The effect of a startle-eliciting device on the foraging success of individual harbor seals (Phoca vitulina).

Pinniped predation on commercially and ecologically important prey has been a source of conflict for centuries. In the Salish Sea, harbor seals (Phoca vitulina) are suspected of impeding the recovery of culturally and ecologically critical Pacific salmon (Oncorhynchus spp.). In Fall 2020, a novel deterrent called Targeted Acoustic Startle Technology (TAST) was deployed at Whatcom Creek to deter harbor seals from preying on fall runs of hatchery chum (O. keta) and Chinook (O. tshawytscha) salmon in Bellingham, Washington, USA. Field observations were conducted in 2020 to compare the presence and foraging success of individual harbor seals across sound exposure (TAST-on) and control (TAST-off) conditions. Observations conducted the previous (2019) and following (2021) years were used to compare the effects observed in 2020 to two control years. Using photo-identification, individual seals were associated with foraging successes across all 3 years of the study. Generalized linear mixed models showed a significant 45.6% reduction in the duration (min) individuals remained at the creek with TAST on, and a significant 43.8% reduction in the overall foraging success of individuals. However, the observed effect of TAST varied across individual seals. Seals that were observed regularly within one season were more likely to return the year after, regardless of TAST treatment. Generalized linear models showed interannual variation in the number of seals present and salmon consumed. However, the effect of TAST in 2020 was greater than the observed variation across years. Our analyses suggest TAST can be an effective tool for managing pinniped predation, although alternate strategies such as deploying TAST longer-term and using multi-unit setups to increase coverage could help strengthen its effects. Future studies should further examine the individual variability found in this study.

Individual variability in foraging success of a marine predator informs predator management.

The complexities of trophic dynamics complicate the management of predator populations. Targeted culling campaigns are one management strategy meant to control predation for the benefit of the prey population. In these campaigns, individual predators are often considered "rogue" based on visitation rates to the site of concern. This definition assumes that all predators impact prey equally. However, individual variability in foraging success may compromise this assumption. To examine this hypothesis, we studied harbor seals preying on adult salmonids during the 2014-2019 fall runs in Whatcom Creek, Bellingham, Washington, USA, and recorded visitation rate and foraging success of individual seals from photographs and field observations. We then used Generalized Linear Mixed-Effects Models to model individual foraging success. Models including harbor seal identity better explained foraging success than models based on visitation rate alone. We concluded that considering intraspecific variability and classifying "rogue individuals" based on foraging success is a more accurate protocol for managing predator populations than relying solely on visitation rate of the predators.

Data on the diets of Salish Sea harbour seals from DNA metabarcoding.

Marine trophic ecology data are in high demand as natural resource agencies increasingly adopt ecosystem-based management strategies that account for complex species interactions. Harbour seal (Phoca vitulina) diet data are of particular interest because the species is an abundant predator in the northeast Pacific Ocean and Salish Sea ecosystem that consumes Pacific salmon (Oncorhynchus spp.). A multi-agency effort was therefore undertaken to produce harbour seal diet data on an ecosystem scale using, 1) a standardized set of scat collection and analysis methods, and 2) a newly developed DNA metabarcoding diet analysis technique designed to identify prey species and quantify their relative proportions in seal diets. The DNA-based dataset described herein contains records from 4,625 harbour seal scats representing 52 haulout sites, 7 years, 12 calendar months, and a total of 11,641 prey identifications. Prey morphological hard parts analyses were conducted alongside, resulting in corresponding hard parts data for 92% of the scat DNA samples. A custom-built prey DNA sequence database containing 201 species (192 fishes, 9 cephalopods) is also provided.

Estimating the abundance of the critically endangered Baltic Proper harbour porpoise (Phocoena phocoena) population using passive acoustic monitoring.

Knowing the abundance of a population is a crucial component to assess its conservation status and develop effective conservation plans. For most cetaceans, abundance estimation is difficult given their cryptic and mobile nature, especially when the population is small and has a transnational distribution. In the Baltic Sea, the number of harbour porpoises (Phocoena phocoena) has collapsed since the mid-20th century and the Baltic Proper harbour porpoise is listed as Critically Endangered by the IUCN and HELCOM; however, its abundance remains unknown. Here, one of the largest ever passive acoustic monitoring studies was carried out by eight Baltic Sea nations to estimate the abundance of the Baltic Proper harbour porpoise for the first time. By logging porpoise echolocation signals at 298 stations during May 2011-April 2013, calibrating the loggers' spatial detection performance at sea, and measuring the click rate of tagged individuals, we estimated an abundance of 71-1105 individuals (95% CI, point estimate 491) during May-October within the population's proposed management border. The small abundance estimate strongly supports that the Baltic Proper harbour porpoise is facing an extremely high risk of extinction, and highlights the need for immediate and efficient conservation actions through international cooperation. It also provides a starting point in monitoring the trend of the population abundance to evaluate the effectiveness of management measures and determine its interactions with the larger neighboring Belt Sea population. Further, we offer evidence that design-based passive acoustic monitoring can generate reliable estimates of the abundance of rare and cryptic animal populations across large spatial scales.

