Ringed and bearded seal densities in the eastern Chukchi Sea, 1999–2000

Aerial surveys were conducted in 1999 and 2000 to estimate the densities of ringed (Phoca hispida) and bearded (Erignathus barbatus) seals in the eastern Chukchi Sea. Survey lines were focused mainly on the coastal zone within 37 km of the shoreline, with additional lines flown 148–185 km offshore to assess how densities of seals changed as a function of distance from shore. Satellite-linked time-depth recorders were attached to ringed seals in both years to evaluate the time spent basking on the ice surface. Haulout patterns indicated that ringed seals transitioned to basking behavior in late May and early June, and that the largest proportion of seals (60–68%) was hauled out between 0830 and 1530 local solar time. Ringed seals were relatively common in nearshore fast ice and pack ice, with lower densities in offshore pack ice. The average density of ringed seals was 1.91 seals km⁻² in 1999 (range 0.37–16.32) and 1.62 seals km⁻² in 2000 (range 0.42–19.4), with the highest densities of ringed seals found in coastal waters south of Kivalina and near Kotzebue Sound. The estimated abundance of ringed seals for the entire study area was similar in 1999 (252,488 seals, SE=47,204) and 2000 (208,857 seals, SE=25,502). Bearded seals were generally more common in offshore pack ice, with the exception of high bearded seal numbers observed near the shore south of Kivalina. Bearded seal densities were not adjusted for haulout behavior, and therefore, abundance was not estimated. Unadjusted average bearded seal density was 0.07 seals km⁻² in 1999 (range 0.011–0.393) and 0.14 seals km⁻² in 2000 (range 0.009–0.652). Levels of primary productivity, benthic biomass, and fast ice distribution may influence the distributions of ringed and bearded seals in the Chukchi Sea. Information on movement and haulout behavior of ringed and bearded seals would be very useful for designing future surveys.     

Density and distribution of the ringed seal in the Bothnian Bay

A modified strip census of basking ringed seals in the Bothnian Bay was carried out during the last week of April and the first week of May 1988. Of the total ice covered area, 23174 km², 3236 km² (14%) was covered by the transects. The mean density in the area was 0.101 ringed seals km^-2, with substantial regional and local variation. Highest densities were found in the compact drift ice area. The estimated total population of ringed seals on the ice was 2093 in the Bothnian Bay proper and 248 in the North Quark. The 95% confidence limits of the estimates are ±24%.     

EFFECTS OF AN OFFSHORE OIL DEVELOPMENT ON LOCAL ABUNDANCE AND DISTRIBUTION OF RINGED SEALS (PHOCA HISPIDA) OF THE ALASKAN BEAUFORT SEA1

This study investigates how densities of ringed seals were affected by construction and oil production activities at Northstar, an artificial island built in the nearshore Alaskan Beaufort Sea. Intensive and replicated aerial surveys of seals on landfast ice were conducted during six spring seasons: for three seasons before island construction began (1997–1999); after a winter of intensive island construction (2000); and after more limited construction plus drilling (2001) and drilling plus oil production (2002). A Poisson regression model was used to examine seal densities relative to distance from Northstar after allowance for environmental covariates. Post hoc power analysis indicated that the study design and Poisson regression approach had high power to detect small‐scale changes in seal densities near Northstar if such changes had occurred. However, seal densities during spring were not significantly affected by proximity to Northstar in 2000–2002. Habitat, temporal, and weather factors did have significant effects on seal densities. This study shows that effects of the Northstar oil development on local distribution of basking ringed seals are no more than slight, and are small relative to the effects of natural environmental factors. An understanding of environmental effects is essential when assessing potential impacts of industrial activity on ringed seals.     

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.     

