Male Sea Otters Research Articles

The Effects of Age and Sex on the Energy Intake of Captive Sea Otters (Enhydra lutris): Implications for Captive Management and Species Conservation

Sea otters (Enhydra lutris) are known ecosystem engineers that have significant impacts on their kelp forest and rocky intertidal communities due to their high levels of food intake. Quantifying sea otter food biomass and energy intake is a valuable way to understand potential ecological impacts of sea otter populations on ecosystems and for predicting future population trends and potential for expansion. While detailed, fine-scale, age-specific food intake is difficult to quantify in wild sea otters, there is a wealth of potential information available from otters under human care. This study used food and energy intake data from husbandry records of 10 sea otters collected over three decades at the Vancouver Aquarium. Within these husbandry records, daily food biomass intake and body mass measurements were recorded and converted to annual average food mass and gross energy intake (GEI). Age-, sex-, and mass-specific trends were also observed. Young sea otters had the highest relative ingested food mass, equivalent to ~26% of body mass, which decreased to ~20% in adult otters. Young otters similarly had the highest mass-specific GEI, where measures from near birth to year 1 were ~40% higher than at year 3, the age of sexual and physical maturity. There were also key differences in trends between sexes. Captive adult male sea otters were 25 to 42% larger than females and their GEI was 23 to 58% higher, although mass-specific GEI was almost identical for male and non-reproductive female otters at all ages, plateauing at ~650 kJ kg-1 d-1. Despite high levels of ingested food mass, GEI was only 5 to 15% higher than for other captive marine mammals and was comparable to previous estimates for wild sea otters. These estimates of ingested food mass and energy intake requirements are valuable when modelling the ecological impact of sea otter populations and for considering the potential effects of future environmental changes.

Do differences in skull morphology and bite performance explain dietary specialization in sea otters?

Intraspecific studies of morphology and performance are essential for understanding the factors that enable resource partitioning within ecological communities. The sea otter (Enhydra lutris) is one of the few mammal species in which individual-level dietary specialization has been documented, making them an ideal system to investigate the morphological basis of food resource partitioning. Here, we test if differences in food resource use within and between sea otter subspecies can be explained by differences in ecologically relevant metrics of bite performance that are mainly the product of variation in size, cranial morphology, or a combination of these traits. We use geometric morphometrics to evaluate variation in cranium size and morphology, and 2-dimensional models to estimate bite performance differences between 2 sea otter subspecies that differ in dietary ecology: the northern sea otter (Enhydra lutris kenyoni, a facultative generalist) and the southern sea otter (E. l. nereis, a specialist). We found significant differences in cranium shape and size between subspecies and between male and female sea otters. These differences were subtle yet consistent with most subspecies classifications and known sexually dimorphic traits. Cranial morphological differences did not translate into differences in estimated bite force between subspecies or sexes, but dentary strength differed significantly between male and female sea otters. Sea otters have short, blunt crania with pronounced sagittal and lambdoidal crests, and strong mandibles. These traits combine to produce high bite forces for their size. We propose that high bite performance capacity in sea otters enables resource-use variation by widening the diversity of available food resources they can procure from their environment; this allows them to behave as either generalists or specialists within different habitats.

Sexual dimorphism in craniomandibular morphology of southern sea otters (Enhydra lutris nereis)

The niche divergence hypothesis suggests that if a species exhibits intersexual differences in diet, selection should favor divergence in the feeding apparatus between the sexes. Recent work revealed that male and female southern sea otters ( Enhydra lutris nereis ) utilize different dietary resources in response to increased population density; females exhibit more specialized diets as a function of smaller home ranges, whereas males exhibit larger home ranges, potentially allowing them to expand their dietary breadths by feeding on prey items that are not found in female home ranges. These dietary differences suggest the potential for sexual dimorphism of the feeding apparatus (i.e., the skull). Here, we tested the hypothesis that male and female southern sea otters exhibit differences in craniomandibular traits directly related to biting ability. Univariate and multivariate analyses of 12 craniomandibular traits showed that size is the primary axis of skull variation, whereas only a handful of craniomandibular traits demonstrated significant shape differences between the sexes. Relative postorbital constriction breadth, masseter in-lever length, and cranial height differed significantly between the sexes. These 3 traits can increase the surface area of jaw muscle attachment sites and thus are directly linked to the mechanics of biting ability. Collectively, these morphological differences indicate that niche divergence may be an important mechanism maintaining sexual dimorphism in southern sea otters.

