An apparent population decrease,or change in distribution,of Weddell seals along the Victoria Land coast

Ground counts during 1959–1968 compared with counts using high resolution (0.6 m²) satellite imagery during 2008–2012 indicated many fewer Weddell seals (Leptonychotes weddellii) at two major molting areas in the western Ross Sea: Edisto Inlet‐Moubray Bay, northern Victoria Land, and McMurdo Sound, southern Victoria Land. Breeding seals have largely disappeared from Edisto‐Moubray, though the breeding population in McMurdo Sound appears to have recovered from harvest in the 1960s. The timing of decline, or perhaps spreading (lower numbers of seals in more places), is unknown but appears unrelated to changes in sea ice conditions. We analyzed both historic and satellite‐derived ice data confirming a large expansion of pack ice mostly offshore of the Ross Sea, and not over the continental shelf (main Weddell seal habitat), and a thinning of fast ice along Victoria Land (conceivably beneficial to seals). Timing of fast ice presence and extent in coves and bays along Victoria Land, remains the same. The reduction in numbers is consistent with an altered food web, the reasons for which are complex. In the context of a recent industrial fishery targeting a seal prey species, a large‐scale seal monitoring program is required to increase understanding of seal population changes.     

Contrasting behavior between two populations of an ice‐obligate predator in East Antarctica

The Austral autumn–winter is a critical period for capital breeders such as Weddell seals that must optimize resource acquisition and storage to provision breeding in the subsequent spring. However, how Weddell seals find food in the winter months remains poorly documented. We equipped adult Weddell seals after their annual molt with satellite‐relayed data loggers at two sites in East Antarctica: Dumont D'Urville (n = 12, DDU) and Davis (n = 20). We used binomial generalized mixed‐effect models to investigate Weddell seals’ behavioral response (i.e., “hunting” vs. “transit”) to physical aspects of their environment (e.g., ice concentration). Weddell seal foraging was concentrated to within 5 km of a breathing hole, and they appear to move between holes as local food is depleted. There were regional differences in behavior so that seals at Davis traveled greater distances (three times more) and spent less time in hunting mode (half the time) than seals at DDU. Despite these differences, hunting dives at both locations were pelagic, concentrated in areas of high ice concentration, and over areas of complex bathymetry. There was also a seasonal change in diving behavior from transiting early in the season to more hunting during winter. Our observations suggest that Weddell seal foraging behavior is plastic and that they respond behaviorally to changes in their environment to maximize food acquisition and storage. Such plasticity is a hallmark of animals that live in very dynamic environments such as the high Antarctic where resources are unpredictable.     

Harbor seal population decline in the Aleutian Archipelago

Populations of Steller sea lions, northern fur seals, and northern sea otters declined substantially during recent decades in the Bering Sea and Aleutian Islands region, yet the population status of harbor seals has not been assessed adequately. We determined that counts obtained during skiff‐based surveys conducted in 1977–1982 represent the earliest estimate of harbor seal abundance throughout the Aleutian Islands. By comparing counts from 106 islands surveyed in 1977–1982 (8,601 seals) with counts from the same islands during a 1999 aerial survey (2,859 seals), we observed a 67% decline over the ～20‐yr period. Regionally, the largest decline of 86% was in the western Aleutians (n= 7 islands), followed by 66% in the central Aleutians (n= 64 islands), and 45% in the eastern Aleutians (n= 35 islands). Harbor seal counts decreased at the majority of islands in each region, the number of islands with >100 seals decreased ～70%, and the number of islands with no seals counted increased ～80%, indicating that harbor seal abundance throughout the Aleutian Islands was substantially lower in the late 1990s than in the 1970s and 1980s.     

Influence of time of day and month on Weddell seal haul-out patterns at the Vestfold Hills, Antarctica

A diurnal pattern in haul-out behaviour has been described for Weddell seals at McMurdo Sound, Antarctica, but regional and seasonal variations were previously unknown. Knowledge of activity patterns is important for standardising census methods and census data. This study quantified the diurnal pattern in haul-out behaviour of Weddell seals at the Vestfold Hills from October 1994 to March 1995. Sequential counts of seals on the ice showed that, between 0900 and 1930 hours, seal abundance differed up to 95%. Fewer seals were hauled out in the morning than in the afternoon. The maximum numbers of seals were hauled out at the warmest time of day. The diurnal cycle was less pronounced in the breeding season than in the moulting season. The findings indicated the importance of censusing Weddell seals after 1430 hours and before 1700 hours local time, especially in the moulting season. Correction factors are given for month and time of day. Received: 27 November 1996 / Accepted: 1 May 1997     

