Sea Otters Research Articles

Pathology, microbiology, and genetic diversity associated with Erysipelothrix rhusiopathiae and novel Erysipelothrix spp. infections in southern sea otters (Enhydra lutris nereis).

Erysipelothrix spp., including E. rhusiopathiae, are zoonotic bacterial pathogens that can cause morbidity and mortality in mammals, fish, reptiles, birds, and humans. The southern sea otter (SSO; Enhydra lutris nereis) is a federally-listed threatened species for which infectious disease is a major cause of mortality. We estimated the frequency of detection of these opportunistic pathogens in dead SSOs, described pathology associated with Erysipelothrix infections in SSOs, characterized the genetic diversity and antimicrobial susceptibility of SSO isolates, and evaluated the virulence of two novel Erysipelothrix isolates from SSOs using an in vivo fish model. From 1998 to 2021 Erysipelothrix spp. were isolated from six of >500 necropsied SSOs. Erysipelothrix spp. were isolated in pure culture from three cases, while the other three were mixed cultures. Bacterial septicemia was a primary or contributing cause of death in five of the six cases. Other pathology observed included suppurative lymphadenopathy, fibrinosuppurative arteritis with thrombosis and infarction, bilateral uveitis and endophthalmitis, hypopyon, petechia and ecchymoses, mucosal infarction, and suppurative meningoencephalitis and ventriculitis. Short to long slender Gram-positive or Gram-variable bacterial rods were identified within lesions, alone or with other opportunistic bacteria. All six SSO isolates had the spaA genotype-four isolates clustered with spaA E. rhusiopathiae strains from various terrestrial and marine animal hosts. Two isolates did not cluster with any known Erysipelothrix spp.; whole genome sequencing revealed a novel Erysipelothrix species and a novel E. rhusiopathiae subspecies. We propose the names Erysipelothrix enhydrae sp. nov. and Erysipelothrix rhusiopathiae ohloneorum ssp. nov. respectively. The type strains are E. enhydrae UCD-4322-04 and E. rhusiopathiae ohloneorum UCD-4724-06, respectively. Experimental injection of tiger barbs (Puntigrus tetrazona) resulted in infection and mortality from the two novel Erysipelothrix spp. Antimicrobial susceptibility testing of Erysipelothrix isolates from SSOs shows similar susceptibility profiles to isolates from other terrestrial and aquatic animals. This is the first description of the pathology, microbial characteristics, and genetic diversity of Erysipelothrix isolates recovered from diseased SSOs. Methods presented here can facilitate case recognition, aid characterization of Erysipelothrix isolates, and illustrate assessment of virulence using fish models.

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 geomorphology1,2, but the evidence for this remains correlational and intensely debated3,4. 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 services5,6. Monitoring of the Elkhorn Slough estuary over several decades suggested top-down control in the system, because the erosion of saltmarsh edges has generally slowed with increasing seaotter abundance, despite the consistently increasing physical stress in the system (that is, nutrient loading, sea-level rise and tidal scour7-9). 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 crabsand 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 seaotter 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.

Sea otter recovery buffers century-scale declines in California kelp forests

The status of kelp forests and their vulnerability to climate change are of global significance. As the foundation for productive and extensive ecosystems, understanding long-term kelp forest trends is critical to coastal ecosystem management, climate resiliency, and restoration programs. In this study, we curate historical US government kelp canopy inventories, develop methods to compare them with contemporary surveys, and use a machine learning framework to evaluate and rank the drivers of change for California kelp forests over the last century. Historical surveys documented Macrocystis and Nereocystis kelp forests covered approximately 120.4 km2 in 1910–1912, which is only slightly above surveys in 2014–2016 (112.0 km2). These statewide comparisons, however, mask dramatic regional changes with increases in Central California (+57.6%, +19.7 km2) and losses along the Northern (-63.0%, -8.1 km2), and Southern (-52.1%, -18.3 km2) mainland coastlines. Random Forest models rank sea otter (Enhydra lutris nereis) population density as the primary driver of kelp changes, with benthic substrate, extreme heat, and high annual variation in primary productivity also significant. This century-scale perspective identifies dramatically different outcomes for California’s kelp forests, providing a blueprint for nature-based solutions that enhance coastal resilience to climate change.

Tissue-specific carbon isotope patterns of amino acids in southern sea otters.

