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Over the past several decades, scientists have constructed bioenergetic models for marine mammals to assess potential population-level consequences following exposure to a disturbance, stressor, or environmental change, such as under the Population Consequences of Disturbance (pCOD) framework. The animal's metabolic rate (rate of energy expenditure) is a cornerstone for these models, yet the cryptic lifestyles of marine mammals, particularly cetaceans, have limited our ability to quantify basal (BMR) and field (FMR) metabolic rates using accepted 'gold standard' approaches (indirect calorimeter via oxygen consumption and doubly labeled water, respectively). Thus, alternate methods have been used to quantify marine mammal metabolic rates, such as extrapolating from known allometric relationships (e.g. Kleiber's mouse to elephant curve) and developing predictive relationships between energy expenditure and physiological or behavioral variables. To understand our current knowledge of marine mammal metabolic rates, we conducted a literature review (1900-2023) to quantify the magnitude and variation of metabolic rates across marine mammal groups. A compilation of data from studies using 'gold standard' methods revealed that BMR and FMR of different marine mammal species ranges from 0.2 to 3.6 and 1.1 to 6.1 x Kleiber, respectively. Mean BMR and FMR varied across taxa; for both measures odontocete levels were intermediate to higher values for otariids and lower values of phocids. Moreover, multiple intrinsic (e.g. age, sex, reproduction, molt, individual) and extrinsic (e.g. food availability, water temperature, season) factors, as well as individual behaviors (e.g. animal at water's surface or submerged, activity level, dive effort and at-sea behaviors) impact the magnitude of these rates. This review provides scientists and managers with a range of reliable metabolic rates for several marine mammal groups as well as an understanding of the factors that influence metabolism to improve the discernment for inputs into future bioenergetic models., Competing Interests: There are no conflicts of interest to report., (© The Author(s) 2023. Published by Oxford University Press and the Society for Experimental Biology.)

The demands on the locomotor muscles at birth are different for cetaceans than terrestrial mammals. Cetacean muscles do not need to support postural costs as the neonate transitions from the womb because water's buoyant force supports body weight. Rather, neonatal cetacean muscles must sustain locomotion under hypoxic conditions as the neonate accompanies its mother swimming underwater. Despite disparate demands at birth, cetaceans like terrestrial mammals require postnatal development to attain mature musculature. Neonatal cetaceans have a low proportion of muscle mass, and their locomotor muscles have lower mitochondrial density, myoglobin content (Mb), and buffering capacity than those found in the adult locomotor muscle. For example, the locomotor muscle of the neonatal bottlenose dolphin has only 10 and 65% of the Mb and buffering capacity, respectively, found in the adult locomotor muscle. The maturation period required to achieve mature Mb and buffering capacity in the locomotor muscle varies across cetacean species from 0.75 to 4 and 1.17 to 3.4 years, respectively. The truncated nursing interval of harbor porpoises and sub-ice travel of beluga whales may be drivers for faster muscle maturation in these species. Despite these postnatal changes in the locomotor muscle, ontogenetic changes in locomotor muscle fiber type seem to be rare in cetaceans. Regardless, the underdeveloped aerobic and anaerobic capacities of the locomotor muscle of immature dolphins result in diminished thrusting capability and swim performance. Size-specific stroke amplitudes (23-26% of body length) of 0-3-month-old dolphins are significantly smaller than those of >10-month-olds (29-30% of body length), and 0-1-month-olds only achieve 37 and 52% of the mean and maximum swim speed of adults, respectively. Until swim performance improves with muscle maturation, young cetaceans are precluded from achieving their pod's swim speeds, which could have demographic consequences when fleeing anthropogenic disturbances., (© The Author(s) 2023. Published by Oxford University Press on behalf of the Society for Integrative and Comparative Biology.)

