in foraging behaviour during the breeding season, their numbers are decreasing globally. Further investigations of their foraging behaviour are needed to assess potential competition with other predators and krill fisheries.     

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     

Diving pattern and performance in the macaroni penguin Eudptes chrysolophus

The pattern and characteristics of diving in two female macaroni penguins Eudyptes chrysolophus was studied, during the brooding period, using continuous-recording time-depth recorders, for a total of I8 days (15 consecutive days) during which the depth, duration and timing of 4876 dives were recorded. Diving in the first 11 days was exclusively diurnal, averaging 244 dives on trips lasting 12 hours. Near the end of the brooding period trips were longer and included diving at night. About half of all trips (except those involving continuous night-time diving) was spent in diving and dive rate averaged 14–25 dives per hour (42 per hour at night). The duration of day time dives varied between trips, and averaged 1.4–1.7 min, with a subsequent surface interval of 0.5–0.9 min. Dive duration was significantly directly related to depth, the latter accounting for 53% of the variation. The average depths of daytime dives were 20–35 m (maximum depth 11 5 m). Dives at night were shorter (average duration 0.9 min) and much shallower (maximum 11 m); depth accounted for only 6% of the variation in duration. Estimates of potential prey capture rates (3–5 krill per dive; one krill every 17–20 s) are made. Daily weight changes in chicks were directly related to number of dives, but not to foraging trip duration nor time spent diving. Of the other species at the same site which live by diving to catch krill, gentoo penguins forage exclusively diurnally, making longer. deeper dives; Antarctic fur seals, which dive to similar depths as macaroni penguins, do so mainly at night.     

Summer diving and haul‐out behavior of leopard seals (Hydrurga leptonyx) near mesopredator breeding colonies at Livingston Island,Antarctic Peninsula

Leopard seals are conspicuous apex predators in Antarctic coastal ecosystems, yet their foraging ecology is poorly understood. Historically, the ecology of diving vertebrates has been studied using high‐resolution time‐depth records; however, to date such data have not been available for leopard seals. Twenty‐one time‐depth recorders were deployed on seasonally resident adult females in January and February between 2008 and 2014. The average deployment length was 13.65 ± 11.45 d and 40,308 postfilter dives were recorded on 229 foraging trips. Dive durations averaged 2.20 ± 1.23 min. Dives were shallow with 90.1% measuring 30 m or less, and a mean maximum dive depth of 16.60 ± 10.99 m. Four dive types were classified using a k‐means cluster analysis and compared with corresponding animal‐borne video data. Dive activity (number of dives/hour) was concentrated at night, including crepuscular periods. Haul‐out probabilities were highest near midday and were positively correlated with available daylight. Visual observations and comparisons of diving activity between and within years suggest individual‐based differences of foraging effort by time of day. Finally, dive and video data indicate that in addition to at‐surface hunting, benthic searching and facultative scavenging are important foraging strategies for leopard seals near coastal mesopredator breeding colonies.     

ANALYSIS OF SWIM VELOCITIES DURING DEEP AND SHALLOW DIVES OF TWO NORTHERN FUR SEALS, CALLORHINUS URSINUS

Swim velocities at 15-sec intervals and maximum depth per dive were recorded by microprocessor units on two "mixed diver" adult female northern fur seals during summer foraging trips. These records allowed comparison of swim velocities of deep (>75 m) and shallow (<75 m) dives.
Deep dives averaged 120 m depth and 3 min duration; shallow dives averaged 30 m and 1.2 min. Mean swim velocities on deep dives were 1.8 and 1.5 m/sec for the two animals; mean swim velocities on shallow dives were 1.5 and 1.2 m/sec. The number of minutes per hour spent diving during the deep and shallow dive patterns were 11 and 27 min, respectively.
Swim velocity, and hence, relative metabolic rate, did not account for the differences in dive durations between deep and shallow dives. The long surface durations associated with deep dives, and estimates of metabolic rates for the observed swim velocities, suggest that deep dives involve significant anaerobic metabolism.     

SUMMER DIVING BEHAVIOR OF MALE WALRUSES IN BRISTOL BAY, ALASKA

Pacific walruses ( Odobenus rosmarus divergens ) make trips from ice or land haul-out sites to forage for benthic prey. We describe dive and trip characteristics from time-depth-recorder data collected over a one-month period during summer from four male Pacific walruses in Bristol Bay, Alaska. Dives were classified into four types. Shallow (4 m), short (2.7 min), square-shaped dives accounted for 11% of trip time, and many were probably associated with traveling. Shallow (2 m) and very short (0.5 min) dives composed only 1% of trip time. Deep (41 m), long (7.2 min), square-shaped dives accounted for 46% of trip time and were undoubtedly associated with benthic foraging. V-shaped dives ranged widely in depth, were of moderate duration (4.7 min), and composed 3% of trip time. These dives may have been associated with navigation or exploration of the seafloor for potential prey habitat. Surface intervals between dives were similar among dive types, and generally lasted 1–2 min. Total foraging time was strongly correlated with trip duration and there was no apparent diel pattern of diving in any dive type among animals. We found no correlation between dive duration and postdive surface interval within dive types, suggesting that diving occurred within aerobic dive limits. Trip duration varied considerably within and among walruses (0.3–9.4 d), and there was evidence that some of the very short trips were unrelated to foraging. Overall, walruses were in the water for 76.6% of the time, of which 60.3% was spent diving.     

