COST OF TRANSPORT IN STELLER SEA LIONS, EUMETOPIAS JUBATUS

The cost of swimming is a key component in the energy budgets of marine mammals. Unfortunately, data to derive predictive allometric equations are limited, and estimates exist for only one other species of otariid. Our study measured the oxygen consumption of three juvenile Steller sea lions ( Eumetopias jubatus ) swimming in a flume tank at velocities up to 2.2 m sec⁻¹. Minimum measured cost of transport ranged from 3.5–5.3 J kg⁻¹ m⁻¹, and was reached at swimming speeds of 1.7–2.1 m s⁻¹. These cost-of-transport values are higher than those reported for other marine mammals. However, once differences in stationary metabolic rate were accounted for, the locomotor costs (LC) for the Steller sea lions were commensurate with those of other marine mammals. Locomotor costs (LC in J m⁻¹) appeared to be directly proportional to body mass (M in kg) such that LC = 1.651M^1.01. These estimates for the cost of locomotion can be incorporated into bioenergetic models and used to determine the energetic consequences of observed swimming behavior in wild marine mammals.     

GEOGRAPHIC VARIATION IN SKULL MORPHOLOGY OF ADULT STELLER SEA LIONS (EUMETOPIAS JUBATUS)

Data from cranial specimens of adult E. jubatus were analyzed to compare intraspecific morphology of skulls. Males and females were grouped separately to avoid bias from sexual dimorphism. Geographic variation was observed in adult male E. jubutus , indicating the potential presence of three morphologically disparate groups: those from Alaska, those from California, and those from Japan and Russia. Although sample sizes were small, results from cluster and discriminant function analyses indicated that specimens from eastern and western Alaska were morphologically similar, and that the most divergent specimens for the species appeared to be those from Japan. Skulls from Alaska possessed a typically longer, less robust skull, whereas those from Japan appeared smaller, yet most robust. Skulls from California were intermediate.     

RANGE-WIDE SURVEY AND ESTIMATION OF TOTAL NUMBER OF STELLER SEA LIONS IN 1989

Abstract: The first range-wide survey of Steller (northern) sea lions ( Eumetopias jubatus ) was completed in 1989 with a total of 68,094 adult and juvenile (nonpup) Steller sea lions counted. This total count includes 10,000 in Russia (15% of the range-wide count), 47,960 in Alaska (70%), 6,109 in British Columbia (9%), 2,261 in Oregon (3%), and 1,764 in California (3%). A range-wide pup count was not obtained. We estimated the 1989 world population based on a calculation for total pups and obtained a range-wide estimate of 116,000 total animals, or about 39–48% of the 240,000-300,000 estimated 30 yr ago.     

A FIELD-BASED METHOD FOR ESTIMATING AGE IN FREE-RANGING STELLER SEA LIONS (EUMETOPIAS JUBATUS) LESS THAN TWENTY-FOUR MONTHS OF AGE

Studies of health, survival, and development of juvenile Alaskan Steller sea lions ( Eumetopias jubatus , SSL) require accurate estimates of age for wild-captured animals. However, the value and accuracy of several potential predictors of age have not been assessed with data from known-age free-ranging animals. During 2001–2005, forty-six individual SSL originally branded or tagged at ≤6 mo of age were recaptured by the Alaska Department of Fish and Game (ADF&G). Using a series of general linear models, we evaluated the ability of morphometrics measurements: permanent canine tooth length (CTL), diastema (DIAS), whisker length (WHIS), and dorsal standard length (DSL) to predict the age of forty-six known-age juveniles ( n = 46 ≤23 mo of age). Permanent CTL was the strongest individual predictor ( r ²= 0.80); followed by DSL, DIAS, and WHIS ( r ²= 0.70, 0.56, and 0.45, respectively). The inclusion of a single sample from a 44-mo-old sea lion suggested quadratic relationships between age and all predictors for older animals. Only models including CTL predicted age to within 6 mo of known age. The equation Age = (−3.0112 +[0.6726 * CTL]+[0.4965 * DIAS]) allows for accurate age estimates of SSL ≤23 mo for both sexes.     

QUANTIFICATION OF TERRESTRIAL HAUL-OUT AND ROOKERY CHARACTERISTICS OF STELLER SEA LIONS

Steller sea lions ( Eumetopias jubatus ) are known to have occupied the same terrestrial haul-out and rookery sites across the North Pacific Rim for centuries, but it is not known why they choose and stay at these locations, or what defines their preferred habitat. Classifying and comparing the shoreline type of haul-outs and rookeries against sites not used by Steller sea lions showed that they preferentially locate their haul-outs and rookeries on exposed rocky shorelines and wave-cut platforms. However, no preference was found for selecting rookeries on sheltered shore types. Shoreline types used less frequently by sea lions included fine-to-medium-grained sand beaches, mixed sand and gravel beaches, gravel beaches, and sheltered rocky shores. Quantifying the shoreline types used by sea lions confirms anecdotal reports of habitat preferences and may prove useful in identifying and protecting sea lion terrestrial habitat, or in forecasting how climate change might affect the distribution of sea lions.     