Some marine birds and mammals can perform dives of extraordinary duration and depth. Such dive performance is dependent on many factors, including total body oxygen (O 2 ) stores. For diving penguins, the respiratory system (air sacs and lungs) constitutes 30-50% of the total body O 2 store. To better understand the role and mechanism of parabronchial ventilation and O 2 utilization in penguins both on the surface and during the dive, we examined air sac partial pressures of O 2 ( P O 2 ) in emperor penguins ( Aptenodytes forsteri ) equipped with backpack P O 2  recorders. Cervical air sac P O 2  values at rest were lower than in other birds, while the cervical air sac to posterior thoracic air sac P O 2  difference was larger. Pre-dive cervical air sac P depletion rate for dives up to 11 min was approximately one-tenth that of the posterior air sacs. Low cervical air sac O 2  values were often greater than those at rest, but had a wide range and were not significantly different from those at rest. The maximum respiratory O 2 store and total body O 2 stores calculated with representative anterior and posterior air sac P O 2  data did not differ from prior estimates. The mean calculated anterior air sac O 2 depletion rate for dives up to 11 min was approximately one-tenth that of the posterior air sacs. Low cervical air sac P O 2  values at rest may be secondary to a low ratio of parabronchial ventilation to parabronchial blood O 2 extraction. During dives, overlap of simultaneously recorded cervical and posterior thoracic air sac P O 2  profiles supported the concept of maintenance of parabronchial ventilation during a dive by air movement through the lungs., Competing Interests: Competing interestsThe authors declare no competing or financial interests., (© 2021. Published by The Company of Biologists Ltd.)

The narwhal ( Monodon monoceros ) is a high-Arctic species inhabiting areas that are experiencing increases in sea temperatures, which together with reduction in sea ice are expected to modify the niches of several Arctic marine apex predators. The Scoresby Sound fjord complex in East Greenland is the summer residence for an isolated population of narwhals. The movements of 12 whales instrumented with Fastloc-GPS transmitters were studied during summer in Scoresby Sound and at their offshore winter ground in 2017-2019. An additional four narwhals provided detailed hydrographic profiles on both summer and winter grounds. Data on diving of the whales were obtained from 20 satellite-linked time-depth recorders and 16 Acousonde™ recorders that also provided information on the temperature and depth of buzzes. In summer, the foraging whales targeted depths between 300 and 850 m where the preferred areas visited by the whales had temperatures ranging between 0.6 and 1.5°C (mean = 1.1°C, SD  = 0.22). The highest probability of buzzing activity during summer was at a temperature of 0.7°C and at depths > 300 m. The whales targeted similar depths at their offshore winter ground where the temperature was slightly higher (range: 0.7-1.7°C, mean = 1.3°C, SD  = 0.29). Both the probability of buzzing events and the spatial distribution of the whales in both seasons demonstrated a preferential selection of cold water. This was particularly pronounced in winter where cold coastal water was selected and warm Atlantic water farther offshore was avoided. It is unknown if the small temperature niche of whales while feeding is because prey is concentrated at these temperature gradients and is easier to capture at low temperatures, or because there are limitations in the thermoregulation of the whales. In any case, the small niche requirements together with their strong site fidelity emphasize the sensitivity of narwhals to changes in the thermal characteristics of their habitats., Competing Interests: There are no competing interests., (© 2020 The Authors. Ecology and Evolution published by John Wiley & Sons Ltd.)

The significance of symbioses between eukaryotic hosts and microbes extends from the organismal to the ecosystem level and underpins the health of Earth's most threatened marine ecosystems. Despite rapid growth in research on host-associated microbes, from individual microbial symbionts to host-associated consortia of significantly relevant taxa, little is known about their interactions with the vast majority of marine host species. We outline research priorities to strengthen our current knowledge of host-microbiome interactions and how they shape marine ecosystems. We argue that such advances in research will help predict responses of species, communities, and ecosystems to stressors driven by human activity and inform future management strategies., Competing Interests: The authors have declared that no competing interests exist.