The measurement of stable isotope values of individual compounds, such as amino acids (AAs), has become a powerful tool in animal ecology and ecophysiology. As with any emerging technique, questions remain regarding thecapabilities and limitations of this approach, including how metabolism and tissue synthesis impact the isotopic values of individual AAs and subsequent multivariate patterns. We measured carbon isotope (δ13C) values of essential (AAESS) and nonessential (AANESS) AAs in bone collagen, whisker, muscle, and liver from ten southern sea otters (Enhydra lutris nereis) thatstranded in Monterey Bay, California. Sea otters in this population exhibit high degrees of individual dietary specialization, making this an excellent dataset to explore differences in AA δ13C values among tissues in a wild population. We found the δ13C values of the AANESS glutamic acid, proline, serine, andglycine and the AAESS threonine differed significantly among tissues, indicating possible isotopic discrimination during tissue synthesis. Threonine δ13C values were higher in liver relative to bone collagen and muscle, which may indicate catabolism of threonine for gluconeogenesis, an interpretation further supported by correlations between the δ13C values of threonine and its gluconeogenic products glycine and serine in liver. This intraindividual isotopic variation yielded different ecological interpretations among tissues; for 6/10 of the sea otter individuals analyzed, at least one tissue indicated reliance on a different primary producer source than the other tissues. Our results highlight the importance of gluconeogenesis in a carnivorous marine mammal and indicate that metabolic processes influence AAESS and AANESS δ13C values and multivariate AA δ13C patterns.

Data needs for sea otter bioenergetics modeling.

Sea otters are keystone predators whose recovery and expansion from historical exploitation throughout their range can serve to enhance local biodiversity, promote community stability, and buffer against habitat loss in nearshore marine systems. Bioenergetics models have become a useful tool in conservation and management efforts of marine mammals generally, yet no bioenergetics model exists for sea otters. Previous research provides abundant data that can be used to develop bioenergetics models for this species, yet important data gaps remain. Here we review the available data that could inform a bioenergetics model, and point to specific open questions that could be answered to more fully inform such an effort. These data gaps include quantifying energy intake through foraging by females with different aged pups in different quality habitats, the influence of body size on energy intake through foraging, and determining the level of fat storage that is possible in sea otters of different body sizes. The more completely we fill these data gaps, the more confidence we can have in the results and predictions produced by future bioenergetics modeling efforts for this species.

High prevalence and diversity of Toxoplasma gondii DNA in feral cat feces from coastal California.

Toxoplasma gondii is a zoonotic parasite that can cause severe morbidity and mortality in warm-blooded animals, including marine mammals such as sea otters. Free-ranging cats can shed environmentally resistant T. gondii oocysts in their feces, which are transported through rain-driven runoff from land to sea. Despite their large population sizes and ability to contribute to environmental oocyst contamination, there are limited studies on T. gondii oocyst shedding by free-ranging cats. We aimed to determine the frequency and genotypes of T. gondii oocysts shed by free-ranging domestic cats in central coastal California and evaluate whether genotypes present in feces are similar to those identified in sea otters that died from fatal toxoplasmosis. We utilized a longitudinal field study of four free-ranging cat colonies to assess oocyst shedding prevalence using microscopy and molecular testing with polymerase chain reaction (PCR). T. gondii DNA was confirmed with primers targeting the ITS1 locus and positive samples were genotyped at the B1 locus. While oocysts were not visualized using microscopy (0/404), we detected T. gondii DNA in 25.9% (94/362) of fecal samples. We genotyped 27 samples at the B1 locus and characterized 13 of these samples at one to three additional loci using multi locus sequence typing (MLST). Parasite DNA detection was significantly higher during the wet season (16.3%, 59/362) compared to the dry season (9.7%; 35/362), suggesting seasonal variation in T. gondii DNA presence in feces. High diversity of T. gondii strains was characterized at the B1 locus, including non-archetypal strains previously associated with sea otter mortalities. Free-ranging cats may thus play an important role in the transmission of virulent T. gondii genotypes that cause morbidity and mortality in marine wildlife. Management of free-ranging cat colonies could reduce environmental contamination with oocysts and subsequent T. gondii infection in endangered marine mammals and people.

Body size evolution in otters distinguished from terrestrial mustelids.

