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Pinnipeds spend large portions of their lives at sea, submerged, or hauled-out on land, often on remote off-shore islands. This fundamentally limits access by researchers to critical parts of pinniped life history and has spurred the development and implementation of a variety of externally attached telemetry devices (ETDs) to collect information about movement patterns, physiology and ecology of marine animals when they cannot be directly observed. ETDs are less invasive and easier to apply than implanted internal devices, making them more widely used. However, ETDs have limited retention times and their use may result in negative short- and long-term consequences including capture myopathy, impacts to energetics, behavior, and entanglement risk. We identify 15 best practice recommendations for the use of ETDs with pinnipeds that address experimental justification, animal capture, tag design, tag attachment, effects assessments, preparation, and reporting. Continued improvement of best practices is critical within the framework of the Three Rs (Reduction, Refinement, Replacement); these best practice recommendations provide current guidance to mitigate known potential negative outcomes for individuals and local populations. These recommendations were developed specifically for pinnipeds; however, they may also be applicable to studies of other marine taxa. We conclude with four desired future directions for the use of ETDs in technology development, validation studies, experimental designs and data sharing.

Wildlife researchers must balance the need to safely capture and handle their study animals to sample tissues, collect morphological measurements, and attach dataloggers while ensuring their results are not confounded by stress artifacts caused by handling. To determine the physiological effects of research activities including chemical immobilization, transport, instrumentation with biologgers, and overnight holding on a model marine mammal species, we collected hormone, blood chemistry, hematology, and heart rate data from 19 juvenile northern elephant seals (Mirounga angustirostris) throughout a translocation experiment. Across our six sampling timepoints, cortisol and aldosterone data revealed a moderate hormonal stress response to handling accompanied by minor changes in hematocrit and blood glucose, but not ketone bodies or erythrocyte sedimentation rate. We also examined heart rate as a stress indicator and found that interval heart rate, standard deviation of heart rate, and apnea-eupnea cycles were influenced by handling. However, when seals were recaptured after several days at sea, all hormonal and hematological parameters had returned to baseline levels. Furthermore, 100 % of study animals were resighted in the wild post-translocation, with some individuals observed over four years later. Together, these findings suggest that while northern elephant seals exhibit measurable physiological stress in response to handling, they recover rapidly and show no observable long-term deleterious effects, making them a robust species for ecological and physiological research., Competing Interests: Declaration of competing interest The authors declare that they have no known competing financial interests or personal relationships that could have appeared to influence the work reported in this paper., (Copyright © 2024 The Authors. Published by Elsevier Inc. All rights reserved.)

Northern elephant seals (Mirounga angustirostris) have been integral to the development and progress of biologging technology and movement data analysis, which continue to improve our understanding of this and other species. Adult female elephant seals at Año Nuevo Reserve and other colonies along the west coast of North America were tracked annually from 2004 to 2020, resulting in a total of 653 instrument deployments. This paper outlines the compilation and curation process of these high-resolution diving and location data, now accessible in two Dryad repositories. The code used for data processing alongside the corresponding workflow is available through GitHub and Zenodo. This data set represents 3,844,927 dives and 596,815 locations collected from 475 individual seals with 178 repeat samplings over 17 years. We anticipate that these data will stimulate further analysis and investigation into elephant seal biology and aid in developing new analytical approaches for large marine predators., Competing Interests: Competing interests: The authors declare no competing interests., (© 2024. The Author(s).)

Despite elite human free divers achieving incredible feats in competitive free diving, there has yet to be a study that compares consummate divers, (i.e. northern elephant seals) to highly conditioned free divers (i.e., elite competitive free-diving humans). Herein, we compare these two diving models and suggest that hematological traits detected in seals reflect species-specific specializations, while hematological traits shared between the two species are fundamental mammalian characteristics. Arterial blood samples were analyzed in elite human free divers ( n = 14) during a single, maximal volitional apnea and in juvenile northern elephant seals ( n = 3) during rest-associated apnea. Humans and elephant seals had comparable apnea durations (∼6.5 min) and end-apneic arterial Po 2 [humans: 40.4 ± 3.0 mmHg (means ± SE); seals: 27.1 ± 5.9 mmHg; P = 0.2]. Despite similar increases in arterial Pco 2 (humans: 33 ± 5%; seals: 16.3 ± 5%; P = 0.2), only humans experienced reductions in pH from baseline (humans: 7.45 ± 0.01; seals: 7.39 ± 0.02) to end apnea (humans: 7.37 ± 0.01; seals: 7.38 ± 0.02; P < 0.0001). Hemoglobin P 50 was greater in humans compared to elephant seals (29.9 ± 1.5 and 28.7 ± 0.6 mmHg, respectively; P = 0.046). Elephant seals overall had higher carboxyhemoglobin (COHb) levels (5.9 ± 2.6%) compared to humans (0.8 ± 1.2%; P < 0.0001); however, following apnea, COHb was reduced in seals (baseline: 6.1 ± 0.3%; end apnea: 5.6 ± 0.3%) and was slightly elevated in humans (baseline: 0.7 ± 0.1%; end apnea: 0.9 ± 0.1%; P < 0.0002, both comparisons). Our data indicate that during static apnea, seals have reduced hemoglobin P 50 , greater pH buffering, and increased COHb levels. The differences in hemoglobin P 50 are likely due to the differences in the physiological environment between the two species during apnea, whereas enhanced pH buffering and higher COHb may represent traits selected for in elephant seals. NEW & NOTEWORTHY This study uses similar methods and protocols in elite human free divers and northern elephant seals. Using highly conditioned divers (elite free-diving humans) and highly adapted divers (northern elephant seals), we explored which hematological traits are fundamentally mammalian and which may have been selected for. We found differences in P 50 , which may be due to different physiological environments between species, while elevated pH buffering and carbon monoxide levels might have been selected for in seals.