ABUNDANCE OF RINGED SEALS (PUSA HISPIDA) IN THE FJORDS OF SPITSBERGEN, SVALBARD, DURING THE PEAK MOLTING PERIOD

Ringed seal (Pusa hispida) abundance in Spitsbergen, Svalbard, was estimated during the peak molting period via aerial, digital photographic surveys. A total of 9,145 images, covering 41.7%–100% of the total fast‐ice cover (1,496 km²) of 18 different fjords and bays, were inspected for the presence of ringed seals. A total of 1,708 seals were counted, and when accounting for ice areas that were not covered by images, a total of 3,254 (95% CI: 3,071–3,449) ringed seals were estimated to be hauled out during the surveys. Extensive behavioral data from radio‐tagged ringed seals (collected in a companion study) from one of the highest density fjords during the molting period were used to create a model that predicts the proportion of seals hauled out on any given date, time of day, and under various meteorological conditions. Applying this model to the count data from each fjord, we estimated that a total of 7,585 (95% CI: 6,332–9,085) ringed seals were present in the surveyed area during the peak molting period. Data on interannual variability in ringed seal abundance suggested higher numbers of seals in Van Keulenfjorden in 2002 compared to 2003, while other fjords with very stable ice cover showed no statistical differences. Poor ice conditions in general in 2002 probably resulted in seals from a wide area coming to Van Keulenfjorden (a large fjord with stable ice in 2002). The total estimated number of ringed seals present in the study area at the time of the survey must be regarded as a population index, or at least a minimum estimate for the area, because it does not account for individuals leaving and arriving, which might account for a considerable number of animals. The same situation is likely the case for many other studies reporting aerial census data for ringed seals. To achieve accurate estimates of population sizes from aerial surveys, more extensive knowledge of ringed seal behavior will be required.     

SEAL RESPONSES TO AIRGUN SOUNDS DURING SUMMER SEISMIC SURVEYS IN THE ALASKAN BEAUFORT SEA

Numbers, sighting distances, and behavior of seals were studied during a nearshore seismic program off northern Alaska in July-September 1996. We observed from the seismic vessel for 885.6 h, including all periods (day and night) when airguns operated and many periods without airguns. Of 422 seals seen, 421 were seen in daylight; 91-8% were ringed seals, 7.3% were bearded seals, and 0.9% were spotted seals. About 79% were first seen within 250 m of the seismic boat, and sighting rate declined rapidly at lateral distances > 50 m. During daylight, seals were seen at nearly identical rates (0.60-0.63/ h) during periods with no airguns firing, one airgun, and a "full-array" of 8-11 120-in³ airguns. However, seals tended to be farther away ( P < 0.0001) during full-array seismic. There was partial avoidance of the zone <150 m from the boat during full-array seismic, but seals apparently did not move much beyond 250 m. "Swimming away" was more common during full-array than no-airgun periods, but relative frequencies of five behaviors did not differ significantly among distance categories. Airgun operations were interrupted 112 times when seals were sighted within safety radii (150–250 m). The National Marine Fisheries Service specified these radii in the Incidental Harassment Authorization issued for the project; they are based on a 190 dB re 1 μPa (rms) criterion for broadband received level. Methods for estimating numbers of seals potentially affected by the seismic program are described, and effectiveness of monitoring and mitigation is discussed. There is an urgent need for more data on effects of strong seismic pulses on seals.     

A population size estimate of the finless porpoise,Neophocaena phocaenoides, from aerial sighting surveys in Ariake Sound and Tachibana Bay, Japan

We estimated the population size of the finless porpoise (Neophocaena phocaenoides) in Ariake Sound and Tachibana Bay of western Kyushu, southwestern Japan, from aerial sighting surveys using line transect methods. All 12 surveys were conducted from May 1993 to May 1994 (8 in Ariake Sound and 4 in Tachibana Bay). In addition to these, 14 surveys were also carried out to obtain information on porpoise occurrence in Tachibana Bay (5 surveys) and in neighboring Sumo Nada (5) and Yatsushiro Sound (4). In Ariake Sound, 225 porpoise groups (369 animals) were detected during all flights totalling 1,694.4 km. In Tachibana Bay, a total of 997.8 km was surveyed and 55 groups (290 animals) were sighted. However, no sightings were recorded in Sumo Nada (distance searched = 148.7 km) and Yatsushiro Sound (208.4 km). In Ariake Sound, few sightings were recorded from waters shallower than 5 m in depth. In Tachibana Bay all animals were detected from waters of less than 50 m depth. The population size was estimated as 1,983 animals in Ariake Sound (95% CI = 1,382-2,847), 1,110 in Tachibana Bay (95% CI = 642-1,920), and 3,093 in the 2 waters (1.3 individuals/km², 95% CI = 2,278-4,201).     

