Food habits of Steller sea lions (Eumetopias jubatus) off Oregon and northern California, 1986–2007

We described the diet of the eastern stock of Steller sea lions (Eumetopias jubatus) from 1416 scat samples collected from five sites in Oregon and northern California from 1986 through 2007. A total of 47 prey types from 30 families were identified. The most common prey was Pacific hake (Merluccius productus), followed by salmonids (Oncorhynchus spp.), skates (Rajidae), Pacific lamprey (Lampetra tridentata), herrings (Clupeidae), rockfish (Sebastes spp.), and northern anchovy (Engraulis mordax). Steller sea lion diet composition varied seasonally, annually, and spatially. Hake and salmonids were the most commonly identified prey in scats collected during the summer (breeding season), whereas hake and skate were most common in the nonbreeding season. Continued research on Steller sea lion diet and foraging behavior in the southern extent of their range is necessary to address issues such as climate change, interaction with competing California sea lions, and predation impacts on valuable or sensitive fish stocks.

Effects of commercial fishing on local abundance of Pacific cod (Gadus macrocephalus) in the Bering Sea

Groundfish fisheries in the southeast Bering Sea in Alaska have been constrained in recent years by management measures to protect the endangered Steller sea lion (Eumetopias jubatus). There is concern that the present commercial harvest may produce a localized depletion of groundfish that would affect the foraging success of Steller sea lions or other predators. A three-year field experiment was conducted to determine whether an intensive trawl fishery in the southeast Bering Sea created a localized depletion in the abundance of Pacific cod (Gadus macrocephalus). This experiment produced strongly negative results; no difference was found in the rate of seasonal change in Pacific cod abundance between stations within a regulatory no-trawl zone and stations in an immediately adjacent trawled area. Corollary studies showed that Pacific cod in the study area were highly mobile and indicated that the geographic scale of Pacific cod movement was larger than the spatial scale used as the basis for current no-trawl zones. The idea of localized depletion is strongly dependent on assumed spatial and temporal scales and contains an implicit assumption that there is a closed local population. The scale of movement of target organisms is critical in determining regional effects of fishery removals.

Diets of Steller sea lions (Eumetopias jubatus) in Southeast Alaska, 1993−1999

The diet of Steller sea lions (Eumetopias jubatus) was determined from 1494 scats (feces) collected at breeding (rookeries) and nonbreeding (haulout) sites in Southeast Alaska from 1993 to 1999. The most common prey of 61 species identified were walleye pollock (Theragra chalcogramma), Pacific herring (Clupea pallasii), Pacific sand lance (Ammodytes hexapterus), Pacific salmon (Salmonidae), arrowtooth flounder (Atheresthes stomias), rockfish (Sebastes spp.), skates (Rajidae), and cephalopods (squid and octopus). Steller sea lion diets at the three Southeast Alaska rookeries differed significantly from one another. The sea lions consumed the most diverse range of prey categories during summer, and the least diverse during fall. Diet was more diverse in Southeast Alaska during the 1990s than in any other region of Alaska (Gulf of Alaska and Aleutian Islands). Dietary differences between increasing and declining populations of Steller sea lions in Alaska correlate with rates of population change, and add credence to the view that diet may have played a role in the decline of sea lions in the Gulf of Alaska and Aleutian Islands.

Survey- and fishery-derived estimates of Pacific cod (Gadus macrocephalus) biomass: implications for strategies to reduce interactions between groundfish fisheries and Steller sea lions (Eumetopias jubatus)

Survey- and fishery-derived biomass estimates have indicated that the harvest indices for Pacific cod (Gadus macrocephalus) within a portion of Steller sea lion (Eumetopias jubatus) critical habitat in February and March 2001 were five to 16 times greater than the annual rate for the entire Bering Sea-Aleutian Islands stock. A bottom trawl survey yielded a cod biomass estimate of 49,032 metric tons (t) for the entire area surveyed, of which less than half (23,329 t) was located within the area used primarily by the commercial fishery, which caught 11,631 t of Pacific cod. Leslie depletion analyses of fishery data yielded biomass estimates of approximately 14,500 t (95% confidence intervals of approximately 9,000–25,000 t), which are within the 95% confidence interval on the fished area survey estimate (12,846–33,812 t). These data indicate that Leslie analyses may be useful in estimating local fish biomass and harvest indices for certain marine fisheries that are well constrained spatially and relatively short in duration (weeks). In addition, fishery effects on prey availability within the time and space scales relevant to foraging sea lions may be much greater than the effects indicated by annual harvest rates estimated from stock assessments averaged across the range of the target spec

