USGS ScienceSearch

Geology topics

Michelle M. Staedler

Publications and source records attributed to Michelle M. Staedler.

At least 19 recordsLinked to original sources

Quantifying southern sea otter (Enhydra lutris nereis) reactions to a quadcopter drone in central California

Drones are useful for wildlife research and management, but they can cause disturbance and harassment to wildlife. Sea otters ( Enhydra lutris ) are candidates for drone-based observation and monitoring but are vulnerable to disturbance. No studies have evaluated drone effects on sea otter behavior, but based on prior disturbance studies, we hypothesized: (1) sea otters would exhibit behaviors indicating higher reactivity in the presence of drones than when drones were absent and (2) drone disturbance to sea otters would be greater when drones were closer. At two sites in Monterey Bay, CA, we conducted 37 observational sessions, recording behavior codes for focal sea otters during a baseline (no drone) period and three consecutive drone flights. Data were analyzed using ANOVA and ordinal logistic regression models. At both locations, focal sea otters had higher behavior codes during drone trials compared to baseline, and behavior codes increased with descending drone altitude. Pup presence, group size, flight trial number, and gull presence were significant covariables. We calculated multipliers to predict drone-mediated behavioral responses at a range of drone altitudes. Our findings can inform best practices for a variety of uses of drones around sea otters, including population monitoring, oil spill response, and drone photography/videography.

California

Keystone interdependence: Sea otter responses to a prey surplus following the collapse of a rocky intertidal predator

The sea star Pisaster ochraceus and sea otters ( Enhydra lutris ) are two predators capable of shaping rocky intertidal and kelp forest community structure and functioning. In 2013, a sea star wasting event decimated populations of Pisaster along the west coast of North America. The collapse of this species in the rocky intertidal revealed an unexpected relationship between two keystone predators. In this study, we show how the loss of Pisaster along the Monterey Peninsula, CA, USA led to an increase in mussel ( Mytilus californianus ) size and expansion into lower tidal zones. Before the sea star wasting event, the local sea otter population fluctuated around a near equilibrium. However, in the absence of Pisaster , sea otters increased their dietary intake on mussels, which contributed in part to a local population-level rise. These results demonstrate how the loss of a keystone predator in one ecosystem may impart population-level changes to another.

California

Top-predator recovery abates geomorphic decline of a coastal ecosystem

The recovery of top predators is thought to have cascading effects on vegetated ecosystems and their geomorphology 1,2 , but the evidence for this remains correlational and intensely debated 3,4 . Here we combine observational and experimental data to reveal that recolonization of sea otters in a US estuary generates a trophic cascade that facilitates coastal wetland plant biomass and suppresses the erosion of marsh edges—a process that otherwise leads to the severe loss of habitats and ecosystem services 5,6 . Monitoring of the Elkhorn Slough estuary over several decades suggested top-down control in the system, because the erosion of salt marsh edges has generally slowed with increasing sea otter abundance, despite the consistently increasing physical stress in the system (that is, nutrient loading, sea-level rise and tidal scour 7,8,9 ). Predator-exclusion experiments in five marsh creeks revealed that sea otters suppress the abundance of burrowing crabs, a top-down effect that cascades to both increase marsh edge strength and reduce marsh erosion. Multi-creek surveys comparing marsh creeks pre- and post-sea otter colonization confirmed the presence of an interaction between the keystone sea otter, burrowing crabs and marsh creeks, demonstrating the spatial generality of predator control of ecosystem edge processes: densities of burrowing crabs and edge erosion have declined markedly in creeks that have high levels of sea otter recolonization. These results show that trophic downgrading could be a strong but underappreciated contributor to the loss of coastal wetlands, and suggest that restoring top predators can help to re-establish geomorphic stability.

California

Tool use increases mechanical foraging success and tooth health in southern sea otters (Enhydra lutris nereis)

Although tool use may enhance resource utilization, its fitness benefits are difficult to measure. By examining longitudinal data from 196 radio-tagged southern sea otters ( Enhydra lutris nereis ), we found that tool-using individuals, particularly females, gained access to larger and/or harder-shelled prey. These mechanical advantages translated to reduced tooth damage during food processing. We also found that tool use diminishes trade-offs between access to different prey, tooth condition, and energy intake, all of which are dependent on the relative prey availability in the environment. Tool use allowed individuals to maintain energetic requirements through the processing of alternative prey that are typically inaccessible with biting alone, suggesting that this behavior is a necessity for the survival of some otters in environments where preferred prey are depleted.

