Using DNA metabarcoding, we examined diet composition, prey resources, and saxitoxin (STX) and parasite exposure in Steller sea lions (SSL) in the Gulf of Alaska in July and October 2022–2023 using 251 scat samples and 45 seawater samples. Using 12S rRNA MiFish gene metabarcoding, diet items in Prince William Sound (PWS), Cook Inlet and near Kodiak ordered by prevalence were salmon, gadids, herring, flatfishes and capelin. Using cytochrome c oxidase subunit I (COI) gene metabarcoding, pink salmon were the most prevalent item, and two cephalopods (giant Pacific octopus and magister armhook squid) were important diet components. Diet was more diverse in October (female and male data) versus July (male only data) and in Kodiak versus other regions. Region*season variation in diet included (1) in July, dominance of pink salmon in PWS versus capelin and gadids in Kodiak, (2) magister armhook squid and pollock in PWS were replaced by giant Pacific octopus and Pacific cod in Kodiak and Cook Inlet; and (3) diverse flatfishes were observed in Cook Inlet. October male diets included more flatfishes, giant Pacific octopus and cods versus more sculpins, herring and salmon for females. Prevalent genera of parasitic intestinal worms included: Anisakis spp. roundworms (range 76.7–100% among region*seasons), Diphyllobothrium spp. tapeworms (34.5–68.0%), and Pseudoterranova spp. roundworms (19.4–50.0%). Regional fish DNA in July seawater mirrored regional July SSL diet (e.g., capelin and sculpin near Kodiak and pink salmon in PWS). STX prevalence (60.8%) and concentrations were higher than previously reported, suggesting SSL in Alaska may be chronically exposed to STX at low concentrations over a wide geographic region. Mean STX concentrations ranged from 9.28–53.32 ng/g among seasons/regions (maximum = 195 ng/g). STX was highest in PWS and Southeast Alaska (SEAK) in July, still below the seafood safety regulatory limit of 800 ng/g. Diversity of fish DNA in seawater was highest in Kodiak and SEAK. STX-producing dinoflagellates (Alexandrium spp.) were highest in July seawater in SEAK and PWS, where the STX in SSL scats were highest. DNA metabarcoding of diet and seawater, coupled with STX studies, allow better monitoring of ecosystem change affecting marine top predators and of the recovery of the endangered western SSL population.
Steller sea lions (SSL) are sentinels for monitoring environmental contaminants in remote areas of the Aleutian Islands, Alaska. Therefore, concentrations of several organochlorines (OCs) were measured in blood from 123 SSL pups sampled from 3 regions; the western Aleutian Islands (WAI), central Aleutian Islands (CAI), and the central Gulf of Alaska. Blood, blubber, and milk from 12 adult female SSL from WAI, CAI and southeast Alaska also were analyzed. Findings included the following. SSL pups had higher concentrations of some OCs and mercury (Hg) on rookeries in the WAI than those more easterly. Pups had significantly higher blood concentrations of many OC classes than adult females sampled within the same region; some pups had PCB concentrations exceeding thresholds of concern (∑PCBs >8600 ng/g lw). ∑PCB concentration in pup whole blood was positively correlated with the trophic marker, δ15N within the regions sampled, along with two PCB congeners (PCB138 and PCB153). This suggests that the dams of pups with higher ∑PCBs, PCB138, and PCB153 concentrations were feeding on more predatory prey. Adult female blubber ∑DDT and hexachlorocyclohexane concentrations were also positively correlated with δ15N values. Several pups (mostly from WAI) had blood Hg concentrations and/or blood PCB concentrations (surrogate for overall OC exposures) of concern. The finding that WAI SSL pups have been exposed to multiple contaminants calls for future investigation of their cumulative exposure to a mixture of contaminants especially their transplacental and then transmammary exposure routes.
