AN ANALYSIS OF CALIFORNIA SEA OTTER (ENHYDRA LUTRIS) PELAGE AND INTEGUMENT

The sea otter is associated with a cold, marine habitat, has no insulating fat layer, and relies on its fur layer for insulation. Soiled pelage provides inadequate insulation and can lead to hypothermia and death. Information on sea otter pelage, the surface and bound lipids found in the pelage, and histology of the integument is thus relevant to the development of rehabilitation and management techniques for sea otters soiled with oil. We present detailed data on the sea otter pelage and integument, including hair bundle density (737–2,465 bundles per cm²), hair density per bundle (19–91.1), total hair density (26,413–164,662 hairs per cm²), guardhair length (8.2–26.9 mm), underfur length (4.6–15.8 mm), guardhair diameter (44.0–106.0 μ), underfur diameter (7.6–11.9 μ), angle of hair with respect to the skin (61.9°- 84.3°) and structure of individual hairs as seen with a scanning electron microscope. The hydrocarbon squalene was found to be the major component of the lipids associated with the pelage. Various layers of skin from eight sites on a single animal are described histologically. With the exception of density of the hair coat, sea otter integument is similar to that of domestic carnivores.     

ATYPICAL PUP REARING STRATEGIES BY SEA OTTERS

Eight tagged sea otter (Enhydra lutris) pups in central Prince William Sound, Alaska, weighed 6–15 kg at the time of separation from their mother. Four pups weighing 15 kg were able to forage successfully on their own. Three pups weighing ≤9 kg had negligible chances of survival and apparently were abandoned by sick females. Abandonment of a pup may reduce the burden on a sick female, enabling recovery and subsequent reproduction. One of the three sick females that abandoned a pup in this study recovered and pupped again. Abandonment of pups should occur most often in populations where females are stressed by poor food resources. Reassociation with a previous offspring, as observed once in this study, also may occur most frequently in food-limited populations where reproductive failures are most common and pup survivorship is significantly increased by additional maternal assistance.     

CAUSES OF MORTALITY IN CALIFORNIA SEA OTTERS DURING PERIODS OF POPULATION GROWTH AND DECLINE

Elevated mortality appears to be the main reason for both sluggish growth and periods of decline in the threatened California sea otter population. We assessed causes of mortality from salvage records of 3,105 beach-cast carcasses recovered from 1968 through 1999, contrasting two periods of growth with two periods of decline. Overall, an estimated 40%-60% of the deaths were not recovered and 70% of the recovered carcasses died from unknown causes. Nonetheless, several common patterns were evident in the salvage records during the periods of population decline. These included greater percentages of (1) prime age animals (3–10 yr), (2) carcasses killed by great white shark attacks, (3) carcasses recovered in spring and summer, and (4) carcasses for which the cause of death was unknown. Neither sex composition nor the proportion of carcasses dying of infectious disease varied consistently between periods of population increase and decline. The population decline from 1976 to 1984 was likely due to incidental mortality in a set-net fishery, and the decline from 1995 to 1999 may be related to a developing live-fish fishery. Long-term trends unrelated to periods of growth and decline included a decrease in per capita pup production and mass/length ratios of adult carcasses over the 31-yr study. The generally high proportion of deaths from infectious disease suggests that this factor has contributed to the chronically sluggish growth rate of the California sea otter population.     

DETECTION OF SEA OTTERS IN BOAT-BASED SURVEYS OF PRINCE WILLIAM SOUND, ALASKA

Boat-based surveys have been commonly used to monitor sea otter populations, but there has been little quantitative work to evaluate detection biases that may affect these surveys. We used ground-based observers to investigate sea otter detection probabilities in a boat-based survey of Prince William Sound, Alaska. We estimated that 30% of the otters present on surveyed transects were not detected by boat crews. Approximately half (53%) of the undetected otters were missed because the otters left the transects, apparently in response to the approaching boat. Unbiased estimates of detection probabilities will be required for obtaining unbiased population estimates from boat-based surveys of sea otters. Therefore, boat-based surveys should include methods to estimate sea otter detection probabilities under the conditions specific to each survey. Unbiased estimation of detection probabilities with ground-based observers requires either that the ground crews detect all of the otters in observed subunits, or that there are no errors in determining which crews saw each detected otter. Ground-based observer methods may be appropriate in areas where nearly all of the sea otter habitat is potentially visible from ground-based vantage points.     