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.     

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.     

CLASSIFYING PREY HARD PART STRUCTURES RECOVERED FROM FECAL REMAINS OF CAPTIVE STELLER SEA LIONS (EUMETOPIAS JUBATUS)

Feces were collected from six Steller sea lions ( Eumetopias jubatus ) that consumed known amounts of Atka mackerel ( Pleurogrammus monopterygius ), Pacific herring ( Clupea barengus ), pink salmon ( Oncorhynchus gorbuscha ), walleye pollock ( Theragra chalcogramma ), and squid ( Loligo opalacens ). The goal was to determine the numbers and types of taxon-specific hard parts that pass through the digestive tract and to develop correction factors for certain abundantly occurring structures. Over 20,000 fish and squid were consumed during 267 d of fecal collection. During this period, over 119,000 taxon-specific hard parts, representing 56 different structures, were recovered. Skeletal structures and non-skeletal structures accounted for 72% and 28% of all hard parts, respectively. The branchiocranium, axial skeleton, and dermocranium regions of the skeletal system accounted for the greatest number of hard parts recovered. Over 70% of all recovered hard parts were represented by one to six taxa specific structures for each prey type. The average number of hard parts (3.1–31.2) and structure types (2.0–17.7) recovered per individual prey varied across taxa and were used to derive correction factors (to reconstruct original prey numbers). A measure of the variability of hard part recovery among sea lions showed no difference for certain herring, pollock, and squid structures, however, there was a significant difference for salmon and Atka mackerel structures. Identifying all taxon- specific prey hard parts increases the likelihood of identifying and estimating the number of prey consumed.     

MASS ESTIMATES USING BODY MORPHOLOGY IN STELLER SEA LIONS

Abstract: Analytical procedures developed from studies on phocids for relating body morphometrics (length and girth) to mass were applied to 390 Steller sea lions from the Gulf of Alaska and the Bering Sea. The equations relating body volume to mass were significant and had very high correlation coefficients for animals from about 60 to over 900 kg with a standard error of ±0.6% in fitted mass. There was a significant difference in these relationships for female sea lions studied in the 1970s and those examined in the 1980s. The latter sea lions were smaller and/or leaner than those collected in the 1970s.     

THE RELIABILITY OF SKINFOLD-CALIPERS FOR MEASURING BLUBBER THICKNESS OF STELLER SEA LION PUPS (EUMETOPIAS JUBATUS)

Twelve dead Steller sea lion pups ( Eumetopias jubatus ) aged 3-14 d were recovered from rookeries in Southeast Alaska. They had a wide range of body sizes and conditions (small to large and fat to no fat). The ability of calipers to estimate the thickness of their blubber layer was assessed with a set of skinfold calipers. Average error of measurement for skin and blubber thickness was an acceptable 5.4%, but the skin and blubber of the pups were highly compressible. Skinfold thickness increased with body mass but did not necessarily reflect the development of blubber, given that pups with no blubber also showed an increase in skinfold thickness with increases in body mass. Skinfold thickness of sea lion pups appears to predict body size better than it predicts blubber thickness, making it difficult if not impossible to develop a simple index of body condition or a calculation of percent body fat for Steller sea lion pups from skinfold caliper measurements.     

REDUCED BODY SIZE OF FEMALE STELLER SEA LIONS FROM A DECLINING POPULATION IN THE GULF OF ALASKA

Nutritional stress is a leading hypothesis behind the decline in numbers of Steller sea lions in the Gulf of Alaska, the Aleutian Islands, and the Bering Sea. To evaluate this hypothesis we compared body growth of female Steller sea lions 1.0–13.9 yr of age collected in the Gulf of Alaska during two time periods, 1975–1978 just prior to or early in the decline and 1985–1986 when the decline was well established. We found that growth, as measured by standard length, axillary girth, and mass, was reduced during the 1980s, supporting the undernutrition hypothesis. We also found a suggestion of reduced growth in our 1970s and 1980s samples when compared to a collection of Steller sea lions obtained from the Gulf of Alaska in 1958. However, no direct link has been demonstrated between undernutrition and the actual decline in numbers.     

