FORAGING STRATEGIES OF SYMPATRIC KILLER WHALE (ORCINUS ORCA) POPULATIONS IN PRINCE WILLIAM SOUND, ALASKA

Killer whales ( Orcinus orca ) feed on a wide variety of fish, cephalopods, and marine mammals throughout their cosmopolitan range; however, the dietary breadth that characterizes the species is not reflected in all populations. Here, we present the findings of a 14-yr study of the diet and feeding habits of killer whales in Prince William Sound, Alaska. Two non-associating forms of killer whale, termed resident and transient (Bigg et al. 1987), were identified. All prey seen taken by transients were marine mammals, including harbor seals ( Phoca vitulina ), Dall's porpoises ( Phocoenoides dalli ), Steller sea lions ( Eumetopias jubatus ), and harbor porpoises ( Phocoena phocoena ). Resident killer whales appeared to prey principally on salmon ( Oncorhynchus spp.), preferring coho salmon ( O. kisutch ) over other, more abundant salmon species. Pacific herring ( Clupea pallasi ) and Pacific halibut ( Hippocampus stenolepis ) were also taken. Resident killer whales frequently were seen to interact in non-predatory ways with Steller sea lions and Dall's porpoises, while transients were not. Differences in the social organization and behavior of the resident and transient killer whales in Prince William Sound are discussed in the light of the dietary differences documented here.     

Total mercury in stranded marine mammals from the Oregon and southern Washington coasts

Muscle samples from 105 marine mammals stranded along the Oregon and Washington coasts (2002–2009) were tested for levels of total mercury (THg) by Cold Vapor Atomic Fluorescence Spectrometry. The THg present is in the form of the highly toxic methylmercury. After normalizing tissue to 75% water weight, Steller sea lions and northern elephant seals exhibited the highest mean concentrations of THg followed by harbor seals, harbor porpoises, and California sea lions, 0.34 ± 0.278, 0.34 ± 0.485, 0.21 ± 0.216, 0.17 ± 0.169, and 0.15 ± 0.126 mg/kg normalized wet weight (ww), respectively. The mean normalized values demonstrate limited muscle methylmercury accumulation in these species in the Pacific Northwest. However, actual ww concentrations in some of the stranded carcasses may pose a risk to scavengers. Normalizing muscle mercury concentrations eliminated the variability from desiccation, and allowed for a clearer indication of the amount of mercury the animal accumulated before stranding.     

Organochlorine exposure and health effects in stranded Steller sea lions (Eumetopias jubatus) and Pacific harbor seals (Phoca vitulina richardii) from Oregon and southern Washington coasts

Organochlorine (OC) pesticides and polychlorinated biphenyls (PCBs) have been detected in a variety of marine mammal species at levels associated with adverse health effects. Little is known about OC levels and impacts on health in pinnipeds with different life histories. We determined the health and levels of 18 OC pesticides and 16 PCB congeners in blubber samples from 20 Steller sea lions and 39 Pacific harbor seals stranded from Oregon and Southern Washington. The most commonly detected OC at the highest concentration was p,p′- dichlorodiphenyldichloroethylene (DDE). PCBs were detected in all samples as well. Hypothesis testing indicated that diseased Steller sea lions (males and females combined) had higher contaminant concentrations than healthy Steller sea lions, and diseased Pacific harbor seals had higher concentrations of total OCs than healthy animals. Differences were also noted between diseased and healthy animals when looking at individual sexes of each species. Diseased Steller sea lions had higher mean contaminant levels than diseased harbor seals and healthy Steller sea lions had higher mean contaminant concentrations than healthy Pacific harbor seals. These results show that species differences exist in both contaminant loads and sensitivity to contaminants, which may be due to differences in life histories and physiology.     

