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Forage fishes are key energy conduits that transfer primary and secondary productivity to higher trophic levels. As novel environmental conditions caused by climate change alter ecosystems and predator–prey dynamics, there is a critical need to understand how forage fish control bottom-up forcing of food web dynamics. In the northeast Pacific, northern anchovy ( Engraulis mordax ) is an important forage species with high interannual variability in population size that subsequently impacts the foraging and reproductive ecology of marine predators. Anchovy habitat suitability from a species distribution model (SDM) was assessed as an indicator of the diet, distribution and reproduction of four predator species. Across 22 years (1998–2019), this anchovy ecosystem indicator (AEI) was significantly positively correlated with diet composition of all species and the distribution of common murres ( Uria aalge ), Brandt's cormorants ( Phalacrocorax penicillatus ) and California sea lions ( Zalophus californianus ), but not rhinoceros auklets ( Cerorhinca monocerata ). The capacity for the AEI to explain variability in predator reproduction varied by species but was strongest with cormorants and sea lions. The AEI demonstrates the utility of forage SDMs in creating ecosystem indicators to guide ecosystem-based management.

Bioenergetics is the study of how animals achieve energetic balance. Energetic balance results from the energetic expenditure of an individual and the energy they extract from their environment. Ingested energy depends on several extrinsic (e.g prey species, nutritional value and composition, prey density and availability) and intrinsic factors (e.g. foraging effort, success at catching prey, digestive processes and associated energy losses, and digestive capacity). While the focus in bioenergetic modelling is often on the energetic costs an animal incurs, the robust estimation of an individual’s energy intake is equally critical for producing meaningful predictions. Here, we review the components and processes that affect energy intake from ingested gross energy to biologically useful net energy (NE). The current state of knowledge of each parameter is reviewed, shedding light on research gaps to advance this field. The review highlighted that the foraging behaviour of many marine mammals is relatively well studied via biologging tags, with estimates of success rate typically assumed for most species. However, actual prey capture success rates are often only assumed, although we note studies that provide approaches for its estimation using current techniques. A comprehensive collation of the nutritional content of marine mammal prey species revealed a robust foundation from which prey quality (comprising prey species, size and energy density) can be assessed, though data remain unavailable for many prey species. Empirical information on various energy losses following ingestion of prey was unbalanced among marine mammal species, with considerably more literature available for pinnipeds. An increased understanding and accurate estimate of each of the components that comprise a species NE intake are an integral part of bioenergetics. Such models provide a key tool to investigate the effects of disturbance on marine mammals at an individual and population level and to support effective conservation and management.

This work was funded by the Marine Mammal Commission (MMC19-173). The Office of Naval Research funded the bioenergetic workshop (N000142012392) that provided support for this work. Bioenergetic approaches are increasingly used to understand how marine mammal populations could be affected by a changing and disturbed aquatic environment. There remain considerable gaps in our knowledge of marine mammal bioenergetics, which hinder the application of bioenergetic studies to inform policy decisions. We conducted a priority-setting exercise to identify high-priority unanswered questions in marine mammal bioenergetics, with an emphasis on questions relevant to conservation and management. Electronic communication and a virtual workshop were used to solicit and collate potential research questions from the marine mammal bioenergetic community. From a final list of 39 questions, 11 were identified as ‘key’ questions because they received votes from at least 50% of survey participants. Key questions included those related to energy intake (prey landscapes, exposure to human activities) and expenditure (field metabolic rate, exposure to human activities, lactation, time-activity budgets), energy allocation priorities, metrics of body condition and relationships with survival and reproductive success and extrapolation of data from one species to another. Existing tools to address key questions include labelled water, animal-borne sensors, mark-resight data from long-term research programs, environmental DNA and unmanned vehicles. Further validation of existing approaches and development of new methodologies are needed to comprehensively address some key questions, particularly for cetaceans. The identification of these key questions can provide a guiding framework to set research priorities, which ultimately may yield more accurate information to inform policies and better conserve marine mammal populations. Publisher PDF

Marine central-place foragers are increasingly faced with altered prey landscapes, necessitating predictions of the impact of such changes on behavior, reproductive success, and population dynamics. We used state-dependent behavioral life history theory implemented via Stochastic Dynamic Programming (SDP) to explore the influence of changes in prey distribution and energy gain from foraging on the behavior and reproductive success of a central place forager during lactation. Our work is motivated by northern fur seals (Callorhinus ursinus) because of the ongoing population decline of the Eastern Pacific stock and projected declines in biomass of walleye pollock (Gadus chalcogrammus), a key fur seal prey species in the eastern Bering Sea. We also explored how changes in female and pup metabolic rates, body size, and lactation duration affected model output to provide insight into traits that might experience selective pressure in response to reductions in prey availability. Simulated females adopted a central-place foraging strategy after an initial extended period spent on land (4.7 - 8.3 days). Trip durations increased as the high energy prey patch moved farther from land or when the energy gain from foraging decreased. Increases in trip duration adversely affected pup growth rates and wean mass despite attempts to compensate by increasing land durations. Metabolic rate changes had the largest impacts on pup wean mass, with reductions in a pup's metabolic rate allowing females to successfully forage at distances of 600+ km from land for up to 15+ days. Our results indicate that without physiological adaptations, a rookery is unlikely to be viable if the primary foraging grounds are 400 km or farther from the rookery. To achieve pup growth rates characteristic of a population experiencing rapid growth, model results indicate the primary foraging grounds need to be150 km from the rookery.

