Contemporary diets of walruses in Bristol Bay, Alaska suggest temporal variability in benthic community structure.

BACKGROUND: Pacific walruses (Odobenus rosmarus divergens) are a conspicuous and important component of the Bristol Bay ecosystem and human social systems, but very little is known about walrus ecology in this region, principally their feeding ecology. The present work provides contemporary data on the diets of walruses at four haulout locations throughout Bristol Bay between 2014 and 2018. METHODS: We analyzed scat and gastrointestinal tract samples from these animals using quantitative polymerase chain reaction to amplify prey DNA, which allowed for diet estimates based on frequencies of prey item occurrence and on the relative importance of dietary items as determined from DNA threshold cycle scores. RESULTS: Diets were highly diverse at all locations, but with some variation in composition that may be related to the time of year that samples were collected (summer vs. autumn), or to spatial variability in the distribution of prey. Overall, polychaetes and tunicates had the highest frequencies of occurrence and relative abundances in 2014-15, but a major change in diet appears to have occurred by 2017-18. While some sample sizes were small, diets in these later years contrasted sharply, with a greater prevalence of sea cucumbers and mollusks, and reduced importance of decapods and fishes compared to the earlier years. Prey identified in scat samples from one collection site also contrasted sharply with those reported from the same location in 1981. The apparent temporal shifts in walrus prey may represent a changing benthic ecosystem due to warming waters in recent decades.

Transhemispheric ecosystem disservices of pink salmon in a Pacific Ocean macrosystem.

Pink salmon (Oncorhynchus gorbuscha) in the North Pacific Ocean have flourished since the 1970s, with growth in wild populations augmented by rising hatchery production. As their abundance has grown, so too has evidence that they are having important effects on other species and on ocean ecosystems. In alternating years of high abundance, they can initiate pelagic trophic cascades in the northern North Pacific Ocean and Bering Sea and depress the availability of common prey resources of other species of salmon, resident seabirds, and other pelagic species. We now propose that the geographic scale of ecosystem disservices of pink salmon is far greater due to a 15,000-kilometer transhemispheric teleconnection in a Pacific Ocean macrosystem maintained by short-tailed shearwaters (Ardenna tenuirostris), seabirds that migrate annually between their nesting grounds in the South Pacific Ocean and wintering grounds in the North Pacific Ocean. Over this century, the frequency and magnitude of mass mortalities of shearwaters as they arrive in Australia, and their abundance and productivity, have been related to the abundance of pink salmon. This has influenced human social, economic, and cultural traditions there, and has the potential to alter the role shearwaters play in insular terrestrial ecology. We can view the unique biennial pulses of pink salmon as a large, replicated, natural experiment that offers basin-scale opportunities to better learn how these ecosystems function. By exploring trophic interaction chains driven by pink salmon, we may achieve a deeper conservation conscientiousness for these northern open oceans.

Carbon sources and trophic relationships of ice seals during recent environmental shifts in the Bering Sea.

Dramatic multiyear fluctuations in water temperature and seasonal sea ice extent and duration across the Bering-Chukchi continental shelf have occurred in this century, raising a pressing ecological question: Do such environmental changes alter marine production processes linking primary producers to upper trophic-level predators? We examined this question by comparing the blubber fatty acid (FA) composition and stable carbon isotope ratios of individual FA (δ¹³CFA) of adult ringed seals (Pusa hispida), bearded seals (Erignathus barbatus), spotted seals (Phoca largha), and ribbon seals (Histriophoca fasciata), collectively known as "ice seals," sampled during an anomalously warm, low sea ice period in 2002-2005 in the Bering Sea and a subsequent cold, high sea ice period in 2007-2010. δ¹³C(FA) values, used to estimate the contribution to seals of carbon derived from sea ice algae (sympagic production) relative to that derived from water column phytoplankton (pelagic production), indicated that during the cold period, sympagic production accounted for 62-80% of the FA in the blubber of bearded seals, 51-62% in spotted seals, and 21-60% in ringed seals. Moreover, the δ¹³CFA values of bearded seals indicated a greater incorporation of sympagic FAs during the cold period than the warm period. This result provides the first empirical evidence of an ecosystem-scale effect of a putative change in sympagic production in the Western Arctic. The FA composition of ice seals showed clear evidence of resource partitioning among ringed, bearded, and spotted seals, and little niche separation between spotted and ribbon seals, which is consistent with previous studies. Despite interannual variability, the FA composition of ringed and bearded seals showed little evidence of differences in diet between the warm and cold periods. The findings that sympagic production contributes significantly to food webs supporting ice seals, and that the contribution apparently is less in warm years with low sea ice, raise an important concern: Will the projected warming and continuing loss of seasonal sea ice in the Arctic, and the associated decline of organic matter input from sympagic production, be compensated for by pelagic production to satisfy both pelagic and benthic carbon and energy needs?