The role of Steller sea lions in a large population decline of harbor seals

We provide the first direct evidence that Steller sea lions will prey on harbor seals. Direct observations of predation on marine mammals at sea are rare, but when observed rates of predation are extrapolated, predation mortality may be found to be significant. From 1992 to 2002, harbor seals in Glacier Bay declined steeply, from 6,200 to 2,500 (∼65%). After documenting that Steller sea lions were preying on seals in Glacier Bay, we investigated increased predation by sea lions as a potential explanation for the large decline. In five independent data sets spanning 21–25 yr and including 14,308 d of observations, 13 predation events were recorded. We conducted a fine-scale analysis for an intensively studied haul-out (Spider Island) and a broader analysis of all of Glacier Bay. At Spider Island, estimated predation by sea lions increased and could account for the entirety of annual pup production in 5 of 8 yr since 1995. The predation rate, however, was not proportional to the number of predators. Predation by Steller sea lions is a new source of mortality that contributed to the seal declines; however, life history modeling indicates that it is unlikely that sea lion predation is the sole factor responsible for the large declines.     

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.     

Prevalence and distribution of serum neutralizing antibodies to Tillamook (bovine) calicivirus in selected populations of marine mammals

Neutralizing antibodies to Tillamook calicivirus (TCV) were found in sera collected from California sea lions (Zalophus c. californianus Lesson) in 1983 and 1984 and in sera collected from Steller sea lions (Eumetopias jubatus Schreber) in 1976 and 1985. The combined prevalence of antibodies for these two species was 10/228 = 4.38%. Titers ranged from 1:20 (five animals), to 1:40 (four animals), to 1:80 (one animal) by standard microtiter neutralization assay. The seropositive pinnipeds were dispersed widely along the margins of the eastern Pacific rim, from the Bering Sea to the Santa Barbara Channel. Antibodies to TCV were not found in sera collected from northern fur seals (Callorhinus ursinus L.), Pacific walruses (Odobenus rosmarus divergens Illiger), seals of the family Phocidae, or several cetacean species. Tillamook calicivirus was isolated originally in 1981 from dairy calves in Oregon; the finding of neutralizing antibodies in two widely distributed species of sea lions suggests the possibility of a marine origin for this agent.     

Diet of Pacific sleeper shark, a potential Steller sea lion predator, in the north-east Pacific Ocean

Pacific sleeper sharks Somniosus pacificus were captured near Steller sea lion Eumetopias jubatus rookeries during the period when Steller sea lion pups are most vulnerable to Pacific sleeper shark predation (first water entrance and weaning). Analysis of stomach contents revealed that teleosts were the dominant prey in August and cephalopods were the dominant prey in May ( n = 198). Marine mammals were found in 15% of stomachs regardless of season, but no Steller sea lion tissues were detected. Molecular genetic analysis identified grey whale Eschrichtius robustus and harbour seal Phoca vitulina remains in some Pacific sleeper shark stomachs. Most mammals were cetacean and at least 70% of the cetaceans were probably scavenged. Although Pacific sleeper shark and Steller sea lion ranges overlapped, so predation could potentially occur, the diet study suggested that predation on Steller sea lions is unlikely, at least when pups first enter the water or during weaning. Harbour seals were infrequent prey and may have been consumed alive. Pacific sleeper sharks consume fast-swimming prey like Pacific salmon Oncorhynchus sp., most likely live animals rather than scavenged animals. Pacific sleeper sharks appeared to be opportunistic consumers of the available prey and carrion, feeding both on the bottom and in the water column, and their diet shifted to teleosts and cetacean carrion as the fish grew larger.     

Characterization and clinical manifestations of Arcanobacterium phocae infections in marine mammals stranded along the central California coast

Between 1994 and 2000, 141 Arcanobacterium phocae isolates were recovered from marine mammals that stranded along the central California coast (USA). Arcanobacterium phocae was cultured from tissue sites with abnormal discharge or evidence of inflammation in 66 California sea lions (Zalophus californianus), 50 Pacific harbor seals (Phoca vitulina richardii), 19 northern elephant seals (Mirounga angustirostris), five southern sea otters (Enhydra lutris nereis), and one common dolphin (Delphinus delphis). The overall prevalence of A. phocae among cultured stranded marine mammals was 8%. This is the first report of A. phocae in animals from the Pacific Ocean. Sequence analysis of a portion of the 16S ribosomal RNA gene confirmed recent isolates as A. phocae. Prior to phylogenetic testing and the routine use of the esculin hydrolysis and motility tests, A. phocae isolates may have been misidentified as Listeria ivanovii. Arcanobacterium phocae was commonly isolated from superficial abscesses, was often present in mixed infections, and was susceptible to all antimicrobial agents tested.     