Acoustic disturbance is a growing conservation concern for wildlife populations because it can elicit physiological and behavioral responses that can have cascading impacts on population dynamics. State-dependent behavioral and life history models implemented via Stochastic Dynamic Programming (SDP) provide a natural framework for quantifying biologically meaningful population changes resulting from disturbance by linking environment, physiology, and metrics of fitness. We developed an SDP model using the endangered western gray whale (Eschrichtius robustus) as a case study because they experience acoustic disturbance on their summer foraging grounds. We modeled the behavior and physiological dynamics of pregnant females as they arrived on the feeding grounds and predicted the probability of female and offspring survival, with and without acoustic disturbance and in the presence/absence of high prey availability. Upon arrival in mid-May, pregnant females initially exhibited relatively random behavior before they transitioned to intensive feeding that resulted in continual fat mass gain until departure. This shift in behavior co-occurred with a change in spatial distribution; early in the season, whales were more equally distributed among foraging areas with moderate to high energy availability, whereas by mid-July whales transitioned to predominate use of the location that had the highest energy availability. Exclusion from energy-rich offshore areas led to reproductive failure and in extreme cases, mortality of adult females that had lasting impacts on population dynamics. Simulated disturbances in nearshore foraging areas had little to no impact on female survival or reproductive success at the population level. At the individual level, the impact of disturbance was unequally distributed across females of different lengths, both with respect to the number of times an individual was disturbed and the impact of disturbance on vital rates. Our results highlight the susceptibility of large capital breeders to reductions in prey availability, and indicate that who, where, and when individuals are disturbed are likely to be important considerations when assessing the impacts of acoustic activities. This model provides a framework to inform planned acoustic disturbances and assess the effectiveness of mitigation strategies for large capital breeders.

RATIONALE Stable isotope analysis of keratinized tissues is an informative tool for quantifying foraging ecology that can address questions related to niche specialization and temporal variation in behavior. Application of this approach relies on an understanding of tissue growth and how isotope ratios relate to physiological and ecological processes, data that are lacking for many species. METHODS We collected paired whisker length measurements from northern elephant seals to estimate growth and shedding patterns (n = 16). A subset of seals (n = 5) carried a satellite tag and time-depth recorder across the 7+ month foraging trip following the annual pelage molt. Stable isotopes of carbon and nitrogen were measured in whisker segments grown across the 6+ week fasting on land and the subsequent foraging trip; profiles were combined with growth parameters to timestamp each segment and investigate relationships with foraging behavior. RESULTS Whisker loss and initial regrowth primarily occurred during the annual pelage molt, but newly grown whiskers exhibited active, nonlinear growth across the foraging trip. The δ13 C and δ15 N values were higher in segments grown on land than at sea and exhibited a characteristic decline upon departure from the rookery. There was a relationship between latitude and longitude and δ15 N values, and individual whisker segments grown at sea could be classified to the correct ecoregion with 81% accuracy. CONCLUSIONS Fasting affected both δ13 C and δ15 N values and the ability to exclude these values from ecological investigations is crucial given the temporal overlap with tissue growth. The rapid decline in isotope ratios upon departure can be used to isolate portions of the whisker with a strong physiological signal, even for whiskers with unknown growth histories. The active growth across the foraging trip combined with the ability to identify differences in foraging behavior validates the utility of this approach for addressing ecological questions.

Food availability is a key concern for the conservation of marine top predators, particularly during a time when they face a rapidly changing environment and continued pressure from commercial fishing activities. Northern fur seals (Callorhinus ursinus) breeding on the Pribilof Islands in the eastern Bering Sea have experienced an unexplained population decline since the late-1990s. Dietary overlap with a large U.S. fishery for walleye pollock (Gadus chalcogrammus) in combination with changes in maternal foraging behavior and pup growth has led to the hypothesis that food limitation may be contributing to the population decline. We developed age- and sex-specific bioenergetic models to estimate fur seal energy intake from May–December in six target years, which were combined with diet data to quantify prey consumption. There was considerable sex- and age-specific variation in energy intake because of differences in body size, energetic costs, and behavior; net energy intake was lowest for juveniles (18.9 MJ sea-day–1, 1,409.4 MJ season–1) and highest for adult males (66.0 MJ sea-day–1, 7,651.7 MJ season–1). Population-level prey consumption ranged from 255,232 t (222,159 – 350,755 t, 95% CI) in 2006 to 500,039 t (453,720 – 555,205 t) in 1996, with pollock comprising between 41.4 and 76.5% of this biomass. Interannual variation in size-specific pollock consumption appeared largely driven by the availability of juvenile fish, with up to 81.6% of pollock biomass coming from mature pollock in years of poor age-1 recruitment. Relationships among metabolic rates, trip durations, pup growth rates, and energy intake of lactating females suggest the most feasible mechanism to increase pup growth rates is by increasing foraging efficiency through reductions in maternal foraging effort, which is unlikely to occur without increases in localized prey density. By quantifying year-specific fur seal consumption of pollock, our study provides a pathway to incorporate fur seals into multispecies pollock stock assessment models, which is critical for fur seal and fishery management given they were a significant source of mortality for both juvenile and mature pollock.