Ontogenetic Scaling of Theoretical Bite Force in Southern Sea Otters (Enhydra lutris nereis).

Sexual dimorphism attributed to niche divergence is often linked to differentiation between the sexes in both dietary resources and characters related to feeding and resource procurement. Although recent studies have indicated that southern sea otters (Enhydra lutris nereis) exhibit differences in dietary preferences as well as sexual dimorphism in skull size and shape, whether these intersexual differences translate to differentiation in feeding performances between the sexes remains to be investigated. To test the hypothesis that scaling patterns of bite force, a metric of feeding performance, differ between the sexes, we calculated theoretical bite forces for 55 naturally deceased male and female southern sea otters spanning the size ranges encountered over ontogeny. We then used standardized major axis regressions to simultaneously determine the scaling patterns of theoretical bite forces and skull components across ontogeny and assess whether these scaling patterns differed between the sexes. We found that positive allometric increases in theoretical bite force resulted from positive allometric increases in physiological cross-sectional area for the major jaw adductor muscle and mechanical advantage. Closer examination revealed that allometric increases in temporalis muscle mass and relative allometric decreases in out-lever lengths are driving these patterns. In our analysis of sexual dimorphism, we found that scaling patterns of theoretical bite force and morphological traits do not differ between the sexes. However, adult sea otters differed in their absolute bite forces, revealing that adult males exhibited greater bite forces as a result of their larger sizes. We found intersexual differences in biting ability that provide some support for the niche divergence hypothesis. Continued work in this field may link intersexual differences in feeding functional morphology with foraging ecology to show how niche divergence has the potential to reinforce sexual dimorphism in southern sea otters.

Characterization of the Temporomandibular Joint of Southern Sea Otters (Enhydra lutris nereis).

The structure–function relationship of the temporomandibular joint (TMJ) of southern sea otter has largely not been described. This study aims to describe the histological, biochemical, and biomechanical features of the TMJ disk in the southern sea otter. The TMJ disks from fresh cadaver heads of southern sea otter adult males (n = 8) and females (n = 8) acquired from strandings were examined. Following macroscopical evaluation, the TMJs were investigated for their histological, mechanical, and biochemical properties. We found that the sea otter TMJ disks are, in general, similar to other carnivores. Macroscopically, the TMJ disk was highly congruent, and the mandibular head was encased tightly by the mandibular fossa with a thin disk separating the joint into two compartments. Histologically, the articular surfaces were lined with dense fibrous connective tissue that gradually transitioned into one to two cell thick layer of hyaline-like cartilage. The disk fibers were aligned primarily in the rostrocaudal direction and had occasional lacuna with chondrocyte-like cells. The disk was composed primarily of collagen type 1. Biochemical analysis indicates sulfated glycosaminoglycan content lower than other mammals, but significantly higher in male sea otters than female sea otters. Finally, mechanical analysis demonstrated a disk that was not only stronger and stiffer in the rostrocaudal direction than the mediolateral direction but also significantly stronger and stiffer in females than males. We conclude that the congruent design of the TMJ, thin disk, biochemical content, and mechanical properties all reflect a structure–function relationship within the TMJ disk that is likely designed for the sea otter’s hard diet and continuous food intake.

Growth and sexual maturation in male northern sea otters (Enhydra lutris kenyoni) from Gustavus, Alaska

Northern sea otters (Enhydra lutris kenyoni) are experiencing rapid population growth and expansion in Southeast Alaska. Fluctuations in population density can produce profound changes in ecological constraints, influencing growth and sexual maturity of animals in a population, and can transform the ecology of newly populated ecosystems. Using morphometric measurements and reproductive tracts collected from 40 sea otters by an Alaska Native subsistence hunter, we evaluated growth rates and age at sexual maturity of male sea otters near Gustavus, Alaska. Standard length of mature otters (asymptotic size = 148.2 ± 2.9 cm [SE]) was greater than previous studies in Alaska. At puberty, mean seminiferous tubule diameter, baculum length, and testes mass increased abruptly at 130 cm body length. Animals in this study reached sexual maturity earlier than previous studies, at age 3–4 years. Baculum length increased rapidly at sexual maturity and was a good indication of maturity status (< 14 cm = immature). Increas...