When predators are removed or suppressed for generations, prey populations tend to increase and when predators are re-introduced, prey densities should fall back to pre-control levels. In cases of apparent competition where there are alternate abundant and rare prey species, rare species may decline further than expected or disappear altogether. Recently, concern about the impact of recovering predator populations on wildlife in Laikipia County, Kenya, has led to questions of whether lions (Panthera leo, IUCN Red List Vulnerable) exert top-down pressure on Grevy's zebra (Equus grevyi, IUCN Red List Endangered). We examined effects of lion predation on Plain's zebra (E. quagga, IUCN Red List Near Threatened) and Grevy's zebra populations in a 2,105 km2 area defined by lion movements. We used line transect surveys to estimate density of Grevy's (0.71/km2) and Plain's (15.9/km2) zebras, and satellite telemetry to measure movements for lions and both zebras. We tracked lions to potential feeding sites to estimate predation rates on zebras. We compared field-based estimates of predation rates on both zebras to random gas models of encounters that result in predation to ask if lions prey preferentially on Grevy's zebra at a sufficient rate to drive population declines. Lions preyed on Grevy's zebra significantly less than expected in 15 of 16 (94%) scenarios considered and lions preyed on Plain's zebras as expected or significantly less than expected in 15 of 16 scenarios. Population trend of Grevy's zebra indicates that the Kenya population may be stabilizing. Recruitment rate to the population has tripled since 2004, making it unlikely that lions are having an impact on Grevy's zebras. In Laikipia County, competitive displacement by livestock (Livestock: Grevy's zebra ratio = 864:1) and interference competition for grass with Plain's zebra (Plain's zebra:Grevy's zebra ratio = 22:1) are most likely the predominant threats to Grevy's Zebra recovery., Competing Interests: Australian 60 Minutes is a commercial funder that provided partial unrestricted financial support to LGF. This support does not alter our adherence to PLOS ONE policies on sharing data and materials. Australian 60 Minutes had no role in study design, data collection and analysis, decision to publish, or preparation of the manuscript.

Humans strongly impact the dynamics of coastal systems, yet surprisingly few studies mechanistically link management of anthropogenic stressors and successful restoration of nearshore habitats over large spatial and temporal scales. Such examples are sorely needed to ensure the success of ecosystem restoration efforts worldwide. Here, we unite 30 consecutive years of watershed modeling, biogeochemical data, and comprehensive aerial surveys of Chesapeake Bay, United States to quantify the cascading effects of anthropogenic impacts on submersed aquatic vegetation (SAV), an ecologically and economically valuable habitat. We employ structural equation models to link land use change to higher nutrient loads, which in turn reduce SAV cover through multiple, independent pathways. We also show through our models that high biodiversity of SAV consistently promotes cover, an unexpected finding that corroborates emerging evidence from other terrestrial and marine systems. Due to sustained management actions that have reduced nitrogen concentrations in Chesapeake Bay by 23% since 1984, SAV has regained 17,000 ha to achieve its highest cover in almost half a century. Our study empirically demonstrates that nutrient reductions and biodiversity conservation are effective strategies to aid the successful recovery of degraded systems at regional scales, a finding which is highly relevant to the utility of environmental management programs worldwide., Competing Interests: The authors declare no conflict of interest.

Little is known about the foraging behavior of top predators in the deep mesopelagic ocean. Elephant seals dive to the deep biota-poor oxygen minimum zone (OMZ) (>800 m depth) despite high diving costs in terms of energy and time, but how they successfully forage in the OMZ remains largely unknown. Assessment of their feeding rate is the key to understanding their foraging behavior, but this has been challenging. Here, we assessed the feeding rate of 14 female northern elephant seals determined by jaw motion events (JME) and dive cycle time to examine how feeding rates varied with dive depth, particularly in the OMZ. We also obtained video footage from seal-mounted videos to understand their feeding in the OMZ. While the diel vertical migration pattern was apparent for most depths of the JME, some very deep dives, beyond the normal diel depth ranges, occurred episodically during daylight hours. The midmesopelagic zone was the main foraging zone for all seals. Larger seals tended to show smaller numbers of JME and lower feeding rates than smaller seals during migration, suggesting that larger seals tended to feed on larger prey to satisfy their metabolic needs. Larger seals also dived frequently to the deep OMZ, possibly because of a greater diving ability than smaller seals, suggesting their dependency on food in the deeper depth zones. Video observations showed that seals encountered the rarely reported ragfish ( Icosteus aenigmaticus ) in the depths of the OMZ, which failed to show an escape response from the seals, suggesting that low oxygen concentrations might reduce prey mobility. Less mobile prey in OMZ would enhance the efficiency of foraging in this zone, especially for large seals that can dive deeper and longer. We suggest that the OMZ plays an important role in structuring the mesopelagic ecosystem and for the survival and evolution of elephant seals.