• Highest Steller sea lion pup lanugo mercury concentrations occur in Western Aleutian Islands • >25% of pups in Western Aleutian Islands had mercury concentrations above risk thresholds • Pups with highest mercury may be at risk of mercury-dependent selenium deficiency • δ 15 N and δ 13 C in pup lanugo reflected isotope values in vibrissa segments grown in late gestation
Mercury (Hg) can be neurotoxic to mammals and impact reproduction, whereas selenium (Se) is an important antioxidant known to ameliorate some adverse effects of Hg. Total Hg concentrations ([THg]) were measured in lanugo (pelage grown in utero) of 812 Steller sea lion (Eumetopias jubatus) pups across Alaska and Russia to assess fetal exposure during late gestation. The molar ratio of total Se to THg (TSe:THg) was determined in whole blood collected from 291 pups. Stable isotope ratios of carbon and nitrogen were measured in sections of vibrissae (whiskers, n = 498) and in lanugo (n = 480) of pups grown during late gestation to track diet variations among adult females that can drive Hg and Se exposure during this critical fetal development period. Lanugo [THg] ranged from 1.4 to 73.7 mu g/g dry weight with the lowest median [THg] in Southeast Alaska. Pups from the Western Aleutian Islands had higher median lanugo [THg] than pups from other metapopulations in Alaska. Over 25% of pups in the Western Aleutian Islands had [THg] above published risk thresholds (20 mu g/g) for other mammals. Whole blood molar TSe:THg was significantly lower in the Western Aleutian Islands and in some parts of the Central Aleutian Islands with higher molar ratios found in the Eastern Aleutian Islands and Central Gulf of Alaska. This suggests a limitation on potential protective functions of Se in the western regions with the highest relative [THg]. The Central Aleutian Island pups with [THg] over 20 mu g/g had higher delta N-15 ratios than pups with lower [THg] suggesting dams consuming higher trophic level prey is a key driver for Hg exposure. However, regional differences likely reflect variability in diet of the dam during gestation and in Hg food web dynamics between oceanic regimes east and west of key passes in the Aleutian Islands. (C) 2020 Published by Elsevier B.V.
Steller sea lions (Eumetopias jubatus) are composed of two genetically distinct metapopulations (an increasing "eastern" and a reduced and endangered "western" population, or stock for management purposes in U.S. waters) that are only recently mixing at new rookeries in northern Southeast Alaska, east of the current stock boundary. We used mark-recapture models and 18 years of resighting data of over 3,500 individuals marked at the new rookeries and at neighboring long-established rookeries in both populations to examine morphology, survival, and movement patterns of pups born at new rookeries based on whether they had mitochondrial DNA haplotypes from the western or eastern population (mtW or mtE); examine survival effects of dispersal to the Eastern Stock region for animals born in the Western Stock region; and estimate minimum proportions of animals with western genetic material in regions within Southeast Alaska. Pups born at new rookeries with mtW had similar mass, but reduced body condition and first-year survival (approximately -10%) compared to pups with mtE. mtE pups ranged more widely than mtW pups, including more to the sheltered waters of Southeast Alaska's Inside Passage. Fitness benefits for western-born females that dispersed to Southeast Alaska were observed as higher female survival (+0.127, +0.099, and +0.032 at ages 1, 2, and 3+) and higher survival of their female offspring to breeding age (+0.15) compared to females that remained west of the boundary. We estimated that a minimum of 38% and 13% of animals in the North Outer Coast-Glacier Bay and Lynn Canal-Frederick Sound regions in Southeast Alaska, respectively, carry genetic information unique to the western population. Despite fitness benefits to western females that dispersed east, asymmetric dispersal costs or other genetic or maternal effects may limit the growth of the western genetic lineage at the new rookeries, and these factors require further study.
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.