Population Trend and Elasticities of Vital Rates for Steller Sea Lions (Eumetopias jubatus) in the Eastern Gulf of Alaska: A New Life-History Table Analysis.

Steller sea lion (Eumetopias jubatus) numbers are beginning to recover across most of the western distinct population segment following catastrophic declines that began in the 1970s and ended around the turn of the century. This study makes use of contemporary vital rate estimates from a trend-site rookery in the eastern Gulf of Alaska (a sub-region of the western population) in a matrix population model to estimate the trend and strength of the recovery across this region between 2003 and 2013. The modeled population trend was projected into the future based on observed variation in vital rates and a prospective elasticity analysis was conducted to determine future trends and which vital rates pose the greatest threats to recovery. The modeled population grew at a mean rate of 3.5% per yr between 2003 and 2013 and was correlated with census count data from the local rookery and throughout the eastern Gulf of Alaska. If recent vital rate estimates continue with little change, the eastern Gulf of Alaska population could be fully recovered to pre-decline levels within 23 years. With density dependent growth, the population would need another 45 years to fully recover. Elasticity analysis showed that, as expected, population growth rate (λ) was most sensitive to changes in adult survival, less sensitive to changes in juvenile survival, and least sensitive to changes in fecundity. A population decline could be expected with only a 6% decrease in adult survival, whereas a 32% decrease in fecundity would be necessary to bring about a population decline. These results have important implications for population management and suggest current research priorities should be shifted to a greater emphasis on survival rates and causes of mortality.

A longitudinal study of Steller sea lion natality rates in the Gulf of Alaska with comparisons to census data.

Steller sea lion (Eumetopias jubatus) numbers in the Western Distinct Population Segment are beginning to recover following the dramatic decline that began in the 1970s and ended around the turn of the century. Low female reproductive rates (natality) may have contributed to the decline and remain an issue of concern for this population. During the 2000s we found high natality among Steller sea lions in the Gulf of Alaska indicating a healthy population. This study extends these previous estimates over an additional three years and tests for interannual variations and long-term trends. We further examine the proportions of pups to adult females observed on the rookery and nearby haulouts during the birthing season to assess whether census data can be used to estimate natality. Open robust design multistate models were built and tested using Program MARK to estimate survival, resighting, and state transition probabilities in addition to other parameters dependent on whether or not a female gave birth in the previous year. Natality was estimated at 70% with some evidence of interannual variation but a long-term increasing or decreasing trend was not supported by the data. Bootstrap and regression comparisons of census data with natality estimates revealed no correlation between the two methods suggesting that census data are not an appropriate proxy for natality in this species. Longitudinal studies of individual animals are an appropriate method for estimating vital rates in species with variable detection over time such as the Steller sea lion. This work indicates that natality remains high in this region and is consistent with a population in recovery.

Climate change, pink salmon, and the nexus between bottom-up and top-down forcing in the subarctic Pacific Ocean and Bering Sea.

Climate change in the last century was associated with spectacular growth of many wild Pacific salmon stocks in the North Pacific Ocean and Bering Sea, apparently through bottom-up forcing linking meteorology to ocean physics, water temperature, and plankton production. One species in particular, pink salmon, became so numerous by the 1990s that they began to dominate other species of salmon for prey resources and to exert top-down control in the open ocean ecosystem. Information from long-term monitoring of seabirds in the Aleutian Islands and Bering Sea reveals that the sphere of influence of pink salmon is much larger than previously known. Seabirds, pink salmon, other species of salmon, and by extension other higher-order predators, are tightly linked ecologically and must be included in international management and conservation policies for sustaining all species that compete for common, finite resource pools. These data further emphasize that the unique 2-y cycle in abundance of pink salmon drives interannual shifts between two alternate states of a complex marine ecosystem.

The sun, moon, wind, and biological imperative-shaping contrasting wintertime migration and foraging strategies of adult male and female northern fur seals (Callorhinus ursinus).