The length of time required for postnatal maturation of the locomotor muscle (longissimus dorsi) biochemistry [myoglobin (Mb) content and buffering capacity] in marine mammals typically varies with nursing duration, but it can be accelerated by species-specific behavioral demands, such as deep-diving and sub-ice transit. We examined how the swimming demands of a pelagic lifestyle influence postnatal maturation of Mb and buffering capacity in spinner dolphins ( Stenella longirostris longirostris ). Mb content of newborn (1.16±0.07 g Mb per 100 g wet muscle mass, n =6) and juvenile (2.77±0.22 g per 100 g, n =4) spinner dolphins were only 19% and 46% of adult levels (6.00±0.74 g per 100 g, n =6), respectively. At birth, buffering capacity was 52.70±4.48 slykes ( n =6) and increased to 78.53±1.91 slykes ( n =6) once a body length of 141 cm was achieved, representing 1.6- to 2.0-year-old dolphins. Based on the age of weaning (1.3-1.6 years post-partum), muscle maturation occurred just after weaning as described for coastal bottlenose dolphins ( Tursiops truncatus ). Thus, a pelagic lifestyle does not promote rapid maturation of muscle biochemistry. Rather, it promotes enhanced muscle biochemistry: newborn and adult spinner dolphins had four- and two-times greater Mb contents than newborn and adult bottlenose dolphins, respectively. Indeed, adult levels rivaled those of deep-diving cetaceans. Nonetheless, the relatively underdeveloped muscle biochemistry of calves likely contributes to documented mother-calf separations for spinner dolphins chased by the tuna purse-seine fishery., Competing Interests: Competing interestsThe authors declare no competing or financial interests., (© 2017. Published by The Company of Biologists Ltd.)

Exponential increases in hydrodynamic drag and physical exertion occur when swimmers move quickly through water, and underlie the preference for relatively slow routine speeds by marine mammals regardless of body size. Because of this and the need to balance limited oxygen stores when submerged, flight (escape) responses may be especially challenging for this group. To examine this, we used open-flow respirometry to measure the energetic cost of producing a swimming stroke during different levels of exercise in bottlenose dolphins ( Tursiops truncatus ). These data were then used to model the energetic cost of high-speed escape responses by other odontocetes ranging in mass from 42 to 2738 kg. The total cost per stroke during routine swimming by dolphins, 3.31±0.20 J kg -1  stroke -1 , was doubled during maximal aerobic performance. A comparative analysis of locomotor costs (LC; in J kg -1  stroke -1 ), representing the cost of moving the flukes, revealed that LC during routine swimming increased with body mass ( M ) for odontocetes according to LC=1.46±0.0005 M ; a separate relationship described LC during high-speed stroking. Using these relationships, we found that continuous stroking coupled with reduced glide time in response to oceanic noise resulted in a 30.5% increase in metabolic rate in the beaked whale, a deep-diving odontocete considered especially sensitive to disturbance. By integrating energetics with swimming behavior and dive characteristics, this study demonstrates the physiological consequences of oceanic noise on diving mammals, and provides a powerful tool for predicting the biological significance of escape responses by cetaceans facing anthropogenic disturbances., (© 2017. Published by The Company of Biologists Ltd.)