HAUL-OUT ACTIVITY OF RINGED SEALS (PHOCA HISPIDA) DETERMINED FROM SATELLITE TELEMETRY

The haul-out activity of 15 ringed seals ( Phoca hispida ) equipped with satellite-linked radio transmitters was studied in NW Greenland ( ca. 73°-78°N). Between 19 June 1997 and 30 June 1999, telemetry data on haulout activity were obtained by the "Land-Sea-Reporter" (LSR), "Time-at-Depth" (TAD), and "Timelines" (TIM) systems housed within the satellite transmitters. The haul-out activity (% of total time hauled out) reported by the TIM system, which is specifically designed for collecting haul-out data, was about 1.4 times higher than that inferred from the LSR, but only about 0.7 of that inferred from TAD data. The TIM were used to describe haul-out activity. A total of 1,011 d with TIM were obtained (64.5% of a total of 1,568 "seal-days" monitored) representing data from nearly an entire annual cycle. No differences were found in percentage of time hauled out per month among various age categories. At all seasons the haul-out time showed considerable individual variation. There were no trends in percentage of time hauled out per month during late summer, fall, and winter (August-February). During the High Arctic winter darkness (November-January) the percentage of haul-out per month ranged between 3.9% in an adult (SD = 2.44, range: 1.1%-5.7%, n = 3 mo) and 15.7% in a subadult (SD = 1.95, range: 13.7%-17.6%, n = 3 mo). From late March there was a significant increase in haul-out time. Between 1 and 30 June, when aerial surveys of basking ringed seals usually are conducted, the haul-out time (% per day) increased from about 25% to about 57%. No tendencies in diel haul-out activity were revealed.     

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     

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.     

DIET OF RINGED SEALS (PHOCA HISPIDA) ON THE EAST AND WEST SIDES OF THE NORTH WATER POLYNYA, NORTHERN BAFFIN BAY

In conjunction with the International North Water Polynya Study (NOW) in northern Baffin Bay, we examined the diets of ringed seals ( Phoca hispida )¹ on the west (Grise Fiord, Nunavut) and east (Qaanaaq, Greenland) sides of the polynya, using conventional stomach content analysis, as well as inferences from stable isotope ratios in seal muscle. Between May and July 1998, stomach and muscle tissue samples were collected from 99 ringed seals taken near Grise Fiord and 100 taken near Qaanaaq. The amphipod Themisto libellula was the dominant prey type in the diet of immature ringed seals from Grise Fiord, whereas arctic cod ( Boreogadus saida ) and polar cod ( Arctogadus glacialis ) predominated in the diet of adults. Both immature and adult seals collected near Qaanaaq fed predominantly on arctic cod. Overall, seals collected near Grise Fiord had significantly higher δ¹³C values than those collected near Qaanaaq ( P < 0.001), but there was no statistical separation in δ¹⁵N values between the two samples ( P = 0.06). Differences in diets of ringed seals from the east and west sides of the North Water Polynya may be due to differences in prey distribution and/or differences in biological productivity and fish biomass within the polynya.     

SPRING HAUL-OUT BEHAVIOR OF RINGED SEALS (PUSA HISPIDA) IN KONGSFJORDEN, SVALBARD

Haul‐out behavior of ringed seals (Pusa hispida) was investigated during the spring molting period of 2003 (May–July) in Kongsfjorden, Svalbard, Norway. Hourly counts were conducted on the land‐fast ice in six spatially defined sectors in the inner fjord, from an elevated land‐based vantage point from early May through until the ice began to break up in June, from 0600 to 2200 daily (total counts n= 478). Concomitantly, measurements were made of a variety of weather parameters. Multiple regression analyses revealed that time of day (P < 0.001) and date (P < 0.001) significantly affected the number of ringed seals hauled out on the ice surface. Other factors influencing the number of seals counted on the ice were air temperature (P= 0.011) and wind speed (P < 0.001). Daily peaks occurred in the early afternoon between 1300 and 1400 and the seasonal high (n= 385) was registered during the first week in June, after which the number of seals on the ice in the fjord declined. In addition to the visual counts, 24 ringed seals were equipped with VHF transmitters, and the haul‐out behavior of individuals was monitored from May through July via an automatic recording station. The VHF‐tagged seals exhibited the same diurnal pattern seen in the total counts, with haul‐out most frequent from 1300 to 1400. Pups exhibited short and frequent haul‐outs, whereas longer haul‐out periods were seen in the older age classes; adult females had the greatest number of haul‐out periods that exceeded 24 h. The seasonal peak of haul‐out for the tagged seals preceded the peak seasonal counts by approximately 3 wk. This may reflect significant out‐ and influx of seals from and to the area, a phenomenon warranting further attention because of its implications for assessment studies.     