AGONISTIC BEHAVIOR IN HARBOR SEALS (PHOCA VITULINA) IN RELATION TO THE AVAILABILITY OF HAUL-OUT SPACE

The hypothesis that haul-out space is a contested resource among harbor seals (Phoca vitulina) was tested by comparing the number of agonistic interactions and the time devoted to them, at a haul-out with unlimited space and a haul-out with limited space, in Humboldt County, California. 798 agonistic interactions were recorded in 160 h of observation. The average number of agonistic interactions per seal per hour and the time spent on these interactions were significantly higher at the site with limited haul-our space (P < 0.05, n= 20). At this restricted site, the frequency of agonistic interactions was positively correlated with seal density (r= 0.61, P < 0.05, n= 20), as was time spent on agonistic interactions (r= 0.84, P < 0.05, n= 20). Size, sex, and age-class of individuals all influenced agonistic interactions. In 72.5% of displacements, smaller seals were displaced by larger ones. This may support claims that some harbor seal populations maintain a size-based dominance hierarchy. The results of this study support the hypothesis that there is competition among harbor seals for suitable haul-out space where this resource is limited. The outcome of these interactions may be predicted, at least in part, by theoretical models on pay-off asymmetries, resource holding power, and the prior residency effect.     

MONITORING THE TREND OF HARBOR SEALS IN PRINCE WILLIAM SOUND,ALASKA, AFTER THE EXXON VALDEZ OIL SPILL

We used aerial counts to monitor the trend in numbers of harbor seals, Phoca vitulina richardsi, in Prince William Sound, Alaska, following the 1989 Exxon Valdez oil spill. Repetitive counts were made at 25 haul-out sites during the annual molt period each year from 1990 through 1997. A generalized linear model indicated that time of day, date, and time relative to low tide significantly affected seal counts. When Poisson regression was used to adjust counts to a standardized set of survey conditions, results showed a highly significant decline of 4.6% per year. Unadjusted counts indicated a slight, but not statistically significant, decline in the number of seals. The number of harbor seals on the trend-count route in eastern and central PWS has been declining since at least 1984, with an overall population reduction of 63% through 1997. Programs to monitor long-term changes in animal population sizes should account for factors that can cause short-term variations in indices of abundance. The inclusion of such factors as covariates in models can improve the accuracy of monitoring programs.     

Predation on cephalopods by Weddell seals, <Emphasis Type="Italic">Leptonychotes weddellii</Emphasis>, at Hope Bay,Antarctic Peninsula

Cephalopods play a key role in marine environments as food resources for top predators such as marine mammals and seabirds. However, detailed information on their trophic relationships with Antarctic seals is scarce. The aim of the present study was to examine the cephalopod portion of the diet of adult and subadult Weddell seals, Leptonychotes weddellii, at Hope Bay, Antarctic Peninsula, through the analysis of scats collected during three consecutive summers (2003, 2004 and 2005). Cephalopods occurred in almost 45% of the 217 samples collected during the whole period of study. A total of 662 beaks (358 upper and 304 lower) were removed from scats containing cephalopod remains (n = 93). Octopods were largely dominant in comparison with teuthoids constituting in numerical abundance over 95% of the cephalopod prey. The octopod Pareledone turqueti was the most frequent and dominant prey species representing, respectively, 57.9 and 71.1% in numbers and biomass of cephalopods consumed. Species belonging to the group of papillated Pareledone were second in importance. The results were compared with information from previous dietary studies of L. weddellii at other localities of Antarctica. Based on the examination of the cephalopod prey taxa identified in this study, it is suggested that during the study period Weddell seal individuals foraged mainly on benthic prey resources close to the coast, in inshore waters where octopods were dominant in comparison with pelagic squid.     

Genetically effective population sizes of Antarctic seals estimated from nuclear genes

We analyzed eight nuclear microsatellite loci in three species of Antarctic seals; Weddell seal (Leptonychotes weddellii; mean N = 163), crabeater seal (Lobodon carcinophaga; 138) and Ross seal (Ommatophoca rossii; 35). We estimated genetic diversity (Θ) and effective population size (N _E) for each species. Autosomal microsatellite based N _E estimates were 151,200 for Weddell seals, 880,200 for crabeater seals, and 254,500 for Ross seals. We screened one X-linked microsatellite (Lw18), which yielded similar N _E estimates to the autosomal loci for all species except the Ross seals, where it was considerably larger (~103 times). Microsatellite N _E estimates were comparable with previously published N _E estimates from mitochondrial DNA, but both are substantially lower than direct estimates of population size in all species except the Ross seals. The ratio of maternally versus biparentally derived estimates of N _E for Ross seals was not consistent with the hypothesis that they are a polygynous species. We found no sign of a recent, sustained genetic bottleneck in any of the species.     

