MANAGING THE EXPLOITATION OF PACIFIC WALRUSES: A TRAGEDY OF DELAYED RESPONSE AND POOR COMMUNICATION1

The Pacific walrus population has been depleted and subsequently allowed to recover three times in the past 150 yr. As we see it, the population has been made to fluctuate like an r-selected species, rather than being maintained at a high, stable level, as befits a K-selected species. The latest depletion began in the 1930s but was not recognized until 25 yr later, by which time the population had been reduced by at least half. Without benefit of communication, the U.S.S.R. and the State of Alaska put similar protective measures into place by 1960, and in the next two decades the walrus population recovered again, at least doubling in size. By 1980, it already was showing density-dependent signs of having approached or reached the carrying capacity of its environment. As productivity and calf survival declined sharply in the late 1970s and early 1980s the catches more than doubled. We believe that the combined effects of natural curtailment and human intervention may be bringing the population down again rather rapidly. With the present, crude monitoring methods, delayed management responses, and poor international communications, however, the downward trend may not be acknowledged for at least another decade, by which time the unilateral Soviet and American corrective measures are likely to be too much, too late. Walrus management needs to be based less on response to immediate crisis and more on long term prediction than it has been in the past. Because the U.S.A. and U.S.S.R. are trying to manage the same walrus population, without sufficient communication or consensus and sometimes to opposite ends, an international joint management program needs to be implemented.     

INVESTIGATING TROPHIC RELATIONSHIPS OF PINNIPEDS IN ALASKA AND WASHINGTON USING STABLE ISOTOPE RATIOS OF NITROGEN AND CARBON

We measured stable-nitrogen (δ¹⁵N) and stable-carbon (δ¹³C) isotope ratios in muscle and hair from 7 northern fur seals (Callorhinus ursinus) from the Pribilof Islands, Alaska, and 27 Steller sea lions (Eumetopias jubatus), and 14 harbor seals (Phoca vitulina) from the Gulf of Alaska and coast of Washington State, in order to contrast dietary information derived from isotopic vs. available conventional dietary studies. Stable-nitrogen-isotope analysis of muscle revealed that harbor seals were enriched over sea lions (mean δ¹⁵N = 18.6‰vs. 17.5‰) which were in turn enriched over northern fur seals (mean δ¹⁵N = 16.6‰). Trophic segregation among these species likely results primarily from differential reliance on herring (Clupea harengus), Atka mackerel (Pleurogrammus monopterygius), and large vs. small walleye pollock (Theregra chalcogramma). According to their δ¹⁵N values, adult male Steller sea lions showed a higher trophic position than adult females (mean δ¹⁵N: 18.0‰vs. 17.2‰), whereas adult female northern fur seals were trophically higher than juvenile male fur seals (mean δ¹⁵N: 16.5‰vs. 15.0‰). Each of these observed differences likely resulted from differential reliance on squid or differences in the size range of pollock consumed. Three northern fur seal pups showed higher δ¹⁵N enrichment over adults (mean 17.7‰vs. 15.8‰) due to their reliance on their mother's milk. Stable-carbon isotope measurements of hair revealed a cline toward more negative values with latitude. Segregation in hair δ¹³C between Steller sea lions and harbor seals off the coast of Washington (mean δ¹³C: ?13.6‰vs.?15.0‰) reflected the greater association of harbor seals with freshwater input from the Columbia River. Our study demonstrates the utility of the stable isotope approach to augment conventional dietary analyses of pinnipeds and other marine mammals.     

SEASONAL DISTRIBUTION OF STELLER'S SEA LIONS AT ROOKERIES AND HAUL-OUT SITES IN ALASKA

The Steller's sea lion population has declined by 60%-70% over much of Alaska since the late 1970s. Overlap in species composition and sizes of fishes consumed by sea lions and harvested by commercial fisheries, particularly during winter, has led to examination of potential interaction between commercial fisheries and Steller's sea lions. Abundance and distribution data for Steller's sea lions in Alaska were derived from aerial surveys conducted during the breeding season, mid-June to early July 1992, 1994, and 1996. To study winter distribution of sea lions, we conducted aerial surveys during March 1993, November-December 1994, and March 1999. We counted about one-half as many sea lions during winter surveys compared to the breeding-season surveys. Numbers of sea lions at rookery sites dropped off considerably during winter, whereas numbers at haul-out sites did not. We found little evidence of large-scale, seasonal movement, at least for the western stock of sea lions. Rather, differences between summer and winter distribution were primarily a function of sea lions dispersing to local haul-out sites during the winter. Terrestrial sites, both rookeries and haul-outs, clearly are important to Steller's sea lions during the entire year. Individual sites may be occupied year-round or only during particular times of year.     