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     

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 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).     

Diving pattern and performance in relation to foraging ecology in the gentoo penguin, Pygoscelis papua

We present data on diving pattern and performance (dive depth, duration, frequency and organization during the foraging trip) in gentoo penguins Pygoscelis papua , obtained using time-depth recorders ( n = 9 birds, 99 foraging trips). These data are used to estimate various parameters of foraging activity, e.g. foraging range, prey capture rates, and are compared in relation to breeding chronology. Foraging trip duration was 6 h and 10 h, and trip frequency 1.0/day and 0.96/day, during the brooding and creche periods, respectively. Birds spent on average 52%of each foraging trip diving. Dive depth and duration were highly bimodal: shallow dives (< 21 m) averaged 4 m and 0.23 min, and deep dives (> 30 m) 80 m and 2.5 min, respectively. Birds spent on average 71%and 25%of total diving time in deep and shallow dives, respectively. For deep dives, dive duration exceeded the subsequent surface interval, but shallow dives were followed by surface intervals 2–3 times dive duration. We suggest that most shallow dives are searching/exploratory dives and most deep dives are feeding dives. Deep dives showed clear diel patterns averaging 40 m at dawn and dusk and 80–90 m at midday. Estimated foraging ranges were 2.3 km and 4.1 km during the brood and creche period, respectively. Foraging trip duration increased by 4 h between the brood and creche periods but total time spent in deep dives (i.e. time spent feeding) was the same (3 h). Of 99 foraging trips, 56%consisted of only one dive bout and 44%of 2–4 bouts delimited by extended surface intervals > 10 min. We suggest that this pattern of diving activity reflects variation in spatial distribution of prey rather than the effect of physiological constraints on diving ability.     

Diving patterns of a Ross seal (Ommatophoca rossii ) near the eastern coast of the Antarctic Peninsula

In January 1987 we documented the diving patterns of a female Ross seal (Ommatophoca rossii) in the marginal pack-ice zone near the eastern coast of the Antarctic Peninsula for 2 days using a microprocessor-based time-depth recorder. The seal hauled out during the day and dived continually when in the water at night. Dives averaged 110 m deep and 6.4 min long; the deepest dive was 212 m and the longest 9.8 min. Dives were deepest near twilight and shallowest at night; this pattern suggests that the seal's prey, presumably mid-water squid and fish, may have been making vertical migrations or changing predator-avoidance behavior in response to diel light patterns. The dives of this Ross seal were substantially deeper, on average, than those of crabeater seals (Lobodon carcinophagus), which forage in the same areas on Antarctic krill (Euphausia superba). Received: 15 August 1996 / Accepted: 22 February 1997     

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.     

Diving and foraging behaviour of Adélie penguins in areas with and without fast sea-ice

The diving and foraging behaviours of Adélie penguins, Pygoscelis adeliae, rearing chiks at Hukuro Cove, Lützow-Holm Bay, where the fast sea-ice remained throughout summer, were compared to those of penguins at Magnetic Island, Prydz Bay, where the fast sea-ice disappeared in early January. Parent penguins at Hukuro Cove made shallower (7.1–11.3 m) but longer (90–111 s) dives than those at Magnetic Island (22.9 m and 62 s). Dive duration correlated with dive depth at both colonies (r ² = 0.001 ∼ 0.90), but the penguins atg Hukuro Cove made longer dives for a given depth. Parents at Hukuro Cove made shorter foraging trips (8.1–14.4 h) with proportionally longer walking/swimming (diving < 1 m) travel time (27–40% of trip duration) and returned with smaller meals (253–293 g) than those at Magnetic Island, which foraged on average for 57.2 h, spent 2% of time walking/swimming ( < 1 m) travel, and with meals averaging 525 g. Trip duration at both colonies correlated to the total time spent diving. Trip duration at Hukuro Cove, but not at Magnetic Island, increased as walking/swimming ( < 1 m) travel time increased. These differences in foraging behaviour between colonies probably reflected differences in sea-ice cover and the availability of foraging sites. Received: 3 November 1995/Accepted: 29 May 1996     