Science

Southern (California) sea otter population status and trends at San Nicolas Island, 2020–2023

The population of southern sea otters ( Enhydra lutris nereis ) at San Nicolas Island, California, has been monitored annually since the translocation of 140 southern sea otters to the island was completed in 1990. Monitoring efforts have varied in frequency and type across years. In 2017, the U.S. Navy and the U.S. Fish and Wildlife Service initiated a southern sea otter monitoring and research plan to determine the effects of military readiness activities on the growth or decline of the southern sea otter population at San Nicolas Island. The southern sea otter is the only subspecies of sea otter in California (hereafter, “sea otter"). The monitoring program, at its basic level, includes seasonal surveys of population abundance, distribution, and foraging activity. From 2020 to 2023, we measured a 10-percent per annum increase in population abundance (95-percent confidence interval =0–20 percent), with 146 total individuals as of April 2023. Coinciding with the recent population growth, the sea otter distribution, which previously tended to concentrate on the island’s west end during 2003–2006 before shifting toward more use in the north and south sides during 2017–2019, appears to have shifted again during 2020–2023 to concentrate at the island’s east end. Forage data were collected between February 2020 and April 2023. There was a total of 773 forage dives in 60 forage bouts, with most of the identified prey on successful dives (n=401) recorded as sea urchins (66 percent), followed by bivalves (15 percent), snails (12 percent), and crabs (5.2 percent). Two lobsters and three abalone also were identified among the sea otter prey. Estimates of energy intake rates averaged 14.0 kilocalories per minute (95-percent confidence interval =10.8–17.2 kilocalories per minute). Monitoring data from the past two decades indicate that sea otters at San Nicolas Island have maintained a steady pattern of energy intake and population growth characteristic of a robust population, including a sixfold growth between 2000 and 2023. There was no conclusive evidence of density-dependent effects based on these patterns; however, estimates of energy intake rates for 2020–2023 were slightly lower than previous estimates from 2017 to 2019. Additionally, subtidal monitoring results at four sites around San Nicolas Island indicated that counts of purple sea urchins ( Strongylocentrotus purpuratus ) have increased between 2003 and 2023, whereas sea otter foraging surveys completed during the same period revealed that some sea otters have shifted toward higher consumption of purple sea urchins and bivalves compared to red sea urchins ( S. fransicanus ), which generally are the preferred larger prey of sea otters. These results contribute to the understanding of population dynamics and to the conservation and planning of future monitoring and research of sea otters at San Nicolas Island.

California

Characterizing the oral and distal gut microbiota of the threatened southern sea otter (Enhydra lutris nereis) to enhance conservation practice

The southern sea otter ( Enhydra lutris nereis ) is a threatened sub-species in coastal ecosystems. To understand better the role of diet, monitor health, and enhance management of this and other marine mammal species, we characterized the oral (gingival) and distal gut (rectal and fecal) microbiota of 158 wild southern sea otters living off the coast of central California, USA, and 12 captive sea otters, some of which were included in a diet shift experiment. We found that the sea otter fecal microbiota was distinct from that of three other otter species, and that captivity does not significantly alter the community structure of the sea otter gingival or distal gut microbiota. Metagenomic analysis unexpectedly revealed that the majority of sea otter fecal DNA is derived from prey, rather than from indigenous bacteria or host cells as with most other mammals. We speculate that a reduced bacterial biomass in the sea otter gut reflects rapid gut transit time and a particular strategy for foraging and energy harvest. This study establishes a reference for the healthy sea otter microbiota, highlights how a marine lifestyle may shape the mammalian microbiota, and may inform future health assessments and conservation management of sea otter populations.