SEA OTTER DIETS IN A DECLINING POPULATION IN ALASKA

Diet of sea otters ( Enhydra lutris ) using a haulout site on the north side of the Alaska Peninsula was determined from 50 scats. Dominant prey species were mussels ( Mytilus edulis ), followed by three species of clams ( Siliqua spp., Spisula polynyma , and Tellina lutea ), sand dollars ( Echinarachnius parma ), and helmet crabs ( Telmessus cheiragonus ). Our results support preliminary findings by Cim-berg et al. (1984) that this sea otter population preys heavily on mussels and that the presence of low caloric value sand dollars in their diet is significant. Coupled with population estimates, our results also provide evidence that this population of sea otters may be declining due, in part, to overdepletion of food resources.     

POTENTIAL IMPACT OF OIL SPILLS ON CALIFORNIA SEA OTTERS: IMPLICATIONS OF THE EXXON VALDEZ SPILL IN ALASKA

Based on the survival of sea otters held at rehabilitation centers during the 1989 Exxon Valdez oil spill in Alaska, we built two models of otter mortality. One was based on the relationship between mortality and distance from spill origin, the other was based on the relationship between mortality and time from the spill origin. These models are simplistic and are meant as first steps in arriving at realistic risk estimates and in providing a conceptual framework for relating oil spills and sea otter mortality. Using the distance model, we simulated the impact of an Exxon Valdez event occurring at different locations along the California coast. A spill at the Monterey Peninsula had the greatest impact, exposing 90% of the California sea otter population to oil and killing at least 50% of the individuals. The time model was used to predict the mortality of otters exposed to oil of various ages and for various periods of time. It suggested that efforts to rehabilitate otters should be discontinued 20-30 d after a spill. The limitations of the data available from the Exxon Valdez spill emphasize the importance of being prepared to conduct appropriate research during the next oil spill in sea otter habitat.     

AN ESTIMATION OF CARRYING CAPACITY FOR SEA OTTERS ALONG THE CALIFORNIA COAST

Carrying capacity (K) for the California sea otter ( Enhydra lutris nereis ) was estimated as a product of the density of sea otters at equilibrium within a portion of their existing range and the total area of available habitat. Equilibrium densities were determined using the number of sea otters observed during spring surveys in 1994, 1995, and 1996 in each of three habitat types where sea otters currently exist. Potential sea otter habitat was defined as from the California coastline to the 40-m isobath and classified as rocky, sandy, or mixed habitat according to the amount of kelp and rocky substrate in the area. The amount of habitat available to sea otters in California was estimated using a Geographic Information Systems (GIS) program. The estimated mean number of sea otters that could be supported by the marine environment to a depth of 40 m in California was 15,941 (95% CI 13,538–18,577). The GIS-based approach incorporated detailed bathymetric contours, produced repeatable and accurate estimates, and served as an innovative method of measuring sea otter habitat. We believe the approach described in this paper represents the best available information on how a sea otter population at equilibrium would be distributed along the California coast.     

REPRODUCTION, SURVIVAL AND TAG LOSS IN CALIFORNIA SEA OTTERS

We observed 40 California sea otters, Enhydra lutris , that were instrumented with implanted radio transmitters and flipper-tagged, and obtained additional data on the reproduction of tagged female otters from the California Department of Fish and Game.
The proportion of instrumented females accompanied by a pup peaked in the spring, with a secondary peak in the fall. Two methods of estimating the annual reproductive rate gave comparable values of 0.90 and 0.94. The average inter-birth interval was 389 d. Two methods of estimating pup survival to weaning gave values of 0.46 and 0.58. Pups either remained with a female less than 80 or more than 120 d. Early mortality of dependent pups appears to be more frequent in California than in Prince William Sound, Alaska.
Two methods of estimation indicated that adult females had the highest survival rates and adult males the lowest. Juvenile females had lower survival rates than adult females but juvenile males had higher survival rates than adult males. The survival rate of juvenile females was lower than that of juvenile males.
The estimated annual loss rate for flipper-tags, based on the instrumented individuals, was 0.26. More individuals lost two tags than would be expected by chance. It is unlikely that accurate estimates of sea otter survival rates can be derived from observations of tagged individuals.     