Some taxa of mammals live in water, all of which evolved from land-dwelling ancestors. In the family Mustelidae (Mammalia: Carnivora), most species live on land, while otters, comprising the subfamily Lutrinae, inhabit aquatic environments, which include the almost exclusively aquatic sea otters (Enhydra lutris). Thus, the transition from a terrestrial to an aquatic lifestyle has occurred within this family. Despite potentially different selection pressures on body size in aquatic and terrestrial habitats, no divergence in the evolutionary pattern of body size between otters and other mustelids has previously been shown using models of trait evolution on a phylogeny. We applied models that explicitly incorporated lineage-specific directional selection to the evolution of body mass in living mustelids. Using a simulation-based likelihood and approximate Bayesian computation approach, we demonstrated lineage-specific directional selection for larger body mass in otters, which is distinct from other mustelids. There was no evidence of a difference between sea otters and other otters in the strength of directional selection for larger body mass. Additionally, our analyses supported no difference in the rate at which body mass evolves in both directions between otters and other mustelids. These findings suggest that the evolution of body mass in otters is associated with selective advantages of larger size rather than the relaxation of constraints on body size in aquatic habitats, like other aquatic mammals such as sirenians, cetaceans, and pinnipeds.

Wolves on the Katmai coast hunt sea otters and harbor seals.

Wolves on the Katmai coast hunt sea otters and harbor seals.

A century of canopy kelp persistence and recovery in the Gulf of Alaska.

Coastal Alaska contains vast kelp habitat that supports diverse marine and human communities. Over the past century, the North Pacific Ocean has undergone oceanographic and ecological regime shifts that have the potential to influence the structure and function of kelp ecosystems strongly. However, the remoteness and complexity of the glacially carved region precludes the regular monitoring efforts that would be necessary to detect such changes. To begin to fill this critical knowledge gap, we drew upon historical and modern surveys to analyse the change in spatial coverage and species composition of canopy kelp between two time points (1913 and the early 2000s to 2010s). We also incorporated decadal surveys on sea otter range expansion following complete extirpation and reintroduction to assess the influence of sea otter recovery on the spatial extent of canopy kelp. We found increases in the spatial extent of canopy kelp throughout the Gulf of Alaska where there was coverage from both surveys. Kelp in Southcentral Alaska showed extensive recovery after the catastrophic Novarupta volcano. Kelp in Southeast Alaska showed persistence and spatial increase that closely matched increases in the range of sea otters. Observations of thermally tolerant kelp species increased more than observations of cold-adapted species between the two surveys. Contrary to trends observed at lower latitudes, the kelp forests that ring the Gulf of Alaska have been remarkably stable and even increased in the past century, despite oceanographic and ecosystem changes. To improve monitoring, we propose identification of sentinel kelp beds for regular monitoring to detect changes to these iconic and foundational canopy kelp species more readily.

Sea Otters (Enhydra lutris) Found in the Stomach of a Stranded Killer Whale (Orcinus orca) in the Commander Islands, Western North Pacific

Sea Otters (Enhydra lutris) Found in the Stomach of a Stranded Killer Whale (Orcinus orca) in the Commander Islands, Western North Pacific

Informing management of recovering predators and their prey with ecological diffusion models

The reintroduction and recovery of predators can be ecologically beneficial as well as socially and economically controversial. However, the growth and expansion of predator populations, and thus their ecological, social, and economic impacts, are not static but rather they vary in space and time. We propose a spatiotemporal statistical modeling framework based on ecological diffusion to better inform the ecology and management of recovering predators and their prey. We demonstrate its utility by applying it to a recovering sea otter (Enhydra lutris) population in Southeast Alaska, where sea otters were reintroduced in the late 1960s and have exhibited unprecedented population growth. Estimated parameters yield inferences about movement and population ecology, and our approach provides useful derived quantities, such as local abundance and carrying capacity as well as a quantity we term the equilibrium differential. We used our model to examine how density dependence and carrying capacity of sea otters vary spatially across a region. The diffusion modeling approach we present can be generalized for use in other instances of (re)colonization across taxa to inform management and conservation efforts.