Large-scale molecular barcoding of prey DNA reveals predictors of intrapopulation feeding diversity in a marine predator.

Predator-prey interactions are critical in understanding how communities function. However, we need to describe intraspecific variation in diet to accurately depict those interactions. Harbor seals (Phoca vitulina) are an abundant marine predator that prey on species of conservation concern. We estimated intrapopulation feeding diversity (variation in feeding habits between individuals of the same species) of harbor seals in the Salish Sea. Estimates of feeding diversity were examined relative to sex, month, and location using a novel approach that combined molecular techniques, repeated cross-sectional sampling of scat, and a specialization metric (within-individual consistency in diet measured by the Proportional Similarity Index ( P S i )). Based on 1,083 scat samples collected from five haul-out sites during four nonsequential years, we quantified diet using metabarcoding techniques and determined the sex of the scat depositor using a molecular assay. Results suggest that intrapopulation feeding diversity was present. Specialization was high over short periods (24-48 hr, P S i  = 0.392, 95% CI = 0.013, R = 100,000) and variable in time and space. Females showed more specialization than males, particularly during summer and fall. Additionally, demersal and benthic prey species were correlated with more specialized diets. The latter finding suggests that this type of prey likely requires specific foraging strategies and that there are trade-offs between pelagic and benthic foraging styles for harbor seals. This differential feeding on prey species, as well as between sexes of harbor seals, indicates that predator-prey interactions in harbor seals are complex and that each sex may have a different impact on species of conservation concern. As such, describing intrapopulation feeding diversity may unravel hitherto unknown complex predator-prey interactions in the community.

Large-scale molecular diet analysis in a generalist marine mammal reveals male preference for prey of conservation concern.

Sex-specific diet information is important in the determination of predator impacts on prey populations. Unfortunately, the diet of males and females can be difficult to describe, particularly when they are marine predators. We combined two molecular techniques to describe haul-out use and prey preferences of male and female harbor seals (Phoca vitulina) from Comox and Cowichan Bay (Canada) during 2012-2013. DNA metabarcoding quantified the diet proportions comprised of prey species in harbor seal scat, and qPCR determined the sex of the individual that deposited each scat. Using 287 female and 260 male samples, we compared the monthly sex ratio with GLMs and analyzed prey consumption relative to sex, season, site, and year with PERMANOVA. The sex ratio between monthly samples differed widely in both years (range = 12%-79% males) and showed different patterns at each haul-out site. Male and female diet differed across both years and sites: Females consumed a high proportion of demersal fish species while males consumed more salmonid species. Diet composition was related to both sex and season (PERMANOVA: R 2 = 27%, p < 0.001; R 2 = 24%, p < 0.001, respectively) and their interaction (PERMANOVA: R 2 = 11%, p < 0.001). Diet differences between males and females were consistent across site and year, suggesting fundamental foraging differences, including that males may have a larger impact on salmonids than females. Our novel combination of techniques allowed for both prey taxonomic and spatiotemporal resolution unprecedented in marine predators.

Competing tradeoffs between increasing marine mammal predation and fisheries harvest of Chinook salmon.

Many marine mammal predators, particularly pinnipeds, have increased in abundance in recent decades, generating new challenges for balancing human uses with recovery goals via ecosystem-based management. We used a spatio-temporal bioenergetics model of the Northeast Pacific Ocean to quantify how predation by three species of pinnipeds and killer whales (Orcinus orca) on Chinook salmon (Oncorhynchus tshawytscha) has changed since the 1970s along the west coast of North America, and compare these estimates to salmon fisheries. We find that from 1975 to 2015, biomass of Chinook salmon consumed by pinnipeds and killer whales increased from 6,100 to 15,200 metric tons (from 5 to 31.5 million individual salmon). Though there is variation across the regions in our model, overall, killer whales consume the largest biomass of Chinook salmon, but harbor seals (Phoca vitulina) consume the largest number of individuals. The decrease in adult Chinook salmon harvest from 1975-2015 was 16,400 to 9,600 metric tons. Thus, Chinook salmon removals (harvest + consumption) increased in the past 40 years despite catch reductions by fisheries, due to consumption by recovering pinnipeds and endangered killer whales. Long-term management strategies for Chinook salmon will need to consider potential conflicts between rebounding predators or endangered predators and prey.

wisepair: a computer program for individual matching in genetic tracking studies.

