Correction: Fin whales of the Great Bear Rainforest: Balaenoptera physalus velifera in a Canadian Pacific fjord system.

[This corrects the article DOI: 10.1371/journal.pone.0256815.].

Acoustic and visual cetacean surveys reveal year-round spatial and temporal distributions for multiple species in northern British Columbia, Canada.

Cetaceans spend most of their time below the surface of the sea, highlighting the importance of passive acoustic monitoring as a tool to facilitate understanding and mapping their year-round spatial and temporal distributions. To increase our limited knowledge of cetacean acoustic detection patterns for the east and west coasts of Gwaii Haanas, a remote protected area on Haida Gwaii, BC, Canada, acoustic datasets recorded off SG̱ang Gwaay (Sep 2009-May 2011), Gowgaia Slope (Jul 2017-Jul 2019), and Ramsay Island (Aug 2018-Aug 2019) were analyzed. Comparing overlapping periods of visual surveys and acoustic monitoring confirmed presence of 12 cetacean species/species groups within the study region. Seasonal patterns were identified for blue, fin, humpback, grey and sperm whale acoustic signals. Killer whale and delphinid acoustic signals occurred year-round on both coasts of Haida Gwaii and showed strong diel variation. Cuvier's, Baird's, beaked whale and porpoise clicks, were identified in high-frequency recordings on the west coast. Correlations between environmental factors, chlorophyll-a and sea surface temperature, and cetacean acoustic occurrence off Gwaii Haanas were also examined. This study is the first to acoustically monitor Gwaii Haanas waters for an extended continuous period and therefore serves as a baseline from which to monitor future changes.

Physical disturbance by recovering sea otter populations increases eelgrass genetic diversity.

Most knowledge regarding the role of predators is ecological in nature. Here, we report how disturbance generated by sea otters (Enhydra lutris) digging for infaunal prey in eelgrass (Zostera marina) meadows increases genetic diversity by promoting conditions for sexual reproduction of plants. Eelgrass allelic richness and genotypic diversity were, respectively, 30 and 6% higher in areas where recovering sea otter populations had been established for 20 to 30 years than in areas where they had been present <10 years or absent >100 years. The influence of sea otter occupancy on the aforementioned measures of genetic diversity was stronger than those of depth, temperature, latitude, or meadow size. Our findings reveal an underappreciated evolutionary process by which megafauna may promote genetic diversity and ecological resilience.

Fin whales of the Great Bear Rainforest: Balaenoptera physalus velifera in a Canadian Pacific fjord system.

Fin whales (Balaenoptera physalus) are widely considered an offshore and oceanic species, but certain populations also use coastal areas and semi-enclosed seas. Based upon fifteen years of study, we report that Canadian Pacific fin whales (B. p. velifera) have returned to the Kitimat Fjord System (KFS) in the Great Bear Rainforest, and have established a seasonally resident population in its intracoastal waters. This is the only fjord system along this coast or elsewhere in which fin whales are known to occur regularly with strong site fidelity. The KFS was also the only Canadian Pacific fjord system in which fin whales were commonly found and killed during commercial whaling, pointing to its long-term importance. Traditional knowledge, whaling records, and citizen science databases suggest that fin whales were extirpated from this area prior to their return in 2005-2006. Visual surveys and mark-recapture analysis documented their repopulation of the area, with 100-120 whales using the fjord system in recent years, as well as the establishment of a seasonally resident population with annual return rates higher than 70%. Line transect surveys identified the central and outer channels of the KFS as the primary fin whale habitat, with the greatest densities occurring in Squally Channel and Caamaño Sound. Fin whales were observed in the KFS in most months of the year. Vessel- and shore-based surveys (27,311 km and 6,572 hours of effort, respectively) indicated regular fin whale presence (2,542 detections), including mother-calf pairs, from June to October and peak abundance in late August-early September. Seasonal patterns were variable year-to-year, and several lines of evidence indicated that fin whales arrived and departed from the KFS repeatedly throughout the summer and fall. Additionally, we report on the population's social network and morphometrics. These findings offer insights into the dynamics of population recovery in an area where several marine shipping projects are proposed. The fin whales of the Great Bear Rainforest represent a rare exception to general patterns in this species' natural history, and we highlight the importance of their conservation.