Distribution and Abundance of Steller Sea Lions, Eumetopias jubatus, on the Asian Coast, 1720's-2005

We analyzed published and archived records for the past 250 years to assess changes in distribution and abundance of Steller sea lions, Eumetopias jubatus, along the Asian coast from the Bering Strait to the Korean Peninsula. We found that the northern extent of Steller sea lion distribution has not changed but that the southern limit has moved north by some 500–900 km (~300–500 n.mi.) over the past 50 years. Additionally, the number of animals and their distribution has changed on the Commander Islands, Kuril Islands, and Kamchatka Peninsula. We found no changes in the number of rookeries in the northern Sea of Okhotsk, but a new rookery was established at Tuleny Island on the eastern coast of Sakhalin Island. We estimate that the total abundance of Steller sea lions along the Asian coast in the late 19th century was about 115,000 animals; during the 1960’s, the total estimate was about 27,000 (including pups), most of which were in the Kuril Islands. The fewest number of Steller sea lions occurred in the northwestern Pacific in the late 1980’s–early 1990’s when only about 13,000 individuals (including pups) were estimated in the entire region. During the 1990’s, and especially in early 2000, an increasing trend in abundance occurred in most areas. Present estimated abundance of Steller sea lions in Asia is about 16,000 individuals (including about 5,000 pups), about half of which occur in the Kuril Islands. Changes in abundance occurred during all time periods but varied by site and period. Specifically, over the past 150 years Steller sea lion abundance at most sites has changed. There were no rookeries on the Commander Islands between 1850 and 1960 and abundance was low, but by 1977, abundance increased to 4,800 individuals and a rookery was established in the mid 1980’s; abundance there has declined since the early 1980’s and in 2004 only 895 individuals (including 221 pups) were counted during the breeding season. Between 1940 and 2004, abundance along the eastern coast of Kamchatka declined from ~7,000 to ~600 individuals, an overall reduction of 90%. Steller sea lion abundance on the Kuril Islands declined by >90% from the 1800’s to 2005; the most severe decline there occurred during 1969–1981. Steller sea lion numbers in the northern part of the Sea of Okhotsk declined during 1930–2002 from 7,200 to 3,100 individuals. Numbers at Tuleny Island have increased since establishment of a rookery there during 1983–2005 and by immigration from other sites.

Estimates of Marine Mammal, Sea Turtle, and Seabird Mortality in the California Drift Gillnet Fishery for Swordfish and Thresher Shark, 1996–2002

Estimates of incidental marine mammal, sea turtle, and seabird mortality in the California drift gillnet fishery for broadbill swordfish, Xiphias gladius, and common thresher shark, Alopias vulpinus, are summarized for the 7-year period, 1996 to 2002. Fishery observer coverage was 19% over the period (3,369 days observed/17,649 days fished). An experiment to test the effectiveness of acoustic pingers on reducing marine mammal entanglements in this fishery began in 1996 and resulted in statistically significant reductions in marine mammal bycatch. The most commonly entangled marine mammal species were the short-beaked common dolphin, Delphinus delphis; California sea lion, Zalophus californianus; and northern right whale dolphin, Lissodelphis borealis. Estimated mortality by species (CV and observed mortality in parentheses) from 1996 to 2002 is 861 (0.11, 133) short-beaked common dolphins; 553 (0.16, 103) California sea lions; 151 (0.25, 31) northern right whale dolphins; 150 (0.21, 27) northern elephant seals, Mirounga angustirostris; 54 (0.41, 10) long-beaked common dolphins, Delphinus capensis; 44 (0.53, 6) Dall’s porpoise, Phocoenoides dalli; 19 (0.60, 5) Risso’s dolphins, Grampus griseus; 11 (0.71, 2) gray whales, Eschrichtius robustus; 7 (0.83, 2) sperm whales, Physeter macrocephalus; 7 (0.96, 1) short-finned pilot whales, Globicephala macrorhychus; 12 (1.06, 1) minke whales, Balaenoptera acutorostrata; 5 (1.05, 1) fin whales, Balaenoptera physalus; 11 (0.68, 2) unidentified pinnipeds; 33 (0.52, 4) leatherback turtles, Dermochelys coriacea; 18 (0.57, 3) loggerhead turtles, Caretta caretta; 13 (0.73, 3) northern fulmars, Fulmarus glacialis; and 6 (0.86, 2) unidentified birds.

