Photo-Identification of Sea Otters Using Nose Scars

Abstract: We evaluated the use of naturally occurring nose scars to identify individual sea otters (Enhydra lutris) in Simpson Bay, Prince William Sound, Alaska, USA. We spent 520 hours over 103 days conducting photo-identification surveys from June to August 2002 and 2003. Altogether, we identified 114 individuals. The number of sightings per individual ranged from 1 to 26, with an average of 3.3. The maximum number of sightings of an individual within a single year was 19. We saw 54 otters (47%)on >1 day, with an average of 8.1 sightings per individual for those seen more than once. We identified 8 individuals (19% of those identified in 2002) in both years. Males and otters of undetermined sex that we first sighted in June had the highest re-sighting rates. We considered 45% of all individuals encountered identifiable from nose scars. Nose scars were present in 63% (n = 19) of males, 45% (n = 45) of females, and 40% (n = 49) of otters of undetermined sex. Our results are similar to the results of photo-identification studies of other marine mammals, suggesting that this technique may be a useful tool for the individual identification of sea otters as well.     

Trends and Carrying Capacity of Sea Otters in Southeast Alaska

Sea otter populations in Southeast Alaska, USA, have increased dramatically from just over 400 translocated animals in the late 1960s to >8,000 by 2003. The recovery of sea otters to ecosystems from which they had been absent has affected coastal food webs, including commercially important fisheries, and thus information on expected growth and equilibrium abundances can help inform resource management. We compile available survey data for Southeast Alaska and fit a Bayesian state-space model to estimate past trends and current abundance. Our model improves upon previous analyses by partitioning and quantifying sources of estimation error, accounting for over-dispersion of aerial count data, and providing realistic measurements of uncertainty around point estimates of abundance at multiple spatial scales. We also provide estimates of carrying capacity (K) for Southeast Alaska, at regional and sub-regional scales, and analyze growth rates, current population status and expected future trends. At the regional scale, the population increased from 13,221 otters in 2003 to 25,584 otters in 2011. The average annual growth rate in southern Southeast Alaska (7.8%) was higher than northern Southeast Alaska (2.7%); however, growth varied at the sub-regional scale and there was a negative relationship between growth rates and the number of years sea otters were present in an area. Local populations vary in terms of current densities and expected future growth; the mean estimated density at K was 4.2 ± 1.58 sea otters/km² of habitat (i.e., the sub-tidal benthos between 0 m and 40 m depth) and current densities correspond on average to 50% of projected equilibrium values (range = 1–97%) with the earliest-colonized sub-regions tending to be closer to K. Assuming a similar range of equilibrium densities for currently un-occupied habitats, the projected value of K for all of Southeast Alaska is 74,650 sea otters. Future analyses can improve upon the precision of K estimates by employing more frequent surveys at index sites and incorporating environmental covariates into the process model to generate more accurate, location-specific estimates of equilibrium density. © 2019 The Authors. The Journal of Wildlife Management Published by Wiley Periodicals, Inc.     

Estimating Carrying Capacity for Sea Otters in British Columbia

ABSTRACT We estimated carrying capacity for sea otters (Enhydra lutris) in the coastal waters of British Columbia, Canada, by characterizing habitat according to the complexity of nearshore intertidal and sub-tidal contours. We modeled the total area of complex habitat on the west coast of Vancouver Island by first calculating the complexity of the Checleset Bay-Kyuquot Sound (CB-KS) region, where sea otters have been at equilibrium since the mid-1990s. We then identified similarly complex areas on the west coast of Vancouver Island (WCVI model), and adapted the model to identify areas of similar complexity along the entire British Columbia coast (BC model). Using survey data from the CB-KS region, we calculated otter densities for the habitat predicted by the 2 models. The density estimates for CB-KS were 3.93 otters/km² and 2.53 otters/km² for the WCVI and BC models, respectively, and the resulting 2 estimates of west coast of Vancouver Island complex habitat carrying capacity were not significantly different (WCVI model: 5,123, 95% CI = 3,337–7,104; BC model: 4,883, 95% CI = 3,223–6,832). The BC model identified the region presently occupied by otters on the central British Columbia coast, but the amount of coast-wide habitat it predicted (5,862 km²) was relatively small, and the associated carrying capacity estimate (14,831, 95% CI = 9,790–20,751) was low compared to historical accounts. We suggest that our model captured a type of high-quality or optimum habitat prevalent on the west coast of Vancouver Island, typified by the CB-KS region, and that suitable sea otter habitat elsewhere on the coast must include other habitat characteristics. We therefore calculated a linear, coast-wide carrying capacity of 52,459 sea otters (95% CI = 34,264–73,489)—a more realistic upper limit to sea otters in British Columbia. Our carrying capacity estimates are helping set population recovery targets for sea otters in Canada, and our habitat predictions represent a first step in Critical Habitat identification. This habitat-based approach to estimating carrying capacity is likely suitable for other nonmigratory, density-dependent species.     