Little is known about the postnatal development of the physiological characteristics that support breath-hold in cetaceans, despite their need to swim and dive at birth. Arctic species have the additional demand of avoiding entrapment while navigating under sea ice, where breathing holes are patchily distributed and ephemeral. This is the first investigation of the ontogeny of the biochemistry of the locomotor muscle in a year-round Arctic-dwelling cetacean (beluga whale, Delphinapterus leucas). Compared with what we know about other cetaceans, belugas are born with high myoglobin content (1.56±0.02 g 100 g -1 wet muscle mass, N=2) that matures rapidly. Myoglobin increased by 452% during the first year after birth and achieved adult levels (6.91±0.35 g 100 g -1 wet muscle mass, N=9) by 14 months postpartum. Buffering capacity was 48.88±0.69 slykes (N=2) at birth; adult levels (84.31±1.38 slykes, N=9) were also achieved by 14 months postpartum. As the oxygen stores matured, calculated aerobic dive limit more than doubled over the first year of life, undoubtedly facilitating the movements of calves under sea ice. Nonetheless, small body size theoretically continues to constrain the diving ability of newly weaned 2 year olds, as they only had 74% and 69% of the aerobic breath-hold capacity of larger adult female and male counterparts. These assessments enhance our knowledge of the biology of cetaceans and provide insight into age-specific flexibility to alter underwater behaviors, as may be required with the ongoing alterations in the Arctic marine ecosystem associated with climate change and increased anthropogenic activities., (© 2016. Published by The Company of Biologists Ltd.)

Foraging by large (>25 kg), mammalian carnivores often entails cryptic tactics to surreptitiously locate and overcome highly mobile prey. Many forms of intermittent locomotion from stroke-and-glide maneuvers by marine mammals to sneak-and-pounce behaviors by terrestrial canids, ursids, and felids are involved. While affording proximity to vigilant prey, these tactics are also associated with unique energetic costs and benefits to the predator. We examined the energetic consequences of intermittent locomotion in mammalian carnivores and assessed the role of these behaviors in overall foraging efficiency. Behaviorally-linked, three-axis accelerometers were calibrated to provide instantaneous locomotor behaviors and associated energetic costs for wild adult Weddell seals (Leptonychotes weddellii) diving beneath the Antarctic ice. The results were compared with previously published values for other marine and terrestrial carnivores. We found that intermittent locomotion in the form of extended glides, burst-and-glide swimming, and rollercoaster maneuvers while hunting silverfish (Pleuragramma antarcticum) resulted in a marked energetic savings for the diving seals relative to continuously stroking. The cost of a foraging dive by the seals decreased by 9.2-59.6%, depending on the proportion of time gliding. These energetic savings translated into exceptionally low transport costs during hunting (COTHUNT) for diving mammals. COTHUNT for Weddell seals was nearly six times lower than predicted for large terrestrial carnivores, and demonstrates the importance of turning off the propulsive machinery to facilitate cost-efficient foraging in highly active, air-breathing marine predators., (© The Author 2015. Published by Oxford University Press on behalf of the Society for Integrative and Comparative Biology. All rights reserved. For permissions please email: journals.permissions@oup.com.)

Physiological constraints dictate animals' ability to exploit habitats. For marine mammals, it is important to quantify physiological limits that influence diving and their ability to alter foraging behaviors. We characterized age-specific dive limits of walruses by measuring anaerobic (acid-buffering capacity) and aerobic (myoglobin content) capacities of the muscles that power hind (longissimus dorsi) and fore (supraspinatus) flipper propulsion. Mean buffering capacities were similar across muscles and age classes (a fetus, five neonatal calves, a 3 month old and 20 adults), ranging from 41.31 to 54.14 slykes and 42.00 to 46.93 slykes in the longissimus and supraspinatus, respectively. Mean myoglobin in the fetus and neonatal calves fell within a narrow range (longissimus: 0.92-1.68 g 100 g(-1) wet muscle mass; supraspinatus: 0.88-1.64 g 100 g(-1) wet muscle mass). By 3 months post-partum, myoglobin in the longissimus increased by 79%, but levels in the supraspinatus remained unaltered. From 3 months post-partum to adulthood, myoglobin increased by an additional 26% in the longissimus and increased by 126% in the supraspinatus; myoglobin remained greater in the longissimus compared with the supraspinatus. Walruses are unique among marine mammals because they are born with a mature muscle acid-buffering capacity and attain mature myoglobin content early in life. Despite rapid physiological development, small body size limits the diving capacity of immature walruses and extreme sexual dimorphism reduces the diving capacity of adult females compared with adult males. Thus, free-ranging immature walruses likely exhibit the shortest foraging dives while adult males are capable of the longest foraging dives., (© 2015. Published by The Company of Biologists Ltd.)