Mercury (Hg) is a well-known toxicant in wildlife and humans. High total Hg concentrations ([THg]) have been reported in central California harbor seals Phoca vitulina richardii. We evaluated the effects of presence/absence of early natal coat (lanugo), year (2012 to 2017), sex, stranding location, and trophic ecology (ẟ

Human activities continue to expand in marine and terrestrial environments, leading to increased interactions with wildlife that can have negative impacts on population dynamics. Approaches for quantifying how these interactions translate to population-level effects are therefore crucial for effective management practices and balancing human-wildlife tradeoffs. We developed a method using state-dependent behavioral theory implemented via Stochastic Dynamic Programming (SDP) for predicting the population consequences of disturbance on the physiology and reproductive behavior of an income-breeding mammal, using California sea lions (Zalophus californianus) as a motivating species. Emergent properties of the model included reproductive characteristics associated with long-lived species, such as variation in the age at first reproduction, early termination of pregnancy, and skipped breeding. In undisturbed model simulations, reproductive rates and the average wean date were consistent with empirically-derived estimates from sea lions and other marine mammals, highlighting the utility of this model for quantifying fecundity estimates of data-deficient species and addressing fundamental ecological processes. In disturbed model simulations, exposure to prolonged, repetitive disturbances negatively impacted population growth; in addition, short, infrequent disturbances had the potential for adverse effects depending on the behavioral response of sea lions and the probability of being disturbed. The adverse effect of disturbance on population dynamics was due to a combination of reduced pup recruitment (survival to age one) resulting from a lower wean mass and increased abortion rates that led to skipped reproductive years, both of which have been documented for marine mammal populations experiencing natural fluctuations in prey availability. The derivation of state- and time-dependent reproductive decisions using an SDP model is an effective approach that links behavioral and energetic effects at the individual level to changes at the population level, and one that serves a dual purpose in the ability to quantify basic biological parameters and address ecological questions irrespective of disturbance.

Intraspecific variability in foraging behavior has been documented across a range of taxonomic groups, yet the energetic consequences of this variation are not well understood for many species. Understanding the effect of behavioral variation on energy expenditure and acquisition is particularly crucial for mammalian carnivores because they have high energy requirements that place considerable pressure on prey populations. To determine the influence of behavior on energy expenditure and balance, we combined simultaneous measurements of at‐sea field metabolic rate (FMR) and foraging behavior in a marine carnivore that exhibits intraspecific behavioral variation, the California sea lion (Zalophus californianus). Sea lions exhibited variability in at‐sea FMR, with some individuals expending energy at a maximum of twice the rate of others. This variation was in part attributable to differences in diving behavior that may have been reflective of diet; however, this was only true for sea lions using a foraging strategy consisting of epipelagic (

Summary 1.Anthropogenic disturbance is of increasing concern for wildlife populations, necessitating the development of models that link behavioural changes at the individual level with biologically meaningful changes at the population level. 2.We developed a general framework for estimating the fitness consequences of disturbance that affects foraging behaviour using state-dependent behavioural theory implemented by Stochastic Dynamic Programming (SDP). We illustrate this framework using generalized examples of pinnipeds, a group of marine carnivores that include both capital- and income-breeding species. We examined how disturbance affected pup recruitment separately for each reproductive strategy, and the impact of foraging decisions and parameter values on model results. 3.The effect of disturbance on pup recruitment was greater for income than capital breeders for all disturbance scenarios. Disturbance had negligible effects on pup recruitment when it occurred within less frequented foraging patches, but moderate to large effects when it occurred within an important foraging patch. Model results were sensitive to values of patch productivity (the energy gained from successful foraging), the probability of disturbance, and individual behavioural choices in the face of disturbance. 4.State-dependent behavioural theory implemented by SDP is a powerful tool for investigating when behavioural changes in response to disturbance may be meaningful at the population level. This approach allows us to incorporate many factors that are known to influence the behavioural and physiological responses of animals to anthropogenic disturbance, and places disturbance within the context of a temporally and spatially variable environment. The general framework we have developed can be used to estimate the consequences of anthropogenic disturbance across a broad range of species. This article is protected by copyright. All rights reserved.

