The eastern Bering is a productive high-latitude ecosystem characterized by high interannual variability in physical environmental conditions that impact biological communities. We investigated how the diet composition of northern fur seals (Callorhinus ursinus) breeding on the Pribilof Islands was influenced by this variation, focusing on water temperatures (surface and bottom) and an index of walleye pollock abundance within foraging areas. We also explored whether interannual variation in diet composition influenced fur seal pup mortality rates or body mass. The frequency of occurrence (FO) of all eight fur seal prey groups detected from hard parts analysis of samples collected from 1987 to 2012 was affected by interannual variation in at least one of the three environmental variables. Pollock was the predominant prey group across the study years, highlighting the importance of this species to Pribilof Island fur seals. Not only was pollock consumed more frequently as it became more abundant within fur seal foraging areas, but its relative abundance also affected how frequently other prey groups were consumed. A considerable amount of variation in FO of almost all prey groups was explained by year effects, suggesting that water temperatures alone were not sufficiently capturing the influences of regional and local physical environmental conditions on prey availability for fur seals. The summed FO of non-pollock prey groups had a small but detectable effect on the mass of male pups, indicating that the availability of prey groups beyond just pollock is somewhat beneficial for female northern fur seals early in lactation. Our results suggest that projected environmental changes in the eastern Bering Sea are likely to influence fur seal diets, but predicting the magnitude and direction of such changes is hampered until the underlying drivers of the observed temporal trends are better resolved.
Understanding the abiotic and biotic drivers of species distribution is critical for climate-informed ecosystem management. We aimed to understand habitat selection of northern fur seals in the eastern Bering Sea, a declining population that is also a key predator of walleye pollock, the target species for the largest U.S. commercial fishery. We developed species distribution models using random forest models by combining satellite telemetry data from lactating female fur seals tagged at different rookery complexes on the Pribilof Islands in the eastern Bering Sea with regional ocean model simulations. We explored how data aggregation at two spatial scales (Pribilof-wide and complex-specific) impacted model performance and predicted distributions. Spatial predictions under hindcasted (1992–2018) and projected (2050–2059) physical and biological conditions were used to identify areas of core habitat, overlap with commercial fishery catches, and potential changes in future habitat suitability. The most important environmental predictor variables across all models were bathymetry, bottom temperature, and surface temperature. The Pribilof-wide model both under- and overrepresented the importance of specific areas, while complex-specific models exhibited considerable variability in transferability performance. The majority of core habitat occurred on the continental shelf in areas that overlapped with commercial catches of walleye pollock during the “B” season (June – October), with an average of 76
Food availability is a key concern for the conservation of marine top predators, particularly during a time when they face a rapidly changing environment and continued pressure from commercial fishing activities. Northern fur seals ( Callorhinus ursinus ) breeding on the Pribilof Islands in the eastern Bering Sea have experienced an unexplained population decline since the late-1990s. Dietary overlap with a large U.S. fishery for walleye pollock ( Gadus chalcogrammus ) in combination with changes in maternal foraging behavior and pup growth has led to the hypothesis that food limitation may be contributing to the population decline. We developed age- and sex-specific bioenergetic models to estimate fur seal energy intake from May–December in six target years, which were combined with diet data to quantify prey consumption. There was considerable sex- and age-specific variation in energy intake because of differences in body size, energetic costs, and behavior; net energy intake was lowest for juveniles (18.9 MJ sea-day –1 , 1,409.4 MJ season –1 ) and highest for adult males (66.0 MJ sea-day –1 , 7,651.7 MJ season –1 ). Population-level prey consumption ranged from 255,232 t (222,159 – 350,755 t, 95% CI) in 2006 to 500,039 t (453,720 – 555,205 t) in 1996, with pollock comprising between 41.4 and 76.5% of this biomass. Interannual variation in size-specific pollock consumption appeared largely driven by the availability of juvenile fish, with up to 81.6% of pollock biomass coming from mature pollock in years of poor age-1 recruitment. Relationships among metabolic rates, trip durations, pup growth rates, and energy intake of lactating females suggest the most feasible mechanism to increase pup growth rates is by increasing foraging efficiency through reductions in maternal foraging effort, which is unlikely to occur without increases in localized prey density. By quantifying year-specific fur seal consumption of pollock, our study provides a pathway to incorporate fur seals into multispecies pollock stock assessment models, which is critical for fur seal and fishery management given they were a significant source of mortality for both juvenile and mature pollock.