DEVELOPMENT AND APPLICATION OF A MASS-ESTIMATION METHOD FOR WEDDELL SEALS

Traditional methods of acquiring mass data limit the ability to collect large samples from across populations of some pinnipeds, or to sample without great disturbance to the animals. In order to collect substantial samples of mass data from the Weddell seal (Leptonychotes weddellii) population in Erebus Bay, Antarctica, we developed the equipment and methods for estimating the mass of Weddell seals using digital photographs. Resulting regression models predict the mass of adult female seals to within ±13.8% of estimated mass, and ±25.9% of estimated mass for pups. We show the protocols developed are repeatable and efficient enough to be applied to a large number of animals in a relatively short period of time and may be useful for studies of other marine mammals. We caution that prediction intervals exist around mass estimates and must be accounted for when estimates are applied to biological questions. In a limited application of the method, differences in mass transfer between experienced and inexperienced maternal females and their pups were detected when prediction error variance around mass estimates was explicitly included. Similar mass‐estimation methods may therefore be useful in consideration of biological questions requiring large samples of mass previously unattainable.     

Mark-resight estimates of seasonal variation in harbor seal abundance and site fidelity

Monitoring trends in abundance of pinnipeds typically involves counting seals at terrestrial haul-out sites during the breeding season. Counts of seals made at other times of the year are typically lower; however, it is often unknown whether this is because fewer animals are present or whether lower counts simply reflect a reduction in haul-out probability. Here we illustrate how photo-identification data from an individual-based study of harbor seals (Phoca vitulina) can be used to estimate seasonal variation in abundance and site fidelity. Monthly data collected over a two-year period were analyzed using a mark-recapture mark-resight model accounting for individuals transitioning between observable and unobservable states. Levels of site fidelity were high throughout the year and abundance estimates showed no seasonal pattern. This suggests that individual seals used haul-out sites to the same extent throughout the year, and that peaks in counts during the breeding season are a result of seasonal changes in haul-out probability. The results of this study have implications for understanding population sub-structuring, gene flow and disease spread.     

Summer diet of Weddell Seals (Leptonychotes weddelli) in the eastern and southern Weddell Sea,Antarctica

Summary Stomach and intestine samples from 21 adult Weddell seals were used to study the diet of these seals from the eastern and southern Weddell Sea coast from January to February 1983 and 1985. Fish occurred in all seals, squid in five, octopods in three and Euphausia crystallorophias in one seal. Pleuragramma antarcticum was the predominant fish in the diet, constituting 61.1% of otoliths in 1983 samples and 93.8% in 1985. Aethotaxis mitopteryx, Dissostichus mawsoni, unidentified Trematomus spp. and channichthyids were also recorded. Size and wet weight of P. antarcticum were calculated from uneroded otoliths, found in 6 seal stomachs with liquid food pulp, collected during early morning hours in 1985. Size distribution of P. antarcticum from individual seals was reasonably constant, ranging between 5.0 and 22.0 cm SL; adult fish from about 14.0 to 19.0 cm SL predominated. P. antarcticum in seals from the southern area had a larger median size (16.5 cm SL), than those from further east (15.5 cm SL). Calculated wet weights of all P. antarcticum from individual seal stomachs ranged between 4.7 and 16.9 kg the mean was 12.8 kg. Comparisons with net-hauls from the southern Gould Bay suggest that Weddell seals feed mainly in deeper water layers (>400 m) where adult P. antarcticum occur at higher densities.     

Variation of annual apparent survival and detection rates with age,year and individual identity in male Weddell seals (Leptonychotes weddellii) from long-term mark-recapture data

Exploring age- and sex-specific survival rates provides insight regarding population behavior and life-history trait evolution. However, our understanding of how age-specific patterns of survival, including actuarial senescence, compare between the sexes remains inadequate. Using 36 years of mark-recapture data for 7,516 male Weddell seals (Leptonychotes weddellii) born in Erebus Bay, Antarctica, we estimated age-specific annual survival rates using a hierarchical model for mark-recapture data in a Bayesian framework. Our male survival estimates were moderate for pups and yearlings, highest for 2-year-olds, and gradually declined with age thereafter such that the oldest animals observed had the lowest rates of any age. Reports of senescence in other wildlife populations of species with similar longevity occurred at older ages than those presented here. When compared to recently published estimates for reproductive Weddell seal females, we found that peak survival rates were similar (males: 0.94, 95% CI = 0.92–0.96; females: 0.92, 95% CI = 0.93–0.95), but survival rates at older ages were lower in males. Age-specific male Weddell seal survival rates varied across years and individuals, with greater variation occurring across years. Similar studies on a broad range of species are needed to contextualize these results for a better understanding of the variation in senescence patterns between the sexes of the same species, but our study adds information for a marine mammal species to a research topic dominated by avian and ungulate species.     