Steller sea lions Eumetopias jubatus in the Glacier Bay region of northern Southeast Alaska experience greater survival and more rapid population growth than sea lions elsewhere in this region. To better understand demographics of sea lions in the region, and to describe the origins and behavior of sea lions and relate these descriptions to previous studies, we studied genetic origins, residency, foraging range, diving behavior, and dispersal of immature sea lions (≤24 mo of age) captured in Glacier Bay. Fifty-two percent of individuals had maternal origins in the distant (550 km) endangered western population rather than in the local recovered eastern population. During winter, 5 mo old pups, dependent on their dams for nutrition, remained within Glacier Bay, diving to shallow depths (≤108 m) mainly during daylight, whereas older (17 mo old) juveniles ranged more widely to areas of known seasonal prey aggregations, performing deep (≥241 m) nocturnal dives. Both pups and juveniles remained within the northern portion of Southeast Alaska, in contrast to farther-ranging pup and juvenile sea lions captured elsewhere in Southeast Alaska. Over the long term, females from Glacier Bay remained within this northern area through maturity and were sighted breeding in this area only. Restricted ranging patterns and natal and breeding philopatry by Steller sea lions of both eastern and western distinct population segment origin in the Glacier Bay region reveal that optimal foraging and breeding conditions likely prevail and help explain the recent colonization, increased survival, and rapid population growth of this species in the region.
We measured individual growth, looked for evidence of weaning, and examined the interaction of these changes with diving behavior in young-of-year Steller sea lion (Eumetopias jubatus (Schreber, 1776)) pups in Alaska, USA, during their first winter. Steller sea lions employ an income breeding strategy, in which females provision their young over an individually variable period of months to years. Thus, we set out to identify whether these young sea lions showed evidence of weaning during the challenging winter months, describe the nature of their growth during this time, and examine their behaviors in light of these changes. Between 2005 and 2008, we captured 71 pups during early winter and recaptured 33 of these pups in early spring. Mark–resight and stable nitrogen isotope ratios in vibrissae indicated most pups remained nutritionally dependent on adult females throughout the winter. All pups increased both mass and lipid mass, with half of growth contributed by lipid mass. Changes in behavior were not correlated with growth excepting a weak but significant effect on rate of vertical travel. This study demonstrated that capture–recapture of Steller sea lion pups is possible, provided seasonal timing, locations, and age classes are carefully considered for their likelihood of capture success.
Background: A utilization distribution quantifies the temporal and spatial probability of space use for individuals or populations. These patterns in movement arise from individuals' internal state and from their response to the external environment, and thus can provide insights for assessing factors associated with the management of threatened populations. The Western Distinct Population Segment of the Steller sea lion (Eumetopias jubatus) has declined to approximately 20% of levels encountered 40 years ago. At the height of the decline, juvenile survival appeared to be depressed and currently there is evidence that juvenile mortality due to predation may be constraining recovery in some regions. Therefore, our objectives were to identify what spaces are biologically important to juvenile Steller sea lions in the Kenai Fjords and Prince William Sound regions of the Gulf of Alaska. Methods: We examined geospatial location data from juvenile sea lions tagged between 2000 and 2014 (n = 84) and derived individual and pooled-population utilization distributions (UDs) from their movements. Core areas were defined from the UDs using an individual-based approach; this quantitatively confirmed that all individuals in our sample exhibited concentrated use within their home range (95% UD). Finally, we explored if variation in UD characteristics were associated with sex, season, age, or region. Results: We found evidence that individual juvenile home ranges were region and sex-specific, with males having larger home ranges on average. Core space characteristics were also sex-specific, and exhibited seasonal patterns of reduced size, increased proximity to haulouts, and increased intensity of use in the summer, but only in the Kenai Fjords-Gulf of Alaska region. Conclusions: This study highlights the areas of biological importance during this vulnerable life history stage, and the demographic, seasonal, and spatial factors associated with variation in movement patterns for a marine mesopredator. This can be useful information for promoting species recovery, and for future efforts to understand ecological patterns such as predator-prey interactions.
