Examination of the foraging habits of Pacific harbor seal (Phoca vitulina richardsi) to describe their use of the Umpqua River, Oregon, and their predation on salmonids

The increase in harbor seal (Phoca vitulina richardsi) abundance, concurrent with the decrease in salmonid (Oncorhynchus spp.) and other fish stocks, raises concerns about the potential negative impact of seals on fish populations. Although harbor seals are found in rivers and estuaries, their presence is not necessarily indicative of exclusive or predominant feeding in these systems. We examined the diet of harbor seals in the Umpqua River, Oregon, during 1997 and 1998 to indirectly assess whether or not they were feeding in the river. Fish otoliths and other skeletal structures were recovered from 651 scats and used to identify seal prey. The use of all diagnostic prey structures, rather than just otoliths, increased our estimates of the number of taxa, the minimum number of individuals and percent frequency of occurrence (%FO) of prey consumed. The %FO indicated that the most common prey were pleuronectids, Pacific hake (Merluccius productus), Pacific stag-horn sculpin (Leptocottus armatus), osmerids, and shiner surfperch (Cymatogaster aggregata). The majority (76%) of prey were fish that inhabit marine waters exclusively and fish found in marine and estuarine areas (e.g. anadromous spp.) which would indicate that seals forage predominantly at sea and use the estuary for resting and opportunistic feeding. Salmonid remains were encountered in 39 samples (6%); two samples contained identifiable otoliths, which were determined to be from chi-nook salmon (O. tshawytscha). Because of the complex salmonid composition in the Umpqua River, we used molecular genetic techniques on salmonid bones retrieved from scat to discern species that were rare from those that were abundant. Of the 37 scats with salmonid bones but no otoliths, bones were identified genetically as chinook or coho (O. kisutch) salmon, or steelhead trout (O. mykiss) in 90% of the samples.

Molecular methods for the genetic identification of salmonid prey from Pacific harbor seal (Phoca vitulina richardsi) scat

Twenty-six stocks of Pacific salmon and trout (Oncorhynchus spp.), representing evolutionary significant units (ESU), are listed as threatened or endangered under the Endangered Species Act (ESA) and six more stocks are currently being evaluated for listing. The ecological and economic consequences of these listings are large; therefore considerable effort has been made to understand and respond to these declining populations. Until recently, Pacific harbor seals (Phoca vitulina richardsi) on the west coast increased an average of 5% to 7% per year as a result of the Marine Mammal Protection Act of 1972 (Brown and Kohlman2). Pacific salmon are seasonally important prey for harbor seals (Roffe and Mate, 1984; Olesiuk, 1993); therefore quantifying and understanding the interaction between these two protected species is important for Morphobiologically sound management strategies. Because some Pacific salmonid species in a given area may be threatened or endangered, while others are relatively abundant, it is important to distinguish the species of salmonid upon which the harbor seals are preying. This study takes the first step in understanding these interactions by using molecular genetic tools for species-level identification of salmonid skeletal remains recovered from Pacific harbor seal scats.

Implantation of Subcutaneous Radio Transmitters in the Harbor Seal (Phoca vitulina)

Radio telemetry has become a standard tool for studying the behavior, physiology, life history traits, and population dynamics of marine mammals. Radio transmitters typically are attached to the hind flippers of pinnipeds or glued to the fur using marine epoxy or other cyanocrylare adhesives (Fedak et al. 1983, Bengtson 1993, Jeffries et al. 1993). Longterm data acquisition is difficult, however, because radio-flipper transmitters commonly tear from the webbing of the flipper and instruments that are glued to the fur are shed during the seasonal molt.

Molecular cloning and expression of leptin in gray and harbor seal blubber, bone marrow, and lung and its potential role in marine mammal respiratory physiology

Leptin is a multifunctional hormone, produced predominantly in adipocytes. It regulates energy balance through its impact on appetite and fat metabolism, and its concentration indicates the size of body fat reserves. Leptin also plays a vital role in stretch-induced surfactant production during alveolar development in the fetus. The structure, expression pattern, and role of leptin have not previously been explored in marine mammals. Phocid seals undergo cyclical changes in body composition as a result of prolonged fasting and intensive foraging bouts and experience rapid, dramatic, and repeated changes in lung volume during diving. Here, we report the tissue-specific expression pattern of leptin in these animals. This is the first demonstration of leptin expression in the lung tissue of a mature mammal, in addition to its expression in the blubber and bone marrow, in common with other animals. We propose a role for leptin in seal pulmonary surfactant production, in addition to its likely role in long-term energy balance. We identify substitutions in the phocine leptin sequence in regions normally highly conserved between widely distinct vertebrate groups, and, using a purified seal leptin antiserum, we confirm the presence of the leptin protein in gray seal lung and serum fractions. Finally, we report the substantial inadequacies of using heterologous antibodies to measure leptin in unextracted gray seal serum.