California

An integrated population model for southern sea otters

Southern sea otters ( Enhydra lutris nereis ) have recovered slowly from their near extinction a century ago, and their continued recovery has been challenged by multiple natural and anthropogenic factors. Development of an integrated population model (IPM) for southern sea otters has been identified as a management priority, to help in evaluating the relative impacts of known threats and guide best management options for species recovery. An IPM represents an analytical modeling framework where various types of data relevant to animal health, population trends, and survival can be evaluated collectively to project future population dynamics under different resource management scenarios. Here, we describe the development of a spatially explicit IPM for southern sea otters that is fit by using Bayesian methods to multiple datasets including a time series of range-wide survey counts, estimated survival rates of tagged animals from telemetry-based population studies, and cause-of-death data from comprehensive necropsies of beach-cast carcasses. The core of the model is a stage-structured matrix, in which survival rates for a given life history stage, year, and location are computed as the outcome of multiple ‘competing risks,’ or hazards, allowing for spatiotemporal variation in each hazard, density-dependence, and stochasticity. The parameterized IPM was used to (1) examine how age and sex-specific hazards vary over space and time, (2) gain insights into density-dependent variation in specific hazards, (3) assess population-level effects of known mortality hazards in the past and in future projections, and (4) evaluate the relative benefits of various potential management actions to address these hazards. Our results indicated that different types of hazards have variable impacts at different life history stages of sea otters; for example, shark-bite mortality had a strong impact on mortality of subadult females but relatively low impacts on aged adult female survival, whereas End Lactation Syndrome showed just the opposite age-based pattern. There also was spatial and temporal variation in exposure to different hazards; for example, shark-bite mortality generally was highest at the north and south ends of the sea otter range, End Lactation Syndrome and cardiac disease were highest in the center part of the range, and harmful algal bloom intoxication and protozoal infection mortalities were highest around Morro Bay. The relative impacts of hazards depended on population density; for example, shark-bite mortality had the greatest effect on male survival when population abundance was low, but as densities increased the impacts of cardiac disease (for aged adults) and acanthocephalan peritonitis (for subadults) exceeded the effects of shark-bite mortality. Sensitivity analyses showed that modifying certain hazard rates can have substantial impacts on future population growth; for example, if the shark-bite hazard rate were to decrease by 20 percent, projected abundance after 50 years is predicted to be 18-percent higher, on average, than under baseline conditions. We used the IPM to evaluate the possible impacts of a potential management action: the reintroduction of sea otters to currently unoccupied parts of their historical range. We found that there were large increases in expected growth potential associated with reintroduction programs to various locations to the north and south of the currently occupied range, although a reintroduction to San Francisco Bay was projected to have the greatest potential impacts on future population growth. The IPM for southern sea otters presented here provides resource managers with a useful tool for evaluating the impacts of specific hazards, forecasting future population dynamics and range expansion, and evaluating alternative management scenarios.

California

Behavioral responses across a mosaic of ecosystem states restructure a sea otter–urchin trophic cascade

Consumer and predator foraging behavior can impart profound trait-mediated constraints on community regulation that scale up to influence the structure and stability of ecosystems. Here, we demonstrate how the behavioral response of an apex predator to changes in prey behavior and condition can dramatically alter the role and relative contribution of top-down forcing, depending on the spatial organization of ecosystem states. In 2014, a rapid and dramatic decline in the abundance of a mesopredator ( Pycnopodia helianthoides ) and primary producer ( Macrocystis pyrifera ) coincided with a fundamental change in purple sea urchin ( Strongylocentrotus purpuratus ) foraging behavior and condition, resulting in a spatial mosaic of kelp forests interspersed with patches of sea urchin barrens. We show that this mosaic of adjacent alternative ecosystem states led to an increase in the number of sea otters ( Enhydra lutris nereis ) specializing on urchin prey, a population-level increase in urchin consumption, and an increase in sea otter survivorship. We further show that the spatial distribution of sea otter foraging efforts for urchin prey was not directly linked to high prey density but rather was predicted by the distribution of energetically profitable prey. Therefore, we infer that spatially explicit sea otter foraging enhances the resistance of remnant forests to overgrazing but does not directly contribute to the resilience (recovery) of forests. These results highlight the role of consumer and predator trait-mediated responses to resource mosaics that are common throughout natural ecosystems and enhance understanding of reciprocal feedbacks between top-down and bottom-up forcing on the regional stability of ecosystems.

PNAS

Exposure to domoic acid is an ecological driver of cardiac disease in southern sea otters