Population growth typically involves range expansion and establishment of new breeding sites, while the opposite occurs during declines. Although density dependence is widely invoked in theoretical studies of emigration and colonization in expanding populations, few empirical studies have documented the mechanisms. Still fewer have documented the direction and mechanisms of individual transfer in declining populations. Here, we screen large numbers of pups sampled on their natal rookeries for variation in mtDNA (n=1106) and 16 microsatellite loci (n=588) and show that new Steller sea lion breeding sites did not follow the typical paradigm and were instead colonized by sea lions from both a declining (Endangered) population and an increasing population. Dispersing individuals colonized rookeries in the distributional hiatus between two evolutionarily distinct ((Phi) over bar (st) = 0.222, (R) over bar (st) = 0.053, K=2) metapopulations recently described as separate subspecies. Hardy-Weinberg, mixed-stock and relatedness analysis revealed levels of interbreeding on the new rookeries that exclude (i) assortative mating among eastern and western forms, and (ii) inbreeding avoidance as primary motivations for dispersal. Positive and negative density dependence is implicated in both cases of individual transfer. Migration distance limits, and conspecific attraction and performance likely influenced the sequence of rookery colonizations. This study demonstrates that resource limitation may trigger an exodus of breeding animals from declining populations, with substantial impacts on distribution and patterns of genetic variation. It also revealed that this event is rare because colonists dispersed across an evolutionary boundary, suggesting that the causative factors behind recent declines are unusual or of larger magnitude than normally occur.
Directly observing the foraging behavior of animals in the marine environment can be extremely challenging, if not impossible, as such behavior often takes place beneath the surface of the ocean and in extremely remote areas. In lieu of directly observing foraging behavior, data from time–depth recorders and other types of behavioral data recording devices are commonly used to describe and quantify the behavior of fish, squid, seabirds, sea turtles, pinnipeds, and cetaceans. Often the definitions of actual behavioral units and analytical approaches may vary substantially which may influence results and limit our ability to compare behaviors of interest across taxonomic groups and geographic regions. A workshop was convened in association with the Fourth International Symposium on Bio-logging in Hobart, Tasmania on 8 March 2011, with the goal of providing a forum for the presentation, review, and discussion of various methods and approaches that are used to describe and analyze time–depth recorder and associated behavioral data records. The international meeting brought together 36 participants from 14 countries from a diversity of backgrounds including scientists from academia and government, graduate students, post-doctoral fellows, and developers of electronic tagging technology and analysis software. The specific objectives of the workshop were to host a series of invited presentations followed by discussion sessions focused on (1) identifying behavioral units and metrics that are suitable for empirical studies, (2) reviewing analytical approaches and techniques that can be used to objectively classify behavior, and (3) identifying cases when temporal autocorrelation structure is useful for identifying behaviors of interest. Outcomes of the workshop included highlighting the need to better define behavioral units and to devise more standardized processing and analytical techniques in order to ensure that results are comparable across studies and taxonomic groups.
P>During the 1990s, the Steller sea lion (Eumetopias jubatus Schreber) Western Alaska stock (WS) suffered steep population decline while the Eastern Alaska stock (ES) steadily increased. One bottom-up forcing hypothesis explaining this decline predicted lactating adult female foraging behavior would be different between stocks. To investigate this effect, we monitored 11 ES females at two breeding rookeries using satellite dive recorders (SDR) during the early breeding seasons of 1992-1993, examined their behavior with respect to prey, physiological limitations, and habitat, and made limited comparisons to observations of WS female behavior reported in the literature. ES females were not operating at the extremes of ability, with most diving within the limits of aerobic metabolism, less than one-quarter of possible foraging time during trips spent submerged and most foraging trips requiring less than one-half the lipid store fasting ability of dependent pups. Thus, females may have some capacity to alter behavior to accommodate future changes in foraging conditions, but the extent of this plasticity is unknown. Because recent work suggests WS recovery is impeded by low natality, future studies should test differences between reproductive and non-reproductive mature females in order to properly assess the contribution of foraging ecology to SSL population dynamics.