Many large terrestrial mammalian predators use energy-intensive, high-risk, high-gain strategies to pursue large, high-quality prey. However, similar-sized marine mammal predators with even higher field metabolic rates (FMRs) consistently target prey three to six orders of magnitude smaller than themselves. Here, we address the question of how these active and expensive marine mammal predators can gain sufficient energy from consistently targeting small prey during breath-hold dives. Using harbor porpoises as model organisms, we show that hunting small aquatic prey is energetically cheap (<20% increase in FMR) for these marine predators, but it requires them to spend a large proportion (>60%) of time foraging. We conclude that this grazing foraging strategy on small prey is viable for marine mammal predators despite their high FMR because they can hunt near continuously at low marginal expense. Consequently, cessation of foraging due to human disturbance comes at a high cost, as porpoises must maintain their high thermoregulation costs with a reduced energy intake.

Deep ocean foraging northern elephant seals (Mirounga angustirostris) consume fish and squid in remote depths of the North Pacific Ocean. Contaminants bioaccumulated from prey are subsequently transferred by adult females to pups during gestation and lactation, linking pups to mercury contamination in mesopelagic food webs (200-1000 m depths). Maternal transfer of mercury to developing seal pups was related to maternal mercury contamination and was strongly correlated with maternal foraging behavior (biotelemetry and isotopes). Mercury concentrations in lanugo (hair grown in utero) were among the highest observed worldwide for young pinnipeds (geometric mean 23.01 μg/g dw, range 8.03-63.09 μg/g dw; n = 373); thus, some pups may be at an elevated risk of sub-lethal adverse health effects. Fetal mercury exposure was affected by maternal foraging geographic location and depth; mercury concentrations were highest in pups of the deepest diving, pelagic females. Moreover, pup lanugo mercury concentrations were strongly repeatable among successive pups of individual females, demonstrating relative consistency in pup mercury exposure based on maternal foraging strategies. Northern elephant seals are biosentinels of a remote deep-sea ecosystem. Our results suggest that mercury within North Pacific mesopelagic food webs may also pose an elevated risk to other mesopelagic-foraging predators and their offspring., (© 2024. This is a U.S. Government work and not under copyright protection in the US; foreign copyright protection may apply.)

Diving is central to the foraging strategies of many marine mammals and seabirds. Still, the effect of dive depth on foraging cost remains elusive because energy expenditure is difficult to measure at fine temporal scales in wild animals. We used depth and acceleration data from eight lactating California sea lions (Zalophus californianus) to model body density and investigate the effect of dive depth and tissue density on rates of energy expenditure. We calculated body density in 5 s intervals from the rate of gliding descent. We modeled body density across depth in each dive, revealing high tissue densities and diving lung volumes (DLVs). DLV increased with dive depth in four individuals. We used the buoyancy calculated from dive-specific body-density models and drag calculated from swim speed to estimate metabolic power and cost of transport in 5 s intervals during descents and ascents. Deeper dives required greater mean power for round-trip vertical transit, especially in individuals with higher tissue density. These trends likely follow from increased mean swim speed and buoyant hinderance that increasingly outweighs buoyant aid in deeper dives. This suggests that deep diving is either a 'high-cost, high-reward' strategy or an energetically expensive option to access prey when prey in shallow waters are limited, and that poor body condition may increase the energetic costs of deep diving. These results add to our mechanistic understanding of how foraging strategy and body condition affect energy expenditure in wild breath-hold divers., Competing Interests: Competing interests The authors declare no competing or financial interests., (© 2023. Published by The Company of Biologists Ltd.)