This study was supported by Office of Naval Research grant N00014‐16‐1‐2858: ‘PCoD+: Developing widely‐applicable models of the population consequences of disturbance’. DPC, MM, EAM and LKS were supported by the E&P Sound and Marine Life Joint Industry Project of the International Association of Oil and Gas Producers. JAG was supported by funding from the Young Investigator Program at the Office of Naval Research (award no. N00014‐16‐1‐2477). VH was funded by European Research Council Grant No. 322814 awarded to A.M. de Roos. Animals make behavioural and reproductive decisions that maximise their lifetime reproductive success, and thus their fitness, in light of periodic and stochastic variability of the environment. Modelling the variation of an individual's energy levels formalises this tradeoff and helps to quantify the population‐level consequences of stressors (e.g. disturbance from human activities and environmental change) that can affect behaviour or physiology. In this study, we develop a dynamic state variable model for the spatially explicit behaviour, physiology and reproduction of a female, long‐lived, migratory marine vertebrate. The model can be used to investigate the spatio‐temporal patterns of behaviour and reproduction that allow an individual to maximise its overall reproductive output. We parametrised the model for eastern North Pacific blue whales Balaenoptera musculus, and used it to predict the effects of changing environmental conditions and increasing human disturbance on the population's vital rates. In baseline conditions, the model output had high fidelity to observed energy dynamics, movement patterns and reproductive strategies. Simulated scenarios suggested that environmental changes could have severe consequences on the population's vital rates, but that individuals could tolerate high levels of anthropogenic disturbance. However, this ability depended on where, when and how often disturbance occurred. In scenarios with both environmental change and anthropogenic disturbance, synergistic interactions caused stronger effects than in isolation. In general, larger body size offered a buffer against stochasticity and disturbance, and, consequently, we predicted juveniles to be more susceptible to disturbance. We also predicted that females prioritise their own survival at the expense of the current reproductive attempt, presumably the result of their long lifespan. Our approach provides a general framework to make predictions of the cumulative and synergistic effects of human disturbance and climate change on migratory populations, which can inform effective management and conservation efforts. Publisher PDF

Mercury (Hg) poses a health risk to wildlife populations and has been documented at relatively high concentrations in many marine mammals, including wild-caught pinnipeds along the central California, US coast. We measured total Hg concentrations ([THg]) in hair and blood of live-stranded harbor seals (HS

Quantifying metabolic rates and the factors that influence them is key to wildlife conservation efforts because anthropogenic activities and habitat alteration can disrupt energy balance, which is critical for reproduction and survival. We investigated the effect of diving behaviour, diet and season on field metabolic rates (FMR) and foraging success of lactating northern fur seals (Callorhinus ursinus) from the Pribilof Islands during a period of population decline. Variation in at-sea FMR was in part explained by season and trip duration, with values that ranged from 5.18 to 9.68 W kg−1 (n = 48). Fur seals experienced a 7.2% increase in at-sea FMR from summer to fall and a 1.9% decrease in at-sea FMR for each additional day spent at sea. There was no effect of foraging effort, dive depth or diet on at-sea FMR. Mass gains increased with trip duration and were greater in the fall compared with summer, but were unrelated to at-sea FMR, diving behaviour and diet. Seasonal increases in at-sea FMR may have been due to costs associated with the annual molt but did not appear to adversely impact the ability of females to gain mass on foraging trips. The overall high metabolic rates in conjunction with the lack of any diet-related effects on at-sea FMR suggests that northern fur seals may have reached a metabolic ceiling early in the population decline. This provides indirect evidence that food limitation may be contributing to the low pup growth rates observed in the Pribilof Islands, as a high metabolic overhead likely results in less available energy for lactation. The limited ability of female fur seals to cope with changes in prey availability through physiological mechanisms is particularly concerning given the recent and unprecedented environmental changes in the Bering Sea that are predicted to have ecosystem-level impacts., Lay summary: We measured metabolic rates of northern fur seals from a declining population and examined the influence of season and foraging behavior on energy expenditure. Our results are relevant for bioenergetic modelling efforts and provide insight into the potential role of prey availability in the population decline.