Assessing Risks to Sea Otters and the Exxon Valdez Oil Spill: New Scenarios,Attributable Risk,and Recovery

The Exxon Valdez oil spill occurred more than two decades ago, and the Prince William Sound ecosystem has essentially recovered. Nevertheless, discussion continues on whether or not localized effects persist on sea otters (Enhydra lutris) at northern Knight Island (NKI) and, if so, what are the associated attributable risks. A recent study estimated new rates of sea otter encounters with subsurface oil residues (SSOR) from the oil spill. We previously demonstrated that a potential pathway existed for exposures to polycyclic aromatic hydrocarbons (PAHs) and conducted a quantitative ecological risk assessment using an individual-based model that simulated this and other plausible exposure pathways. Here we quantitatively update the potential for this exposure pathway to constitute an ongoing risk to sea otters using the new estimates of SSOR encounters. Our conservative model predicted that the assimilated doses of PAHs to the 1-in-1000th most-exposed sea otters would remain 1–2 orders of magnitude below the chronic effects thresholds. We re-examine the baseline estimates, post-spill surveys, recovery status, and attributable risks for this subpopulation. We conclude that the new estimated frequencies of encountering SSOR do not constitute a plausible risk for sea otters at NKI and these sea otters have fully recovered from the oil spill.     

A Quantitative Ecological Risk Assessment of the Toxicological Risks from Exxon Valdez Subsurface Oil Residues to Sea Otters at Northern Knight Island,Prince William Sound,Alaska

A comprehensive, quantitative risk assessment is presented of the toxicological risks from buried Exxon Valdez subsurface oil residues (SSOR) to a subpopulation of sea otters (Enhydra lutris) at Northern Knight Island (NKI) in Prince William Sound, Alaska, as it has been asserted that this subpopulation of sea otters may be experiencing adverse effects from the SSOR. The central questions in this study are: could the risk to NKI sea otters from exposure to polycyclic aromatic hydrocarbons (PAHs) in SSOR, as characterized in 2001–2003, result in individual health effects, and, if so, could that exposure cause subpopulation-level effects? We follow the U.S. Environmental Protection Agency (USEPA) risk paradigm by: (a) identifying potential routes of exposure to PAHs from SSOR; (b) developing a quantitative simulation model of exposures using the best available scientific information; (c) developing scenarios based on calculated probabilities of sea otter exposures to SSOR; (d) simulating exposures for 500,000 modeled sea otters and extracting the 99.9% quantile most highly exposed individuals; and (e) comparing projected exposures to chronic toxicity reference values. Results indicate that, even under conservative assumptions in the model, maximum-exposed sea otters would not receive a dose of PAHs sufficient to cause any health effects; consequently, no plausible toxicological risk exists from SSOR to the sea otter subpopulation at NKI.     

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.     

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.     

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.     

Status,trends, and equilibrium abundance estimates of the translocated sea otter population in Washington State