The Threatened Status of Steller Sea Lions, Eumetopias jubatus, under the Endangered Species Act: Effects on Alaska Groundfish Fisheries Management

In April 1990, the Steller sea lion, Eumetopias jubatus, was listed as threatened under the U.S. Endangered Species Act by emergency action. Competitive interactions with the billion-dollar Alaska commercial groundfish fisheries have been suggested as one of the possible contributing factors to the Steller sea lion population decline. Since the listing, fisheries managers have attempted to address the potential impacts of the groundfish fisheries on Steller sea lion recovery. In this paper, we review pertinent Federal legislation, biological information on the Steller sea lion decline, changes in the Alaska trawl fishery for walleye pollock, Theragra chalcogramma, since the late 1970's, andpossible interactions between fisheries and sea lions. Using three cases, we illustrate how the listing of Steller sea lions has affected Alaska groundfish fisheries through: I) actions taken at the time of listing designed to limit the potential for directhuman-related sea lion mortality, 2) actions addressing spatial and temporal separation of fisheries from sea lions, and 3) introduction of risk-adverse stock assessment methodologies and Steller sea lion conservation considerations directly in the annual quota-setting process. This discussion shows some of the ways that North Pacific groundfish resource managers have begun to explicitly consider the conservation ofmarine mammal and other nontarget species.

Sizes of walleye pollock (Theragra chalcogramma) consumed by the eastern stock of Steller sea lions (Eumetopias jubatus) in Southeast Alaska from 1994 to 1999

Lengths of walleye pollock (Theragra chalcogramma) consumed by Steller sea lions (Eumetopias jubatus) were estimated by using allometric regressions applied to seven diagnostic cranial structures recovered from 531 scats collected in Southeast Alaska between 1994 and 1999. Only elements in good and fair condition were selected. Selected structural measurements were corrected for loss of size due to erosion by using experimentally derived condition-specific digestion correction factors. Correcting for digestion increased the estimated length of fish consumed by 23%, and the average mass of fish consumed by 88%. Mean corrected fork length (FL) of pollock consumed was 42.4 ±11.6 cm (range=10.0−78.1 cm, n=909). Adult pollock (FL>45.0 cm) occurred more frequently in scats collected from rookeries along the open ocean coastline of Southeast Alaska during June and July (74% adults, mean FL=48.4 cm) than they did in scats from haul-outs located in inside waters between October and May (51% adults, mean FL=38.4 cm). Overall, the contribution of juvenile pollock (≤20 cm) to the sea lion diet was insignificant; whereas adults contributed 44% to the diet by number and 74% by mass. On average, larger pollock were eaten in summer at rookeries throughout Southeast Alaska than at rookeries in the Gulf of Alaska and the Bering Sea. Overall it appears that Steller sea lions are capable of consuming a wide size range of pollock, and the bulk of fish fall between 20 and 60 cm. The use of cranial hard parts other than otoliths and the application of digestion correction factors are fundamental to correctly estimating the sizes of prey consumed by sea lions and determining the extent that these sizes overlap with the sizes of pollock caught by commercial fisheries.

Diving behavior of immature Steller sea lions (Eumetopias jubatus)

Understanding the ontogenetic relationship between juvenile Steller sea lions (Eumetopias jubatus) and their foraging habitat is key to understanding their relationship to available prey and ultimately their survival. We summarize dive and movement data from 13 young-of-the-year (YOY) and 12 yearling Steller sea lions equipped with satellite dive recorders in the Gulf of Alaska and Aleutian Islands (n=18), and Washington (n=7) from 1994 to 2000. A total of 1413 d of transmission (x =56.5 d, range: 14.5–104.1 d) were received. We recorded 222,073 dives, which had a mean depth of 18.4 m (range of means: 5.8−67.9 m; SD=16.4). Alaska YOY dived for shorter periods and at shallower depths (mean depth=7.7 m, mean duration=0.8 min, mean maximum depth=25.7 m, and maximum depth=252 m) than Alaska yearlings (x =16.6 m, 0=1.1 min, x = 63.4 m, 288 m), whereas Washington yearlings dived the longest and deepest (mean depth=39.4 m, mean duration=1.8 min, mean maximum depth=144.5 m, and maximum depth=328 m). Mean distance for 564 measured trips was 16.6 km; for sea lions ≤10 months of age, trip distance (7.0 km) was significantly less than for those >10 months of age (24.6 km). Mean trip duration for 10 of the 25 sea lions was 12.1 h; for sea lions ≤10 months of age, trip duration was 7.5 h and 18.1 h for those >10 months of age. We identified three movements types: long-range trips (>15 km and >20 h), short-range trips (<15 km and <20 h) during which the animals left and returned to the same site, and transits to other haul-out sites. Long-range trips started around 9 months of age and occurred most frequently around the assumed time of weaning, whereas short-range trips happened almost daily (0.9 trips/day, n=426 trips). Transits began as early as 7 months of age, occurred more often after 9 months of age, and ranged between 6.5 and 454 km. The change in dive characteristics coincided with the assumed onset of weaning. These yearling sea lion movement patterns and dive characteristics suggest that immature Steller sea lions are as capable of making the same types of movements as adults.