Rationale Mixing models are a common method for quantifying the contribution of prey sources to the diet of an individual using stable isotope analysis; however, these models rely upon a known trophic discrimination factor (hereafter, TDF) that results from fractionation between prey and animal tissues. Quantifying TDFs in captive animals is ideal, because diet is controlled and the proportional contributions and isotopic values of all prey items are known. Methods To calculate TDFs for the Hawaiian monk seal, northern elephant seal, bearded seal, ringed seal, spotted seal, harbor seal, and California sea lion, we obtained whiskers, serum, plasma, red blood cells, and prey items from nine captive individuals. We obtained δ13C and δ15N values using continuous-flow isotope-ratio mass spectrometry. The average δ13C and δ15N values from bulk and lipid-corrected prey from the diet were subtracted from the δ13C and δ15N values of each blood and whisker sample to calculate tissue-specific TDFs for each individual (∆13C or ∆15N). Results The ∆13C values ranged from +1.7 to +3.2‰ (bulk prey) and from +0.8 to +1.9‰ (lipid-corrected prey) for the various blood components, and from +3.9 to +4.6‰ (bulk prey) or +2.6 to +3.9‰ (lipid-corrected prey) for whiskers. The ∆15N values ranged from +2.2 to +4.3‰ for blood components and from +2.6 to +4.0‰ for whiskers. The TDFs tended to group by tissue, with whiskers having greater ∆13C values than blood components. In contrast, the ∆15N values were greater in serum and plasma than in red blood cells and whiskers. Conclusions By providing the first TDF values for five seal species (family Phocidae) and one otariid species (family Otariidae), our study facilitates more accurate mixing models for these species. These values are particularly important for critically endangered Hawaiian monk seals and the three Arctic seal species (bearded, ringed, and spotted) that are faced with a rapidly changing environment. Copyright © 2016 John Wiley & Sons, Ltd.

Marine predators frequently exhibit consistency in foraging behaviors despite the dynamic nature of marine ecosystems, which has the potential for ecological and evolutionary implications depending on the timescale at which it persists. We examined behavioral consistency in movements and diving behavior of adult female California sea lions (Zalophus californianus), which are abundant, generalist central-place foragers inhabiting an ecosystem characterized by small- and broad-scale oceanographic variability. We used biologging devices to measure repeatability of behavior within a season and stable isotope analysis of whiskers to quantify behavior across a 2-year period associated with anomalous environmental conditions that affected prey availability. Sea lions were significantly repeatable in all variables across multiple timescales (Radj = 0.26–0.82), although repeatability estimates were generally higher for variables related to characteristics of individual dives (e.g., dive depth) than those that described dive bouts (e.g., bout duration) or spatial use (e.g., volume of 3D utilization distribution). These differences may result from the fact that diving behaviors vary with prey type, whereas spatial use and bout variables may reflect the foraging success within prey patches or movement among patches. There was variation in how predictable individual sea lions were in their diving behaviors, which was largely unrelated or negatively related to foraging site fidelity. The strength of behavioral consistency decreased with time yet persisted across the 2-year period, suggesting that while sea lions alter their behavior in response to environmental change, the behavioral flexibility of individuals may ultimately be constrained by consistency.

Integrating behavior and physiology is critical to formulating new hypotheses on the evolution of animal life-history strategies. Migratory capital breeders acquire most of the energy they need to sustain migration, gestation, and lactation before parturition. Therefore, when predicting the impact of environmental variation on such species, a mechanistic understanding of the physiology of their migratory behavior is required. Using baleen whales as a model system, we developed a dynamic state variable model that captures the interplay among behavioral decisions, energy, reproductive needs, and the environment. We applied the framework to blue whales (Balaenoptera musculus) in the eastern North Pacific Ocean and explored the effects of environmental and anthropogenic perturbations on female reproductive success. We demonstrate the emergence of migration to track prey resources, enabling us to quantify the trade-offs among capital breeding, body condition, and metabolic expenses. We predict that periodic climatic oscillations affect reproductive success less than unprecedented environmental changes do. The effect of localized, acute anthropogenic impacts depended on whales' behavioral response to the disturbance; chronic, but weaker, disturbances had little effect on reproductive success. Because we link behavior and vital rates by modeling individuals' energetic budgets, we provide a general framework to investigate the ecology of migration and assess the population consequences of disturbance, while identifying critical knowledge gaps.