Social structure and underwater behavior of harbor seals in southern Monterey Bay, California

To understand harbor seal social and mating strategies, I examined site fidelity, seasonal abundance and distribution, herd integrity, and underwater behavior of individual harbor seals in southern Monterey Bay. Individual harbor seals (n = 444) were identified by natural markings and represented greater than 80% of an estimated 520 seals within this community. Year to year fidelity of individual harbor seals to southern Monterey Bay coastline was 84% (n = 388), and long-term associations (>2 yrs) among individuals were common (>40%). Consistent with these long-term associations, harbor seals were highly social underwater throughout the year. Underwater social behavior included three primary types: (1) visual and acoustic displays, such as vocalizing, surface splashing, and bubble-blowing; (2) playful or agonistic social behavior such as rolling, mounting, attending, and biting; and (3) signal gestures such as head-thrusting, fore-flipper scratch~ng, and growling. Frequency of these types of behavior was related to seal age, gender, season, and resource availability. Underwater behavior had a variety of functions, including promotion of learning and social development, reduction of aggression and preservation of social bonds by maintaining social hierarchy, and facilitation of mate selection during breeding season. Social behavior among adult males was significantly correlated with vocalization characteristics (r = 0.99, X2 = 37.7, p = 0.00087), indicating that seals may assess their competition based on underwater vocalization displays and adopt individual strategies for attracting females during breeding season based on social status. Individual mating strategies may include defending underwater territories, using scramble tactics, and developing social alliances. (PDF contains 105 pages)

Spatial and seasonal relationships between Pacific harbor seals (Phoca vitulina richardii) and their prey, at multiple scales

Knowing where pinnipeds forage is vital to managing and protecting their populations, and for assessing potential interactions with fisheries. We assessed the spatial relationship between the seasonal distribution of Pacific harbor seals (Phoca vitulina richardii) outfitted with satellite transmitters and the seasonal distributions of potential harbor seal prey species in San Francisco Bay, California. Pearson’s correlation coefficients were calculated between the number of harbor seal locations in an area of the San Francisco Bay and the abundance of specific prey species in the same area. The influence of scale on the analyses was assessed by varying the scale of analysis from 1 to 10 km. There was consistency in the prey species targeted by harbor seals year-round, although there were seasonal differences between the most important prey species. The highest correlations between harbor seals and their prey were found for seasonally abundant benthic species, located within about 10 km of the primary haul-out site. Probable foraging habitat for harbor seals was identified, based on areas with high abundances of prey species that were strongly correlated with harbor seal distribution. With comparable local data inputs, this approach has potential application to pinniped management in other areas, and to decisions about the location of marine reserves designed to protect these species.

New insights into the diets of harbor seals (Phoca vitulina) in the Salish Sea revealed by analysis of fatty acid signatures

Harbor seals (Phoca fvitulina) are an abundant predator along the west coast of North America, and there is considerable interest in their diet composition, especially in regard to predation on valued fish stocks. Available informationon harbor seal diets, primarily derived from scat analysis, suggests that adult salmon (Oncorhynchus spp.), Pacific Herring (Clupea pallasii), and gadids predominate. Because diet assessments based on scat analysis may be biased, we investigated diet composition through quantitative analysis of fatty acid signatures. Blubber samples from 49 harbor seals captured in western North America from haul-outs within the area of the San Juan Islands and southern Strait of Georgia in the Salish Sea were analyzed for fatty acid composition, along with 269 fish and squid specimens representing 27 potential prey classes. Diet estimates varied spatially, demographically, and among individual harbor seals. Findings confirmed the prevalence of previously identified prey species in harbor seal diets, but other species also contributed significantly. In particular, Black (Sebastes melanops) and Yellowtail (S. flavidus) Rockfish were estimated to compose up to 50% of some individual seal diets. Specialization and high predation rates on Black and Yellowtail Rockfish by a subset of harbor seals may play a role in the population dynamics of these regional rockfish stocks that is greater than previously realized.