Mercury bioaccumulation from deep-ocean prey and the extreme life history strategies of adult female northern elephant seals ( Mirounga angustirostris ) provide a unique system to assess the interactive effects of mercury and stress on animal health by quantifying blood biomarkers in relation to mercury (skeletal muscle and blood mercury) and cortisol concentrations. The thyroid hormone thyroxine (tT4) and the antibody immunoglobulin E (IgE) were associated with mercury and cortisol concentrations interactively, where the magnitude and direction of the association of each biomarker with mercury or cortisol changed depending on the concentration of the other factor. For example, when cortisol concentrations were lowest, tT4 was positively related to muscle mercury, whereas tT4 had a negative relationship with muscle mercury in seals that had the highest cortisol concentrations. Additionally, we observed that two thyroid hormones, triiodothyronine (tT3) and reverse triiodothyronine (rT3), were negatively (tT3) and positively (rT3) associated with mercury concentrations and cortisol in an additive manner. As an example, tT3 concentrations in late breeding seals at the median cortisol concentration decreased by 14% across the range of observed muscle mercury concentrations. We also observed that immunoglobulin M (IgM), the pro-inflammatory cytokine IL-6 (IL-6), and a reproductive hormone, estradiol, were negatively related to muscle mercury concentrations but were not related to cortisol. Specifically, estradiol concentrations in late molting seals decreased by 50% across the range of muscle mercury concentrations. These results indicate important physiological effects of mercury on free-ranging apex marine predators and interactions between mercury bioaccumulation and extrinsic stressors. Deleterious effects on animals' abilities to maintain homeostasis (thyroid hormones), fight off pathogens and disease (innate and adaptive immune system), and successfully reproduce (endocrine system) can have significant individual- and population-level consequences.

The impressive breath-hold capabilities of marine mammals are facilitated by both enhanced O 2 stores and reductions in the rate of O 2 consumption via peripheral vasoconstriction and bradycardia, called the dive response. Many studies have focused on the extreme role of the dive response in maximizing dive duration in marine mammals, but few have addressed how these adjustments may compromise the capability to hunt, digest and thermoregulate during routine dives. Here, we use DTAGs, which record heart rate together with foraging and movement behaviour, to investigate how O 2 management is balanced between the need to dive and forage in five wild harbour porpoises that hunt thousands of small prey daily during continuous shallow diving. Dive heart rates were moderate (median minimum 47-69 bpm) and relatively stable across dive types, dive duration (0.5-3.3 min) and activity. A moderate dive response, allowing for some perfusion of peripheral tissues, may be essential for fuelling the high field metabolic rates required to maintain body temperature and support digestion during diving in these small, continuously feeding cetaceans. Thus, despite having the capacity to prolong dives via a strong dive response, for these shallow-diving cetaceans, it appears to be more efficient to maintain circulation while diving: extreme heart rate gymnastics are for deep dives and emergencies, not everyday use.

Anthropogenic noise can alter marine mammal behaviour and physiology, but little is known about cetacean cardiovascular responses to exposures, despite evidence that acoustic stressors, such as naval sonars, may lead to decompression sickness. Here, we measured heart rate and movements of two trained harbour porpoises during controlled exposure to 6-9 kHz sonar-like sweeps and 40 kHz peak-frequency noise pulses, designed to evoke acoustic startle responses. The porpoises initially responded to the sonar sweep with intensified bradycardia despite unaltered behaviour/movement, but habituated rapidly to the stimuli. In contrast, 40 kHz noise pulses consistently evoked rapid muscle flinches (indicative of startles), but no behavioural or heart rate changes. We conclude that the autonomous startle response appears decoupled from, or overridden by, cardiac regulation in diving porpoises, whereas certain novel stimuli may motivate oxygen-conserving cardiovascular measures. Such responses to sound exposure may contribute to gas mismanagement for deeper-diving cetaceans., Competing Interests: Competing interests The authors declare no competing or financial interests., (© 2021. Published by The Company of Biologists Ltd.)

The aerobic dive limit (ADL) and the hypothesis that most dives are aerobic in nature have become fundamental to the understanding of diving physiology and to the interpretation of diving behavior and foraging ecology of marine mammals and seabirds. An ADL, the dive duration associated with the onset of post-dive blood lactate accumulation, has only been documented with blood lactate analyses in five species. Applications to other species have involved behavioral estimates or use of an oxygen store / metabolic rate formula. Both approaches have limitations, but have proved useful to the evaluation of the dive behavior and ecology of many species., (Copyright © 2020. Published by Elsevier Inc.)

Southern Ocean ecosystems are under pressure from resource exploitation and climate change 1,2 . Mitigation requires the identification and protection of Areas of Ecological Significance (AESs), which have so far not been determined at the ocean-basin scale. Here, using assemblage-level tracking of marine predators, we identify AESs for this globally important region and assess current threats and protection levels. Integration of more than 4,000 tracks from 17 bird and mammal species reveals AESs around sub-Antarctic islands in the Atlantic and Indian Oceans and over the Antarctic continental shelf. Fishing pressure is disproportionately concentrated inside AESs, and climate change over the next century is predicted to impose pressure on these areas, particularly around the Antarctic continent. At present, 7.1% of the ocean south of 40°S is under formal protection, including 29% of the total AESs. The establishment and regular revision of networks of protection that encompass AESs are needed to provide long-term mitigation of growing pressures on Southern Ocean ecosystems.

The Retrospective Analysis of Antarctic Tracking Data (RAATD) is a Scientific Committee for Antarctic Research project led jointly by the Expert Groups on Birds and Marine Mammals and Antarctic Biodiversity Informatics, and endorsed by the Commission for the Conservation of Antarctic Marine Living Resources. RAATD consolidated tracking data for multiple species of Antarctic meso- and top-predators to identify Areas of Ecological Significance. These datasets and accompanying syntheses provide a greater understanding of fundamental ecosystem processes in the Southern Ocean, support modelling of predator distributions under future climate scenarios and create inputs that can be incorporated into decision making processes by management authorities. In this data paper, we present the compiled tracking data from research groups that have worked in the Antarctic since the 1990s. The data are publicly available through biodiversity.aq and the Ocean Biogeographic Information System. The archive includes tracking data from over 70 contributors across 12 national Antarctic programs, and includes data from 17 predator species, 4060 individual animals, and over 2.9 million observed locations.