The cost of reproduction: differential resource specialization in female and male California sea otters.

Intraspecific variation in behavior and diet can have important consequences for population and ecosystem dynamics. Here, we examine how differences in reproductive investment and spatial ecology influence individual diet specialization in male and female southern sea otters (Enhydra lutris nereis). We hypothesize that greater reproductive constraints and smaller home ranges of females lead to more pronounced intraspecific competition and increased specialization. We integrate stable carbon (δ(13)C) and nitrogen (δ(15)N)isotope analysis of sea otter vibrissae with long-term observational studies of five subpopulations in California. We define individual diet specialization as low ratios of within-individual variation (WIC) to total population niche width (TNW). We compare isotopic and observational based metrics of WIC/TNW for males and females to data on population densities, and movement patterns using both general linear and linear mixed-effects models. Consistent with our hypothesis, increasing population density is associated with increased individual diet specialization by females but not by males. Additionally, we find the amount of coastline in a sea otter's home range positively related with individual dietary variability, with increased range span resulting in weaker specialization for both males and females. We attribute our results to sex-based differences in movement, with females needing to specialize in their small ranges to maximize energy gain, and posit that the paradigm of individual prey specialization in sea otters with increased intraspecific competition may be a pattern driven largely by females. Our work highlights a potentially broader role of sex in the mechanistic pressures promoting and maintaining dietspecialization.

Temporal patterns in the foraging behavior of sea otters in Alaska

ABSTRACTActivity time budgets in apex predators have been proposed as indicators of population status relative to resource limitation or carrying capacity. We used archival time‐depth recorders implanted in 15 adult female and 4 male sea otters (Enhydra lutris) from the northernmost population of the species, Prince William Sound, Alaska, USA, to examine temporal patterns in their foraging behavior. Sea otters that we sampled spent less time foraging during summer (females 8.8 hr/day, males 7.9 hr/day) than other seasons (females 10.1–10.5 hr/day, males 9.2–9.5 hr/day). Both sexes showed strong preferences for diurnal foraging and adjusted their foraging effort in response to the amount of available daylight. One exception to this diurnal foraging mode occurred after females gave birth. For approximately 3 weeks post‐partum, females switched to nocturnal foraging, possibly in an effort to reduce the risk of predation by eagles on newborn pups. We used multilevel mixed regression models to assess the contribution of several biological and environmental covariates to variation in the daily foraging effort of parous females. In the random effects only model, 87% of the total variation in foraging effort was within‐otter variation. The relatively small among‐otter variance component (13%) indicates substantial consistency in the foraging effort of sea otters in this northern population. In the top 3 models, 17% of the within‐otter variation was explained by reproductive stage, day length, wind speed, air temperature and a wind speed × air temperature interaction. This study demonstrates the potential importance of environmental and reproductive effects when using activity budgets to assess population status relative to carrying capacity. Published 2014. This article is a U.S. Government work and is in the public domain in the USA.

MODIFIED-CLOSED CASTRATION: A NOVEL TECHNIQUE FOR SEA OTTER (ENHYDRA LUTRIS NEREIS) ORCHIECTOMIES

A novel surgical technique was used in the routine castrations of two intact male southern sea otters, Enhydra lutris nereis, housed at the Georgia Aquarium (Atlanta, Georgia, USA). This technique involved incising the parietal vaginal tunic to allow placement of double ligation of the ductus deferens, testicular artery, and pampiniform plexus en masse. After ligating and transecting these structures, they were introduced back into the tunic, which was closed with a circumferential ligature. The incision site was closed in a routine manner. Both otters recovered well from the procedure. One otter had mild cutaneous dehiscence postoperatively, and the other had no obvious complications. Benefits of this procedure include reduced risk of ligature slippage or loosening and resultant hemorrhage, as provided by the traditional open portion of the castration, and decreased postoperative swelling, as provided by the closed part of the castration. To the authors' knowledge, this is the first time this technique has been described for use in sea otters.