VARIATION IN THE FORAGING LOCATION OF ANTARCTIC FUR SEALS (ARCTOCEPHALUS GAZELLA) AND THE EFFECTS ON DIVING BEHAVIOR

This study investigated how female Antarctic fur seals adapt their foraging behavior, over time scales of days, to spatial unpredictability in the distribution of their food. Lactating Antarctic fur seals are central-place foragers that feed on highly patchy but spatially and temporally dynamic food. We measured the foraging distribution of 28 fur seals to test whether variation in foraging trip durations was reflected in variation in the location of foraging and the diving behavior of seals at sea. Based on the maximum distance travelled from the breeding beach, three categories of foraging trips were denned: those to the continental shelf area ( n = 12, median = 71 km), to oceanic water ( n = 11, median =164 km), and to farther offshore oceanic waters ( n = 5, median = 260 km). Trip duration and mean surface speed were positively correlated with the maximum distance travelled from the breeding beach. Seals on longer trips spent proportionally less of their time submerged, but there was no significant difference in the total number of dives or the total time spent foraging by seals in relation to trip duration. Evidence from this study and previous work investigating energy gain suggests that an animal on a longer foraging trip could potentially have a higher mean energy return per dive than a similar animal on a shorter foraging trip. Evidence presented suggests that the type of foraging trip (near or far) is not predetermined by the animal but may be a simple response to the stochastic distribution of the resources available.     

Dive types and circadian behaviour patterns of Saimaa ringed seals<Emphasis Type="Italic">Phoca hispida saimensis</Emphasis> during the open-water season

Diving and circadian behaviour patterns of 7 free-ranging Saimaa ringed sealsPhoca hispida saimensis Nordquist, 1899 were examined by VHF-radiotelemetry during open-water seasons between May and November in Lake Saimaa, eastern Finland. The mean recorded dive duration ranged from 2.8 to 6.5 min, with a maximum of 21 min. The mean dive depth ranged from 9.8 to 15.7 m, with maximum of 39.6 m. The maximum dive depth of each seal was limited by water depth in the study area. The dive depths were positively correlated with dive duration and body mass of the seal. Five different dive types were defined, as based on their depth-time characteristics, each falling into one of the three functional categories: travelling, feeding, and resting. Long duration diving bouts occurred mostly at night and were presumed to be resting dives. Saimaa ringed seals exhibited a circadian pattern of haul-out behaviour that shifted seasonally. During molting (May–June) the seals hauled-out both day and night, but later in summer haul-out was more frequent at night.     

MONITORING THE PREY-FIELD OF MARINE PREDATORS: COMBINING DIGITAL IMAGING WITH DATALOGGING TAGS

There is increasing interest in the diving behavior of marine mammals. However, identifying foraging among recorded dives often requires several assumptions. The simultaneous acquisition of images of the prey encountered, together with records of diving behavior will allow researchers to more fully investigate the nature of subsurface behavior. We tested a novel digital camera linked to a time-depth recorder on Antarctic fur seals ( Arctocephalus gazella ). During the austral summer 2000–2001, this system was deployed on six lactating female fur seals at Bird Island, South Georgia, each for a single foraging trip. The camera was triggered at depths greater than 10 m. Five deployments recorded still images (640 × 480 pixels) at 3-sec intervals (total 8,288 images), the other recorded movie images at 0.2-sec intervals (total 7,598 frames). Memory limitation (64 MB) restricted sampling to approximately 1.5 d of 5–7 d foraging trips. An average of 8.5% of still pictures (2.4%-11.6%) showed krill ( Euphausia sulperba ) distinctly, while at least half the images in each deployment were empty, the remainder containing blurred or indistinct prey. In one deployment krill images were recorded within 2.5 h (16 km, assuming 1.8 m/sec travel speed) of leaving the beach. Five of the six deployments also showed other fur seals foraging in conjunction with the study animal. This system is likely to generate exciting new avenues for interpretation of diving behavior.     

Deep-diving of Atlantic salmon (<Emphasis Type="Italic">Salmo salar</Emphasis>) during their marine feeding migrations

Data from seven data storage tags recovered from Atlantic salmon marked as smolts were analyzed for depth movements and patterns of deep diving during the marine migration. The salmon mostly stayed at the surface and showed diurnal activity especially from autumn until spring. During the first months at sea the salmon stayed at shallower depths (<100 m). The salmon took short deep dives (>100 m), that were rare or absent during the first summer at sea but increased in frequency and duration especially in late winter. The maximum depth of the dives varied from 419 to 1187 m. Most of dives were short, (<5 h) but could last up to 33 h. The duration of dives increased in late winter until spring and the overall depth and maximum depth per dive increased exponentially over time. The initiation of the dives was more common in evenings and at night, suggesting nocturnal diving. We hypothesized that deep diving is related to feeding of salmon as mesopelagic fish can be important food for salmon during winter.