Temporal variation in distribution and density of ice-obligated seals in western Hudson Bay, Canada

Recent unidirectional climatic trends and changes in top predator population ecology suggest that long-term modifications may be happening in Hudson Bay, Canada. Effects of such changes on ice-obligated seal populations are expected but long-term studies are required to differentiate climate-induced changes from natural variation. We conducted strip-transect surveys in late spring in 1995–1997, 1999–2000 and 2007–2008 to estimate distribution, density and abundance of ice-obligated ringed (Phoca hispida) and bearded (Erignathus barbatus) seals in western Hudson Bay. When hauled out, ringed seals preferred land-fast and consolidated pack ice, whereas bearded seals preferred unconsolidated pack ice. Bearded and ringed seal density estimates varied from 0.0036 to 0.0229 seals/km² of ice and from 0.46 to 1.60 seals/km² of ice, respectively. Strong inter-annual variations were recorded in the abundance estimates of both species, with the largest abundance estimates in 1995 (104,162 and 1,494 ringed and bearded seals, respectively) and the lowest in 2008 for ringed seals (33,701) and 1997 for bearded seals (278). A sine function best described seal density estimates in western Hudson Bay and suggested a decadal cycle. Previous studies that reported low ringed seal demographic parameters in the 1990s and a recovery in the 2000s supported our interpretation of the survey results. We discuss our results in the context of climate warming and suggest that a long-term decline in ice-obligated seal density estimates may overlay a possible natural decadal cycle.     

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.     

Habitat use by harbour seals (Phoca vitulina) in a seasonally ice-covered region, the western Hudson Bay

Over the last few decades, the period of ice cover in Hudson Bay has decreased, owing to climate warming, with breakup occurring approximately 3 weeks earlier than it did 30 years ago. The trend towards lengthening of the open water season has led to speculation that ringed seal numbers would decline, but then harbour seals might become numerous enough to replace ringed seals in the diet of polar bears. The movement patterns of 18 harbour seals equipped with satellite-linked transmitters in the Churchill River estuary (western Hudson Bay) were examined, as well as the dive behaviour of 11 of these seals. During the ice-free period, seals followed a general central place-foraging strategy, making repeated trips between their haul-out site in the Churchill River estuary and nearshore areas (<20 km) near the river mouth and estuary. Seal behaviour changed significantly as ice started to form along the coast of western Hudson Bay: animals remained significantly farther from the Churchill River haul-out site and from the coast and performed longer and deeper dives. However, throughout the entire tracking period, whether ice was present or not, all animals restricted their movements to a narrow band of shallow coastal waters (<50 m depth) along a 600-km stretch of the western Hudson Bay coastline, centred on the Churchill River estuary haul-out site. This natural self-limitation to nearshore shallow waters could restrict the potential for the population to increase in size and replace ringed seals as a primary energy resource for polar bears.     

The origin and genetic relationships of the Baikal seal, Phoca sibirica, by restriction analysis of mitochondrial DNA

The origin and genetic relationships of the Baikal seal, Phoca sibirica, were studied by restriction fragment length polymorphism analysis of mitochondrial DNA (mtDNA). Using 17 different six-base recognition restriction endonucleases, we examined 98 Baikal seals, and two other related species, the ringed seal, P. hispida, (n=87), and the Caspian seal, P. caspica, (n=94). Analysis revealed the existence of 87 mtDNA haplotypes in the total of 279 specimens. The haplotypes of each species were divided into different clusters on a dendrogram obtained by UPGMA based on haplotype frequency and mtDNA base substitution. No common haplotypes were found among the species examined. The Baikal seal is much more closely related to the ringed seal than the Caspian seal. The amount of divergence suggested that an ancestor of the Baikal seal came down to the lake approximately 0.4 million years ago as was previously indicated by paleontological studies. The seals examined here showed lower variabilities.     