Climate change-driven alterations in Arctic environments can influence habitat availability, species distributions and interactions, and the breeding, foraging, and health of marine mammals. Phocine distemper virus (PDV), which has caused extensive mortality in Atlantic seals, was confirmed in sea otters in the North Pacific Ocean in 2004, raising the question of whether reductions in sea ice could increase contact between Arctic and sub-Arctic marine mammals and lead to viral transmission across the Arctic Ocean. Using data on PDV exposure and infection and animal movement in sympatric seal, sea lion, and sea otter species sampled in the North Pacific Ocean from 2001–2016, we investigated the timing of PDV introduction, risk factors associated with PDV emergence, and patterns of transmission following introduction. We identified widespread exposure to and infection with PDV across the North Pacific Ocean beginning in 2003 with a second peak of PDV exposure and infection in 2009; viral transmission across sympatric marine mammal species; and association of PDV exposure and infection with reductions in Arctic sea ice extent. Peaks of PDV exposure and infection following 2003 may reflect additional viral introductions among the diverse marine mammals in the North Pacific Ocean linked to change in Arctic sea ice extent.
In species exhibiting differential migration by sex and age, understanding what differences exist, and the adaptive reasons for these differences is critical for determining how demographic groups will respond to environmental variability and anthropogenic perturbations. We used satellite-telemetered movement and diving data to investigate differential migration and its ontogeny in a highly migratory North Pacific Ocean predator, the northern fur seal ( Callorhinus ursinus ; NFS), with a focus on understudied juvenile (1- to 2-year-old) animals. We instrumented 71 juvenile NFS in two years (2006–07 and 2007–08) at three major North American breeding sites and compared their migratory strategies with pups and adults. Although sexual dimorphism is strong in adult NFS, only weak differences in body mass between sexes were found in juveniles, which had similar body mass to pups (~3–4 months). However, unlike widely-dispersed pups, juvenile male and female NFS dispersed in different directions, and used different habitats characterized by distinct hydrography and prey assemblages during migration, similar to breeding adults. Juvenile diving behavior differed only modestly among habitats and between sexes, consistent with weak differences in body mass. Evidence of habitat sexual segregation by juvenile NFS contradicts previous hypotheses that physiological differences predominantly drive the ontogeny of differential migration.
Environmental microplastics are widely documented in marine life and bioaccumulation may present risks to marine predators. Investigations of microplastics in marine mammals are increasing, though none have examined animals routinely consumed by humans. Here, we investigate microplastic exposure in the northern fur seal (Callorhinus ursinus), a species consumed by humans, using fecal material. We examined 44 feces (scat) at sites encompassing the seals' eastern Pacific range. Multiple contamination control measures were implemented, including field and laboratory controls. Fragments were the most common microplastic recovered, in 55% (24/44) of scat and no controls (range 1 to 86 fragments/scat, mean 16.6, sd 19.1). Microplastic fibers were recovered from 41% of scats (18/44), though some controls contained fibers confounding fiber results. Fecal analysis documented northern fur seal exposure to microplastics throughout their eastern Pacific range.