Temporal records of δ13C and δ15N in North Pacific pinnipeds: inferences regarding environmental change and diet

Sea lion and seal populations in Alaskan waters underwent various degrees of decline during the latter half of the twentieth century and the cause(s) for the declines remain uncertain. The stable carbon (¹³C/¹²C) and nitrogen (¹⁵N/¹⁴N) isotope ratios in bone collagen from wild Steller sea lions (Eumetopias jubatus), northern fur seals (Callorhinus ursinus) and harbor seals (Phoca vitulina) from the Bering Sea and Gulf of Alaska were measured for the period 1951-1997 to test the hypothesis that a change in trophic level may have occurred during this interval and contributed to the population declines. A significant change in '¹⁵N in pinniped tissues over time would imply a marked change in trophic level. No significant change in bone collagen '¹⁵N was found for any of the three species during the past 47 years in either the Bering Sea or the Gulf of Alaska. However, the ¹⁵N in the Steller sea lion collagen was significantly higher than both northern fur seals and harbor seals. A significant decline in '¹³C (almost 2 ‰ over the 47 years) was evident in Steller sea lions, while a declining trend, though not significant, was evident in harbor seals and northern fur seals. Changes in foraging location, in combination with a trophic shift, may offer one possible explanation. Nevertheless, a decrease in '¹³C over time with no accompanying change in '¹⁵N suggests an environmental change affecting the base of the foodweb rather than a trophic level change due to prey switching. A decline in the seasonal primary production in the region, possibly resulting from decreased phytoplankton growth rates, would exhibit itself as a decline in '¹³C. Declining production could be an indication of a reduced carrying capacity in the North Pacific Ocean. Sufficient quantities of optimal prey species may have fallen below threshold sustaining densities for these pinnipeds, particularly for yearlings and subadults who have not yet developed adequate foraging skills.     

Beneath the surface: Profiling blubber depth in pinnipeds with infrared imaging

Infrared thermography (IRT) was assessed as a non-invasive tool to evaluate body condition in juvenile female harbor seals (Phoca vitulina), (n=6) and adult female Steller sea lions (Eumetopias jubatus), (n=2). Surface temperature determined by IRT and blubber depth assessed with portable imaging ultrasound were monitored concurrently at eight body sites over the course of a year in long-term captive individuals under controlled conditions. Site-specific differences in surface temperature were noted between winter and summer in both species. Overall, surface temperature was slightly higher and more variable in harbor seals (9.8±0.6 °C) than Steller sea lions (9.1±0.5 °C). Limited site-specific relationships were found between surface temperature and blubber thickness, however, insulation level alone explained a very small portion of the variance. Therefore, while validated IRT data collection can potentially provide valuable information on the health, condition and metabolic state of an animal, it cannot provide a generalized proxy for blubber depth.     

Mercury Concentrations in Hair from Neonatal and Juvenile Steller Sea Lions (Eumetopias jubatus): Implications Based on Age and Region in this Northern Pacific Marine Sentinel Piscivore

Mercury is a global contaminant of concern for the fetus and the neonate of piscivores. Methylmercury, produced within marine ecosystems, is of particular concern as a readily absorbed neurotoxicant transported across the blood brain barrier and transplacentally. In the North Pacific Ocean, Steller sea lions are broadly distributed apex predators and, as such, integrate complex food webs and the associated exposure and possible adverse effects of toxic and infectious agents. Hair, including lanugo, was examined using regional and age groupings to assess mercury concentrations in young Alaskan Steller sea lions. The highest concentrations of mercury occurred in the youngest animals, likely via in utero exposure. Based on the adverse developmental outcomes of methylmercury toxicity this specific cohort is of concern. Regionally, higher concentrations of mercury were observed in the endangered western population of Steller sea lions and mirrored patterns observed in human biomonitoring studies of Alaskan coastal communities. These data have broader implications with respect to human and ecosystem health as Steller sea lions rely on similar prey species and foraging areas as those targeted by commercial fisheries and subsistence users and are therefore valuable sentinels of marine ecosystem health.     