Fish consumption by harbor seals (Phoca vitulina) in the San Juan Islands, Washington

The harbor seal (Phoca vitulina) is a large-bodied and abundant predator in the Salish Sea ecosystem, and its population has recovered since the 1970s after passage of the Marine Mammal Protection Act and the cessation of bounties. Little is known about how this large predator population may affect the recovery of fish stocks in the Salish Sea, where candidate marine protected areas are being proposed. We used a bioenergetics model to calculate baseline consumption rates in the San Juan Islands, Washington. Salmonids (Oncorhynchus spp.) and herring (Clupeidae) were the 2 most energetically important prey groups for biomass consumed by harbor seals. Estimated consumption of salmonids was 783 (±380 standard deviation [SD]) metric tons (t) in the breeding season and 675 (±388 SD t in the nonbreeding season. Estimated consumption of herring was 646 (±303 SD) t in the breeding season and 2151 (±706 SD) t in the nonbreeding season. Rockfish, a depressed fish stock currently in need of population recovery, composed one of the minor prey groups consumed by harbor seals (84 [±26 SD] t in the nonbreeding season). The variables of seal body mass and proportion of prey in seal diet explained >80% of the total variation in model outputs. Prey groups, such as rockfish, that are targeted for recovery may still be affected by even low levels of predation. This study highlights the importance of salmonids and herring for the seal population and provides a framework for refining consumption estimates and their confidence intervals with future data.

Space–time zero-inflated count models of Harbor seals

Environmental data are spatial, temporal, and often come with many zeros. In this paper, we included space–time random effects in zero-inflated Poisson (ZIP) and ‘hurdle’ models to investigate haulout patterns of harbor seals on glacial ice. The data consisted of counts, for 18 dates on a lattice grid of samples, of harbor seals hauled out on glacial ice in Disenchantment Bay, near Yakutat, Alaska. A hurdle model is similar to a ZIP model except it does not mix zeros from the binary and count processes. Both models can be used for zero-inflated data, and we compared space–time ZIP and hurdle models in a Bayesian hierarchical model. Space–time ZIP and hurdle models were constructed by using spatial conditional autoregressive (CAR) models and temporal first-order autoregressive (AR(1)) models as random effects in ZIP and hurdle regression models. We created maps of smoothed predictions for harbor seal counts based on ice density, other covariates, and spatio-temporal random effects. For both models predictions around the edges appeared to be positively biased. The linex loss function is an asymmetric loss function that penalizes overprediction more than underprediction, and we used it to correct for prediction bias to get the best map for space–time ZIP and hurdle models.

The brain of the fur seal, Callorhinus ursinus : with a comparative description of those of Zalophus californianus, Phoca vitulina, Ursus americanus and Monachus tropicalis /

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A Predictive Thermal Habitat Model for Harbor Seals in the Northwest Atlantic

We analyzed projections of current and future ambient temperatures along the eastern United States in relationship to the thermal tolerance of harbor seals in air. Using the earth systems model (HadGEM2-ES) and representative concentration pathways (RCPs) 4.5 and 8.5, which are indicative of two different atmospheric CO2 concentrations, we were able to examine possible shifts in distribution based on three metrics: current preferences, the thermal limit of juveniles, and the thermal limits of adults. Our analysis focused on average ambient temperatures because harbor seals are least effective at regulating their body temperature in air, making them most susceptible to rising air temperatures in the coming years. Our study focused on the months of May, June, and August from 2041-2060 (2050) and 2061-2080 (2070) as these are the historic months in which harbor seals are known to annually come ashore to pup, breed, and molt. May, June, and August are also some of the warmest months of the year. We found that breeding colonies along the eastern United States will be limited by the thermal tolerance of juvenile harbor seals in air, while their foraging range will extend as far south as the thermal tolerance of adult harbor seals in air. Our analysis revealed that in 2070, harbor seal pups should be absent from the United States coastline nearing the end of the summer due to exceptionally high air temperatures.

