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Reliable age estimation is an essential tool to assess the status of wildlife populations and inform successful management. Aging methods, however, are often limited by too few data, skewed demographic representation, and by single or uncertain morphometric relationships. In this study, we synthesize age estimates in southern sea otters Enhydra lutris nereis from 761 individuals across 34 years of study, using multiple noninvasive techniques and capturing all life stages from 0 to 17 years of age. From wild, stranded, and captive individuals, we describe tooth eruptions, tooth wear, body length, nose scarring, and pelage coloration across ontogeny and fit sex‐based growth functions to the data. Dental eruption schedules provided reliable and identifiable metrics spanning 0.3–9 months. Tooth wear was the most reliable predictor of age of individuals aged 1–15 years, which when combined with total length, explained >93% of observed age. Beyond age estimation, dental attrition also indicated the maximum lifespan of adult teeth is 13‒17 years, corresponding with previous estimates of life expectancy. Von Bertalanffy growth function model simulations of length at age gave consistent estimates of asymptotic lengths (male Loo = 126.0‒126.8 cm, female Loo = 115.3‒115.7 cm), biologically realistic gestation periods (t 0 = 115 days, SD = 10.2), and somatic growth (male k = 1.8, SD = 0.1; female k = 2.1, SD = 0.1). Though exploratory, we describe how field radiographic imaging of epiphyseal plate development or fusions may improve aging of immature sea otters. Together, our results highlight the value of integrating information from multiple and diverse datasets to help resolve conservation problems., By synthesizing more than two decades of southern sea otter physical examination data from strandings, captive‐rearing for release, and field studies, we provide strong support for adopting minimally invasive aging techniques based on metrics obtainable during brief field capture. These techniques provide the means to gather age‐specific information from a living, wild population to inform management decisions focused on restoring sea otters and their ecosystems throughout California. Our results also highlight the value of integrating information from multiple and diverse datasets to help resolve conservation problems.

The sea otter (Enhydra lutris) is a marine mammal hunted to near extinction during the 1800s. Despite their well‐known modern importance as a keystone species, we know little about historical sea otter ecology. Here, we characterize the ecological niche of ancient southern sea otters (E. lutris nereis) using δ13C analysis and δ15N analysis of bones recovered from archaeological sites spanning ~7,000 to 350 years before present (N = 112 individuals) at five regions along the coast of California. These data are compared with previously published data on modern animals (N = 165) and potential modern prey items. In addition, we analyze the δ15N of individual amino acids for 23 individuals to test for differences in sea otter trophic ecology through time. After correcting for tissue‐specific and temporal isotopic effects, we employ nonparametric statistics and Bayesian niche models to quantify differences among ancient and modern animals. We find ancient otters occupied a larger isotopic niche than nearly all modern localities; likely reflecting broader habitat and prey use in prefur trade populations. In addition, ancient sea otters at the most southerly sites occupied an isotopic niche that was more than twice as large as ancient otters from northerly regions. This likely reflects greater invertebrate prey diversity in southern California relative to northern California. Thus, we suggest the potential dietary niche of sea otters in southern California could be larger than in central and northern California. At two sites, Año Nuevo and Monterey Bay, ancient otters had significantly higher δ15N values than modern populations. Amino acid δ15N data indicated this resulted from shifting baseline isotope values, rather than a change in sea otter trophic ecology. Our results help in better understanding the contemporary ecological role of sea otters and exemplify the strength of combing zooarchaeological and biological information to provide baseline data for conservation efforts., We measured the dietary niche of modern and ancient Southern sea otters in California using stable isotope analysis. We find the size of the realized niche in southern California has decreased over time. These results should be considered for effective management and conservation efforts of this threatened species.

Sea otters (Enhydra lutris) have the highest mass-specific metabolic rate of any marine mammal, which is superimposed on the inherently high costs of reproduction and lactation in adult females. These combined energetic demands have been implicated in the poor body condition and increased mortality of female sea otters nearing the end of lactation along the central California coast. However, the cost of lactation is unknown and currently cannot be directly measured for this marine species in the wild. Here, we quantified the energetic demands of immature sea otters across five developmental stages as a means of assessing the underlying energetic challenges associated with pup rearing that may contribute to poor maternal condition. Activity-specific metabolic rates, daily activity budgets and field metabolic rates (FMR) were determined for each developmental stage. Mean FMR of pre-molt pups was 2.29±0.81 MJ day−1 and increased to 6.16±2.46 and 7.41±3.17 MJ day−1 in post-molt pups and dependent immature animals, respectively. Consequently, daily energy demands of adult females increase 17% by 3 weeks postpartum and continue increasing to 96% above pre-pregnancy levels by the average age of weaning. Our results suggest that the energetics of pup rearing superimposed on small body size, marine living and limited on-board energetic reserves conspire to make female sea otters exceptionally vulnerable to energetic shortfalls. By controlling individual fitness, maternal behavior and pup provisioning strategies, this underlying metabolic challenge appears to be a major factor influencing current population trends in southern sea otters (Enhydra lutris nereis).