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.     

Parasites of forage fishes in the vicinity of Steller sea lion (Eumetopias jubatus) habitat in Alaska

Fish serve as intermediate hosts for a number of larval parasites that have the potential of maturing in marine mammals such as Steller sea lions (Eumetopias jubatus). We examined the prevalence of parasites from 229 fish collected between March and July 2002 near two islands used by Steller sea lions in Southeast Alaska and island habitats in the Aleutian Islands. Sea lion populations have remained steady in Southeast Alaska but have been declining over the last 30 yr in the Aleutian Islands. Even though the fish samples near the Southeast Alaska haul-outs were composed of numerous small species of fish and the Aleutian Islands catch was dominated by juveniles of commercially harvested species, the parasite fauna was similar at all locations. Eleven of the 20 parasite taxa identified were in their larval stage in the fish hosts, several of which have been described from mammalian final hosts. Four species of parasite were more prevalent in Southeast Alaska fish samples, and seven parasite species, including several larval forms capable of infecting marine mammals, were more prevalent in fish from the Aleutian Islands. Nevertheless, parasites available to Steller sea lions from common fish prey are not likely to be a major factor in the decline of this marine mammal species.     

KILLER WHALES, WHALING, AND SEQUENTIAL MEGAFAUNAL COLLAPSE IN THE NORTH PACIFIC: A COMPARATIVE ANALYSIS OF THE DYNAMICS OF MARINE MAMMALS IN ALASKA AND BRITISH COLUMBIA FOLLOWING COMMERCIAL WHALING

The hypothesis that commercial whaling caused a sequential megafaunal collapse in the North Pacific Ocean by forcing killer whales to eat progressively smaller species of marine mammals is not supported by what is known about the biology of large whales, the ecology of killer whales, and the patterns of ecosystem change that took place in Alaska, British Columbia, and elsewhere in the world following whaling. A comparative analysis shows that populations of seals, sea lions, and sea otters increased in British Columbia following commercial whaling, unlike the declines noted in the Gulf of Alaska and Aleutian Islands. The declines of seals and sea lions that began in western Alaska around 1977 were mirrored by increases in numbers of these species in British Columbia. A more likely explanation is that the seal and sea lion declines and other ecosystem changes in Alaska stem from a major oceanic regime shift that occurred in 1977. Killer whales are unquestionably a significant predator of seals, sea lions, and sea otters—but not because of commercial whaling.     

High aerobic capacities in the skeletal muscles of pinnipeds: adaptations to diving hypoxia.

The objective was to assess the aerobic capacity of skeletal muscles in pinnipeds. Samples of swimming and nonswimming muscles were collected from Steller sea lions (Eumetopias jubatus, n = 27), Northern fur seals (Callorhinus ursinus, n = 5), and harbor seals (Phoca vitulina, n = 37) by using a needle biopsy technique. Samples were either immediately fixed in 2% glutaraldehyde or frozen in liquid nitrogen. The volume density of mitochondria, myoglobin concentration, citrate synthase activity, and beta-hydroxyacyl-CoA dehydrogenase was determined for all samples. The swimming muscles of seals had an average total mitochondrial volume density per volume of fiber of 9.7%. The swimming muscles of sea lions and fur seals had average mitochondrial volume densities of 6.2 and 8.8%, respectively. These values were 1.7- to 2.0-fold greater than in the nonswimming muscles. Myoglobin concentration, citrate synthase activity, and beta-hydroxyacyl-CoA dehydrogenase were 1.1- to 2. 3-fold greater in the swimming vs. nonswimming muscles. The swimming muscles of pinnipeds appear to be adapted for aerobic lipid metabolism under the hypoxic conditions that occur during diving.     