Lesions and Behavior Associated with Forced Copulation of Juvenile Pacific Harbor Seals (&lt;I&gt;Phoca vitulina richardsi&lt;/I&gt;) by Southern Sea Otters (&lt;I&gt;Enhydra lutris nereis&lt;/I&gt;)

Nineteen occurrences of interspecific sexual behavior between male southern sea otters (Enhydra lutris nereis) and juvenile Pacific harbor seals (Phoca vitulina richardsi) were reported in Monterey Bay, California, between 2000 and 2002. At least three different male sea otters were observed harassing, dragging, guarding, and copulating with harbor seals for up to 7 d postmortem. Carcasses of 15 juvenile harbor seals were recovered, and seven were necropsied in detail by a veterinary pathologist. Necropsy findings from two female sea otters that were recovered dead from male sea otters exhibiting similar behavior are also presented to facilitate a comparison of lesions. The most frequent lesions included superficial skin lacerations; hemorrhage around the nose, eyes, flippers, and perineum; and traumatic corneal erosions or ulcers. The harbor seals sustained severe genital trauma, ranging from vaginal perforation to vagino-cervical transection, and colorectal perforations as a result of penile penetration. One harbor seal developed severe pneumoperitoneum subsequent to vaginal perforation, which was also observed in both female sea otters and has been reported as a postcoital lesion in humans. This study represents the first description of lesions resulting from forced copulation of harbor seals by sea otters and is also the first report of pneumoperitoneum secondary to forced copulation in a nonhuman animal. Possible explanations for this behavior are discussed in the context of sea otter biology and population demographics.

Interannual assessment of territory quality for male sea otters (Enhydra lutris) in Simpson Bay, Prince William Sound, Alaska

Territory quality and site fidelity were assessed for male sea otters ( Enhydra lutris (L., 1758)) in Simpson Bay, Prince William Sound, Alaska, during the summer months (May–August) from 2003 to 2006. Territory quality was coded with a score of 0–2 for four attributes: territory size, shoreline enclosure, territory accessibility, and number of females observed feeding in each territory. Total quality scores ranged from 0.14 to 1.96 (0.9 ± 0.61, mean ± SD), with the highest quality territories having large areas, moderate to little shoreline enclosure, high accessibility, and many foraging females. A total of 13.1 km2 (62.5%) of Simpson Bay was occupied at least once by a territorial male during the 4-year study, and 7.9 km2 (37.5%) was never occupied by a territorial male. There were 7 ± 2.4 (mean ± SD) territorial males in Simpson Bay each year. Five males maintained territories for two consecutive years, with their respective territories having a 16%–34% overlap between years, indicating that repeat territory holders returned to the same area. The number of territorial sea otters, the size of territories, and whether or not a sea otter was a repeat territory holder did not significantly influence the size or quality of male territories.

PERSISTENT ORGANIC POLLUTANTS IN THE BLOOD OF FREE-RANGING SEA OTTERS (ENHYDRA LUTRIS SSP.) IN ALASKA AND CALIFORNIA

As part of tagging and ecologic research efforts in 1997 and 1998, apparently healthy sea otters of four age-sex classes in six locations in Alaska and three in California were sampled for persistent organic pollutants (POPs) and other chemicals of ecologic or environmental concern (COECs). Published techniques for the detection of POPs (specifically ∑polychlorinated biphenyls [PCBs], ∑DDTs, ∑hexachlorocyclohexanes [HCHs], ∑polycyclic aromatic hydrocarbons [PAHs], ∑chlordanes [CHLs], hexachlorobenzene [HCB], dieldrin, and mirex) in the tissue of dead otters were modified for use with serum from live sea otters. Toxic equivalencies (TEQs) were calculated for POPs with proven bioactivity. Strong location effects were seen for most POPs and COECs; sea otters in California generally showed higher mean concentrations than those in Alaska. Differences in contaminant concentrations were detected among age and sex classes, with high levels frequently observed in subadults. Very high levels of ∑DDT were detected in male sea otters in Elkhorn Slough, California, where strong freshwater outflow from agricultural areas occurs seasonally. All contaminants except mirex differed among Alaskan locations; only ∑DDT, HCB, and chlorpyrifos differed within California. High levels of ∑PCB (particularly larger, more persistent congeners) were detected at two locations in Alaska where associations between elevated PCBs and military activity have been established, while higher PCB levels were found at all three locations in California where no point source of PCBs has been identified. Although POP and COEC concentrations in blood may be less likely to reflect total body burden, concentrations in blood of healthy animals may be more biologically relevant and less influenced by state of nutrition or perimortem factors than other tissues routinely sampled.