The same genotype of Sarcocystis neurona responsible for mass mortality in marine mammals induced a clinical outbreak in raccoons (Procyon lotor) 10 years later

Here, we report the first known outbreak of clinical protozoal myeloencephalitis in naturally infected raccoons by the parasite Sarcocystis neurona. The North American opossum (Didelphis virginiana) and the South American opossum (Didelphis albiventris) are its known definitive hosts. Several other animal species are its intermediate or aberrant hosts. The raccoon (Procyon lotor) is considered the most important intermediate host for S. neurona in the USA. More than 50% of raccoons in the USA have sarcocysts in their muscles, however clinical sarcocystosis in raccoons is rare. In 2014, 38 free-living raccoons were found dead or moribund on the grounds of the Saint Louis Zoo, Missouri, USA. Moribund individuals were weak, lethargic, and mildly ataxic; several with oculo-nasal discharge. Seven raccoons were found dead and 31 were humanely euthanized. Postmortem examinations were conducted on nine raccoons. Neural lesions compatible with acute sarcocystosis were detected in eight raccoons. The predominant lesions were meningoencephalitis and perivascular mononuclear cells. Histologic evidence for the Canine Distemper Virus was found in one raccoon. Schizonts and merozoites were present in the encephalitic lesions of four raccoons. Mature sarcocysts were present within myocytes of five raccoons. In six raccoons, S. neurona schizonts and merozoites were confirmed by immunohistochemical staining with S. neurona-specific polyclonal antibodies. Viable S. neurona was isolated from the brains of two raccoons by bioassay in interferon gamma gene knockout mice and in cell cultures seeded directly with raccoon brain homogenate. Molecular characterization was based on raccoon no. 68. Molecular characterization based on multi-locus typing at five surface antigens (SnSAG1-5-6, SnSAG3 and SnSAG4) and the ITS-1 marker within the ssrRNA locus, using DNA isolated from bradyzoites released from sarcocysts in a naturally infected raccoon (no. 68), confirmed the presence of S. neurona antigen type I, the same genotype that caused a mass mortality event in which 40 southern sea otters stranded dead or dying within a 3 week period in April 2004 with S. neurona-associated disease. An expanded set of genotyping markers was next applied. This study reports the following new genotyping markers at 18S rRNA, 28S rRNA, COX1, ITS-1, RON1, RON2, GAPDH1, ROP20, SAG2, SnSRS21 and TUBA1 markers. The identity of Sarcocystis spp. infecting raccoons is discussed.

Ontogenetic changes in southern sea otter (Enhydra lutris nereis) fur morphology.

Many animals exhibit morphological changes across ontogeny associated with adaptations to their environment. Sea otters (Enhydra lutris) have the densest fur of any animal, which is composed of guard hairs, intermediate hairs, and underhairs. Sea otters live in cold water environments, and their fur traps a layer of air to remain properly insulated, due to morphological adaptations that allow the hairs to trap air when submerged. When a sea otter is born, it has a natal pelage which it will eventually molt and replace with a pelt resembling the adult pelage. Past studies have investigated the morphology and hair density of adult sea otter fur, but these characteristics have not been measured for other age classes, including for the natal pelage. This study quantified ontogenetic changes in hair morphology of southern sea otter (E. lutris nereis) pelts. We measured guard hair length and circularity, shape of cuticular scales on guard hairs and underhairs, and overall hair density for sea otter pelts across six age classes: neonate (<1 month), small pup (1-2 months), large pup (3-5 months), juvenile (6 months-1 year), subadult (1-3 years), and adult (4-9 years). Neonate and small pup pelts had significantly longer guard hairs than older age classes. Natal pelage guard hairs were similarly shaped but smaller in diameter than adult guard hairs. Hairs of the natal pelage had similar cuticular scale patterns as adult hairs, indicating the importance of this structure for the function of the fur. Natal pelage had a lower hair density than the pelage of older age classes, with the adult pelage exhibiting the highest hair density. Overall, the morphological differences between natal and adult pelage in sea otters suggest functional differences that may make sea otter pups more vulnerable to heat loss.

Molecular and morphological confirmation of Profilicollis altmani as the cause of acanthocephalan peritonitis in California sea otters (Enhydra lutris nereis)

Acanthocephalan peritonitis (AP; trans-intestinal migration of acanthocephalan parasites into the peritoneal cavity resulting in severe peritonitis), is a common cause of mortality in southern sea otters (Enhydra lutris nereis). Although Profilicollis spp. acanthocephalans have been implicated in these infections, the species causing AP has been an important unresolved question for decades. We used morphological and molecular techniques to characterize acanthocephalans from the gastrointestinal (GI) tract and peritoneal omentum of eighty necropsied southern sea otters. Only P. altmani was found to have perforated through the intestinal wall and migrated into the peritoneal cavity of examined sea otters, resulting in AP. Morphological and molecular criteria confirmed that Profilicollis kenti was synonymous with P. altmani. A second Profilicollis sp., likely P. botulus, was present only in the intestinal lumen, did not penetrate through the intestinal wall, and was not associated with AP.