Harbor seal population decline in the Aleutian Archipelago

Populations of Steller sea lions, northern fur seals, and northern sea otters declined substantially during recent decades in the Bering Sea and Aleutian Islands region, yet the population status of harbor seals has not been assessed adequately. We determined that counts obtained during skiff‐based surveys conducted in 1977–1982 represent the earliest estimate of harbor seal abundance throughout the Aleutian Islands. By comparing counts from 106 islands surveyed in 1977–1982 (8,601 seals) with counts from the same islands during a 1999 aerial survey (2,859 seals), we observed a 67% decline over the ～20‐yr period. Regionally, the largest decline of 86% was in the western Aleutians (n= 7 islands), followed by 66% in the central Aleutians (n= 64 islands), and 45% in the eastern Aleutians (n= 35 islands). Harbor seal counts decreased at the majority of islands in each region, the number of islands with >100 seals decreased ～70%, and the number of islands with no seals counted increased ～80%, indicating that harbor seal abundance throughout the Aleutian Islands was substantially lower in the late 1990s than in the 1970s and 1980s.     

Linking seasonal distribution patterns with prey availability in a central-place forager, the Steller sea lion

Aim We used a novel approach to infer foraging areas of a central‐place forager, the Steller sea lion (Eumetopias jubatus), by assessing changes in the temporal and spatial distribution patterns of sea lions at terrestrial sites. Specifically, our objectives were (1) to classify seasonal distribution patterns of Steller sea lions and (2) to determine to what extent the seasonal distribution of Steller sea lions is explained by seasonal concentrations of prey. Location Southeast Alaska, USA. Methods Steller sea lions of all age classes were counted monthly (2001–04) by aerial surveys at 28 terrestrial sites. Hierarchical cluster analysis and principal components analysis were used to classify seasonal distribution patterns of Steller sea lions at these terrestrial sites. We estimated the proportion of sea lions in the study area that were associated with each seasonal distribution pattern. Results Multivariate ordination techniques revealed four distinct seasonal distributional patterns. During December, 55% of the sea lions in the study area were found at Type 1 sites, located near over‐wintering herring aggregations. During May, 56% of sea lions were found at Type 2 sites, near aggregations of spring‐spawning forage fish. In July, 78% of sea lions were found at Type 3 sites, near summer migratory corridors of salmon. During September, 44% of sea lions were found at Type 4 sites, near autumn migratory corridors of salmon. Main conclusions Seasonal attendance patterns of sea lions were commonly associated with the seasonal availability of prey species near terrestrial sites and reflected seasonal foraging patterns of Steller sea lions in Southeast Alaska. A reasonable annual foraging strategy for Steller sea lions is to forage on herring (Clupea pallasii) aggregations in winter, spawning aggregations of forage fish in spring, salmon (Oncorhynchus spp.) in summer and autumn, and pollock (Theragra chalcogramma) and Pacific hake (Merluccius productus) throughout the year. The seasonal use of haulouts by sea lions and ultimately haulout‐specific foraging patterns of Steller sea lions depend in part upon seasonally available prey species in each region.     

The decline of Steller sea lions Eumetopias jubatus in Alaska: a review of the nutritional stress hypothesis