Cascading social-ecological costs and benefits triggered by a recovering keystone predator.

Predator recovery often leads to ecosystem change that can trigger conflicts with more recently established human activities. In the eastern North Pacific, recovering sea otters are transforming coastal systems by reducing populations of benthic invertebrates and releasing kelp forests from grazing pressure. These changes threaten established shellfish fisheries and modify a variety of other ecosystem services. The diverse social and economic consequences of this trophic cascade are unknown, particularly across large regions. We developed and applied a trophic model to predict these impacts on four ecosystem services. Results suggest that sea otter presence yields 37% more total ecosystem biomass annually, increasing the value of finfish [+9.4 million Canadian dollars (CA$)], carbon sequestration (+2.2 million CA$), and ecotourism (+42.0 million CA$). To the extent that these benefits are realized, they will exceed the annual loss to invertebrate fisheries (-$7.3 million CA$). Recovery of keystone predators thus not only restores ecosystems but can also affect a range of social, economic, and ecological benefits for associated communities.

Spatial epidemiological patterns suggest mechanisms of land-sea transmission for Sarcocystis neurona in a coastal marine mammal.

Sarcocystis neurona was recognised as an important cause of mortality in southern sea otters (Enhydra lutris nereis) after an outbreak in April 2004 and has since been detected in many marine mammal species in the Northeast Pacific Ocean. Risk of S. neurona exposure in sea otters is associated with consumption of clams and soft-sediment prey and is temporally associated with runoff events. We examined the spatial distribution of S. neurona exposure risk based on serum antibody testing and assessed risk factors for exposure in animals from California, Washington, British Columbia and Alaska. Significant spatial clustering of seropositive animals was observed in California and Washington, compared with British Columbia and Alaska. Adult males were at greatest risk for exposure to S. neurona, and there were strong associations with terrestrial features (wetlands, cropland, high human housing-unit density). In California, habitats containing soft sediment exhibited greater risk than hard substrate or kelp beds. Consuming a diet rich in clams was also associated with increased exposure risk. These findings suggest a transmission pathway analogous to that described for Toxoplasma gondii, with infectious stages traveling in freshwater runoff and being concentrated in particular locations by marine habitat features, ocean physical processes, and invertebrate bioconcentration.

Sex and occupation time influence niche space of a recovering keystone predator.

Predators exert strong effects on ecological communities, particularly when they re-occupy areas after decades of extirpation. Within species, such effects can vary over time and by sex and cascade across trophic levels. We used a space-for-time substitution to make foraging observations of sea otters (Enhydra lutris) across a gradient of reoccupation time (1-30 years), and nonmetric multidimensional scaling (nMDS) analysis to ask whether (a) sea otter niche space varies as a function of occupation time and (b) whether niche space varies by sex. We found that niche space varied among areas of different occupation times. Dietary niches at short occupation times were dominated by urchins (Mesocentrotus and Strongylocentrotus spp; >60% of diets) in open habitats at 10-40 m depths. At longer occupation times, niches were dominated by small clams (Veneroida; >30% diet), mussels (Mytilus spp; >20% diet), and crab (Decapoda; >10% diet) in shallow (<10 m) kelp habitats. Diet diversity was lowest (H' = 1.46) but energy rich (~37 kcal/min) at the earliest occupied area and highest, but energy poor (H' = 2.63, ~9 kcal/min) at the longest occupied area. A similar transition occurred through time at a recently occupied area. We found that niche space also differed between sexes, with bachelor males consuming large clams (>60%), and urchins (~25%) from deep waters (>40 m), and females and territorial males consuming smaller, varied prey from shallow waters (<10 m). Bachelor male diets were less diverse (H' = 2.21) but more energy rich (~27 kcal/min) than territorial males (H' = 2.54, ~13 kcal/min) and females (H' = 2.74, ~11 kcal/min). Given recovering predators require adequate food and space, and the ecological interactions they elicit, we emphasize the importance of investigating niche space over the duration of recovery and considering sex-based differences in these interactions.