The existence of individual prey specializations has been reported for an ever-growing number of taxa, and has important ramifications for our understanding of predator–prey dynamics. We use the California sea otter population as a case study to validate the use of archival time–depth data to detect and measure differences in foraging behaviour and diet. We collected observational foraging data from radio-tagged sea otters that had been equipped with Mk9 time depth recorders (TDRs, Wildlife Computers, Redmond, WA). After recapturing the study animals and retrieving the TDRs it was possible to compare the two data types, by matching individual dives from the TDR record with observational data and thus examining behavioural correlates of capture success and prey species. Individuals varied with respect to prey selection, aggregating into one of three distinct dietary specializations. A number of TDR-derived parameters, particularly dive depth and post-dive surface interval, differed predictably between specialist types. A combination of six dive parameters was particularly useful for discriminating between specialist types, and when incorporated into a multivariate cluster analysis, these six parameters resulted in classification of 13 adult female sea otters into three clusters that corresponded almost perfectly to the diet-based classification (1 out of 13 animals was misclassified). Thus based solely on quantifiable traits of time–depth data that have been collected over an appropriate period (in this case 1 year per animal), it was possible to assign female sea otters to diet type with >90% accuracy. TDR data can thus be used as a tool to measure the degree of individual specialization in sea otter populations, a conclusion that will likely apply to other diving marine vertebrates as well. Our ultimate goals must be both to understand the causes of individual specialization, and to incorporate such variation into models of population- and community-level food web dynamics.

Summary 1 Growth models for body mass and length were fitted to data collected from 1842 sea otters Enhydra lutris shot or live-captured throughout south-west Alaska between 1967 and 2004. Growth curves were constructed for each of two main year groups: 1967–71 when the population was at or near carrying capacity and 1992–97 when the population was in steep decline. Analyses of data collected from animals caught during 2004, when the population density was very low, were precluded by a small sample size and consequently only examined incidentally to the main growth curves. 2 Growth curves demonstrated a significant increase in body mass and body length at age in the 1990s. Asymptotic values of body mass were 12–18% higher in the 1990s than in the 1960s/70s, and asymptotic values for body length were 10–11% higher between the same periods. Data collected in 2004 suggest a continued increase in body size, with nearly all data points for mass and length falling significantly above the 1990s growth curves. 3 In addition to larger asymptotic values for mass and length, the rate of growth towards asymptotic values was more rapid in the 1990s than in the 1960s/70s: sea otters reached 95% of asymptotic body mass and body length 1–2 years earlier in the 1990s. 4 Body condition (as measured by the log mass/log length ratio) was significantly greater in males than in females. There was also an increasing trend from the 1960s/70s through 2004 despite much year-to-year variation. 5 Population age structures differed significantly between the 1960s/70s and the 1990s with the latter distribution skewed toward younger age classes (indicating an altered lx function) suggesting almost complete relaxation of age-dependent mortality patterns (i.e. those typical of food-limited populations). 6 This study spanned a period of time over which the population status of sea otters in the Aleutian archipelago declined precipitously from levels at or near equilibrium densities at some islands in the 1960s/70s to

Recently published data demonstrated a sea otter population collapse across theAleutian archipelago (Doroff et al. 2003). This assessment, undertaken in responseto earlier reports of precipitous declines at several islands (Estes et al. 1998), wasbased on archipelago-wide aerial surveys conducted in 1965, 1992, and 2000, andskiff surveys of particular islands or small island groups during various years fromthe early 1990s through 2000. To summarize the main findings: (1) aerial countsdeclined by about 70% from 1992 to 2000, (2) populations at those islandsthought by Kenyon (1969) to be at or near carrying capacity in 1965 had declined88% by 2000, (3) most of the overall decline occurred after the late 1980s or early1990s, and (4) sea otters had reached a uniformly low population density (numbersper unit length of shoreline) throughout the Aleutian archipelago by 2000. Doroffet al. (2003) interpreted this latter finding to mean that the factors responsible forthe decline were density-dependent, thus implying that a steady state may havebeen attained between killer whale predation, the purported cause of the decline(Estes et al. 1998), and the reduced availability of sea otters. If that conjecture is

Summary 1. Longitudinal records of prey selection by 10 adult female sea otters on the Monterey Peninsula, California, from 1983 to 1990 demonstrate extreme inter-individual variation in diet. Variation in prey availability cannot explain these differences as the data were obtained from a common spatial-temporal area. 2. Individual dietary patterns persisted throughout our study, thus indicating that they are life-long characteristics. 3. Individual dietary patterns in sea otters appear to be transmitted along matrilines, probably by way of learning during the period of mother‐young association. 4. Efficient utilization of different prey types probably requires radically different sensory/motor skills, each of which is difficult to acquire and all of which may exceed the learning and performance capacities of any single individual. This would explain the absence of generalists and inertia against switching, but not the existence of alternative specialists. 5. Such individual variation might arise in a constant environment from frequencydependent effects, whereby the relative benefit of a given prey specialization depends on the number of other individuals utilizing that prey. Additionally, many of the sea otter’s prey fluctuate substantially in abundance through time. This temporal variation, in conjunction with matrilineal transmission of foraging skills, may act to mediate the temporal dynamics of prey specializations. 6. Regardless of the exact cause, such extreme individual variation in diet has broad ramifications for population and community ecology. 7. The published literature indicates that similar patterns occur in many other species.

After nearly a century of recovery from overhunting, sea otter populations are in abrupt decline over large areas of western Alaska. Increased killer whale predation is the likely cause of these declines. Elevated sea urchin density and the consequent deforestation of kelp beds in the nearshore community demonstrate that the otter's keystone role has been reduced or eliminated. This chain of interactions was probably initiated by anthropogenic changes in the offshore oceanic ecosystem.