Sea otters (Enhydra lutris kenyoni) historically occurred in Washington State, USA, until their local extinction in the early 1900s as a result of the maritime fur trade. Following their extirpation, 59 sea otters were translocated from Amchitka Island, Alaska, USA, to the coast of Washington, with 29 released at Point Grenville in 1969 and 30 released at La Push in 1970. The Washington Department of Fish and Wildlife has outlined 2 main objectives for sea otter recovery: a target population level and a target geographic distribution. Recovery criteria are based on estimates of population abundance, equilibrium abundance (K), and geographic distribution; therefore, estimates of these parameters have important management implications. We compiled available survey data for sea otters in Washington State since their translocation (1977–2019) and fit a Bayesian state-space model to estimate past and current abundance, and equilibrium abundance at multiple spatial scales. We then used forward projections of population dynamics to explore potential scenarios of range recolonization and as the basis of a sensitivity analysis to evaluate the relative influence of movement behavior, frontal wave speed, intrinsic growth, and equilibrium density on future population recovery potential. Our model improves upon previous analyses of sea otter population dynamics in Washington by partitioning and quantifying sources of estimation error to estimate population dynamics, by providing robust estimates of K, and by simulating long-term population growth and range expansion under a range of realistic parameter values. Our model resulted in predictions of population abundance that closely matched observed counts. At the range-wide scale, the population size in our model increased from an average of 21 independent sea otters (95% CI = 13–29) in 1977 to 2,336 independent sea otters (95% CI = 1,467–3,359) in 2019. The average estimated annual growth rate was 12.42% and varied at a sub-regional scale from 6.42–14.92%. The overall estimated mean K density of sea otters in Washington was 1.71 ± 0.90 (SD) independent sea otters/km² of habitat (1.96 ± 1.04 sea otters/km², including pups), and estimated densities within the current range correspond on average to 87% of mean sub-regional equilibrium values (range = 66–111%). The projected value of K for all of Washington was 5,287 independent sea otters (95% CI = 2,488–8,086) and 6,080 sea otters including pups (95% CI = 2,861–9,300), assuming a similar range of equilibrium densities in currently un-occupied habitats. Sensitivity analysis of simulations of sea otter population growth and range expansion suggested that mean K density estimates in currently occupied sub-regions had the largest impact on predicted future population growth (r² = 0.52), followed by the rate of southward range expansion (r² = 0.26) and the mean K density estimate of currently unoccupied sub-regions to the south of the current range (r² = 0.04). Our estimates of abundance and sensitivity analysis of simulations of future population abundance and geographic range help determine population status in relation to population recovery targets and identify the most influential parameters affecting future population growth and range expansion for sea otters in Washington State.     

Monitoring population productivity in the saiga antelope

Effective conservation requires a good understanding of factors causing variation in population growth rate. We here analyse the relationship between female age and fecundity in the saiga antelope Saiga tatarica tatarica , a critically endangered ungulate of the Eurasian steppes and semideserts, at both individual and population levels. Annual variation in age structure and twinning rates was investigated using long-term datasets, sampling a total of 3308 females in four populations over more than 40 years. Further, a new non-invasive method is presented, estimating twinning rates from both calves and placentas encountered during calving aggregation transects. At an individual level, the most parsimonious model for twinning rates included three age classes (1, 2 and ≥3 years); however, the model with only two classes (1 and ≥2 years) was competitive and particularly useful for monitoring because these two age classes can reliably be determined by direct observation in the field. Among yearlings, 77.4% were fecund and 11.7% twinned, whereas among older females 94.6% were fecund and 72.6% twinned. At a population level, annual variation in age structure (proportion ≥2 years) correlated well with annual variation in twinning rate except in the north-west Pre-Caspian population. Our results suggest that the recent poaching-driven collapse in saiga numbers has potentially resulted in reductions in fecundity, which will have an impact on population growth rate. Our results highlight the potential for monitoring of twinning rate using non-invasive calving aggregation transects as a cost-effective additional tool to population counts for monitoring the status of this critically endangered species. These monitoring methods are also potentially transferable to other ungulate species.     

A Rapid Method to Estimate Population Variables for African Elephants

Abstract: We developed a noninvasive method to estimate reproductive and survival parameters for free-ranging African savannah elephants (Loxodonta africana africana) and used these to estimate finite population growth rates. We used published data from 2 populations with known growth rates and birth and survival histories to validate our technique. Based on body measurements, our method yielded estimates of age at first and last calving, calving interval, and age-specific survival rates that were similar to those determined during long-term studies at both Addo Elephant National Park and Amboseli National Park. Our technique generated population data required to estimate population growth rates. The method may be particularly useful where censuses yield imprecise estimates or where long-term population data are unavailable. (JOURNAL OF WILDLIFE MANAGEMENT 72(3):822–829; 2008)     