PROJECTING THE TREND OF STELLER SEA LION POPULATIONS IN WESTERN ALASKA

This paper attempts to project the trends of Steller sea lion ( Eumetopias jubatus ) populations in six subdivisions of the western Alaska population. The overall Western Alaska population has declined dramatically since the 1970s. Trends in half of the areas appear to have leveled-off and possibly to be on the increase. Bootstrapping has been used to provide confidence intervals on predictions for the 2004 counts. For the three areas in which we expect increases, the 95% confidence intervals on predictions were: Eastern Gulf (2,430–3,740), Central Gulf (3,260–3,660) and Central Aleutians (5,160–6,580). The Western Gulf counts have been somewhat erratic, with a gradual rate of decrease (about 2% per year) and wide confidence limits on a linear prediction (logarithmic scale) of 2,690–3,240. Trends in the Eastern Aleutians have been even more erratic, so that about all that can be inferred is that the population may be roughly stabilized. Only the Western Aleutians appear to be rapidly declining at about 10% per year, with a 95% confidence interval on a linear trend of 610–1,100. The predictions were made before the 2004 counts and are in reasonable accord with the 2004 counts. Age structure changes do not appeat to provide a viable explanation for the changing trends.     

STABLE ISOTOPE ANALYSES OF TOOTH ANNULI REVEAL TEMPORAL DIETARY RECORDS: AN EXAMPLE USING STELLER SEA LIONS

Stable isotope analysis of teeth of marine mammals can provide valuable information on trophic level and source of feeding. However, the isotopic analysis of whole teeth presents only an average dietary estimate for individuals across the period of growth of that tooth. While such analyses can be valuable, particularly in the case of fossil material, in contrast, isotopic analysis of individual annuli of teeth can provide dietary information for each year of tooth growth, in some cases representing the whole of the animal's life. We measured stable-carbon isotope ratios (¹³C/¹²C) in the inorganic (hydroxyapatite) and stable-nitrogen isotope ratios (¹⁵N/¹⁴N) in the organic (primarily collagenous) components of individual tooth annuli of 18 male Steller sea lions (Eumetopias jubatus) obtained from archived collections from the Bering Sea and Gulf of Alaska and from single northern fur seals (Callorbinus ursinus) and northern elephant seals (Mirounga angustirostris) from the central Aleutian Islands and eastern Gulf of Alaska, respectively. In several individuals, we detected considerable variation in stable isotope values among annuli, up to 6.1%_O for δ¹⁵N and 5.1%_O for δ¹³C values. Enrichment in δ¹⁵N and depletion of δ¹³C values in the first annulus may correspond to dietary inputs from mother's milk during the period of suckling. Other variations among years may be caused by dietary changes or movements of individuals between regions differing in isotopic signatures of foodweb primary production. Our study indicates that the isotopic signatures of foodweb primary production. Our study indicates that the isotopic analysis of individual tooth annuli represents a fine-seale tool for dietary reconstructions involving marine mammals, and cautions against the use of whole-tooth material averaged over several annuli.     

DISPERSAL, ROOKERY FIDELITY, AND METAPOPULATION STRUCTURE OF STELLER SEA LIONS (EUMETOPIAS JUBATUS) IN AN INCREASING AND A DECREASING POPULATION IN ALASKA

Over the past 24 yr, 8,596 Steller sea lion ( Eumetopias jubatus ) pups were branded on their natal rookeries throughout Alaska with the objectives of determining survival rates, recruitment, movements, and site fidelity. Our objectives here were to examine the extent of dispersal of Steller sea lions away from their natal rookeries, movements between stocks, and degree of natal rookery fidelity. Pups (<1 yr old) usually remained within 500 km of their natal rookery. Branded juveniles dispersed widely and were resighted at distances up to 1,785 km from their natal rookeries. Adults generally remained within 500 km of their natal rookeries. No interchange of breeding animals between the ES (eastern stock) and WS (western stock) was observed. Although natal rookery fidelity was prevalent, 33% of the 12 observations of females branded in the WS during 1987–1988 and 19% of the 29 observations of females branded in the ES during 1994–1995 were observed with newly born pups at sites other than their natal rookeries. Steller sea lions generally conformed to the metapopulation concept as depicted by Hanski and Simberloff (1997), with local breeding populations (rookeries) and movements among these local populations having the potential of affecting local dynamics.