Non-invasive detection of Orthohalarachne attenuata (Banks, 1910) and Orthohalarachne diminuata (Doetschman, 1944) (Acari: Halarachnidae) in free-ranging synanthropic South American sea lions Otaria flavescens (Shaw, 1800).

Respiratory mites of the genera Orthohalarachne and Halarachne (Acari: Halarachnidae) are causative agents of nasopharyngeal/nasopulmonary acariasis in pinnipeds and sea otters. Until now, these endoparasitic mites were mainly diagnosed via necropsies and invasive procedures. So far, non-invasive diagnostic techniques have neither been developed nor applied in free-ranging pinnipeds. In the current study, we aimed to evaluate the halarachnid mite infestation status of free-ranging "urban" South American sea lions Otaria flavescens in the city of Valdivia, Chile. Therefore, non-invasive sampling methods were used in the current study, e. g. by observation-based sampling of freshly expectorated nasal mucus in the animal environment. Further, collection devices were developed for target-oriented sampling of sneezed nasal mucus, including sterile petri dishes and stretched clingfilms mounted on telescopic rods. Applying these techniques, 26 individual sputum samples were collected. 11.5% of sputum samples proved positive for halarachnid larvae (in total, n=7), which were morphologically identified as Orthohalarachne attenuata (n=2) or Orthohalarachne diminuata (n=5). In one of the individual sea lion mucus samples, both Orthohalarachne species were detected, thereby confirming a patent co-infestation in vivo. 16S rDNA-based molecular identification of individual Orthohalarachne spp. larvae confirmed morphological findings. For the first time, we here molecularly characterized Orthohalarachne spp. on the basis of three gene regions [18S, 28S and the internal transcribed spacer 1 (ITS1)]. Overall, current data include the successful application of non-invasive techniques to sample halarachnid mites from free-ranging synanthropic pinnipeds and contribute to the current knowledge on respiratory mites infesting South American sea lions by combining morphological and molecular methods to overcome challenges in species identification. This study should further serve as baseline study and calls for more research on occurrence, biology and health implications of orthohalarachnosis in free-living as well as captive pinnipeds.

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.

“Soft Gold” Before the Gold Rush: Sea Otter Pelts in the “Competitive Expansion” of Merchant Capitalism and the Creation of a Pacific Ocean Economy

Objective/Context: In this article, we examine how Chinese demand for pelts—of sea otters and other marine mammals—fueled the eighteenth-century competitive expansion of European and later American merchants and explorers in the Pacific islands and along the Pacific coast of North America. Sea otter pelts were a commodity directly linked to the creation of a Pacific Ocean economy, and California’s colonization resulted from imperial dynamics to which this trade gave rise. Methodology: Using both primary and secondary sources, we examine how these merchants’ quest to supply the lucrative Chinese luxury market with furs—“soft gold”—brought them into contact with indigenous peoples whose livelihoods and commercial networks would also be recruited into this global market but on quite unequal terms and with devastating consequences. Originality: The growth of this Pacific trade throughout the eighteenth century fueled geopolitical rivalries that led to the colonization of California with a system of missions and military garrisons (presidios) and, eventually, to a new ecology as a result of plants and animals brought from New Spain before the Gold Rush and the Treaty of Guadalupe Hidalgo. Many, including Marx and Engels, have claimed that the Gold Rush contributed significantly to the dynamics of industrial capitalism; nevertheless, we argue that its conditions of possibility were laid out centuries earlier via the complex geopolitical and ecological connections through which this Pacific economy was articulated. Conclusions: During the eighteenth century, the Pacific’s products were extracted and commodified in circuits whose demand center was China. It is precisely the prior existence of these global markets—centered on silver and “soft gold,” or sea otter furs—that explains the presence of Europeans and Americans in California eager to prospect for the yellow metal in 1848.