REPRODUCTIVE CHARACTERISTICS OF FEMALE SEA OTTERS

Abstract: Several important aspects of reproduction in the female sea otter, such as gestation, pupping frequency, period of pup dependency, and annual pupping rate, were unclear when this study was begun. We present data from 75 tagged adult females that indicate gestation is variable, but on average is about 6 months, the length of pup dependency is 6 mo, thus the pupping interval is usually 12–13 mo. Most females breed for the first time in their fifth year of life. About 85–90% of adult females pup in a given year.     

MOVEMENTS, HOME RANGE, AND TERRITORIES OF MALE SEA OTTERS OFF CENTRAL CALIFORNIA

Sixty male sea otters ( Enhydra lutris ) were tagged on the rear flippers with colored tags. Of these, 46 (77%) were resighted. Movements of 127 km were documented for adults and 187 km for subadults. Adults maintained breeding territories that averaged 40.3 ha ( n = 10, SE = 4.0). They returned to the same territory seasonally for up to seven consecutive years. Territorial males moved from areas of high male abundance to areas of high female abundance on a seasonal basis. During the winter, 74% of adult males left breeding areas and joined concentrations of males located near the ends of the range. Thirty percent of the subadult males were observed in male groups near the extremities of the range. During the summer and fall, the density of adult males (15/1,000 ha) and adult male to independent otter (non-pup) ratio (1:5) in female areas was highest. The number of adult males in areas of female abundance was inversely related to the number of dependent pups, perhaps because when pup numbers are low (late summer and fall) the number of estrous females is high. Subadult males may remain in female areas on a year round basis until their second or third year. However, they were not generally associated with adult females.     

FORAGING DEPTHS OF SEA OTTERS AND IMPLICATIONS TO COASTAL MARINE COMMUNITIES

We visually observed 1,251 dives, of 14 sea otters instrumented with TDRs in southeast Alaska, and used attribute values from observed dives to classify 180,848 recorded dives as foraging (0.64), or traveling (0.36). Foraging dives were significantly deeper, with longer durations, bottom times, and postdive surface intervals, and greater descent and ascent rates, compared to traveling dives. Most foraging occurred in depths between 2 and 30 m (0.84), although 0.16 of all foraging was between 30 and 100 m. Nine animals, including all five males, demonstrated bimodal patterns in foraging depths, with peaks between 5 and 15 m and 30 and 60 m, whereas five of nine females foraged at an average depth of 10 m. Mean shallow foraging depth was 8 m, and mean deep foraging depth was 44 m. Maximum foraging depths averaged 61 m (54 and 82 for females and males, respectively) and ranged from 35 to 100 m. Female sea otters dove to depths ≤20 m on 0.85 of their foraging dives while male sea otters dove to depths ≥45 m on 0.50 of their foraging dives. Less than 0.02 of all foraging dives were >55 m, suggesting that effects of sea otter foraging on nearshore marine communities should diminish at greater depths. However, recolonization of vacant habitat by high densities of adult male sea otters may result in initial reductions of some prey species at depths >55 m.     

DIET AND FORAGING BEHAVIOR OF SEA OTTERS IN SOUTHEAST ALASKA

Abstract: Direct observations of feeding sea otters ( Enhydra lutris ) at 11 sites in southeast Alaska showed infaunal clams to be the primary prey utilized by otters throughout the region. Foraging dive times associated with clam and sea urchin prey were significantly longer than those for more easily captured prey (crabs and mussels). Dive times and surface intervals were also generally correlated with water depth or apparent difficulty in obtaining buried prey. Male otters, which fed more extensively on clams than females, made significantly longer foraging dives than females. Foraging success remained high, even at sites where prey numbers were found to be very low during a related study. The very deeply burrowing geoduck clam ( Panope abrupta ), while common at several otter feeding sites, was rarely captured by otters. These results, combined with those of a companion study on prey numbers, indicate that butter clams ( Saxidomus giganteus ) account for the majority of the sea otter diet in southeast Alaska, and that sea urchins may represent relatively short-term prey in comparison to infaunal bivalves in regions where both prey types co-exist. Furthermore, the importance of butter clams in the sea otter diet and the tendency for this bivalve to retain chronically high levels of paralytic shellfish poisoning toxins in southeast Alaska increases the probability that toxic phytoplankton blooms influence sea otter distribution in this region.     