Harmful algal blooms produce toxins that bioaccumulate in the food web and adversely affect humans, animals, and entire marine ecosystems. Blooms of the diatom Pseudo-nitzschia can produce domoic acid (DA), a toxin that most commonly causes neurological disease in endothermic animals, with cardiovascular effects that were first recognized in southern sea otters. Over the last 20 years, DA toxicosis has caused significant morbidity and mortality in marine mammals and seabirds along the west coast of the USA. Identifying DA exposure has been limited to toxin detection in biological fluids using biochemical assays, yet measurement of systemic toxin levels is an unreliable indicator of exposure dose or timing. Furthermore, there is little information regarding repeated DA exposure in marine wildlife. Here, the association between long-term environmental DA exposure and fatal cardiac disease was investigated in a longitudinal study of 186 free-ranging sea otters in California from 2001 – 2017, highlighting the chronic health effects of a marine toxin. A novel Bayesian spatiotemporal approach was used to characterize environmental DA exposure by combining several DA surveillance datasets and integrating this with life history data from radio-tagged otters in a time-dependent survival model. In this study, a sea otter with high DA exposure had a 1.7-fold increased hazard of fatal cardiomyopathy compared to an otter with low exposure. Otters that consumed a high proportion of crab and clam had a 2.5- and 1.2-times greater hazard of death due to cardiomyopathy than otters that consumed low proportions. Increasing age is a well-established predictor of cardiac disease, but this study is the first to identify that DA exposure affects the risk of cardiomyopathy more substantially in prime-age adults than aged adults. A 4-year-old otter with high DA exposure had 2.3 times greater risk of fatal cardiomyopathy than an otter with low exposure, while a 10-year old otter with high DA exposure had just 1.2 times greater risk. High Toxoplasma gondii titers also increased the hazard of death due to heart disease 2.4-fold. Domoic acid exposure was most detrimental for prime-age adults, whose survival and reproduction are vital for population growth, suggesting that persistent DA exposure will likely impact long-term viability of this threatened species. These results offer insight into the pervasiveness of DA in the food web and raise awareness of under-recognized chronic health effects of DA for wildlife at a time when toxic blooms are on the rise.

California

Robust age estimation of southern sea otters from multiple morphometrics

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 L oo = 126.0‒126.8 cm, female L oo = 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.

Ecology and Evolution

Emergence of a zoonotic pathogen in a coastal marine sentinel: Capillaria hepatica (syn. Calodium hepaticum)-associated hepatitis in southern sea otters (Enhydra lutris nereis)

Capillaria hepatica is a globally distributed zoonotic nematode parasite that most commonly infects feral and native rats. Soil contact, pica, and living in close proximity to rat populations are risk factors for human infection. Larval nematodes and eggs that were morphologically consistent with C. hepatica were observed microscopically in livers of stranded southern sea otters ( Enhydra lutris nereis ) from California. Large (90–100 × 45–55 μm), barrel-shaped non-embryonated parasite eggs with large polar prominences and a roughened or striated surface, or 105–120 μm diameter larval aphasmid nematode profiles with a prominent stichosome and hypodermal bands were observed in the livers of three otters. The liver of a fourth animal exhibited serpiginous tracts of necrosis, micro-cavitation and pleocellular inflammation, with intralesional linear eosinophilic material that resembled cuticle from degenerating metazoan parasites. Capillaria hepatica -associated hepatitis and capsular adhesions were the cause of death for one otter, and parasite-associated liver lesions were observed in all cases. All infected otters were adult females that stranded from 2006 through 2016 at multiple sites along the central California coast. All cases stranded from December through May; during and soon after peak seasonal precipitation and land-sea runoff in California. This same seasonal pattern has been reported for other land-based parasites infecting southern sea otters. Neither C. hepatica , nor any similar nematodes have been reported from marine mammals, and southern sea otters are not typical hosts for C. hepatica or any other nematode parasites. The most likely route of exposure was via freshwater runoff containing embryonated eggs liberated from predated or decomposing terrestrial hosts, especially rats. Similar to the land-based parasites Toxoplasma gondii and Sarcocystis neurona , C. hepatica eggs may be concentrated and transmitted through filter-feeding marine invertebrates that serve as southern sea otter prey, which may also pose an unrecognized public health risk for people who consume these species.

California

Species recovery and recolonization of past habitats: Lessons for science and conservation from sea otters in estuaries