The dive response, bradycardia (decreased heart rate) and peripheral vasoconstriction, is the key mechanism allowing breath-hold divers to perform long-duration dives while actively swimming and hunting prey. This response is variable and modulated by factors such as dive duration, depth, exercise and cognitive control. This study assessed the potential role of exercise and relative lung volume in the regulation of heart rate ( f H ) during dives of adult female California sea lions instrumented with electrocardiogram (ECG), depth and tri-axial acceleration data loggers. A positive relationship between activity (minimum specific acceleration) and f H throughout dives suggested increased muscle perfusion associated with exercise. However, apart from late ascent, f H during dives was still less than or equal to resting f H (on land). In addition, the activity- f H relationship was weaker in long, deep dives consistent with prioritization of blood oxygen conservation over blood oxygen delivery to muscle in those dives. Pulmonary stretch receptor reflexes may also contribute to f H regulation as f H profiles generally paralleled changes in relative lung volume, especially in shallower dives and during early descent and late ascent of deeper dives. Overall, these findings support the concept that both exercise and pulmonary stretch receptor reflexes may influence the dive response in sea lions., Competing Interests: Competing interestsThe authors declare no competing or financial interests., (© 2020. Published by The Company of Biologists Ltd.)

Pronounced dive responses through peripheral vasoconstriction and bradycardia enable prolonged apnoea in marine mammals. For most vertebrates, the dive response is initiated upon face immersion, but little is known about the physical drivers of diving and surfacing heart rate in cetaceans whose faces are always mostly submerged. Using two trained harbour porpoises instrumented with an ECG-measuring sound-and-movement tag (DTAG-3), we investigated the initiation and progression of bradycardia and tachycardia during apnoea and eupnoea for varying levels of immersion. We show that paranasal wetting drives bradycardia initiation and progression, whereas apnoea leads to dive-level bradycardia eventually, but not instantly. At the end of dives, heart rate accelerates independently of lung expansion, perhaps in anticipation of surfacing; however, full tachycardia is only engaged upon inhalation. We conclude that breathing drives surface tachycardia, whereas blowhole wetting is an important driver of bradycardia; however, anticipatory/volitional modulation can overrule such responses to sensory inputs., Competing Interests: Competing interestsThe authors declare no competing or financial interests., (© 2019. Published by The Company of Biologists Ltd.)

Reliable estimates of field metabolic rates (FMRs) in wild animals are essential for quantifying their ecological roles, as well as for evaluating fitness consequences of anthropogenic disturbances. Yet, standard methods for measuring FMR are difficult to use on free-ranging cetaceans whose FMR may deviate substantially from scaling predictions using terrestrial mammals. Harbour porpoises ( Phocoena phocoena ) are among the smallest marine mammals, and yet they live in cold, high-latitude waters where their high surface-to-volume ratio suggests high FMRs to stay warm. However, published FMR estimates of harbour porpoises are contradictory, with some studies claiming high FMRs and others concluding that the energetic requirements of porpoises resemble those of similar-sized terrestrial mammals. Here, we address this controversy using data from a combination of captive and wild porpoises to estimate the FMR of wild porpoises. We show that FMRs of harbour porpoises are up to two times greater than for similar-sized terrestrial mammals, supporting the hypothesis that small, carnivorous marine mammals in cold water have elevated FMRs. Despite the potential cost of thermoregulation in colder water, harbour porpoise FMRs are stable over seasonally changing water temperatures. Varying heat loss seems to be managed via cyclical fluctuations in energy intake, which serve to build up a blubber layer that largely offsets the extra costs of thermoregulation during winter. Such high FMRs are consistent with the recently reported high feeding rates of wild porpoises and highlight concerns about the potential impact of human activities on individual fitness and population dynamics., Competing Interests: Competing interestsThe authors declare no competing or financial interests., (© 2018. Published by The Company of Biologists Ltd.)

Long and deep dives in marine mammals are enabled by high mass-specific oxygen stores and the dive response, which reduces oxygen consumption in concert with increased peripheral vasoconstriction and a lowered heart rate during dives. Diving heart rates of pinnipeds are highly variable and modulated by many factors, such as breath holding (apnea), pressure, swimming activity, temperature and even cognitive control. However, the individual effects of these factors on diving heart rate are poorly understood because of the difficulty of parsing their relative contributions in diving pinnipeds. Here, we examined the effects of apnea and external sensory inputs as autonomic drivers of bradycardia. Specifically, we hypothesized that (1) water stimulation of facial receptors would - as is the case for terrestrial mammals - enhance the dive response, (2) increasing the facial area stimulated would lead to a more intense bradycardia, and (3) cold water would elicit a more pronounced bradycardia than warm water. Three harbor seals ( Phoca vitulina ) and a California sea lion ( Zalophus californianus ) were trained to breath hold in air and with their heads submerged in a basin with variable water level and temperature. We show that bradycardia occurs during apnea without immersion. We also demonstrate that bradycardia is strengthened by both increasing the area of facial submersion and colder water. Thus, we conclude that the initiation of the dive response in pinnipeds is more strongly related to breath holding than in terrestrial mammals, but the degree of the dive response is potentiated autonomically via stimulation of facial mechano- and thermo-receptors upon submergence., Competing Interests: Competing interestsThe authors declare no competing or financial interests., (© 2018. Published by The Company of Biologists Ltd.)