The extent of increasing anthropogenic impacts on large marine vertebrates partly depends on the animals’ movement patterns. Effective conservation requires identification of the key drivers of movement including intrinsic properties and extrinsic constraints associated with the dynamic nature of the environments the animals inhabit. However, the relative importance of intrinsic versus extrinsic factors remains elusive. We analyze a global dataset of ∼2.8 million locations from >2,600 tracked individuals across 50 marine vertebrates evolutionarily separated by millions of years and using different locomotion modes (fly, swim, walk/paddle). Strikingly, movement patterns show a remarkable convergence, being strongly conserved across species and independent of body length and mass, despite these traits ranging over 10 orders of magnitude among the species studied. This represents a fundamental difference between marine and terrestrial vertebrates not previously identified, likely linked to the reduced costs of locomotion in water. Movement patterns were primarily explained by the interaction between species-specific traits and the habitat(s) they move through, resulting in complex movement patterns when moving close to coasts compared with more predictable patterns when moving in open oceans. This distinct difference may be associated with greater complexity within coastal microhabitats, highlighting a critical role of preferred habitat in shaping marine vertebrate global movements. Efforts to develop understanding of the characteristics of vertebrate movement should consider the habitat(s) through which they move to identify how movement patterns will alter with forecasted severe ocean changes, such as reduced Arctic sea ice cover, sea level rise, and declining oxygen content., Workshop funding was granted by the University of Western Australia (UWA) Oceans Institute, the Australian Institute of Marine Science (AIMS), and King Abdullah University of Science and Technology (KAUST). A.M.M.S. was supported by Australian Research Council Grant DE170100841 and an Indian Ocean Ocean Marine Research Centre (UWA, AIMS, Commonwealth of Scientific and Industrial Research Organisation) fellowship. J.P.R., V.M.E., and J.F.G. were supported by Agencia Estatal de Investigación (AEI, Spain) and Fondo Europeo de Desarrollo Regional (FEDER) through project Spatiotemporality in Sociobological Interactions, Models and Methods (SPASIMM) (FIS2016-80067-P AEI/FEDER, European Union), and by research funding from KAUST. J.P.R. was supported by Ministerio de Educación, Cultura y Deporte (Formación de Profesorado Universitario Grant, Spain). D.W.S. was supported by the UK Natural Environment Research Council and Save Our Seas Foundation. N.Q. was supported by Fundação para a Ciência e Tecnologia (Portugal). M.M.C.M. was supported by a Coordenação de Aperfeiçoamento de pessoal de Nível Superior fellowship (Ministry of Education).

Feeding ecology is a primary driver of contaminant exposure for mammalian species, thus tools to quantify diet and habitat use are a key component of many ecotoxicological studies. In this chapter, we discuss the fate and transport of contaminants to coastal ecosystems and review the feeding ecology tools available for coastal mammals, a group comprised of marine mammals and terrestrial mammals that rely on marine-derived resources. We highlight these ecological connections related to contaminants, focusing on stable isotopes of carbon, nitrogen, and sulfur for elucidating trophic and nontrophic interactions with contaminant exposure and tissue concentrations. In addition, we discuss tools that can be used alone or in conjunction with the aforementioned stable isotopes to describe the feeding ecology of individuals, populations, or species. Our mission is to expose readers to a nonlinear view of contaminant sources and transport to coastal ecosystems and to facilitate the use of cross-disciplinary techniques and tools to further the fields of ecotoxicology and ecology.

We measured total mercury (THg) concentrations in California sea lions (Zalophus californianus) and examined how concentrations varied with age class, colony, and sex. Because Hg exposure is primarily via diet, we used nitrogen (δ (15)N) and carbon (δ (13)C) stable isotopes to determine if intraspecific differences in THg concentrations could be explained by feeding ecology. Blood and hair were collected from 21 adult females and 57 juveniles from three colonies in central and southern California (San Nicolas, San Miguel, and Año Nuevo Islands). Total Hg concentrations ranged from 0.01 to 0.31 μg g(-1) wet weight (ww) in blood and 0.74 to 21.00 μg g(-1) dry weight (dw) in hair. Adult females had greater mean THg concentrations than juveniles in blood (0.15 vs. 0.03 μg(-1) ww) and hair (10.10 vs. 3.25 μg(-1) dw). Age class differences in THg concentrations did not appear to be driven by trophic level or habitat type because there were no differences in δ (15)N or δ (13)C values between adults and juveniles. Total Hg concentrations in adult females were 54 % (blood) and 24 % (hair) greater in females from San Miguel than females from San Nicolas Island, which may have been because sea lions from the two islands foraged in different areas. For juveniles, we detected some differences in THg concentrations with colony and sex, although these were likely due to sampling effects and not ecological differences. Overall, THg concentrations in California sea lions were within the range documented for other marine mammals and were generally below toxicity benchmarks for fish-eating wildlife.

Mercury (Hg) biomonitoring of pinnipeds increasingly utilizes nonlethally collected tissues such as hair and blood. The relationship between total Hg concentrations ([THg]) in these tissues is not well understood for marine mammals, but it can be important for interpretation of tissue concentrations with respect to ecotoxicology and biomonitoring. We examined [THg] in blood and hair in multiple age classes of four pinniped species. For each species, we used paired blood and hair samples to quantify the ability of [THg] in hair to predict [THg] in blood at the time of sampling and examined the influence of varying ontogenetic phases and life history of the sampled animals. Overall, we found that the relationship between [THg] in hair and blood was affected by factors including age class, weaning status, growth, and the time difference between hair growth and sample collection. Hair [THg] was moderately to strongly predictive of current blood [THg] for adult female Steller sea lions (Eumetopias jubatus), adult female California sea lions (Zalophus californianus), and adult harbor seals (Phoca vitulina), whereas hair [THg] was poorly predictive or not predictive (different times of year) of blood [THg] for adult northern elephant seals (Mirounga angustirostris). Within species, except for very young pups, hair [THg] was a weaker predictor of blood [THg] for prereproductive animals than for adults likely due to growth, variability in foraging behavior, and transitions between ontogenetic phases. Our results indicate that the relationship between hair [THg] and blood [THg] in pinnipeds is variable and that ontogenetic phase and life history should be considered when interpreting [THg] in these tissues.