Weak polygyny in California sea lions and the potential for alternative mating tactics

Female aggregation and male territoriality are considered to be hallmarks of polygynous mating systems. The development of genetic parentage assignment has called into question the accuracy of behavioral traits in predicting true mating systems. In this study we use 14 microsatellite markers to explore the mating system of one of the most behaviorally polygynous species, the California sea lion (Zalophus californianus). We sampled a total of 158 female-pup pairs and 99 territorial males across two breeding rookeries (San Jorge and Los Islotes) in the Gulf of California, Mexico. Fathers could be identified for 30% of pups sampled at San Jorge across three breeding seasons and 15% of sampled pups at Los Islotes across two breeding seasons. Analysis of paternal relatedness between the pups for which no fathers were identified (sampled over four breeding seasons at San Jorge and two at Los Islotes) revealed that few pups were likely to share a father. Thirty-one percent of the sampled males on San Jorge and 15% of the sampled males on Los Islotes were assigned at least one paternity. With one exception, no male was identified as the father of more than two pups. Furthermore, at Los Islotes rookery there were significantly fewer pups assigned paternity than expected given the pool of sampled males (p<0.0001). Overall, we found considerably lower variation in male reproductive success than expected in a species that exhibits behavior associated with strongly polygynous mating. Low variation in male reproductive success may result from heightened mobility among receptive females in the Gulf of California, which reduces the ability of males to monopolize groups of females. Our results raise important questions regarding the adaptive role of territoriality and the potential for alternative mating tactics in this species.

Regional differences in the spatial and temporal heterogeneity of oceanographic habitat used by Steller sea lions

Over the past three decades, the decline and altered spatial distribution of the western stock of Steller sea lions (Eumetopias jubatus) in Alaska have been attributed to changes in the distribution or abundance of their prey due to the cumulative effects of fisheries and environmental perturbations. During this period, dietary prey occurrence and diet diversity were related to population decline within metapopulation regions of the western stock of Steller sea lions, suggesting that environmental conditions may be variable among regions. The objective of this study, therefore, was to examine regional differences in the spatial and temporal heterogeneity of oceanographic habitat used by Steller sea lions within the context of recent measures of diet diversity and population trajectories. Habitat use was assessed by deploying satellite-depth recorders and satellite relay data loggers on juvenile Steller sea lions (n = 45) over a five-year period (2000–2004) within four regions of the western stock, including the western, central, and eastern Aleutian Islands, and central Gulf of Alaska. Areas used by sea lions during summer months (June, July, and August) were demarcated using satellite telemetry data and characterized by environmental variables (sea surface temperature [SST] and chlorophyll a [chl a]), which possibly serve as proxies for environmental processes or prey. Spatial patterns of SST diversity and Steller sea lion population trends among regions were fairly consistent with trends reported for diet studies, possibly indicating a link between environmental diversity, prey diversity, and distribution or abundance of Steller sea lions. Overall, maximum spatial heterogeneity coupled with minimal temporal variability of SST appeared to be beneficial for Steller sea lions. In contrast, these patterns were not consistent for chl a, and there appeared to be an ecological threshold. Understanding how Steller sea lions respond to measures of environmental heterogeneity will ultimately be useful for implementing ecosystem management approaches and developing additional conservation strategies.

Differential Expression of Immune Response Genes in Steller Sea Lions (Eumetopias jubatus): An Indicator of Ecosystem Health?