1. The decline of Steller sea lions Eumetopias jubatus in the Gulf of Alaska and Aleutian Islands between the late 1970s and 1990s may have been related to reduced availability of suitable prey. Many studies have shown that pinnipeds and other mammals suffering from nutritional stress typically exhibit reduced body size, reduced productivity, high mortality of pups and juveniles, altered blood chemistry and specific behavioural modifications. 2. Morphometric measurements of Steller sea lions through the 1970s and 1980s in Alaska indicate reduced body size. Reduced numbers of pups born and an apparent increase in juvenile mortality rates also appear to be nutritionally based. Blood chemistry analyses have further shown that Steller sea lions in the Gulf of Alaska and Aleutian Islands area exhibited signs of an acute phase reaction, or immune reaction, in response to unidentified physical and/or environmental stress. Behavioural studies during the 1990s have not noted any changes that are indicative of an overall shortage in the quantity of prey available to lactating female sea lions. 3. The data collected in Alaska are consistent with the hypothesis that Steller sea lions in the declining regions were nutritionally compromised because of the relative quality of prey available to them (chronic nutritional stress), rather than because of the overall quantity of fish per se (acute nutritional stress). This is further supported by captive studies that indicate the overall quality of prey that has been available to Steller sea lions in the declining population could compromise the health of Steller sea lions and hinder their recovery.     

Depth and movement behaviour of the Pacific sleeper shark in the north-east Pacific Ocean

Satellite-linked radio telemetry was used to study the geographic movements and vertical movement behaviour of the Pacific sleeper shark Somniosus pacificus . The fish were tagged near Steller sea lion Eumetopias jubatus rookeries in the Gulf of Alaska during periods when Steller sea lions pups were most vulnerable to predation; when Steller sea lion pups first enter the water (July to August) and when Steller sea lion pups are weaned (April to May). Final locations recovered from most Pacific sleeper sharks (76%) were within 100 km of release locations, 16% were within 100–250 km and 8% were within 250–500 km. The most striking behavioural feature was their extensive, nearly continuous vertical movements. Median daily depth range was 184 m; the most time (61%) was spent between 150 and 450 m, but ascents above 100 m were common (58% of days). Median vertical movement rate was 6 km day⁻¹ and steady. The longest period of continuous vertical movement (> 60 m h⁻¹) was 330 h. Systematic vertical oscillations were most common (60%), followed by diel vertical migrations (25%) and irregular vertical movements (15%). The Pacific sleeper sharks travelled below the photic zone during the day and approached the surface at night. Pacific sleeper sharks appear to employ a stealth and ambush hunting strategy that incorporates slow vertical oscillations to search for prey, and cryptic colouration and cover of darkness to avoid detection by potential prey. The depth and geographic range of Pacific sleeper shark and Steller sea lions overlap near four important Steller sea lion rookeries in the northern Gulf of Alaska, so the potential exists for predation to occur. None of the tissues in the stomachs of the 198 Pacific sleeper sharks collected during a companion diet study, however, were identified as Steller sea lion.     

PINNIPED POPULATION DYNAMICS IN CENTRAL CALIFORNIA: CORRELATIONS WITH SEA SURFACE TEMPERATURE AND UPWELLING INDICES

We investigated effects of marine climate variability on pinniped populations and assessed the initial stages of recovery following implementation of the U. S. Marine Mammal Protection Act (MMPA) based on long-term (1973-1997) population surveys at the South Farallon Islands and Point Reyes Peninsula, central California. California sea lions increased over the study period, with peak numbers observed during and after major El Niño events. The rate of increase for California sea lions appears to have decreased in recent years. Steller sea lions decreased at the South Farallon Islands and remain depleted at Point Reyes Peninsula. Harbor seal populations increased in a logistic and non-linear fashion at Point Reyes Peninsula and the South Farallon Islands, respectively. Harbor seals were more abundant at the South Farallon Islands during years of relatively high sea-surface temperature, which may be related to their inability to find sufficient prey in coastal waters under these conditions. Northern elephant seal abundance increased in a logistic fashion over the study period at both the South Farallon Islands and Point Reyes Peninsula; however, productivity at the South Farallon Islands decreased in recent years. Maximum haulout numbers for elephant seals at the South Farallon Islands increased in the 1970s, maintained an asymptote throughout the 1980s and early 1990s, but recently declined; additional studies are needed to investigate which age classes are associated with this decline. Protection afforded by the MMPA has facilitated partial to full recovery of all populations except for Steller sea lion. Oceanographic relationships do not appear to confound interpretations of population recovery and may help to explain changes in the Steller sea lion population.