Sea growth, smolt age and age at sexual maturation in Atlantic salmon

Relationships between growth at sea, smolt size and age at sexual maturation of Atlantic salmon Salmo salar were tested. The fish were offspring of brood stocks sampled in eight Norwegian rivers at latitudes between 59° and 70° N, hatchery reared and released at smolting at the mouth of the River Imsa (59° N). Smolt size influenced the subsequent growth rate of Atlantic salmon. The larger the fish were at release, the slower the yearly length increment at sea. Mean sea age at sexual maturity, measured as proportion of the returning adults attaining sexual maturity at sea age 2 years, was significantly correlated with mean growth rate during the first year at sea and mean smolt size ( r ²= 0·74, P < 0·001). Fish attaining maturity at a relatively high sea age were more fast growing during their first year at sea than those maturing at a younger age. The results indicate that high sea age at sexual maturation is a population-specific characteristic and associated with high early growth rate at sea.     

Fitness measures in selection analyses: sensitivity to the overall number of offspring produced in a lifetime

Age at first (α) and last (ω) breeding are important life‐history traits; however, the direction and strength of selection detected on traits may vary depending on the fitness measure used. We provide the first estimates of lifetime breeding success (LBS) and λ_ind (the population growth rate of an individual) of European badgers Meles meles, by genotyping 915 individuals, sampled over 18 years, for 22 microsatellites. Males are slightly larger than females, and the opportunity for selection was slightly greater for males, as predicted. λ_ind and LBS both performed well in predicting the number of grand‐offspring, and both detected selection for a late ω, until the age of eight. Differential selection (S′_α) for an early α, however, was only detected using LBS, not with λ_ind. In declining populations (λ_ind < 1) selection favours reproduction later in life, whereas early reproduction is selected in increasing populations (λ_ind > 1). As 41% of badgers were assigned only one offspring (λ_ind < 1), whereas 40% were assigned more than two (λ_ind > 1), this cancelled out S′_α measured by λ_ind.     

Strength of density feedback in census data increases from slow to fast life histories

Life-history theory predicts an increasing rate of population growth among species arranged along a continuum from slow to fast life histories. We examine the effects of this continuum on density-feedback strength estimated using long-term census data from >700 vertebrates, invertebrates, and plants. Four life-history traits (Age at first reproduction, Body size, Fertility, Longevity) were related statistically to Gompertz strength of density feedback using generalized linear mixed-effects models and multi-model inference. Life-history traits alone explained 10 to 30% of the variation in strength across species (after controlling for time-series length and phylogenetic nonindependence). Effect sizes were largest for body size in mammals and longevity in birds, and density feedback was consistently stronger for smaller-bodied and shorter-lived species. Overcompensatory density feedback (strength <-1) occurred in 20% of species, predominantly at the fast end of the life-history continuum, implying relatively high population variability. These results support the idea that life history leaves an evolutionary signal in long-term population trends as inferred from census data. Where there is a lack of detailed demographic data, broad life-history information can inform management and conservation decisions about rebound capacity from low numbers, and propensity to fluctuate, of arrays of species in areas planned for development, harvesting, protection, and population recovery.     

Food availability as a major driver in the evolution of life‐history strategies of sibling species

Life‐history theory predicts trade‐offs between reproductive and survival traits such that different strategies or environmental constraints may yield comparable lifetime reproductive success among conspecifics. Food availability is one of the most important environmental factors shaping developmental processes. It notably affects key life‐history components such as reproduction and survival prospect. We investigated whether food resource availability could also operate as an ultimate driver of life‐history strategy variation between species. During 13 years, we marked and recaptured young and adult sibling mouse‐eared bats (Myotis myotis and Myotis blythii) at sympatric colonial sites. We tested whether distinct, species‐specific trophic niches and food availability patterns may drive interspecific differences in key life‐history components such as age at first reproduction and survival. We took advantage of a quasi‐experimental setting in which prey availability for the two species varies between years (pulse vs. nonpulse resource years), modeling mark‐recapture data for demographic comparisons. Prey availability dictated both adult survival and age at first reproduction. The bat species facing a more abundant and predictable food supply early in the season started its reproductive life earlier and showed a lower adult survival probability than the species subjected to more limited and less predictable food supply, while lifetime reproductive success was comparable in both species. The observed life‐history trade‐off indicates that temporal patterns in food availability can drive evolutionary divergence in life‐history strategies among sympatric sibling species.