Characterization of the polygenic and polymorphic features of the Steller sea lion major histocompatibility complex (MHC) provides an ideal window for evaluating immunologic vigor of the population and identifying emergence of new genotypes that reflect ecosystem pressures. MHC genotyping can be used to measure the potential immunologic vigor of a population. However, since ecosystem-induced changes to MHC genotype can be slow to emerge, measurement of differential expression of these genes can potentially provide real-time evidence of immunologic perturbations. MHC DRB genes were cloned and sequenced using peripheral blood mononuclear leukocytes derived from 10 Steller sea lions from Southeast Alaska, Prince William Sound, and the Aleutian Islands. Nine unique DRB gene sequences were represented in each of 10 animals. MHC DRB gene expression was measured in a subset of six sea lions. Although DRB in genomic DNA was identical in all individuals, relative levels of expressed DRB mRNA was highly variable. Selective suppression of MHC DRB genes could be indicative of geographically disparate environmental pressures, thereby serving as an immediate and sensitive indicator of population and ecosystem health.

Sea lions, sacrifice, and ritual : The symbolism of Moche Island imagery

The Mochica, or Moche (c. 100-800 AD) culture, flourished along the northern Peruvian coast. The Moche did not have a formal written language; as such, contemporary scholars base their analysis on Moche iconography and archaeological burial remains. Especially renown for their ceramic artistry, Moche vessels exhibit a wide range of subject matter, including animal and enigmatic figural representations that evoke terrestrial, marine and possibly, spiritual realms. While research has focused on political organization and the interrelationship between sacrifice and warfare, many marine themes have not been fully explored in the discourse. An exploration of sea lion imagery and sacrifice themes suggests that the marine mammals were ritually hunted. A careful iconographic analysis of island scenes demonstrates ritual and gender affiliations held by the Moche about sea lions. In a multi-disciplinary approach, scientific, archaeological and ethnographic resources substantiate this claim.

Incidental Catch of Marine Mammals by Foreign and Joint Venture Trawl Vessels in the U.S. EEZ of the North Pacific, 1973-88

During 1973-88, 3,661 marine mammals of 17 species were reported as incidental catch by U.S. fishery observers aboard foreign and joint venture trawl vessels in the U.S. Exclusive Economic Zone in the North Pacific Ocean and the Bering Sea. Northern sea lions (Eumetopias jubatus) accounted for 90% of the reported incidental mortality in the Gulf of Alaska and eastern Bering Sea. Nearly half of these sea lions were taken in trawl nets in the Shelikof Strait, Alaska, joint venture fishery during 1982-84. However, high incidental mortality rates (>25 sea lions per 10,000 metric tons of groundfish catch) also occurred in the foreign fisheries near Kodiak Island and in the Aleutian Islands area in earlier years. Estimated annual mortality of incidentally caught northern sea lions in Alaska declined from 1,000 to 2,000 animals per year during the early 1970s and 1982 to fewer than 100 animals in 1988. In the Bering Sea most sea lions incidentally caught were males, while in the Gulf of Alaska females were more frequently caught. Females may also have been dominant in the incidental catch of sea lions in the Aleutian Islands area, but age and sex composition data are limited. Incidental mortality of adult female sea lions by foreign trawl fisheries in these areas could have partially contributed to the reported declines in northern sea lion populations in Alaska during the 1970s, but it cannot alone account for the present decline in population size. (PDF file contains 64 pages.)

Changes over time in the spatial distribution of walleye pollock (Theragra chalcogramma) in the Gulf of Alaska, 1984-1996

Triennial bottom trawl survey data from 1984 to 1996 were used to evaluate changes in the summer distribution of walleye pollock in the western and central Gulf of Alaska. Differences between several age groups of pollock were evaluated. Distribution was examined in relation to several physical characteristics, including bottom depth and distance from land. Interspecies associations were also analyzed with the Bray-Curtis clustering technique to better understand community structure. Our results indicated that although the population numbers decreased, high concentrations of pollock remained in the same areas during 1984–96. However, there was an increase in the number of stations where low-density pollock concentrations of all ages were observed, which resulted in a decrease in mean population density of pollock within the GOA region. Patterns emerging from our data suggested an alternative to Mac-Call’s “basin hypothesis” which states that as population numbers decrease, there should be a contraction of the population range to optimal habitats. During 1984–96 there was a concurrent precipitous decline in Steller sea lions in the Gulf of Alaska. The results of our study suggest that decreases in the mean density of adult pollock, the main food in the Steller sea lion diet, combined with slight changes in the distribution of pollock (age-1 pollock in particular) in the mid-1980s, may have contributed to decreased foraging efficiency in Steller sea lions. Our results support the prevailing conceptual model for pollock ontogeny, although there is evidence that substantial spawning may also occur outside of Shelikof Strait.