Recovering species are often limited to much smaller areas than they historically occupied. Conservation planning for the recovering species is often based on this limited range, which may simply be an artifact of where the surviving population persisted. Southern sea otters ( Enhydra lutris nereis ) were hunted nearly to extinction but recovered from a small remnant population on a remote stretch of the California outer coast, where most of their recovery has occurred. However, studies of recently-recolonized estuaries have revealed that estuaries can provide southern sea otters with high quality habitats featuring shallow waters, high production and ample food, limited predators, and protected haul-out opportunities. Moreover, sea otters can have strong effects on estuarine ecosystems, fostering seagrass resilience through their consumption of invertebrate prey. Using a combination of literature reviews, population modeling, and prey surveys we explored the former estuarine habitats outside the current southern sea otter range to determine if these estuarine habitats can support healthy sea otter populations. We found the majority of studies and conservation efforts have focused on populations in exposed, rocky coastal habitats. Yet historical evidence indicates that sea otters were also formerly ubiquitous in estuaries. Our habitat-specific population growth model for California’s largest estuary—San Francisco Bay—determined that it alone can support about 6,600 sea otters, more than double the 2018 California population. Prey surveys in estuaries currently with (Elkhorn Slough and Morro Bay) and without (San Francisco Bay and Drakes Estero) sea otters indicated that the availability of prey, especially crabs, is sufficient to support healthy sea otter populations. Combining historical evidence with our results, we show that conservation practitioners could consider former estuarine habitats as targets for sea otter and ecosystem restoration. This study reveals the importance of understanding how recovering species interact with all the ecosystems they historically occupied, both for improved conservation of the recovering species and for successful restoration of ecosystem functions and processes.

California

Surrogate rearing a keystone species to enhance population and ecosystem restoration

Translocation and rehabilitation programs are critical tools for wildlife conservation. These methods achieve greater impact when integrated in a combined strategy for enhancing population or ecosystem restoration. From 2002-2016, we reared 37 orphaned southern sea otter (Enhydra lutris nereis) pups, using captive sea otters as surrogate mothers, then released them into a degraded coastal estuary. As a keystone species, observed increases in the local sea otter population unsurprisingly brought many ecosystem benefits. The role that surrogate-reared otters played in this success story, however, remained uncertain. To resolve this question, we developed an individual-based model (IBM) of the local population using surveyed individual fates (survival and reproduction) of surrogate-reared and wild-captured otters, and modeled estimates of immigration. Estimates derived from a decade of population monitoring indicated that surrogate-reared and wild sea otters experienced similar reproductive and survival rates. This was true for males and females, across all ages (1-13 years) and locations evaluated. The IBM simulations indicated that reconstructed counts of the wild population are best explained by surrogate-reared otters combined with low levels of unassisted immigration. In addition, the model shows that 55% of observed population growth over this period is attributable to surrogate-reared otters and their wild progeny. Together, our results indicate that the integration of surrogacy methods and reintroduction of juvenile sea otters helped establish a biologically successful population and restore a once-impaired ecosystem.

California

Drift and beaching patterns of sea otter carcasses and car tire dummies

Enumerating and examining marine animal carcasses is important for quantifying mortality rates and determining causes of mortality. Drifter experiments are one tool for estimating at‐sea mortality and determining factors affecting carcass drift, but they require validation to confirm drifters accurately replicate the drift characteristics of the species of interest. The goal of this study was to determine whether dummies constructed from car tires were appropriate substitutes for sea otter ( Enhydra lutris ) carcasses. We released 33 sets of targets (carcasses and dummies) in a one‐to‐one ratio on 15 randomly chosen dates between January 1995 and December 1996. They were telemetrically tracked until they beached or were no longer detected. Beaching rates were similar between carcasses (69.7%) and dummies (66.7%). Our results indicated that there was no statistical difference in the drifting pattern, as measured by distance traveled and location, between carcasses and dummies, and that cumulative wind speed, days since release, and release month were predictors of drift patterns. We concluded that dummies constructed from car tires do imitate sea otter carcasses and could be used to estimate at‐sea mortality of sea otters, or, if released during or after an oil spill, could be used to direct search efforts for carcasses.

California

Lactation and resource limitation affect stress responses, thyroid hormones, immune function, and antioxidant capacity of sea otters (Enhydra lutris)