We hypothesized that: (1) Steller sea lion Eurmetopias jubatus diet choice is a function of prey availability, (2) sea lions move to take advantage of times and locations of seasonal prey concentrations and (3) the number present depends on the amount of prey available (numerical response). Over 3 yr, typically on a quarterly basis, in Frederick Sound, SE Alaska, multiple measurements were taken of Steller sea lion abundance (aerial surveys), diet (scats), dive behavior (satellite telemetry) and prey availability and caloric density (nearshore, pelagic and demersal fish surveys). We found that Steller sea lions shifted diet composition in response to changes in prey availability of pollock Theragra chalcogramma, hake Merluccius productus, herring Clupea pallasi and salmon Oncorhynchus spp. They selected intermediate-sized fish and avoided small (<10 cm) and large (>60 cm) fish, and moved between areas as prey became available seasonally. The number of sea lions present depended on the amount of prey available; a standing biomass of 500 to 1700 t of prey in a non-breeding area such as Frederick Sound, depending on species composition, can attract and sustain about 500 sea lions. Pollock was more frequent in sea lion diet in inside waters of SE Alaska-including Frederick Sound, Stephens Passage and Lynn Canal-than anywhere else in Alaska and contributed similar to 1/3 of the dietary energy in Frederick Sound. This finding implies that a diet with substantial year-round contributions from less nutritious, but abundant prey such as pollock can form part of a healthy diet as long as more nutritious prey such as herring, salmon or eulachon Thaleichthys pacificus also are consumed. Our study supports the conclusion that the Steller sea lion is an opportunistic marine predator with a flexible foraging strategy that selects abundant, accessible prey and shifts among seasonally available species.
Reduced juvenile survival caused by prey depletion is one hypothesis for the decline in the western Alaska population of Steller sea lions ( Eumetopias jubatus (Schreber, 1776)). To understand the exposure of young sea lions to these depletions, the swimming and diving behavior of pups, juveniles, and subadults was evaluated relative to prey behavior. Pups made shorter and shallower dives (13 m, 0.9 min) than juveniles or subadults, as expected based on physiological limitations, but juveniles and subadults dived to similar depths and durations (29 m, 1.7 min and 38 m, 2.0 min, respectively). Activity patterns of juveniles and subadults reflected diurnal prey migrations, while pup activity did not. Longitudinal trends in pup dive behavior reflected both physiological and behavioral development, while juvenile dive behaviors reflected seasonal changes in prey availability. Results suggest that adult females must continue to provide nutritional support to pups during winter because of the limited diving ability of these young animals. For this reason, the flexible lactation strategies that allow for longer nursing periods during periods of low prey availability and reduce female fecundity may improve juvenile survival.
We calculated the durations of time on-shore and at-sea for juvenile Steller sea lions (Eumetopias jubatus) using satellite dive recorders deployed between 2000 and 2002, and compared two genetically distinct populations; one increasing (eastern stock; n=42) and one that experienced an 80% decline in population since the mid-1970s (western stock; n=89). Data represented 24-h periods divided into 72 20-min increments indicating whether an animal was on-shore (dry) or at-sea (wet). Time apportioned between land and sea was described on a per-trip basis (rather than a 24-h cycle) and durations ranged from 20min to several days. We tested differences in the durations of on-shore and at-sea events among sex, geographic region, year, and age at capture using mixed-effects models. Animal identifier was included as a random effect to account for repeated measures on the same individual. Sea lions from the eastern Aleutian Islands, central Aleutian Islands, and central Gulf of Alaska hauled out just after sunrise, and departure times coincided with dusk. For Prince William Sound and Southeast Alaska animals, arrivals and departures occurred throughout the day and were not related to crepuscular period. Mean duration on-shore did not differ among sex, region, year or age, and was unrelated to previous trip duration. This may suggest a minimum rest period for juvenile Steller sea lions or that dependant animals are maximizing their time on-shore suckling. Time spent at-sea varied among individuals from both populations and development of maternal independence, inferred from significant increases in time spent at sea, occurred approximately 10 months later in individuals from Prince William Sound and Southeast Alaska than in the other regions, suggesting environmental and developmental differences among regions.