Summer activity pattern and Field Metabolic Rate of adult male sea otters ( Enhydra lutris) in a soft sediment habitat in Alaska

Individual follows and instantaneous sampling were used to examine the behavior of adult male sea otters ( Enhydra lutris) in Simpson Bay, Prince William Sound, Alaska, during the summer (May to August) of 2005 and 2006. Six behaviors (foraging, grooming, interacting with other otters, patrolling, resting, and swimming at the surface) were observed during four time periods (dawn, day, dusk, and night) to create 24-h activity budgets. Adult male sea otters were observed during 190 focal follows, representing 98 h of observation. Male otters allocated 27% of their time (over a 24-h period) to resting, 26% to swimming, 19% to grooming, 14% foraging, 9% to patrolling and 5% to interacting with other otters. Field Metabolic Rate (FMR) was estimated by combining the energetic costs for foraging, grooming, resting, and swimming behavior of captive otters from Yeates et al. (2007) with our activity budgets. Our study considered ‘patrolling’ to be energetically similar to ‘swimming’ and therefore the two categories were combined. This combined category accounted for the greatest percentage (43%) of energy expended each day followed by grooming (23%), resting (15%), feeding (13%) and other (5%). The estimated weight specific FMR for all activities was 686.7 kJ day − 1 kg − 1 and the total FMR was 19.04 MJ day − 1 . The FMR was 6.6 times the resting metabolic rate and 2.2 times greater than the allometric prediction for terrestrial mammals of similar size but similar to other marine mammals. With a peak summer sea otter density in 5.6 otters km − 2 , the low percentage of time spent foraging (even after correction for possible sampling biases) indicates that Simpson Bay is still below equilibrium density.

Diving and foraging energetics of the smallest marine mammal, the sea otter (Enhydra lutris)

As the smallest and one of the most recently evolved marine mammals, sea otters face physiological challenges rarely encountered by larger, more derived aquatic species. To examine the effect of these challenges on foraging costs and resultant daily energy budgets, we measured the energetics of resting, grooming, diving and foraging for adult, male sea otters. The energy expended for these different behaviors as determined from open flow respirometry was then standardized across activity budgets measured for wild sea otters to estimate field metabolic rates (FMR). We found that the metabolic rate of captive otters performing single dives ranging in duration from 40 to 192 s was 17.6+/-0.5 ml O(2) kg(-1) min(-1) and only 1.3 times resting rates. This rate increased significantly if the animals foraged during submergence. The cost of a foraging dive for sea otters was nearly twice that predicted for phocid seals, which was attributed in part to elevated locomotor costs associated with buoyancy and swimming style. Our behavioral studies indicate that wild sea otters spend the greatest proportion of the day feeding and resting, with the largest daily energy expenditure (6.1+/-1.1 MJ day(-1)) associated with foraging. The resulting mean FMR for wild sea otters based on the energy expended for all behaviors was 15.7+/-2.7 MJ day(-1) and matched predicted FMR values based upon a regression of known FMR values for other marine mammals across a range of body sizes. This was achieved by counterbalancing elevated foraging costs with prolonged periods of rest on the water surface.

Association between Perfluorinated Compounds and Pathological Conditions in Southern Sea Otters

Concentrations of four perfluorinated contaminants, including perfluorooctanesulfonate (PFOS) and perfluorooctanoic acid (PFOA), were measured in liver tissue from 80 adult female sea otters collected from the California coast during 1992-2002. Concentrations of PFOS and PFOA were in the ranges of <1-884 and <5-147 ng/g, wet wt, respectively. Concentrations of PFOA in the livers of these sea otters were among the highest values reported for marine mammals to date. Liver tissue from 6 male sea otters also was analyzed and contained significantly higher concentrations of PFOS than did tissues from female otters. To examine the association between exposures and potential effects, concentrations of PFOS and PFOA were compared among the adult female otters that died from infectious diseases, noninfectious causes, and from apparent emaciation. Concentrations of both PFOA and PFOS were significantly higher in sea otters in the infectious disease category than in the noninfectious category. Concentrations of PFOS and PFOA were not significantly different between noninfectious and emaciated otters, suggesting that the poor nutritive (body) status of emaciated otters did not affectthe concentrations of perfluorochemicals in livers. Concentrations of PFOA increased significantly from 1992 to 2002, whereas PFOS concentrations increased from 1992to 1998 and then decreased after 2000. Significant association between infectious diseases and elevated concentrations of PFOS/PFOA in the livers of sea otters is a cause for concern and suggests the need for further studies.