Adult male and female northern fur seals (Callorhinus ursinus) are sexually segregated in different regions of the North Pacific Ocean and Bering Sea during their winter migration. Explanations for this involve interplay between physiology, predator-prey dynamics, and ecosystem characteristics, however possible mechanisms lack empirical support. To investigate factors influencing the winter ecology of both sexes, we deployed five satellite-linked conductivity, temperature, and depth data loggers on adult males, and six satellite-linked depth data loggers and four satellite transmitters on adult females from St. Paul Island (Bering Sea, Alaska, USA) in October 2009. Males and females migrated to different regions of the North Pacific Ocean: males wintered in the Bering Sea and northern North Pacific Ocean, while females migrated to the Gulf of Alaska and California Current. Horizontal and vertical movement behaviors of both sexes were influenced by wind speed, season, light (sun and moon), and the ecosystem they occupied, although the expression of the behaviors differed between sexes. Male dive depths were aligned with the depth of the mixed layer during daylight periods and we suspect this was the case for females upon their arrival to the California Current. We suggest that females, because of their smaller size and physiological limitations, must avoid severe winters typical of the northern North Pacific Ocean and Bering Sea and migrate long distances to areas of more benign environmental conditions and where prey is shallower and more accessible. In contrast, males can better tolerate often extreme winter ocean conditions and exploit prey at depth because of their greater size and physiological capabilities. We believe these contrasting winter behaviors 1) are a consequence of evolutionary selection for large size in males, important to the acquisition and defense of territories against rivals during the breeding season, and 2) ease environmental/physiological constraints imposed on smaller females.

High natality rates of endangered Steller sea lions in Kenai Fjords, Alaska and perceptions of population status in the Gulf of Alaska.

Steller sea lions experienced a dramatic population collapse of more than 80% in the late 1970s through the 1990s across their western range in Alaska. One of several competing hypotheses about the cause holds that reduced female reproductive rates (natality) substantively contributed to the decline and continue to limit recovery in the Gulf of Alaska despite the fact that there have been very few attempts to directly measure natality in this species. We conducted a longitudinal study of natality among individual Steller sea lions (n = 151) at a rookery and nearby haulouts in Kenai Fjords, Gulf of Alaska during 2003-2009. Multi-state models were built and tested in Program MARK to estimate survival, resighting, and state transition probabilities dependent on whether or not a female gave birth in the previous year. The models that most closely fit the data suggested that females which gave birth had a higher probability of surviving and giving birth in the following year compared to females that did not give birth, indicating some females are more fit than others. Natality, estimated at 69%, was similar to natality for Steller sea lions in the Gulf of Alaska prior to their decline (67%) and much greater than the published estimate for the 2000s (43%) which was hypothesized from an inferential population dynamic model. Reasons for the disparity are discussed, and could be resolved by additional longitudinal estimates of natality at this and other rookeries over changing ocean climate regimes. Such estimates would provide an appropriate assessment of a key parameter of population dynamics in this endangered species which has heretofore been lacking. Without support for depressed natality as the explanation for a lack of recovery of Steller sea lions in the Gulf of Alaska, alternative hypotheses must be more seriously considered.

Fatty acid signatures of stomach oil and adipose tissue of northern fulmars (Fulmarus glacialis) in Alaska: implications for diet analysis of Procellariiform birds.

Procellariiforms are unique among seabirds in storing dietary lipids in both adipose tissue and stomach oil. Thus, both lipid sources are potentially useful for trophic studies using fatty acid (FA) signatures. However, little is known about the relationship between FA signatures in stomach oil and adipose tissue of individuals or whether these signatures provide similar information about diet and physiology. We compared the FA composition of stomach oil and adipose tissue biopsies of individual northern fulmars (N = 101) breeding at three major colonies in Alaska. Fatty acid signatures differed significantly between the two lipid sources, reflecting differences in dietary time scales, metabolic processing, or both. However, these signatures exhibited a relatively consistent relationship between individuals, such that the two lipid sources provided a similar ability to distinguish foraging differences among individuals and colonies. Our results, including the exclusive presence of dietary wax esters in stomach oil but not adipose tissue, are consistent with the notion that stomach oil FA signatures represent lipids retained from prey consumed during recent foraging and reflect little metabolic processing, whereas adipose tissue FA signatures represent a longer-term integration of dietary intake. Our study illustrates the potential for elucidating short- versus longer-term diet information in Procellariiform birds using different lipid sources.