The dive response, a decrease in heart rate ( f H ) and peripheral vasoconstriction, is the key mechanism allowing breath-hold divers to perform long-duration dives. This pronounced cardiovascular response to diving has been investigated intensely in pinnipeds, but comparatively little is known for cetaceans, in particular in ecologically relevant settings. Here, we studied the dive f H response in one of the smallest cetaceans, the harbour porpoise ( Phocoena phocoena ). We used a novel multi-sensor data logger to record dive behaviour, f H , ventilations and feeding events in three trained porpoises, providing the first evaluation of cetacean f H regulation while performing a variety of natural behaviours, including prey capture. We predicted that tagged harbour porpoises would exhibit a decrease in f H in all dives, but the degree of bradycardia would be influenced by dive duration and activity, i.e. the dive f H response would be exercise modulated. In all dives, f H decreased compared with surface rates by at least 50% (mean maximum surface f H =173 beats min -1 , mean minimum dive f H =50 beats min -1 ); however, dive f H was approximately 10 beats min -1 higher in active dives as a result of a slower decrease in f H and more variable f H during pursuit of prey. We show that porpoises exhibit the typical breath-hold diver bradycardia during aerobic dives and that the f H response is modulated by exercise and dive duration; however, other variables such as expectations and individual differences are equally important in determining diving f H ., Competing Interests: Competing interestsThe authors declare no competing or financial interests., (© 2018. Published by The Company of Biologists Ltd.)

The depletion rate of the blood oxygen store, development of hypoxemia and dive capacity are dependent on the distribution and rate of blood oxygen delivery to tissues while diving. Although blood oxygen extraction by working muscle would increase the blood oxygen depletion rate in a swimming animal, there is little information on the relationship between muscle workload and blood oxygen depletion during dives. Therefore, we examined flipper stroke rate, a proxy of muscle workload, and posterior vena cava oxygen profiles in four adult female California sea lions ( Zalophus californianus ) during foraging trips at sea. Flipper stroke rate analysis revealed that sea lions minimized muscle metabolism with a stroke-glide strategy when diving, and exhibited prolonged glides during the descent of deeper dives (>100 m). During the descent phase of these deep dives, 55±21% of descent was spent gliding, with the longest glides lasting over 160 s and covering a vertical distance of 340 m. Animals also consistently glided to the surface from 15 to 25 m depth during these deeper dives. Venous hemoglobin saturation ( S O 2 ) profiles were highly variable throughout dives, with values occasionally increasing during shallow dives. The relationship between S O 2 and flipper stroke rate was weak during deeper dives, while this relationship was stronger during shallow dives. We conclude that (1) the depletion of oxygen in the posterior vena cava in deep-diving sea lions is not dependent on stroke effort, and (2) stroke-glide patterns during dives contribute to a reduction of muscle metabolic rate., Competing Interests: Competing interestsThe authors declare no competing or financial interests., (© 2017. Published by The Company of Biologists Ltd.)

Recent publications have emphasized the potential generation of morbid cardiac arrhythmias secondary to autonomic conflict in diving marine mammals. Such conflict, as typified by cardiovascular responses to cold water immersion in humans, has been proposed to result from exercise-related activation of cardiac sympathetic fibers to increase heart rate, combined with depth-related changes in parasympathetic tone to decrease heart rate. After reviewing the marine mammal literature and evaluating heart rate profiles of diving California sea lions ( Zalophus californianus ), we present an alternative interpretation of heart rate regulation that de-emphasizes the concept of autonomic conflict and the risk of morbid arrhythmias in marine mammals. We hypothesize that: (1) both the sympathetic cardiac accelerator fibers and the peripheral sympathetic vasomotor fibers are activated during dives even without exercise, and their activities are elevated at the lowest heart rates in a dive when vasoconstriction is maximal, (2) in diving animals, parasympathetic cardiac tone via the vagus nerve dominates over sympathetic cardiac tone during all phases of the dive, thus producing the bradycardia, (3) adjustment in vagal activity, which may be affected by many inputs, including exercise, is the primary regulator of heart rate and heart rate fluctuations during diving, and (4) heart beat fluctuations (benign arrhythmias) are common in marine mammals. Consistent with the literature and with these hypotheses, we believe that the generation of morbid arrhythmias because of exercise or stress during dives is unlikely in marine mammals., Competing Interests: Competing interestsThe authors declare no competing or financial interests., (© 2017. Published by The Company of Biologists Ltd.)