Individual-based data sets tracking organisms over space and time are fundamental to answering broad questions in ecology and evolution. A 'permanent' genetic tag circumvents a need to invasively mark or tag animals, especially if there are little phenotypic differences among individuals. However, genetic tracking of individuals does not come without its limits; correctly matching genotypes and error rates associated with laboratory work can make it difficult to parse out matched individuals. In addition, defining a sampling design that effectively matches individuals in the wild can be a challenge for researchers. Here, we combine the two objectives of defining sampling design and reducing genotyping error through an efficient Python-based computer-modelling program, wisepair. We describe the methods used to develop the computer program and assess its effectiveness through three empirical data sets, with and without reference genotypes. Our results show that wisepair outperformed similar genotype matching programs using previously published from reference genotype data of diurnal poison frogs (Allobates femoralis) and without-reference (faecal) genotype sample data sets of harbour seals (Phoca vitulina) and Eurasian otters (Lutra lutra). In addition, due to limited sampling effort in the harbour seal data, we present optimal sampling designs for future projects. wisepair allows for minimal sacrifice in the available methods as it incorporates sample rerun error data, allelic pairwise comparisons and probabilistic simulations to determine matching thresholds. Our program is the lone tool available to researchers to define parameters a priori for genetic tracking studies.

The social structure of Golfo Dulce bottlenose dolphins (Tursiops truncatus) and the influence of behavioural state.

Ecological factors such as habitat and food availability affect the social structure of bottlenose dolphins (Tursiops spp.). Here, we describe the social structure of bottlenose dolphins (T. truncatus) in Golfo Dulce, Costa Rica, a semi-enclosed, fjord-like tropical embayment resembling a pelagic system. We also examine behaviour-linked social strategies by comparing social structure relative to behavioural state: feeding versus non-feeding. We analysed 333 sightings over 210 days from boat-based surveys. Despite the uniqueness of the area, the 47 analysed adults had a social structure similar to other populations: a well-differentiated fission-fusion society with sex-specific patterns of associations and aggression. These results indicate that differences in social structure relative to other populations were a matter of degree. Association strength of dyads was highly correlated across behavioural states, indicating constraints on social fluidity. Males displayed a marked difference in lagged association rate and females displayed a small difference in association homogeneity between states. We suggest this difference in population-wide social connections between behavioural states, particularly for males, was due to mating strategies, a pressure which is strongest during non-feeding behaviour and relaxed during feeding. This finding highlights the importance of considering behavioural state when examining individual bonds and the behavioural plasticity for which the bottlenose dolphin is well known.

Quantitative classification of harbor seal breeding calls in Georgia Strait, Canada.

During breeding season, male harbor seals (Phoca vitulina) produce underwater calls used in sexual competition and advertisement. Call characteristics vary among populations, and within-population differences are thought to represent individual variation. However, vocalizations have not been described for several populations of this widely-distributed and genetically diverse species. This study describes the vocal repertoire of harbor seals from British Columbia, Canada. Underwater recordings were made near Hornby Island during the summer of 2014 using a single hydrophone. A wide variability was detected in breeding vocalizations within this single breeding site. Four candidate call types were identified, containing six subtypes. Linear discriminant analysis showed 88% agreement with subjective classification of call types, and 74% agreement for call subtypes. Classification tree analysis gave a 92% agreement with candidate call types, with all splits made on the basis of call duration. Differences in duration may have reflected individual differences among seals. This study suggests that the vocal repertoire of harbor seals in this area comprises a vocal continuum rather than discrete call types. Further work with the ability to localize calls may help to determine whether this complexity represents variability due to propagation conditions, animal orientation, or differences among individual seals.

Fine-scale variability in harbor seal foraging behavior.

Understanding the variability of foraging behavior within a population of predators is important for determining their role in the ecosystem and how they may respond to future ecosystem changes. However, such variability has seldom been studied in harbor seals on a fine spatial scale (<30 km). We used a combination of standard and Bayesian generalized linear mixed models to explore how environmental variables influenced the dive behavior of harbor seals. Time-depth recorders were deployed on harbor seals from two haul-out sites in the Salish Sea in 2007 (n = 18) and 2008 (n = 11). Three behavioral bout types were classified from six dive types within each bout; however, one of these bout types was related to haul-out activity and was excluded from analyses. Deep foraging bouts (Type I) were the predominant type used throughout the study; however, variation in the use of bout types was observed relative to haul-out site, season, sex, and light (day/night). The proportional use of Type I and Type II (shallow foraging/traveling) bouts differed dramatically between haul-out sites, seasons, sexes, and whether it was day or night; individual variability between seals also contributed to the observed differences. We hypothesize that this variation in dive behavior was related to habitat or prey specialization by seals from different haul-out sites, or individual variability between seals in the study area. The results highlight the potential influence of habitat and specialization on the foraging behavior of harbor seals, and may help explain the variability in diet that is observed between different haul-out site groups in this population.