Transformation of metabolism with age and lifestyle in Antarctic seals: a case study of systems biology approach to cross-species microarray experiment

Background  

The metabolic transformation that changes Weddell seal pups born on land into aquatic animals is not only interesting for the study of general biology, but it also provides a model for the acquired and congenital muscle disorders which are associated with oxygen metabolism in skeletal muscle. However, the analysis of gene expression in seals is hampered by the lack of specific microarrays and the very limited annotation of known Weddell seal (Leptonychotes weddellii) genes.     

Spatial utilisation of fast-ice by Weddell seals Leptonychotes weddelli during winter

This study describes the distribution of Weddell seals Leptonychotes weddelli in winter (May–September 1999) at the Vestfold Hills, in Prydz Bay, East Antarctica. Specifically, we describe the spatial extent of haul-out sites in shore–fast sea-ice, commonly referred to as fast-ice. As winter progressed, and the fast-ice grew thick (ca 2 m), most of the inshore holes closed over, and the seals' distribution became restricted to ocean areas beyond land and islands. Using observations from the end of winter only, we fitted Generalised Additive Models (GAMs) to generate resource selection functions, which are models that yield values proportional to the probability of use. The models showed that seal distribution was defined mainly by distance to ice-edge and distance to land. Distance to ice-bergs was also selected for models of some regions. We present the results as maps of the fitted probability of seal presence, predicted by the binomial GAM for offshore regions, both with and without autocorrelation terms. The maps illustrate the expected distribution encompassing most of the observed distribution. On this basis, we hypothesise that propensity for the fast-ice to crack is the major determinant of Weddell seal distribution in winter. Proximity to open water and pack-ice habitats could also influence the distribution of haul-out sites in fast-ice areas. This is the first quantitative study of Weddell seal distribution in winter. Potential for regional variation is discussed.     

The World’s Northernmost Harbour Seal Population–How Many Are There?

This study presents the first abundance estimate for the world’s northernmost harbour seal (Phoca vitulina) population, which resides in Svalbard, Norway, based on three digital stereoscopic photographic surveys conducted in 2009 and 2010. The counts from these high resolution 3D images were combined with a novel method for estimating correction factors for animals that were in the water at the time of the surveys, in which extensive behavioural data from radio-tagged harbour seals were used together with age distribution data to estimate the proportion of seals of various age and sex classes hauled out at the times of the surveys. To detect possible seasonal shifts in age distribution between surveys, lengths of hauled out seals were measured from the stereoscopic images. No body-length differences were detected between the surveys; but, this may be due to a high degree of sexual dimorphism exhibited in this population. Applying the modelled correction factors, a total of 1888 (95% CI: 1660–3023), 1742 (1381–3549) and 1812 (1656–4418) harbour seals were estimated for the surveys flown on 01 August 2009, 01 August 2010 and 19 August 2010, respectively. The similarity between the three survey estimates (despite significant differences in the number of animals actually counted on the photos from each survey effort) suggests that the variation in numbers of hauled out seals is reasonably accurately adjusted for by the haul-out probability model. The low population size, the limited spatial distribution of the population and its reduced genetic diversity make this population vulnerable to chance events, such as disease epidemics.     

Underwater calling rates of harp and Weddell seals as a function of hydrophone location

Stereo recordings of harp seal (Pagophilus groenlandicus) and Weddell seal (Leptonychotes weddellii) underwater vocalizations were made near breeding groups using separations between two hydrophones of 159–339 m. Within a large herd, harp seal call numbers varied slightly between channels. Counts of Weddell seal calls were higher near a small herd on the ice than 159–180 m away. Repeat counts of harp seal calls by a single observer differed significantly. Source levels of Weddell seal calls varied and higher amplitude calls would be detected up to 2 orders of magnitude farther away than quieter calls. Hydrophone location (especially near small groups), observer variability and call source level differences will bias the use of monitoring underwater seal vocalizations to index locations, population size or underwater behaviours. Using hydrophone arrays and multiple observers may mitigate these problems. Accepted: 15 September 2000     

Dinucleotide microsatellite markers from the Antarctic seals and their use in other Pinnipeds

Twenty‐four microsatellite loci were isolated from three species of Antarctic seals (Subfamily Monachinae, Tribe Lobodontini). Eleven loci were cloned from Weddell seal, Leptonychotes weddellii, seven from leopard seal, Hydrurga leptonyx, and six from crabeater seal, Lobodon carcinophagus. Variability was assessed in Weddell seals collected in McMurdo Sound, leopard seals from Bird Island, South Georgia, and crabeater seals sampled in the eastern Ross Sea. All loci were variable in the three species used for cloning and 22 of these loci amplified variable products in the Ross seal, Ommatophoca rossii. Cross‐species amplification was largely successful, with an average of 19 loci amplifying products in other phocids.     