RISK OF EXTIRPATION OF STELLER SEA LIONS IN THE GULF OF ALASKA AND ALEUTIAN ISLANDS: A POPULATION VIABILITY ANALYSIS BASED ON ALTERNATIVE HYPOTHESES FOR WHY SEA LIONS DECLINED IN WESTERN ALASKA

We estimated the risk that the Steller sea lion will be extirpated in western Alaska using a population viability analysis (PVA) that combined simulations with statistically fitted models of historical population dynamics. Our analysis considered the roles that density‐dependent and density‐independent factors may have played in the past, and how they might influence future population dynamics. It also established functional relationships between population size, population growth rate and the risk of extinction under alternative hypotheses about population regulation and environmental variability. These functional relationships can be used to develop recovery criteria and guide research and management decisions. Life table parameters (e.g., birth and survival rates) operating during the population decline (1978–2002) were estimated by fitting simple age‐structured models to time‐series of pup and non‐pup counts from 33 rookeries (subpopulations). The PVA was carried out by projecting all 33 subpopulations into the future using these estimated site‐specific life tables (with associated uncertainties) and different assumptions about carrying capacities and the presence or absence of density‐dependent population regulation. Results suggest that the overall predicted risk of extirpation of Steller sea lions as a species in western Alaska was low in the next 100 yr under all scenarios explored. However, most subpopulations of Steller sea lions had high probabilities of going extinct within the next 100 yr if trends observed during the 1990s were to continue. Two clusters of contiguous subpopulations occurring in the Unimak Pass area in the western Gulf of Alaska/eastern Aleutian Islands and the Seguam–Adak region in the central Aleutian Islands had relatively lower risks of extinction. Risks of extinction for a number of subpopulations in the Gulf of Alaska were reduced if the increases observed since the late 1990s continue into the future. The risks of subpopulations going extinct were small when density‐dependent compensation in birth and survival rates was assumed, even when random stochasticity in these vital rates was introduced.     

ASSESSING KILLER WHALE PREDATION ON STELLER SEA LIONS FROM FIELD OBSERVATIONS IN KENAI FJORDS, ALASKA

The behavioral and predatory patterns of Gulf of Alaska (GOA) transient killer whales ( Orcinus orca ) were studied between 2000 and 2005 using remote video and vessel-based observations near the Chiswell Island Steller sea lion ( Eumetopias jubatus ) rookery and in the broader Kenai Fjords (KF) region of the northern GOA. GOA transient killer whales were observed on 118 d over the 6-yr period; the median group size was two (range: 1–9). Nine predation events were observed from vessels and an additional sixteen were inferred from remote video studies; all involved Steller sea lions. Estimates from field observations suggest that fifty-nine sea lions were consumed over the summer seasons of 2002–2005; whereas estimates based on published caloric requirements of transient killer whales would suggest a loss of 103 sea lions over the same time period. GOA transients spent a large proportion (43%) of their time resting which may be a strategy for conserving energy. Predation on sea lion pups at the Chiswell Island rookery was greatest during years when a single killer whale was foraging alone and when a 1.5-yr-old calf was evidently being trained to handle prey. Predation on pups was low during years when killer whales were foraging in groups and were observed and presumed to be taking mostly juvenile sea lions. Our study suggests that GOA transients are having a minor effect on the recovery of Steller sea lions in the GOA.     

DEVELOPMENT OF DISPERSAL, MOVEMENT PATTERNS, AND HAUL-OUT USE BY PUP AND JUVENILE STELLER SEA LIONS (EUMETOPIAS JUBATUS) IN ALASKA

Population 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.     

STELLER SEA LION BODY CONDITION INDICES

We evaluated various measurements of mass, morphology, and blubber thickness as indices of fatness for Steller sea lions by correlation with the percentage of total body mass comprised by the sculp (%SCULP). We concluded LMD-index was the best index evaluated because it had a relatively high r ² (0.58), had a linear relationship with %SCULP, and the intercept term was not different from O. We suggest the development of a LMD-index for otariids would likely reduce the unexplained variation in the index. We developed a multiple regression model ( r ²= 0.745, P < 0.001) for predicting %SCULP with LMD-index and functions of sex, age, and season as predictor variables. Steller sea lions < 5 yr of age had higher %SCULP values than those ≥ 5 yr. %SCULP declined with age for sea lions < 5 yr. Both younger and older males were fatter during the winter/spring period than during summer/ fall. Females of both age classes had similar %SCULP values throughout the year. Steller sea lions are relatively lean pinnipeds; estimates of blubber and total body lipids ranged from 5% to 17% of total body mass.     