by M. M. Muto, V. T. Helker, R. P. Angliss, B. A. Allen, P. L. Boveng, J. M. Breiwick, M. F. Cameron, P. J. Clapham, S. P. Dahle, M. E. Dahlheim, B. S. Fadely, M. C. Ferguson, L. W. Fritz, R. C. Hobbs, Y. V. Ivashchenko, A. S. Kennedy, J. M. London, S. A. Mizroch, R. R. Ream, E. L. Richmond, K. E. W. Shelden, R. G. Towell, P. R. Wade, J. M. Waite, and A. N. Zerbini Alaska Marine Mammal Stock Assessments, 2016
STOCK DEFINITION AND GEOGRAPHIC RANGE Harbor seals inhabit coastal and estuarine waters off Baja California, north along the western coasts of the United States, British Columbia, and Southeast Alaska, west through the Gulf of Alaska and Aleutian Islands, and in the Bering Sea north to Cape Newenham and the Pribilof Islands. They haul out on rocks, reefs, beaches, and drifting glacial ice and feed in marine, estuarine, and occasionally fresh waters. Harbor seals generally are non-migratory, with local movements associated with such factors as tides, weather, season, food availability, and reproduction (Scheffer and Slipp 1944; Fisher 1952; Bigg 1969, 1981; Hastings et al. 2004). The results of past and recent satellitetagging studies in Southeast Alaska, Prince William Sound, Kodiak Island, and Cook Inlet are also consistent with the conclusion that harbor seals are non-migratory (Swain et al. 1996, Lowry et al. 2001, Small et al. 2003, Boveng et al. 2012). However, some long-distance movements of tagged animals in Alaska have been recorded (Pitcher and McAllister 1981, Lowry et al. 2001, Small et al. 2003, Womble 2012, Womble and Gende 2013). Strong fidelity of individuals for haul-out sites during the breeding season has been documented in several populations (Härkönen and Harding 2001), including some harbor seal stocks in Alaska such as South Kodiak Island, Prince William Sound, Glacier Bay/Icy Strait, and Cook Inlet (Pitcher and McAllister 1981, Small et al. 2005, Boveng et al. 2012, Womble 2012, Womble and Gende 2013). Local or regional trends in harbor seal numbers have been monitored at various time intervals since the 1970s, revealing diverse spatial patterns in apparent population trends. Where declines have been observed, they seem Figure 1. Approximate distribution of harbor seals in Alaska waters (shaded coastline area).
We developed models to predict foraging habitat of adult female northern fur seals (Callorhinus ursinus) using stable carbon (δ13C) and nitrogen (δ15N) isotope values from plasma and red blood cells. Binomial generalized linear mixed models were developed using blood isotope samples collected from 35 adult female fur seals on three breeding colonies in Alaska during July-October 2006. Satellite location and dive data were used to define habitat use in terms of the proportion of time spent or dives made in different oceanographic/bathymetric domains. For both plasma and red blood cells, the models accurately predicted habitat use for animals that foraged exclusively off or on the continental shelf. The models did not perform as well in predicting habitat use for animals that foraged in both on- and off-shelf habitat; however, sample sizes for these animals were small. Concurrently collected scat, fatty acid, and dive data confirmed that the foraging differences predicted by isotopes were associated with diet differences. Stable isotope samples, dive data, and GPS location data collected from an additional 15 females during August-October 2008 validated the effective use of the models across years. Little within year variation in habitat use was indicated from the comparison between stable isotope values from plasma (representing 1-2 weeks) and red blood cells (representing the prior few months). Constructing predictive models using stable isotopes provides an effective means to assess habitat use at the population level, is inexpensive, and can be applied to other marine predators.
Behavioral responses by top marine predators to oceanographic features such as eddies, river plumes, storms, and coastal topography suggest that biophysical interactions in these zones affect predators' prey, foraging behaviors, and potentially fitness. However, examining these pathways is challenged by the obstacles inherent in obtaining simultaneous observations of surface and subsurface environmental fields and predator behavior. In this study, migratory movements and, in some cases, diving behavior of 40 adult female northern fur seals (NFS; Callorhinus ursinus) were quantified across their range and compared to remotely-sensed environmental data in the Gulf of Alaska and California Current ecosystems, with a particular focus off the coast of Washington State (USA)--a known foraging ground for adult female NFS and where autonomous glider sampling allowed opportunistic comparison of seal behavior to subsurface biophysical measurements. The results show that in these ecosystems, adult female habitat utilization was concentrated near prominent coastal topographic, riverine, or inlet features and within 200 km of the continental shelf break. Seal dive depths, in most ecosystems, were moderated by surface light level (solar or lunar), mirroring known behaviors of diel vertically-migrating prey. However, seal dives differed in the California Current ecosystem due to a shift to more daytime diving concentrated at or below the surface mixed layer base. Seal movement models indicate behavioral responses to season, ecosystem, and surface wind speeds; individuals also responded to mesoscale eddies, jets, and the Columbia River plume. Foraging within small scale surface features is consistent with utilization of the inner coastal transition zone and habitats near coastal capes, which are known eddy and filament generation sites. These results contribute to our knowledge of NFS migratory patterns by demonstrating surface and subsurface behavioral responses to a spatially and temporally dynamic ocean environment, thus reflecting its influence on associated NFS prey species.