Heart rate and peripheral blood flow distribution are the primary determinants of the rate and pattern of oxygen store utilisation and ultimately breath-hold duration in marine endotherms. Despite this, little is known about how otariids (sea lions and fur seals) regulate heart rate (fH) while diving. We investigated dive fH in five adult female California sea lions (Zalophus californianus) during foraging trips by instrumenting them with digital electrocardiogram (ECG) loggers and time depth recorders. In all dives, dive fH (number of beats/duration; 50±9 beats min(-1)) decreased compared with surface rates (113±5 beats min(-1)), with all dives exhibiting an instantaneous fH below resting (<54 beats min(-1)) at some point during the dive. Both dive fH and minimum instantaneous fH significantly decreased with increasing dive duration. Typical instantaneous fH profiles of deep dives (>100 m) consisted of: (1) an initial rapid decline in fH resulting in the lowest instantaneous fH of the dive at the end of descent, often below 10 beats min(-1) in dives longer than 6 min in duration; (2) a slight increase in fH to ~10-40 beats min(-1) during the bottom portion of the dive; and (3) a gradual increase in fH during ascent with a rapid increase prior to surfacing. Thus, fH regulation in deep-diving sea lions is not simply a progressive bradycardia. Extreme bradycardia and the presumed associated reductions in pulmonary and peripheral blood flow during late descent of deep dives should (a) contribute to preservation of the lung oxygen store, (b) increase dependence of muscle on the myoglobin-bound oxygen store, (c) conserve the blood oxygen store and (d) help limit the absorption of nitrogen at depth. This fH profile during deep dives of sea lions may be characteristic of deep-diving marine endotherms that dive on inspiration as similar fH profiles have been recently documented in the emperor penguin, another deep diver that dives on inspiration.

Widely ranging marine predators often adopt stereotyped, energy-saving behaviours to minimize the energetic cost of transport while maximizing energy gain. Environmental and anthropogenic disturbances can disrupt energy balance by prompting avoidance behaviours that increase transport costs, thereby decreasing foraging efficiency. We examined the ability of 12 free-ranging, juvenile northern elephant seals (Mirounga angustirostris) to mitigate the effects of experimentally increased transport costs by modifying their behaviour and/or energy use in a compensatory manner. Under normal locomotion, elephant seals had low energy requirements (106.5±28.2 kJ kg(-1) day(-1)), approaching or even falling below predictions of basal requirements. Seals responded to a small increase in locomotion costs by spending more time resting between dives (149±44 s) compared with matched control treatments (102±11 s; P<0.01). Despite incurred costs, most other dive and transit behaviours were conserved across treatments, including fixed, rhythmic swimming gaits. Because of this, and because each flipper stroke had a predictable effect on total costs (P<0.001), total energy expenditure was strongly correlated with time spent at sea under both treatments (P<0.0001). These results suggest that transiting elephant seals have a limited capacity to modify their locomotory behaviour without increasing their transport costs. Based on this, we conclude that elephant seals and other ocean predators occupying similar niches may be particularly sensitive to increased transport costs incurred when avoiding unanticipated disturbances.

The management and depletion of O2 stores underlie the aerobic dive capacities of marine mammals. The California sea lion (Zalophus californianus) presumably optimizes O2 store management during all dives, but approaches its physiological limits during deep dives to greater than 300 m depth. Blood O2 comprises the largest component of total body O2 stores in adult sea lions. Therefore, we investigated venous blood O2 depletion during dives of California sea lions during maternal foraging trips to sea by: (1) recording venous partial pressure of O2 (P(O2)) profiles during dives, (2) characterizing the O2-hemoglobin (Hb) dissociation curve of sea lion Hb and (3) converting the P(O2) profiles into percent Hb saturation (S(O2)) profiles using the dissociation curve. The O2-Hb dissociation curve was typical of other pinnipeds (P50=28±2 mmHg at pH 7.4). In 43% of dives, initial venous S(O2) values were greater than 78% (estimated resting venous S(O2)), indicative of arterialization of venous blood. Blood O2 was far from depleted during routine shallow dives, with minimum venous S(O2) values routinely greater than 50%. However, in deep dives greater than 4 min in duration, venous S(O2) reached minimum values below 5% prior to the end of the dive, but then increased during the last 30-60 s of ascent. These deep dive profiles were consistent with transient venous blood O2 depletion followed by partial restoration of venous O2 through pulmonary gas exchange and peripheral blood flow during ascent. These differences in venous O2 profiles between shallow and deep dives of sea lions reflect distinct strategies of O2 store management and suggest that underlying cardiovascular responses will also differ.

Stressors associated with human activities interact in complex ways to affect marine ecosystems, yet we lack spatially explicit assessments of cumulative impacts on ecologically and economically key components such as marine predators. Here we develop a metric of cumulative utilization and impact (CUI) on marine predators by combining electronic tracking data of eight protected predator species (n=685 individuals) in the California Current Ecosystem with data on 24 anthropogenic stressors. We show significant variation in CUI with some of the highest impacts within US National Marine Sanctuaries. High variation in underlying species and cumulative impact distributions means that neither alone is sufficient for effective spatial management. Instead, comprehensive management approaches accounting for both cumulative human impacts and trade-offs among multiple stressors must be applied in planning the use of marine resources.