Characterizing habitat suitability for a marine predator requires an understanding of the environmental heterogeneity and variability over the range in which a population moves during a particular life cycle. Female California sea lions (Zalophus californianus) are central-place foragers and are particularly constrained while provisioning their young. During this time, habitat selection is a function of prey availability and proximity to the rookery, which has important implications for reproductive and population success. We explore how lactating females may select habitat and respond to environmental variability over broad spatial and temporal scales within the California Current System. We combine near-real-time remotely sensed satellite oceanography, animal tracking data (n = 72) from November to February over multiple years (2003-2009) and Generalized Additive Mixed Models (GAMMs) to determine the probability of sea lion occurrence based on environmental covariates. Results indicate that sea lion presence is associated with cool (

We measured total selenium and total mercury concentrations ([TSe] and [THg]) in hair ( n = 138) and blood ( n = 73) of harbor seals ( Phoca vitulina ) from California to assess variation by geography and sex, and inferred feeding relationships based on carbon, nitrogen, and sulfur stable isotopes. Harbor seals from Hg-contaminated sites had significantly greater [THg], and lesser [TSe] and TSe:THg molar ratios than seals from a relatively uncontaminated site. Males had significantly greater [THg] than females at all locations. Sulfur stable isotope values explained approximately 25% of the variability in [THg], indicating increased Hg exposure for seals with a greater use of estuarine prey species. Decreased [TSe] in harbor seals from Hg-contaminated regions may indicate a relative Se deficiency to mitigate the toxic effects of Hg. Further investigation into the Se status and the potential negative impact of Hg on harbor seals from Hg-contaminated sites is warranted.

Intraspecific variability is increasingly recognized as an important component of foraging behavior that can have implications for both population and community dynamics. We used an individual-level approach to describe the foraging behavior of an abundant, generalist predator that inhabits a dynamic marine ecosystem, focusing specifically on the different foraging strategies used by individuals in the same demographic group. We collected data on movements and diving behavior of adult female California sea lions (Zalophus californianus) across multiple foraging trips to sea. Sea lions (n=35) used one of three foraging strategies that primarily differed in their oceanic zone and dive depth: a shallow, epipelagic strategy, a mixed epipelagic/benthic strategy, and a deep-diving strategy. Individuals varied in their degree of fidelity to a given strategy, with 66% of sea lions using only one strategy on all or most of their foraging trips across the two-month tracking period. All foraging strategies were present in each of the sampling years, but there were inter-annual differences in the population-level importance of each strategy that may reflect changes in prey availability. Deep-diving sea lions traveled shorter distances and spent a greater proportion of time at the rookery than sea lions using the other two strategies, which may have energetic and reproductive implications. These results highlight the importance of an individual-based approach in describing the foraging behavior of female California sea lions and understanding how they respond to the seasonal and annual changes in prey availability that characterize the California Current System.

Mercury (Hg) poses a health risk to wildlife populations and has been documented at relatively high concentrations in many marine mammals, including wild-caught pinnipeds along the central...

On 16 June 1979, a herd of 41 sperm whales stranded near the mouth of the Siuslaw River in Florence, Oregon. The stomach contents from 32 whales were collected, identified to the lowest taxonomic level possible, enumerated, and measured. A total of 20,247 cephalopod lower beaks that represented 24 species from 14 different families were recovered. The most numerous species were Histioteuthis hoylei (25.9%), Taonius borealis (12.9%), Galiteuthis phyllura (11.2%), Gonatopsis/Berryteuthis type (10.9%), and Moroteuthis robusta (10.7%). Reconstructed estimates of mass indicated that M. robusta contributed almost 50% of the total mass of cephalopods consumed, followed by H. hoylei (19.3%), and T. borealis (7.0%). The most important species in the diet of stranded whales were M. robusta, H. hoylei, T. borealis, G. phyllura, Octopoteuthis deletron, and Gonatopsis/Berryteuthis type. There were significant differences in the diet of males and females, but no differences between sperm whales of different age groups. Overall, sperm whales primarily consumed small cephalopods that were likely eaten south of 45oN in or near the California Current System. This study provides new estimates of the food habits of sperm whales in the northeast Pacific from one of the largest strandings of this species.

Skin biopsies were collected from free-ranging harbor seals (Phoca vitulina richardii) from central California (n = 53). Microscopic examination of hematoxylin and eosin-stained tissue sections revealed the presence of tightly coiled nematode larvae within the ostia of numerous hair follicles of four seals. Parasites were characterized by paired lateral alae, platymyarian musculature, and an indistinct, uninucleate digestive tract. Mild chronic superficial dermatitis and perifolliculitis were evident microscopically in association with the intrafollicular parasites. Histomorphologic features of the larvae and their presence within hair follicles are consistent with previous reports of the facultative nematode parasite Pelodera strongyloides. This is the first published report of P. strongyloides infection in any marine mammal. This parasite may be acquired by marine mammals through close contact with soil or decaying organic material and should be considered as a potential differential diagnosis for dermatitis in marine mammals that use terrestrial resting sites.