SURVIVAL RATES OF SEA OTTER PUPS IN ALASKA AND CALIFORNIA

Adult female sea otters ( Enhydra lutris ) were instrumented with implanted radio-transmitters in Prince William Sound (PWS), Alaska, and survival rates were estimated for their dependent pups. Overall, 94 of 141 (67%) of the pups studied survived to a minimum age of 120 d and were assumed to have successfully weaned. Survival of pups in six cohorts ranged from 53% to 88%. The mean interval between successive visual observations was 12.5 d. For these calculations, the assumption was made that pups were successfully weaned if they accompanied mothers for at least 120 d. Estimated survival rates were different when this assumption was changed to either 90 d or 150 d (73% and 52%, respectively).
Females were palpated for pregnancy when instrumented. Of 19 believed to be pregnant, 17 were subsequently seen with young pups giving a detection rate for births of 89.5%. When the above observed survival rate of pups was adjusted for undetected births, the estimated overall survival rate for the study population was 60% (120 d minimum dependency).
Survival rates of pups in PWS and a population at Kodiak Archipelago (KOD) (Monson and DeGange 1995) were compared with that of pups in the population in California (CA, four studies). These data did not support the hypothesis that survival rates were lower in California (CA: 103/160, P_surv. 0.64; PWS: 94/141, P_surv. = 0.67; KOD: 19/23, P_surv. = 0.83; pairwise comparisons, X², P > 0.05). Comparison of pup survival rates among studies was hindered by small sample sizes, methodological differences, and lack of detail about assumptions underlying estimates.     

MORTALITY OF SEA OTTERS IN PRINCE WILLIAM SOUND FOLLOWING THE EXXON VALDEZ OIL SPILL

Abstract: This paper presents an estimate of the total number of sea otters that died as a direct consequence of the oil spill that occurred when the T/V Exxon Valdez grounded in Prince William Sound, Alaska on 24 March 1989. We compared sea otter counts conducted from small boats throughout the Sound during the summers of 1984 and 1985 to counts made after the spill during the summer of 1989. We used ratio estimators, corrected for sighting probability, to calculate otter densities and population estimates for portions of the Sound affected by the oil spill. We estimated the otter population in the portion of Prince William Sound affected by the oil was 6,546 at the time of the spill and that the post-spill population in the summer of 1989 was 3,898, yielding a loss estimate of approximately 2,650. Bootstrapping techniques were used to approximate confidence limits on the loss estimate of about 500–5,000 otters. The wide confidence limits are a result of the complex scheme required to estimate losses and limitations of the data. Despite the uncertainty of the loss estimate it is clear that a significant fraction of the otters in the spill zone survived. We observed otters persisting in relatively clean embayments throughout the oil spill zone suggesting that the highly convoluted coastline of Prince William Sound produced refuges that allowed some sea otters in the oil spill area to survive.     

LENGTH-MASS AND TOTAL BODY LENGTH OF ADULT FEMALE SEA OTTERS IN PRINCE WILLIAM SOUND BEFORE AND AFTER THE EXXON VALDEZ OIL SPILL

After the 1989 T/V Exxon Valdez oil spill (EVOS), the body condition of non-pregnant female sea otters ( Enhydra lutris ) ages 4 yr and older in the EVOS-affected region of western Prince William Sound, Alaska (WPWS), was significantly poorer than that of individuals captured in the same or adjacent habitat in WPWS approximately a decade earlier, and than that of individuals inhabiting unoiled habitat in eastern PWS (EPWS) between 1984 and 1990. However, the body condition of females of this age category captured in WPWS prior to EVOS was not significantly different from that of pre-and postspill EPWS females. The mean total body length (TBL) of non-pregnant females captured prespill in WPWS was significantly less than that of pre-and postspill EPWS and postspill WPWS females. Evidence from this and other studies suggests that the body condition of at least some classes of sea otters was negatively affected by one or more EVOS-related factors.     