COUNTING STELLER SEA LION PUPS IN ALASKA: AN EVALUATION OF MEDIUMFORMAT, COLOR AERIAL PHOTOGRAPHY

Estimates of Steller sea lion ( Eumetopias jubatus ) pup production are valuable for estimating population trend and size. Currently in Alaska, pups are counted by visiting rookeries, driving older animals into the water, then walking through the rookeries and counting the pups, a highly disruptive procedure. At smaller rookeries, with good vantage points, pups are occasionally counted from the periphery of rookeries without disturbing the sea lions. We evaluated counts made from medium-format, color, aerial photographs as an alternative to drive counts and peripheral counts. Neither the peripheral counts nor the aerial photographic counts disturbed animals on the rokeries. There were strong 1:1 linear relationships between photographic counts and drive counts ( r ²= 0.966, P < 0.001) and between photographic counts and peripheral counts ( r ²= 0.999, P < 0.001). Precision was similar for all three methods of counting. We suggest that medium-format, color, aerial photographs is appropriate for routine surveys of Steller sea lion pups in Alaska because it is not disruptive to the hauled-out sea lions and provides comparable estimates with similar precision to drive and peripheral counts. Large areas canbe rapidly surveyed during periods of good weather with a minimum of manpower.     

Neonatal survival of Steller sea lions (Eumetopias jubatus)

Neonatal survival of Steller sea lions ( Eumetopias jubatus ) are often considered inconsequential to their population dynamics. However, observations of dead animals on rookeries and in surrounding waters suggest that early mortality is not uncommon. This study used the natural markings of adult females in a mark and resighting framework to estimate the apparent survival (φ) of pups with the Cormack–Jolly–Seber model at two sites on Lowrie Island, Alaska from birth to 3 wk old. Estimates varied greatly by site and year; 2002 Area 5:     (95% CI: 0.199, 0.684; n = 21), 2002 Area 1:     (0.437, 0.916; n = 21), 2003 Area 5:     (0.414, 0.738; n = 56), and 2003 Area 1:     (0.695, 0.997; n = 32). The mean estimate across the four area × year combinations was     (0.569, 0.772). Survival was lowest on the first day of life and then leveled off at a higher rate. None of the four environmental covariates we considered (swell height, interaction of tide and swell heights, density, or birth date) were significantly related to neonatal survival. Our results suggest that estimates of first-year survival that do not account for mortality prior to dispersal from the natal rookery may significantly overestimate survival rate.     

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.     

STELLER SEA LION STATUS AND TREND IN SOUTHEAST ALASKA: 1979–1997

Steller sea lion (Eumetopias jubatus) numbers in the United States declined by about 75% over the past 20+ yr. They are classified, under the U. S. Endangered Species Act, as “threatened” in the eastern portion of their range and as “endangered” in the western portion. We analyzed trends in numbers of pup and non-pup Steller sea lions counted in Southeast Alaska between 1979 and 1997. Sea lion numbers, based on counts of pups on rookeries, increased by an average of 5.9% per year between 1979 and 1997. However, numbers of pups increased at a much slower rate (+ 1.7% per year) between 1989 and 1997. For counts of non-pup Steller sea lions we used models that controlled for the effects of date, time, and tide at the time of the survey to analyze trends. This technique reduced bias and increased precision of the resulting trend estimates. Numbers of sea lions were stable (+0.5%) between 1989 and 1996, based on counts of non-pups. We estimated the Southeast Alaska breeding population of Steller sea lions at about 19,000 animals of all ages in 1997, a level that is probably near the highest in recorded history.     

BEHAVIORAL AND PHYSIOLOGICAL MEASUREMENTS OF MATERNAL INVESTMENT IN THE STELLER SEA LION, EUMETOPIAS JUBATUS

The onshore and at-sea cycles of females, suckling behavior of pups and their milk intake were studied in Steller sea lions ( Eumetopias jubatus ) during 1983 at Año Nuevo Island, California. Females averaged approximately 21 h ashore and 36 h at sea. The trips to sea lengthened as pups aged, resulting in an overall decline in female time ashore to 30% by the sixth week following parturition. Activity budgets of pups showed no significant differences among suckling time, age and sex. Milk intake, estimated using labeled water studies, revealed that heavier pups consumed more milk than lighter ones (milk ingestion in ml/d = 4.26 + 0.0687 [Pup Mass in kg]). Mean milk intake was 1.78 ± 0.33 liters/d. Mean pup growth rate was 0.38 ± 0.1 kg/d. The results suggest that female attendance patterns are shaped by the increasing nutritional demands of growing pups and their increasing efficiency at suckling.