Development, validation, and utilization of a competitive enzyme-linked immunosorbent assay for the detection of antibodies against Brucella species in marine mammals

A competitive enzyme-linked immunosorbent assay (cELISA) was developed by using a whole-cell antigen from a marine Brucella sp. isolated from a harbor seal (Phoca vitulina). The assay was designed to screen sera from multiple marine mammal species for the presence of antibodies against marine-origin Brucella. Based on comparisons with culture-confirmed cases, specificity and sensitivity for cetacean samples tested were 73% and 100%, respectively. For pinniped samples, specificity and sensitivity values were 77% and 67%, respectively. Hawaiian monk seal (Monachus schauinslandi; n = 28) and bottlenose dolphin (Tursiops truncatus; n = 48) serum samples were tested, and the results were compared with several other assays designed to detect Brucella abortus antibodies. The comparison testing revealed the marine-origin cELISA to be more sensitive than the B. abortus tests by the detection of additional positive serum samples. The newly developed cELISA is an effective serologic method for detection of the presence of antibodies against marine-origin Brucella sp. in marine mammals.

Improving pinniped diet analyses through identification of multiple skeletal structures in fecal samples

Marine mammal diet is typically characterized by identifying fish otoliths and cephalopod beaks retrieved from stomachs and fecal material (scats). The use and applicability of these techniques has been the matter of some debate given inherent biases associated with the method. Recent attempts to identify prey using skeletal remains in addition to beaks and otoliths are an improvement; however, difficulties incorporating these data into quantitative analyses have limited results for descriptive analyses such as frequency of occurrence. We attempted to characterize harbor seal (Phoca vitulina) diet in an area where seals co-occur with several salmon species, some endangered and all managed by state or federal agencies, or both. Although diet was extremely variable within sampling date, season, year, and between years, the frequency and number of individual prey were at least two times greater for most taxa when prey structures in addition to otoliths were identified. Estimating prey mass in addition to frequency and number resulted in an extremely different relative importance of prey in harbor seal diet. These data analyses are a necessary step in generating estimates of the size, total number, and annual biomass of a prey species eaten by pinnipeds for inclusion in fisheries management plans.

Pinniped diet composition: a comparison of estimation models

Along the west coast of the United States, the potential impact of increasing pinniped populations on declining salmonid (Oncorhynchus spp.) stocks has become an issue of concern. Fisheries managers need species-specific estimates of consumption by pinnipeds to evaluate their impact on salmonid stocks. To estimate consumption, we developed a model that estimates diet composition by reconstructing prey biomass from fecal samples. We applied the model to data collected from harbor seals (Phoca vitulina) that are present year-round in the lower Columbia River where endangered stocks of salmonids pass as returning adults and as seaward-migrating smolts. Using the same data, we applied the split-sample frequency of occurrence model, which avoids reconstructing biomass by assuming that each fecal sample represents an equal volume of consumption and that within each sample each prey item represents an equal proportion of the volume. The two models for estimating diet composition yielded size-specific differences in consumption estimates that were as large as tenfold for the smallest and largest prey. Conclusions about the impact of harbor seal predation on adult salmonids, some of their largest prey species, remain uncertain without some appropriate rationale or further information (e.g. empirical captive studies) to discriminate between these models.

Characterization of TRPC2, an Essential Genetic Component of VNS Chemoreception, Provides Insights into the Evolution of Pheromonal Olfaction in Secondary-Adapted Marine Mammals

Pheromones are chemical cues released and sensed by individuals of the same species, which are of major importance in regulating reproductive and social behaviors of mammals. Generally, they are detected by the vomeronasal system (VNS). Here, we first investigated and compared an essential genetic component of vomeronasal chemoreception, that is, TRPC2 gene, of four marine mammals varying the degree of aquatic specialization and related terrestrial species in order to provide insights into the evolution of pheromonal olfaction in the mammalian transition from land to water. Our results based on sequence characterizations and evolutionary analyses, for the first time, show the evidence for the ancestral impairment of vomeronasal pheromone signal transduction pathway in fully aquatic cetaceans, supporting a reduced or absent dependence on olfaction as a result of the complete adaptation to the marine habitat, whereas the amphibious California sea lion was found to have a putatively functional TRPC2 gene, which is still under strong selective pressures, reflecting the reliance of terrestrial environment on chemical recognition among the semiadapted marine mammals. Interestingly, our study found that, unlike that of the California sea lion, TRPC2 genes of the harbor seal and the river otter, both of which are also semiaquatic, are pseudogenes. Our data suggest that other unknown selective pressures or sensory modalities might have promoted the independent absence of a functional VNS in these two species. In this respect, the evolution of pheromonal olfaction in marine mammals appears to be more complex and confusing than has been previously thought. Our study makes a useful contribution to the current understanding of the evolution of pheromone perception of mammals in response to selective pressures from an aquatic environment.