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.     

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.     

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.     

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.     

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.     

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.     

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.     

AN INTRAPERITONEAL RADIO TRANSMITTER FOR SEA OTTERS

A small intraperitoneal radio transmitter was developed and used successfully in California sea otters. The transmitter weighs about 120 g and has an internal antenna. Thirty-five implanted transmitters were located for an average of 526 d. The carcasses of five animals that died were recovered; there were no complications associated with the transmitters in these individuals. Reproductive performance of the adult females with implanted transmitters appeared normal. The main advantages of this transmitter are its reliability and long life. It has enabled the collection of significant new information on California sea otters.     

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.     

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.     

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.     

STOCK STRUCTURE OF SEA OTTERS (ENHYDRA LUTRIS KENYONI) IN ALASDA

Sea otters in Alaska are recognized as a single subspecies ( Enhydra lutris kenyoni ) and currently managed as a single, interbreeding population. However, geographic and behavioral mechanisms undoubrably constrain sea otter movements on much smaller scales. This paper applies the phylogeographic method (Dizon et al . 1992) and considers distribution, population response, phenotype and genotype data to identify stocks of sea otters within Alaska. The evidence for separate stock identity is genotypic (all stocks), phenotypic (Southcentral and Southwest stocks), and geographic distribution (Southeast stock), whereas population response data are equivocal (all stocks). Differences in genotype frequencies and the presence of unique genotypes among areas indicate restricted gene flow. Genetic exchange may be limited by little or no movement across proposed stock boundaries and discontinuities in distribution at proposed stock boundaries. Skull size differences (phenotypic) between Southwest and Southcentral Alaska populations further support stock separation. Population response information was equivocal in either supporting or refuting stock identity. On the basis of this review, we suggest the following: (1) a Southeast stock extending from Dixon Entrance to Cape Yakataga; (2) a Southcentral stock extending from Cape Yakataga to Cape Douglas including Prince William Sound and Kenai peninsula coast; and (3) a Southwest stock including Alaska Peninsula coast, the Aleutians to Attu Island, Barren, Kodiak, Pribilof Islands, and Bristol Bay.     

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.     

A GRADIENT IN BENTHIC INTERTIDAL ALGAL ASSEMBLAGES ALONG THE SOUTHERN CALIFORNIA COAST1

Based on an analysis floristic data, a gradient appears to exist in the composition of intertidal algae along the 450 km of southern California coastline immediately south of Point Conception. Reciprocal averaging ordination of the algal flora at 51 sites in this area suggests that the gradient is not strictly latitudinal. Variation from a latitudinal gradient occurs ca. 60-80 km south of Point Conception. An ordination of frequently occurring species indicated that compositional changes are substantial when sites at the extreme north and south are compared, but that the changes are gradual and continuous. Possible explanations for the gradient are discussed, and these include: sea temperature, upwelling, sand movement, human disturbance (i.e., pollution, trampling), and wave action.     

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.     

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.     

中国农业生态系统的生产潜力和人口承载力

中国农业生态系统的生产潜力和人口承载力是全球关注的问题。本研究将农业生态系统的生产潜力定义为由生物遗传特性和4大宏观生态条件(太阳辐射、温度、水资源和土地资源)共同决定的生产力上限;以无机环境-第一性生产-第二性生产之间的结构适应性和能量-物质流平衡为主线,发展了对农业生态系统生产潜力和人口承载力的综合评价模型,它包括第一性生产潜力子模型和第二性生产潜力与人口承载力优化子模型;并应用该模型把中国农业生态系统分为603个区域单元,进行第一性生产潜力,第二性生产潜力和人口承载力的评价。     

PHOTOPERIOD DEFINES THE PHENOLOGY OF BIRTH IN CAPTIVE CALIFORNIA SEA LIONS

A population-based analysis was used to evaluate the role of photoperiod in regulating the precise annual birth timing of captive California sea lions. Latitudinal variation of 463 birth dates was explained by a response to a common photoperiod, 11.48 h/d, occurring 242 d pre-partum and immediately before blastocyst implantation. The lengths of the pupping seasons at the 18 largest captive colonies were significantly correlated to those estimated by the photoperiod model. Hence, the phenology of birth in this species is defined by photoperiod.