Analytical Validation and Assessment of Baseline Fecal Glucocorticoid Metabolites in Northern Sea Otters (Enhydra lutris kenyoni) in Human Care

Fecal glucocorticoid metabolites (FGMs) have been used as a non-invasive and indirect measurement of the complex stress response in a variety of species. Animals in facilities under managed care allow for the longitudinal study of FGMs in a controlled environment. Animal histories often include environmental, husbandry, and medical notes that can be matched to FGM concentrations to aid in the physiological validation of adrenal stimulation and response. The goal of this study was to demonstrate analytical validations using two enzyme-linked immunosorbent assays (EIA) to measure FGMs from northern sea otters (Enhydra lutris kenyoni) under human care (Seattle Aquarium, Seattle, WA, USA) and to determine baseline and stress response spike levels for individual sea otters. Individual variation was found among the four subjects in the study with fecal baseline levels ranging from 20.2 to 83.7 ng/g for cortisol-immunoreactive metabolites and 52.3 to 102 ng/g for corticosterone-immunoreactive metabolites. As a retrospective study, 39 percent of hormone peaks were associated with notes and most FGM spikes were associated with veterinary procedures or days in which enrichment items were provided and produced an excitatory response. Monitoring baseline FGMs levels and events associated with hormone peak values may provide insight into effective husbandry management to improve the overall welfare of sea otters and other marine mammals.

The role of spatial structure in at-risk metapopulation recoveries.

Metapopulations are often managed as a single contiguous population despite the spatial structure underlying their local and regional dynamics. Disturbances from human activities can also be spatially structured with mortality impacts concentrated to just a few local populations among the aggregate. Scale transitions between local and regional processes can generate emergent properties whereby the whole system can fail to recover as quickly as expected for an equivalent single population. Here, we draw on theory and empirical case studies to ask: what is the consequence of spatially structured ecological and disturbance processes on metapopulation recoveries? We suggest that exploring this question could help address knowledge gaps for managing metapopulations including: Why do some metapopulations recover quickly while others remain collapsed? And, what risks are unaccounted for when metapopulations are managed at aggregate scales? First, we used model simulations to examine how scale transitions among ecological and disturbance conditions interact to generate emergent metapopulation recovery outcomes. In general, we found that the spatial structure of disturbance was a strong determinant of recovery outcomes. Specifically, disturbances that unevenly impacted local populations consistently generated the slowest recoveries and highest conservation risks. Ecological conditions that dampened metapopulation recoveries included low dispersal, variable local demography, sparsely connected habitat networks, and spatially and temporally correlated stochastic processes. Second, we illustrate the unexpected challenges of managing metapopulations by examining the recoveries of three USA federally listed endangered species: Florida Everglade snail kites, California and Alaska sea otters, and Snake River Chinook salmon. Overall, our results show the pivotal role of spatial structure in metapopulation recoveries whereby the interplay between local and regional processes shapes the resilience of the whole system. With this understanding, we provide guidelines for resource managers tasked with conserving and managing metapopulations and identify opportunities for research to support the application of metapopulation theory to real-world challenges.

The Economic Value of Sea Otters and Recreational Tourism in a California Estuary

The recovery of marine megafauna can lead to improved ecosystem function and services, but not all stakeholders may benefit equally. Quantifying the local economic value of a species’ presence may appeal to broader range of stakeholders when developing conservation strategies. This study aims to examine the economic effect recreational activities can have on a local region, and to determine what role the presence of southern sea otters (Enhydra lutris nereis) had on the value visitors placed on the visit and the area’s preservation. We surveyed visitors to Elkhorn Slough, a small estuary in California known for diverse wildlife, modeled direct and indirect economic contributions to local economics, and evaluated perceptions and value placed on the area and wildlife. Annually, visitors contributed approximately $3.2 million (USD) in direct spending, with an additional $1.85 million in indirect economic gains, which could support over 300 full-time, part-time, and seasonal jobs to the region. Whether sea otters were observed during a trip influenced how visitors ranked their importance, and the perceived value of the estuary and sea otters. Combined, this study quantified what recreational visitors could contribute to local economies and that sea otters play a role in what visitor’s value about their visit. These results provide additional support to the benefits of species presence at a local scale. We discuss how these types of studies can be used as part of larger species and ecosystem management plans, particularly considering species recovery and range expansion.