Territory quality of male sea otters in Prince William Sound, Alaska: relation to body and territory maintenance behaviors

Based on optimality models, lekking males holding higher quality territories should spend more effort on territory maintenance and less effort on body maintenance. We tested the hypothesis that benefits are correlated with costs for male sea otters, Enhydra lutris (L., 1758). Activity state was recorded during focal follows of 10 individuals (n = 127). Higher quality territories had larger area, more food resources attractive to females, a higher ratio of protective shoreline edge, and higher accessibility for females evading male harassment. Contrary to our prediction, territory quality was uncorrelated with measures of cost: territory maintenance (patrolling, interacting) and body maintenance (feeding, grooming). We rejected the hypothesis that proximate benefits would be correlated with costs and suggested the following alternative working hypotheses: (i) given the high metabolic rate of sea otters, male breeding success may depend as much on maintaining body condition as maintaining a territory; (ii) higher quality territories with shoreline edge may not require additional patrolling effort; (iii) males may not expend extra effort in territory maintenance until more females come into estrus; or (iv) our seasonal measures of the benefits and costs of territoriality may not have accurately reflected factors influencing the switch between territorial and non-territorial tactics.

Behavior of Territorial Male Sea Otters (&lt;I&gt;Enhydra lutris&lt;/I&gt;) in Prince William Sound, Alaska

Photo-identification and focal animal sampling were used to examine the daytime behavior of territorial male sea otters (Enhydra lutris) in Simpson Bay, Prince William Sound, Alaska, during the summer (June to August) of 2003. The average number of otters (all age classes of males and females) in the study area was 121 ± 12.1 SD (n = 5 surveys). The bout duration of six behaviors (resting, grooming, foraging, interacting with other otters, swimming at the surface, and patrolling), activity time budgets, and interactions with females were determined for territorial males. Ten males were observed during 183 focal follows (i.e., observation periods), representing 92 h of observation. More time was spent foraging (30%) than on any other activity, and foraging bouts were longer than all other activities. Males interacted with females with pups (59%) and with single females (41%). Two of three consortships (i.e., mating associations lasting ca. three days) were formed with single females. Sixty-seven percent of interactions between territorial males were aggressive and were longer than one min.

FORAGING DEPTHS OF SEA OTTERS AND IMPLICATIONS TO COASTAL MARINE COMMUNITIES

AbstractWe 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 &gt;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 &gt;55 m.

AGE‐ AND SEX‐SPECIFIC MORTALITY AND POPULATION STRUCTURE IN SEA OTTERS

AbstractWe used 742 beach‐cast carcasses to characterize age‐ and sex‐specific sea otter mortality during the winter of 1990‐1991 at Bering Island, Russia. We also examined 363 carcasses recovered after the 1989 grounding of the T/V Exxon Valdez, to characterize age and sex composition in the living western Prince William Sound (WPWS) sea otter population. At Bering Island, mortality was male‐biased (81%), and 75% were adults. The WPWS population was female‐biased (59%) and most animals were subadult (79% of the males and 45% of the females). In the decade prior to 1990‐1991 we found increasing sea otter densities (particularly among males), declining prey resources, and declining weights in adult male sea otters at Bering Island. Our findings suggest the increased mortality at Bering Island in 1990‐1991 was a density‐dependent population response. We propose male‐maintained breeding territories and exclusion of juvenile females by adult females, providing a mechanism for maintaining densities in female areas below densities in male areas and for potentially moderating the effects of prey reductions on the female population. Increased adult male mortality at Bering Island in 1990‐1991 likely modified the sex and age class structure there toward that observed in Prince William Sound.

ATYPICAL INTERACTIONS BETWEEN MALE SOUTHERN SEA OTTERS and PINNIPEDS

ATYPICAL INTERACTIONS BETWEEN MALE SOUTHERN SEA OTTERS and PINNIPEDS