Lacaziosis-like disease among bottlenose dolphins Tursiops truncatus photographed in Golfo Dulce, Costa Rica.

Lacaziosis (also known as lobomycosis) is a chronic dermal disease caused by the fungal agent Lacazia loboi, which affects both humans and dolphins. Photographic data have been used to identify lacaziosis-like disease (LLD) among dolphins in the waters of North and South America, and here we report LLD in bottlenose dolphins Tursiops truncatus off the coast of Costa Rica, the first reporting in Central American waters. During the periods of 1991 to 1992 and 2010 to 2011, 3 research teams conducted separate dolphin surveys in the Pacific tropical fiord Golfo Dulce, and each documented skin lesions in the resident population of bottlenose dolphins. Photo-ID records were used to identify LLD-affected bottlenose dolphins and to assess their lesions. Findings showed between 13.2 and 16.1% of the identified dolphins exhibited lesions grossly resembling lacaziosis. By combining efforts and cross-referencing photographic data, the teams explored the presence of LLD in Golfo Dulce over a time gap of approximately 20 yr. Our findings expand the geographical range of the disease and offer insight into its longevity within a given population of dolphins.

Long distance movements and disjunct spatial use of harbor seals (Phoca vitulina) in the inland waters of the Pacific Northwest.

BACKGROUND: Worldwide, adult harbor seals (Phoca vitulina) typically limit their movements and activity to <50 km from their primary haul-out site. As a result, the ecological impact of harbor seals is viewed as limited to relatively small spatial scales. Harbor seals in the Pacific Northwest are believed to remain <30 km from their primary haul-out site, one of several contributing factors to the current stock designation. However, movement patterns within the region are not well understood because previous studies have used radio-telemetry, which has range limitations. Our objective was to use satellite-telemetry to determine the regional spatial scale of movements. METHODOLOGY/PRINCIPAL FINDINGS: Satellite tags were deployed on 20 adult seals (n=16 males and 4 females) from two rocky reefs and a mudflat-bay during April-May 2007. Standard filtering algorithms were used to remove outliers, resulting in an average (± SD) of 693 (± 377) locations per seal over 110 (± 32) days. A particle filter was implemented to interpolate locations temporally and decrease erroneous locations on land. Minimum over-water distances were calculated between filtered locations and each seal's capture site to show movement of seals over time relative to their capture site, and we estimated utilization distributions from kernel density analysis to reflect spatial use. Eight males moved >100 km from their capture site at least once, two of which traveled round trip to and from the Pacific coast, a total distance >400 km. Disjunct spatial use patterns observed provide new insight into general harbor seal behavior. CONCLUSIONS/SIGNIFICANCE: Long-distance movements and disjunct spatial use of adult harbor seals have not been reported for the study region and are rare worldwide in such a large proportion of tagged individuals. Thus, the ecological influence of individual seals may reach farther than previously assumed.

Effects of the presence of official-looking volunteers on harassment of New Zealand fur seals.

An increased number of tourists viewing animals in the wild have increased stress on these animals (hereafter wildlife). Many wildlife-viewing locations rely on voluntary compliance with posted regulations to protect animals from tourists because of the expense of employing on-site enforcement personnel. Voluntary compliance, however, is ineffective. The presence of official-looking volunteers may decrease the incidence of wildlife harassment by tourists. To test this possibility, we observed tourists interacting with 5- to 12-month-old New Zealand fur seals (Arctocephalus forsteri) at the popular Ohau Stream waterfall while in the absence or presence of a young woman in plain sight wearing a neon vest (i.e., observer) and when an observer was not present. We observed 254 tourist groups at the waterfall when young seals were present. The percentage of groups in which at least one person harassed (approached, touched, or threw objects) a young seal was two-thirds lower when the official-looking observer was present. Frequency of harassment was inversely related to observer presence. Programs in which volunteers work at tourist sites are popular in countries with high tourism rates, such as New Zealand. Our results show that a relatively inexpensive and effective tourism-management strategy may be to post such volunteers as observers at sites where tourists view wildlife.