Feeding ecology of phocid seals and some walrus in the Alaskan and Canadian Arctic as determined by stomach contents and stable isotope analysis

Feeding habits of ringed (Phoca hispida), bearded (Erignathus barbatus), spotted (Phoca largha) and ribbon (Phoca fasciata) seals and walrus (Odobenus rosmarus) were studied using stomach contents and stable carbon and nitrogen isotopes. Bearded seals fed benthically, primarily crustaceans and mollusks. Both zooplankton and fish were significant prey for ringed seals, while fish was principal spotted seal prey. Few gastric contents were available from ribbon seals. δ¹⁵N was positively correlated with age in ribbon seals and δ¹³C was positively correlated with age in ringed and ribbon seals. δ¹⁵N was highest in spotted seals, in agreement with their fish-dominated diet. δ¹⁵N was not different between Alaskan-harvested ringed and bearded seals, while δ¹⁵N was lowest in ribbon seals and walrus. Carbon-13 was most enriched in bearded seals and walrus reflecting benthic ecosystem use. Canadian ringed seals were depleted in ¹³C compared to Alaskan pinnipeds, likely because of Beaufort Sea versus Chukchi and Bering seas influence.     

Habitat selection by ice-associated pinnipeds near St. Lawrence Island,Alaska in March 2001

Aerial surveys of ice-associated pinnipeds were conducted south of St. Lawrence Island in March 2001. The observed distributions of bearded seals (Erignathus barbatus), ribbon seals (Phoca fasciata), ringed seals (P. hispida), spotted seals (P. largha), and walruses (Odobenus rosmarus) were compared to the distributions of ice habitat types and benthic communities. Randomization tests were used to investigate habitat selection for each species. Both ringed seals and walruses preferred large ice floes (>48 m in diameter) that were common in the interior ice pack. Spotted seals favored smaller ice floes (<20 m in diameter) common near the ice edge, and bearded seals avoided large floes and preferred transitional habitat between small and large floes. Ringed seals also seemed to prefer areas with greater than 90% sea ice coverage, and bearded seals preferred 70–90% sea ice coverage while avoiding areas with greater than 90% coverage. All species, except spotted seals, were seen most frequently in a region of high benthic biomass, and randomization tests suggested that bearded seals actively selected that region.     

Polar bear predatory behaviour reveals seascape distribution of ringed seal lairs

Ringed seal (Pusa hispida) breeding distribution has been extensively studied across near-shore habitats, but has received limited attention at a seascape scale due to the difficulty in accessing offshore sea ice environments. Employing highly visible predation attempts by polar bears (Ursus maritimus) on ringed seals in subnivean lairs observed by helicopter, the spatial relationship between predatory behaviour and ringed seal breeding habitat was examined. Resource selection functions were used to determine the relative probability of predation attempts on ringed seals in lairs as a function of habitat during a period of low ringed seal natality (2004–2006). Ringed seal pup kill locations were compared between years of low (2003–2006) and high (2007–2011) natality to assess the effect of reproductive output on habitat use. During years of low natality, polar bear hunting attempts were more likely in near-shore fast ice, and pup kills were observed predominately in fast ice (fast = 65 %, pack = 29 %, P = 0.002) at a median distance of 36 km from shore. In years of high natality, pup kills were observed farther from shore (median = 46 km, P = 0.03), and there was no difference in the proportion of observations in fast ice and pack ice (fast = 43 %, pack = 52 %, P = 0.29). These results suggest that the facultative use of adjacent offshore pack ice by breeding ringed seals may be influenced by natality. This study illustrates how documenting the behaviour of a predator can facilitate insight into the distribution of a cryptic prey.     

A survey of the Van Mijen fiord, Svalbard, as habitat for ringed seals, Phoca hispida

The ringed seal breeding habitat of the Van Mijen fiord, Svalbard, Norway, was studied from 30 March to 22 May 1986. A Siberian husky dog was used to detect ringed seal birth lairs and breathing holes. Fifteen percent of the total fiord areas was sampled. We estimated the densitites of birth lairs and breathing holes to be 0.04 km^-2 and 1.30 km^-2, respectively. The Van Mijen fiord was a poor ringed seal breeding habitat compared with breeding habitats investigated both in Canada and other parts of Svalbard. The main reason is probably shallow snow depth and lack of structures such as pressure ridges around which enough snow could accumulate for the ringed seal females to be able to dig out their lairs. The number of seals inhabiting the area during the breeding period 1986 was approximately 125 animals.