Lactation is the most energetically demanding stage of reproduction in female mammals. Increased energetic allocation toward current reproduction may result in fitness costs, although the mechanisms underlying these trade‐offs are not well understood. Trade‐offs during lactation may include reduced energetic allocation to cellular maintenance, immune response, and survival and may be influenced by resource limitation. As the smallest marine mammal, sea otters (Enhydra lutris) have the highest mass‐specific metabolic rate necessitating substantial energetic requirements for survival. To provide the increased energy needed for lactation, female sea otters significantly increase foraging effort, especially during late‐lactation. Caloric insufficiency during lactation is reflected in the high numbers of maternal deaths due to End‐Lactation Syndrome in the California subpopulation. We investigated the effects of lactation and resource limitation on maternal stress responses, metabolic regulation, immune function, and antioxidant capacity in two subspecies of wild sea otters (northern: E. l. nereis and southern: E. l. kenyoni) within the California, Washington, and Alaska subpopulations. Lactation and resource limitation were associated with reduced glucocorticoid responses to acute capture stress. Corticosterone release was lower in lactating otters. Cortisol release was lower under resource limitation and suppression during lactation was only evident under resource limitation. Lactation and resource limitation were associated with alterations in thyroid hormones. Immune responses and total antioxidant capacity were not reduced by lactation or resource limitation. Southern sea otters exhibited higher concentrations of antioxidants, immunoglobulins, and thyroid hormones than northern sea otters. These data provide evidence for allocation trade‐offs during reproduction and in response to nutrient limitation but suggest self‐maintenance of immune function and antioxidant defenses despite energetic constraints. Income‐breeding strategists may be especially vulnerable to the consequences of stress and modulation of thyroid function when food resources are insufficient to support successful reproduction and may come at a cost to survival, and thereby influence population trends.

California

Gaps in kelp cover may threaten the recovery of California sea otters

Despite more than a century of federal protection, the California sea otter Enhydra lutris nereis remains threatened under the U.S. Endangered Species Act (ESA), and the population has not appreciably expanded its range in two decades. Here, we examine a novel dataset of 725 sea otter live strandings from 1984–2015 to gain insights into demographic and environmental factors underlying threats to sea otter recovery. Using multinomial logistic regression to evaluate spatiotemporal patterns of stranding causes, we demonstrate that increases in stranding rates, particularly outside the range center, are related to a substantial increase in shark bites. By contrast, trauma linked to human activities has declined dramatically, and now accounts for less than 5% of stranding cases. Within the range core, where the sea otter population seems regulated by prey availability, symptoms of energetic stress represent more than 63% of all strandings and are strongly associated with high sea otter density. Conversely, in range peripheries, the majority of strandings are caused by shark bite and neurological disease. Notably, these threats are virtually absent where nearshore habitat is characterized by at least 10% kelp canopy cover. Our analyses reveal that declining kelp cover may therefore constrain the population's spatial expansion and recovery in two key ways. Absence of kelp intensifies density‐independent threats in the range peripheries, and likely limits dispersal of reproductive females, which depend on kelp canopy for nursery habitat. These results highlight the significance of both top‐down and bottom‐up processes in population dynamics, and inform an ecosystem‐based approach to conservation planning.

California

The high cost of motherhood: End-lactation syndrome in southern sea otters ( Enhydra lutris nereis ) on the central California, USA, coast

Sea otters ( Enhydra lutris ) have exceptionally high energetic requirements, which nearly double during lactation and pup care. Thus, females are extremely vulnerable to caloric insufficiency. Despite a number of compensatory strategies, the metabolic challenge of reproduction culminates in numerous maternal deaths annually. Massive depletion of energy reserves results in a case presentation that we define as end-lactation syndrome (ELS), characterized by moderate to severe emaciation not attributable to a concurrent, independent disease process in females dying during late pup care or postweaning. We compiled detailed data for 108 adult female southern sea otters ( Enhydra lutris nereis ) examined postmortem that stranded in California, US, 2005–12, and assessed pathology, reproductive status, and the location and timing of stranding. We introduce simple, grossly apparent, standardized physical criteria to assess reproductive stage for female sea otters. We also describe ELS, examine associated risk factors, and highlight female life history strategies that likely optimize reproduction and survival. Our data suggest that females can reset both the timing and energetic demands of reproduction through fetal loss, pup abandonment, or early weaning as part of specific physiologic checkpoints during each reproductive cycle. Females appear to preload nutritionally during delayed implantation and gestation to increase fitness and reproductive success. We found that ELS was a major cause of death, affecting 56% of enrolled adult females. Peak ELS prevalence occurred in late spring, possibly reflecting the population trend toward fall/winter pupping. Increasing age and number of pregnancies were associated with a higher risk of ELS. Although the proportion of ELS females was highest in areas with dense sea otter populations, cases were recovered throughout the range, suggesting that death from ELS is associated with, but not solely caused by, population resource limitation.

California

Lesions and behavior associated with forced copulation of juvenile Pacific harbor seals (Phoca vitulina richardsi) by southern sea otters (Enhydra lutris nereis)

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.

California