A B S T R A C T Domoic acid (DA) is a potent neurotoxin that has caused strandings and mortality of seabirds and marine mammals off the California coast. Pacific harbor seals (Phoca vitulina richardii) are an abundant, nearshore species in California; however, DA exposure and toxicosis have not been documented for harbor seals in this region. To investigate DA exposure in harbor seals, samples were collected from freeranging and stranded seals off California to assess exposure, clinical signs of toxicosis, and brain lesions in harbor seals exposed to DA. Domoic acid was detected in 65% (17/26) of urine samples collected from apparently healthy free-ranging seals, with concentrations of 0.4–11.7 ng/ml. Domoic acid also was detected in feces (2.4–2887 ng/g), stomach contents (1.4 ng/g; stranded only), milk (2.2 ng/ml; stranded only), amniotic fluid (9.7 ng/ml; free-ranging only), fetal meconium (14.6–39.8 ng/g), and fetal urine (2.0–10.2 ng/ml). Clinical signs indicative of DA toxicosis were observed in two live-stranded seals, and included disorientation, seizures, and uncoordinated movements. Histopathology revealed the presence of brain lesions consistent with DA toxicosis in two live-stranded seals, and one free-ranging seal that died during capture. Results indicated that harbor seals were exposed to DA, exhibited clinical signs and histological lesions associated with DA exposure, and that pups were exposed to DA in utero and during lactation via milk. Future investigation is required to determine the magnitude of impact that DA has on the health and mortality of harbor seals.

Whisker growth dynamics in two North Pacific pinnipeds: implications for determining foraging ecology from stable isotope analysis McHuron, EA. 1* , Walcott, S.M. 2,3 , Zeligs, J. 4 , Skrovan, S. 4 , Costa, D.P 1 , Reichmuth, C. 2 Department of Ecology & Evolutionary Biology, University of California, Santa Cruz, CA 95060, USA Institute of Marine Sciences, Long Marine Laboratory, Santa Cruz, CA 95060, USA Department of Biological Sciences, University of Alaska, Anchorage, AK 99508, USA Moss Landing Marine Laboratories, Moss Landing, CA 95039, USA *Corresponding author, emchuron@ucsc.edu Abstract Stable isotope analysis (SIA) of whiskers is increasingly used to investigate the foraging ecology of pinnipeds. An understanding of whisker growth dynamics is lacking for most species, yet is necessary for study design and interpretation of isotope data. Here we present measurements of whisker growth in five California sea lions (Zalophus californianus) and two spotted seals (Phoca largha) obtained using photogrammetry. Data were collected from captive individuals for at least one year, resulting in serial measurements of 321 sea lion and 153 spotted seal whiskers. The sea lion whiskers exhibited linear growth, with growth rates that ranged from

Since the introduction of the population consequences of disturbance (PCoD) conceptual model, lost energy has become a central component of modelling biologically meaningful disturbance. Long-term datasets on northern elephant seals of Ano Nuevo, CA and bottlenose dolphins of Sarasota Bay, FL span several environmental disturbances, allowing us to quantify how behavior, physiology, and vital rates change with natural reductions in prey. While complete PCoD models are possible with large datasets, health metrics in particular can be difficult or impossible to collect for many species. In addition, PCoD models to date have assumed the worst-case scenario, in that animals exposed to a disturbance cannot compensate for lost foraging. Combined with knowledge of energetic requirements of individuals, Stochastic Dynamic Programming (SDP) models examine potential compensatory behavioral mechanisms that individuals can employ in the presence of a disturbance. However, SDP bioenergetic models require data on the energetic landscape, time-activity budgets, movement probabilities between habitats, and metabolic rates on a timescale appropriate for energy acquisition. This approach is currently being implemented for California sea lions and gray whales with the potential to ground-truth SDP models by comparing them with current elephant seal and bottlenose dolphin models.

Hair is used to determine trace elements exposure and status of pinnipeds because it is an excretory route for many elements and can be collected non-lethally. Despite increased use, there have been few studies on how sampling designs and procedures (e.g., hair type, collection site) affect results. The objective of this study was to determine whether concentrations of an essential (selenium; Se) and non-essential element (mercury; Hg) differed between hair samples collected from two body locations on harbor seals (Phoca vitulina). Concentrations of Se and total Hg (THg) differed between mid-dorsal midline and neck samples, and although the absolute differences were relatively small (Δ(absolute) Se = 0.69 μg g(-1), Δ(absolute) THg = 2.86 μg g(-1)), the relative differences were large (Δ(relative) Se = 49%, Δ(relative) THg = 17%). These differences highlight the need to standardize the collection site for trace element determination in pinnipeds.