Lung collapse is considered the primary mechanism that limits nitrogen absorption and decreases the risk of decompression sickness in deep-diving marine mammals. Continuous arterial partial pressure of oxygen profiles in a free-diving female California sea lion (Zalophus californianus) revealed that (i) depth of lung collapse was near 225 m as evidenced by abrupt changes in during descent and ascent, (ii) depth of lung collapse was positively related to maximum dive depth, suggesting that the sea lion increased inhaled air volume in deeper dives and (iii) lung collapse at depth preserved a pulmonary oxygen reservoir that supplemented blood oxygen during ascent so that mean end-of-dive arterial was 74 ± 17 mmHg (greater than 85% haemoglobin saturation). Such information is critical to the understanding and the modelling of both nitrogen and oxygen transport in diving marine mammals.

Identifying individuals' foraging strategies is critical to understanding the ecology of a species, and can provide the means to predict possible ecological responses to environmental change. Our study combines stable isotope analysis and satellite telemetry to study the variability in individual foraging strategies of adult female southern elephant seals (Mirounga leonina). Our hypothesis is that female elephant seals from the Western Antarctica Peninsula (WAP) display individual specialization in their diets. We captured adult female elephant seals (n = 56, 2005-2009) at Livingston Island (Antarctica), and instrumented them with SMRU-CTD satellite tags. We collected blood, fur, and vibrissae samples for δ(13)C and δ(15)N analyses. The mean values for all vibrissae were -21.0 ± 0.7‰ for δ(13)C, and 10.4 ± 0.8‰, for δ(15)N. The individual variability of δ(13)C (60%) was more important than the within-individual variability (40%) in explaining the total variance observed in our data. For δ(15)N, the results showed the opposite trend, with the within-individual variability (64%) contributing more to the total variance than the individual variability (36%), likely associated with the effect that the fasting periods have on δ(15)N values. Most individuals were specialists, as inferred from the low intra-individual variability of δ(13)C values with respect to the population variability, with half the individuals utilizing 31% or less of their available niche. We found eight different foraging strategies for these animals. Female elephant seals from the WAP are a diverse group of predators with individuals utilizing only a small portion of the total available niche, with the consequent potential to expand their foraging habits to exploit other resources or environments in the Southern Ocean.

Life-history theory predicts that selection will favor optimal levels of parental effort that balance benefits of current reproduction with costs to survival and future reproduction. The optimal level of effort depends on parental traits, offspring traits, and provisioning strategy. Additionally, how these factors influence effort may differ depending on the stage of reproduction. The relative importance of maternal and offspring traits on energy allocation to offspring was investigated in known-age Antarctic fur seals Arctocephalus gazella across four stages of reproduction, using birth mass and milk-consumption measurements. Maternal traits were important during three of the four stages investigated, with larger females giving birth to larger pups and investing more in pups during perinatal and molt stages. Pup mass influenced maternal effort during the premolt stage, and provisioning strategy influenced postnatal maternal effort at all stages. Energy provided to the offspring during an attendance visit was positively related to the duration of the foraging-trip/visit cycle; however, when investment was controlled for trip/visit cycle duration, the overall rate of energy transfer was similar across trip durations. In addition to strong effects of maternal mass, pup traits affected energy allocation, suggesting that pup demand is important in determining maternal care. These findings emphasize the importance of considering state variables in life-history studies and suggest that timing of measurements of effort in species with long provisioning periods may influence conclusions and our ability to make comparisons of reproductive effort among species.

Little is known about how variation in the pattern and magnitude of parental effort influences allocation decisions in offspring. We determined the energy budget of Antarctic fur seal pups and examined the relative importance of timing of provisioning, pup traits (mass, condition, sex), and weather (wind chill and solar radiation) on allocation of energy obtained in milk by measuring milk energy intake, field metabolic rate (FMR), and growth rate in 48 Antarctic fur seal pups over three developmental stages (perinatal, premolt, and molt). The relative amount of milk energy used for growth was 59.1% ± 8.1% during the perinatal period but decreased to 23.4% ± 15.5% and 26.0% ± 13.9% during the premolt and molt. This decrease was associated with a greater amount of time spent fasting, along with an increase in pup activity while the mother was at sea foraging. Average daily milk intake, pup mass, and condition were all important in determining how much energy was available for growth, but the amount of energy obtained as milk was the single most important factor determining pup growth. While mean mass-specific FMR did not change with developmental stage (range = 1.74-1.77 mL O(2)/g/h), the factors that accounted for variation in FMR did. Weather (wind chill and solar radiation) and pup traits (mass and condition) influenced mass-specific FMR, but these impacts varied across development. This study provides information about the factors influencing how offspring allocate energy toward growth and maintenance and improves our predictions about how a changing environment may affect energy allocation in pups.