Development of competent diving ability is critical to obtaining nutritional independence in marine mammals such as Steller sea lions (SSLs), Eumetopias jubatus (Schreber, 1776). We studied diving performance in pup (75) and juvenile (36) SSLs using satellite data recorders. In general, dives by SSLs were brief and shallow. Overall, 82.3% of dives were < 2 min long and 86.9% of dives were < 10 m deep. Long ( > 5 min) and deep dives ( > 100 m) constituted only 2.49% and 0.77%, respectively, of total dives. We used linear mixed-effects models to investigate the relationships between the response variables maximum-daily-depth, time-at-depth, mean-dive-duration, dive rate, and time-at-sea and the predictor variables age, sex, population (eastern and western Alaska populations), time-of-day, and month-of-year. All response variables except dive rate were positively related (P < 0.05) to age. Dive rate declined (P < 0.001) with age. Time-of-day, month, population, sex, and some first-order interactions were all significantly (P < 0.05) related to some measure of diving performance. With large samples we were able to identify significant relationships between the response variables and the predictor variables, even though the total amount of variation explained by the models was low, because most dives were short and shallow regardless of age, sex, population, time-of-day, or month-of-year. Depths and durations of dives by juvenile animals increased throughout the range of ages studied and were similar to or greater than those previously reported for juveniles and adult females. We expect maximum depths and durations to continue to increase with age until body mass plateaus at about 10 years of age. Therefore, we expect older animals to be more efficient foragers, as they would have greater aerobic dive limits as well as more experience locating and capturing prey.
Movement patterns of Alaska harbor seal pups were studied using satellite telemetry during 1997-2000. Mean tracking duration was 277.3 d (SD = 105.8) for Tugidak Island pups (n = 26) and 171.2 d (108.3) for Prince William Sound (PWS) pups (n = 27). Movements were similar for males and females and were largely restricted to the continental shelf. Multiple return trips of > 75 km from the natal area and up to similar to 3 wk duration were most common, followed by movements restricted to < 25 km from the natal area; one way movements from the natal site were rare. Distances moved and home range sizes remained relatively stable or increased gradually from July through winter, then decreased markedly through spring. Monthly movements (maximum distance from tagging location, mean distance from haul-outs to at-sea locations, and home range size) were significantly greater for Tugidak vs. PWS pups. Six of seven pups from each region that traveled farthest and were tracked the longest had returned to their tagging site when their last location was recorded, indicating philopatry or limited dispersal during their first year of life. Seal pups exhibited similar movement patterns in the distinct habitats of the two regions, but differed in the spatial extent of their movements.
AbstractPopulation declines of Steller sea lions (Eumetopias juhatus) in western Alaska (west of 144°W) may be a result of reduced juvenile survival. We used satellite telemetry to study the at‐sea distribution and movement patterns of pup (1.6–11.9 mo) and juvenile (12.0–35.1 mo) Steller sea lions. We studied trip distance, duration, and interhaul‐out movements of sea lions in relation to age, sex, and month of year in the decreasing western population (WP; Prince William Sound, Kodiak, Aleutian Islands, Alaska) and the increasing eastern population (EP; Southeast Alaska). We deployed 103 satellite transmitters (29 WP; 74 EP) on sea lions between 1998 and 2001. Round trip distance and duration increased with age, trip distance was greater in the WP than the EP, trip duration was greater for females than males, and haul‐out use was clustered. Changes in round trip distance and duration occurred from April to June for all age classes studied indicating that the annual timing of weaning may be less variable than the age of weaning. Overall, 90% of round trips were ≤ 15 km from haul‐outs and 84% were <20 h, indicating nearshore areas adjacent to haulouts are critical to the developing juvenile.