Loss of genetic diversity in sea otters (Enhydra lutris) associated with the fur trade of the 18th and 19th centuries

Sea otter (Enhydra lutris) populations experienced widespread reduction and extirpation due to the fur trade of the 18th and 19th centuries. We examined genetic variation within four microsatellite markers and the mitochondrial DNA (mtDNA) d-loop in one prefur trade population and compared it to five modern populations to determine potential losses in genetic variation. While mtDNA sequence variability was low within both modern and extinct populations, analysis of microsatellite allelic data revealed that the prefur trade population had significantly more variation than all the extant sea otter populations. Reduced genetic variation may lead to inbreeding depression and we believe sea otter populations should be closely monitored for potential associated negative effects.     

Activity Budgets Derived From Time-Depth Recorders in a Diving Mammal

Abstract: We describe a method to convert continuously collected time-depth data from archival time-depth recorders (TDRs) into activity budgets for a benthic-foraging marine mammal. We used data from 14 TDRs to estimate activity-specific time budgets in sea otters (Enhydra lutris) residing near Cross Sound, southeast Alaska, USA. From the TDRs we constructed a continuous record of behavior for each individual over 39-46 days during summer of 1999. Behaviors were classified as foraging (diving to the bottom), other diving (traveling, grooming, interacting), and nondiving (assumed resting). The overall average activity budget (proportion of 24-hr/d) was 0.37 foraging (8.9 hr/d), 0.11 in other diving (2.6 hr/d), and 0.52 nondiving time (12.5 hr/d). We detected significant differences in activity budgets among individuals and between groups within our sample. Historically, the sea otter population in our study area had been expanding and sequentially reoccupying vacant habitat since their reintroduction to the area in the 1960s, and our study animals resided in 2 adjacent yet distinct locations. Males (n = 5) and individuals residing in recently occupied habitat (n = 4) spent 0.28-0.30 of their time foraging (6.7-7.2 hr/d), 0.17-0.18 of their time in other diving behaviors (4.1-4.3 hr/d), and 0.53-0.54 of their time resting (12.7-13.0 hr/d). In contrast, females (n = 9) and individuals residing in longer occupied habitat (n = 10) spent 0.40 of their time foraging (9.6 hr/d), 0.08-0.09 of their time in other diving behaviors (1.9-2.2 hr/d), and 0.51-0.52 of their time resting (12.2-12.5 hr/d). Consistent with these differences, sea otters residing in more recently occupied habitat captured more and larger clams (Saxidomus spp., Protothaca spp., Macoma spp., Mya spp., Clinocardium spp.) and other prey, and intertidal clams were more abundant and larger in this area. We found that TDRs provided data useful for measuring activity time budgets and behavior patterns in a diving mammal over long and continuous time periods. Fortuitous contrasts in time budgets between areas where our study animals resided suggest that activity time budgets estimated from TDRs may be a sensitive indicator of population status, particularly in relation to prey availability.     

Evaluating Cementum to Determine Past Reproduction in Northern Sea Otters

Abstract: Age at first reproduction (AFR) has been difficult to quantify in mammals, as the most commonly used methods require reproductive tracts or direct observations. However, work in several large mammal species suggests that the width of cementum light bands in teeth decline once females begin to reproduce, suggesting that teeth structures might provide a new tool to examine AFR. To determine if changes in cementum light band width could be used to calculate AFR for the northern sea otter (Enhydra lutris kenyoni), we measured cementum light band widths on sectioned premolar teeth and compared them to reproductive tracts. We classified otters as parous if any single light band was narrower than a threshold value, selected as the value that minimized error rates. At a threshold value of 0.32, we correctly identified otters as parous or nulliparous in 83% of cases (n = 92) as compared to reproductive tracts, and the AFR estimated from teeth samples (3.52 ± 0.032 yr) was not different from that determined by reproductive tract analysis (3.45 ± 0.031 yr; t-test, P > 0.05). These data support the use of cementum as an indicator of past reproduction in individual female otters, which can then be used to estimate average AFR. Given that declines in cementum width have been described for other mammal species, the same quantitative approach used here could be applied to other species. (JOURNAL OF WILDLIFE MANAGEMENT 72(3):618–624; 2008)