The mesopelagic zone of the northeast Pacific Ocean is an important foraging habitat for many predators, yet few studies have addressed the factors driving basin-scale predator distributions or inter-annual variability in foraging and breeding success. Understanding these processes is critical to reveal how conditions at sea cascade to population-level effects. To begin addressing these challenging questions, we collected diving, tracking, foraging success, and natality data for 297 adult female northern elephant seal migrations from 2004 to 2010. During the longer post-molting migration, individual energy gain rates were significant predictors of pregnancy. At sea, seals focused their foraging effort along a narrow band corresponding to the boundary between the sub-arctic and sub-tropical gyres. In contrast to shallow-diving predators, elephant seals target the gyre-gyre boundary throughout the year rather than follow the southward winter migration of surface features, such as the Transition Zone Chlorophyll Front. We also assessed the impact of added transit costs by studying seals at a colony near the southern extent of the species' range, 1,150 km to the south. A much larger proportion of seals foraged locally, implying plasticity in foraging strategies and possibly prey type. While these findings are derived from a single species, the results may provide insight to the foraging patterns of many other meso-pelagic predators in the northeast Pacific Ocean.

Top predators integrate resources over time and space, and depending on the particular species they represent, different components of the marine environment. The habitat utilization of top predators has been studied using electronic tags to follow their movements and foraging behavior. In addition, these tags provide information on the physical characteristics of the water column (temperature and salinity) at a scale and resolution that is coincident with the animals' behavior. In addition to data on the animals' behavior, these tags provide physical oceanographic data in regions or at times they cannot be collected using other currently available technologies. These data inform us on how these important top predators are likely to respond to climatic change, as well as about how the Southern Ocean is changing.

The range of foraging behaviors available to deep-diving, air-breathing marine vertebrates is constrained by their physiological capacity to breath-hold dive. We measured body oxygen stores (blood volume and muscle myoglobin) and diving behavior in adult female northern elephant seals, Mirounga angustirostris, to investigate age-related effects on diving performance. Blood volume averaged 74.4+/-17.0 liters in female elephant seals or 20.2+/-2.0% of body mass. Plasma volume averaged 32.2+/-7.8 liters or 8.7+/-0.7% of body mass. Absolute plasma volume and blood volume increased independently with mass and age. Hematocrit decreased weakly with mass but did not vary with age. Muscle myoglobin concentration, while higher than previously reported (7.4+/-0.7 g%), did not vary with mass or age. Pregnancy status did not influence blood volume. Mean dive duration, a proxy for physiological demand, increased as a function of how long seals had been at sea, followed by mass and hematocrit. Strong effects of female body mass (range, 218-600 kg) on dive duration, which were independent of oxygen stores, suggest that larger females had lower diving metabolic rates. A tendency for dives to exceed calculated aerobic limits occurred more frequently later in the at-sea migration. Our data suggest that individual physiological state variables and condition interact to determine breath-hold ability and that both should be considered in life-history studies of foraging behavior.

The efficiency with which mothers convert acquired energy into milk is a key determinant of the magnitude of parental investment in mammals; however, the mechanisms underlying lactation efficiency are poorly understood. Investigations on northern elephant seals have shown lactation efficiency, measured as the proportion of total energy expenditure that goes to the pup as milk, increases with age. In a cross-sectional study the physiological and behavioral determinants of lactation efficiency were investigated in eight young and seven prime (older) elephant seals by conducting behavioral observations and collecting milk, blood, and tissue on days 3 and 22 of lactation. Milk composition, circulating fatty acid and triglyceride concentrations, and mammary and blubber lipoprotein lipase activity were determined. Prime females had significantly greater percent milk fat and circulating fatty acids on day 3 than did young females, but these differences disappeared by day 22. The ability for prime females to produce higher-energy milk early in lactation may allow them to increase lactation efficiency by increasing the rate of energy transfer. In addition, prime females spent significantly more time resting. A combination of reduced activity and more rapid energy delivery likely explains the increase in lactation efficiency with age found in a previous study.

Interpolation of geolocation or Argos tracking data is a necessity for habitat use analyses of marine vertebrates. In a fluid marine environment, characterized by curvilinear structures, linearly interpolated track data are not realistic. Based on these two facts, we interpolated tracking data from albatrosses, penguins, boobies, sea lions, fur seals and elephant seals using six mathematical algorithms. Given their popularity in mathematical computing, we chose Bézier, hermite and cubic splines, in addition to a commonly used linear algorithm to interpolate data. Performance of interpolation methods was compared with different temporal resolutions representative of the less-precise geolocation and the more-precise Argos tracking techniques. Parameters from interpolated sub-sampled tracks were compared with those obtained from intact tracks. Average accuracy of the interpolated location was not affected by the interpolation method and was always within the precision of the tracking technique used. However, depending on the species tested, some curvilinear interpolation algorithms produced greater occurrences of more accurate locations, compared with the linear interpolation method. Total track lengths were consistently underestimated but were always more accurate using curvilinear interpolation than linear interpolation. Curvilinear algorithms are safe to use because accuracy, shape and length of the tracks are either not different or are slightly enhanced and because analyses always remain conservative. The choice of the curvilinear algorithm does not affect the resulting track dramatically so it should not preclude their use. We thus recommend using curvilinear interpolation techniques because of the more realistic fluid movements of animals. We also provide some guidelines for choosing an algorithm that is most likely to maximize track quality for different types of marine vertebrates.