For central place foragers, forming colonies can lead to extensive competition for prey around breeding areas and a zone of local prey depletion. As populations grow, this area of reduced prey can expand impacting foraging success and forcing animals to alter foraging behaviour. Here, we examine a population of marine predators, the northern fur seal (Callorhinus ursinus), which colonized a recently formed volcanic island, and assess changes in foraging behaviour associated with increasing population density. Specifically, we measured pup production and adult foraging behaviour over a 15-year period, during which the population increased 4-fold. Using measures of at-sea movements and dive behaviour, we found clear evidence that as the population expanded, animals were required to allot increasing effort to obtain resources. These changes in behaviour included longer duration foraging trips, farther distances travelled, a larger foraging range surrounding the island and deeper maximum dives. Our results suggest that as the northern fur seal population increased, local prey resources were depleted as a result of increased intraspecific competition. In addition, the recent slowing of population growth indicates that this population may be approaching carrying capacity just 31 years after a natural colonization event. Our study offers insight into the dynamics of population growth and impacts of increasing population density on a large marine predator. Such data could be vital for understanding future population fluctuations that occur in response to the dynamic environment, as natural and anthropogenic factors continue to modify marine habitats.
Central place foraging by colonial breeders can lead to depleted prey resources around breeding areas. Segregation of foraging areas both within and between large colonies may act as a mechanism to reduce competition for prey resulting in increased foraging success. We reassessed horizontal (spatial) foraging habitat segregation for northern fur seals (Callorhinus ursinus (L., 1758)) within and between colonies on the Pribilof Islands, Alaska (St. Paul and St. George islands), after the population declined by approximately 40%. Additionally, we examined vertical habitat segregation, where foraging ranges overlapped, and describe the influence of different foraging habitats on northern fur seal dive behavior. Spatial habitat segregation in northern fur seal foraging areas occurred between islands but was variable within islands, which is similar to the pattern previously described. There was no evidence for vertical habitat segregation when fur seals from different rookeries on St. George Island used the same foraging area. Additionally, fur seals from St. Paul Island rookeries that foraged in similar habitats showed fewer differences in dive behavior, indicating that foraging habitat plays a significant role in shaping dive behavior. The use of multiple foraging strategies within the Pribilof Island fur seal population could indicate that a complex management and conservation strategy may be necessary to stop the continuing decline of this population.
We consider a model-based clustering approach to examining abundance trends in a metapopulation. When examining trends for an animal population with management goals in mind one is often interested in those segments of the population that behave similarly to one another with respect to abundance. Our proposed trend analysis incorporates a clustering method that is an extension of the classic Chinese Restaurant Process, and the associated Dirichlet process prior, which allows for inclusion of distance covariates between sites. This approach has two main benefits: (1) nonparametric spatial association of trends and (2) reduced dimension of the spatio-temporal trend process. We present a transdimensional Gibbs sampler for making Bayesian inference that is efficient in the sense that all of the full conditionals can be directly sampled from save one. To demonstrate the proposed method we examine long term trends in northern fur seal pup production at 19 rookeries in the Pribilof Islands, Alaska. There was strong evidence that clustering of similar year-to-year deviation from linear trends was associated with whether rookeries were located on the same island. Clustering of local linear trends did not seem to be strongly associated with any of the distance covariates. In the fur seal trends analysis an overwhelming proportion of the MCMC iterations produced a 73–79 % reduction in the dimension of the spatio-temporal trend process, depending on the number of cluster groups.
During the breeding season northern fur seals ( Callorhinus ursinus ) congregate on the Pribilof Islands in large numbers creating the potential for intraspecific competition. Due to the declining trend in the Pribilof Islands population of fur seals, it is important to understand how prey resources are partitioned among the population. Fur seals exhibit a high degree of sexual dimorphism resulting in energetic differences among age and sex classes. Therefore, we hypothesized that subadult male and adult female fur seals would differ in the type and size of prey consumed. We examined the diets of subadult male (age 2–8; mean mass 28–176 kg) and adult female (age ≥ 3 yr; mean mass 13–50 kg) seals on St. Paul Island from 1992 to 2000. Prey remains found in fecal samples were compared using niche overlap indices. There was nearly complete dietary niche overlap between subadult male and adult female fur seals. Walleye pollock ( Theragra chalcogramma ), Pacific salmon ( Oncorhynchus spp.), Pacific herring ( Clupeia pallasi ), and cephalopods were common prey items found in the diets of both groups. We found differences in the size of pollock consumed and that geographic location of sample collection may be important in determining diet differences. Our results indicate high levels of dietary overlap among subadult male and adult female fur seals.