Pleistocene population expansions of Antarctic seals

We sequenced a portion ( c . 475 bp) of the mitochondrial control region of three species of Antarctic phocid carnivores (Weddell seal, Leptonychotes weddellii , N  = 181; crabeater seal, Lobodon carcinophaga , N  = 143; and Ross seal, Ommatophoca rossii , N  = 41) that live seasonally or permanently in the fast ice and seasonal pack ice of the western Amundsen and Ross seas of western Antarctica. We resolved 251 haplotypes with a haplotype diversity of 0.98 to 0.99. Bayesian estimates of Θ from the program LAMARC ranged from 0.075 for Weddell seals to 0.576 for crabeater seals. We used the values of theta to estimate female effective population sizes ( N_EF ), which were 40 700 to 63 000 for Weddell seals, 44 400 to 97 800 for Ross seals, and 358 500 to 531 900 for crabeater seals. We used mismatch distributions to test for historical population size expansions. Weddell seals and crabeater seals had significant, unimodal mean pairwise difference distributions ( P  = 0.56 and 0.36, respectively), suggesting that their populations expanded suddenly around 731 000 years ago (Weddell seals) and around 1.6 million years ago (crabeater seals). Both of these expansions occurred during times of intensified glaciations and may have been fostered by expanding pack ice habitat.     

Influence of age and individual identity in the use of breeding colony habitat by male Weddell seals in Erebus Bay,Antarctica

Male–male contest behavior can contribute to spatial distributions of male pinnipeds during breeding seasons. To maximize breeding opportunities, the most competitive males would be expected to be surrounded by the highest numbers of reproductive-age females. As information regarding fine-scale spatial ecology of Weddell seals is lacking, we performed an exploratory study using kernel density analyses to evaluate age-specific habitat use of male Weddell seals in Erebus Bay, Antarctica. Additionally, we investigated the relationship between age and number of surrounding reproductive-age females using a competing set of regression models in a Bayesian framework that considered different functional forms of age while incorporating individual heterogeneity. As male adult Weddell seals aged, to at least 20 years, they were more likely to be found in areas associated with the greatest densities of reproductive-age females, but individual heterogeneity also influenced the number of reproductive-age female neighbors. The youngest males tended to haul out in offshore areas associated with better hunting, and older males tended to settle in more nearshore areas associated with more pup production. Our findings from this preliminary investigation indicate that male Weddell seal spatial behavior during the breeding season varies with age and individual and might be related to reproductive activity.     

Histological,histochemical, and ultrastructural investigations on the gastrointestinal system of antarctic seals: Weddell seal (Leptonychotes weddellii) and crabeater seal (Lobodon carcinophagus)

The morphology of the principal sections of the gastrointestinal system of two Antarctic seals with different dietary habits, namely, the Weddell seal (Leptonychotes weddellii) and the crabeater seal (Lobodon carcinophagus), has been investigated. Histologically examined by light microscopy, the tissue layers of the gastrointestinal tract of both seals are almost identical to those observed in most other mammals and no major differences in principle organization could be found between the two seal species. The ultrastructure of the gastric and intestinal epithelial cells has been examined and is also closely comparable to that of these cells in other mammals; however, Paneth cells have not been found in our material. In general, therefore, adaptations of the gastrointestinal tract to the aquatic environment or the diet are not obvious at the morphological levels of organization studied. Histochemical differences are found between the two closely related species; mucins of the surface epithelium in the stomach of Weddell seals are highly sulfated, while those in the crabeater seal are not. Mucous neck cells in Weddell seals contain acid mucosubstances, while those of crabeater seals contain neutral ones. Goblet cells in the small and large intestine in Weddell seals contain both neutral and acid mucosubstances. Both mucin types are detected in the crabeater seal; however, the mucins of the colon in the crabeater seal are more highly sulfated than those in the Weddell seal. The ratio of globet cells to enterocytes in the large intestine of crabeater seals is higher than that in Weddell seals. © 1995 Wiley-Liss, Inc.