Defining the risk landscape in the context of pathogen pollution: Toxoplasma gondii in sea otters along the Pacific Rim.

Pathogens entering the marine environment as pollutants exhibit a spatial signature driven by their transport mechanisms. The sea otter (Enhydra lutris), a marine animal which lives much of its life within sight of land, presents a unique opportunity to understand land-sea pathogen transmission. Using a dataset on Toxoplasma gondii prevalence across sea otter range from Alaska to California, we found that the dominant drivers of infection risk vary depending upon the spatial scale of analysis. At the population level, regions with high T. gondii prevalence had higher human population density and a greater proportion of human-dominated land uses, suggesting a strong role for population density of the felid definitive host of this parasite. This relationship persisted when a subset of data were analysed at the individual level: large-scale patterns in sea otter T. gondii infection prevalence were largely explained by individual exposure to areas of high human housing unit density, and other landscape features associated with anthropogenic land use, such as impervious surfaces and cropping land. These results contrast with the small-scale, within-region analysis, in which age, sex and prey choice accounted for most of the variation in infection risk, and terrestrial environmental features provided little variation to help in explaining observed patterns. These results underscore the importance of spatial scale in study design when quantifying both individual-level risk factors and landscape-scale variation in infection risk.

The interaction of intraspecific competition and habitat on individual diet specialization: a near range-wide examination of sea otters.

The quantification of individuality is a common research theme in the fields of population, community, and evolutionary ecology. The potential for individuality to arise is likely context-dependent, and the influence of habitat characteristics on its prevalence has received less attention than intraspecific competition. We examined individual diet specialization in 16 sea otter (Enhydra lutris) populations from southern California to the Aleutian Islands in Alaska. Because population histories, relative densities, and habitat characteristics vary widely among sites, we could examine the effects of intraspecific competition and habitat on the prevalence of individual diet specialization. Using observed diet data, we classified half of our sites as rocky substrate habitats and the other half containing a mixture of rocky and unconsolidated (soft) sediment substrates. We used stable isotope data to quantify population- and individual-level diet variation. Among rocky substrate sites, the slope [±standard error (SE)] of the positive significant relationship between the within-individual component (WIC) and total isotopic niche width (TINW) was shallow (0.23 ± 0.07) and negatively correlated with sea otter density. In contrast, the slope of the positive WIC/TINW relationship for populations inhabiting mixed substrate habitats was much higher (0.53 ± 0.14), suggesting a low degree of individuality, irrespective of intraspecific competition. Our results show that the potential for individuality to occur as a result of increasing intraspecific competition is context-dependent and that habitat characteristics, which ultimately influence prey diversity, relative abundance, and the range of skillsets required for efficient prey procurement, are important in determining when and where individual diet specialization occurs in nature.

Hydrocarbon concentrations and patterns in free-ranging sea otters (Enhydra lutris) from British Columbia, Canada.

With oil pollution recognized as a major threat to British Columbia's recovering sea otter (Enhydra lutris) population, it is important to distinguish acute from chronic exposures to oil constituent groups in this marine mammal. Concentrations and patterns of alkanes and polycyclic aromatic hydrocarbons (PAHs) were determined in blood samples from 29 live-captured sea otters in two coastal areas of British Columbia, as well as in representative samples of their invertebrate prey. Hydrocarbon concentrations in sea otters were similar between areas and among age and sex classes, suggesting that metabolism dominates the fate of these compounds in sea otters. Biomagnification factors derived from PAH ratios in otter:prey supported this notion. Although some higher alkylated three- and four-ring PAHs appeared to biomagnify, the majority of PAHs did not. The apparent retention of alkyl PAHs was reflected in the composition of estimated sea otter body burdens, which provided an alternative way of evaluating hydrocarbon exposure. Alkyl PAHs made up 86 ± 9% of estimated body burdens (4,340 ± 2,950 µg), with no differences between males and females (p = 0.18). The importance of measuring both parent and alkyl PAHs is underscored by their divergent dynamics in sea otters, with ready depuration of parent PAHs (metabolized or excreted) by sea otters on the one hand and biomagnification of alkyl PAHs on the other.