Background: The energy requirements of free-ranging marine mammals are challenging to measure due to cryptic and far-ranging feeding habits, but are important to quantify given the potential impacts of high-level predators on ecosystems. Given their large body size and carnivorous lifestyle, we would predict that northern elephant seals (Mirounga angustirostris) have elevated field metabolic rates (FMRs) that require high prey intake rates, especially during pregnancy. Disturbance associated with climate change or human activity is predicted to further elevate energy requirements due to an increase in locomotor costs required to accommodate a reduction in prey or time available to forage. In this study, we determined the FMRs, total energy requirements, and energy budgets of adult, female northern elephant seals. We also examined the impact of increased locomotor costs on foraging success in this species., Results: Body size, time spent at sea and reproductive status strongly influenced FMR. During the short foraging migration, FMR averaged 90.1 (SE = 1.7) kJ kg(-1)d(-1) - only 36 % greater than predicted basal metabolic rate. During the long migration, when seals were pregnant, FMRs averaged 69.4 (±3.0) kJ kg(-1)d(-1) - values approaching those predicted to be necessary to support basal metabolism in mammals of this size. Low FMRs in pregnant seals were driven by hypometabolism coupled with a positive feedback loop between improving body condition and reduced flipper stroking frequency. In contrast, three additional seals carrying large, non-streamlined instrumentation saw a four-fold increase in energy partitioned toward locomotion, resulting in elevated FMRs and only half the mass gain of normally-swimming study animals., Conclusions: These results highlight the importance of keeping locomotion costs low for successful foraging in this species. In preparation for lactation and two fasting periods with high demands on energy reserves, migrating elephant seals utilize an economical foraging strategy whereby energy savings from reduced locomotion costs are shuttled towards somatic growth and fetal gestation. Remarkably, the energy requirements of this species, particularly during pregnancy, are 70-80 % lower than expected for mammalian carnivores, approaching or even falling below values predicted to be necessary to support basal metabolism in mammals of this size.

Use of complete mitochondrial genomes (mitogenomes) can greatly increase the resolution achievable in phylogeographic and historical demographic studies. Using next-generation sequencing methods, it is now feasible to efficiently sequence mitogenomes of large numbers of individuals once a reference mitogenome is available. However, assembling the initial mitogenomes of nonmodel organisms can present challenges, for example, in birds, where mtDNA is often subject to gene rearrangements and duplications. We developed a workflow based on Illumina paired-end, whole-genome shotgun sequencing, which we used to generate complete 19-kilobase mitogenomes for each of three species of North Pacific albatross, a group of birds known to carry a tandem duplication. Although this duplication had been described previously, our procedure did not depend on this prior knowledge, nor did it require a closely related reference mitogenome (e.g. a mammalian mitogenome was sufficient). We employed an iterative process including de novo assembly, reference-guided assembly and gap closing, which enabled us to detect duplications, determine gene order and identify sequence for primer positioning to resolve any mitogenome ambiguity (via minimal targeted Sanger sequencing). We present full mtDNA annotations, including 22 tRNAs, 2 rRNAs, 13 protein-coding genes, a control region and a duplicated feature for all three species. Pairwise comparisons supported previous hypotheses regarding the phylogenetic relationships within this group and occurrence of a shared tandem duplication. The resulting mitogenome sequences will enable rapid, high-throughput NGS mitogenome sequencing of North Pacific albatrosses via direct reference-guided assembly. Moreover, our approach to assembling mitogenomes should be applicable to any taxon., (© 2014 John Wiley & Sons Ltd.)

Globally, small-scale fisheries are influenced by dynamic climate, governance, and market drivers, which present social and ecological challenges and opportunities. It is difficult to manage fisheries adaptively for fluctuating drivers, except to allow participants to shift effort among multiple fisheries. Adapting to changing conditions allows small-scale fishery participants to survive economic and environmental disturbances and benefit from optimal conditions. This study explores the relative influence of large-scale drivers on shifts in effort and outcomes among three closely linked fisheries in Monterey Bay since the Magnuson-Stevens Fisheries Conservation and Management Act of 1976. In this region, Pacific sardine (Sardinops sagax), northern anchovy (Engraulis mordax), and market squid (Loligo opalescens) fisheries comprise a tightly linked system where shifting focus among fisheries is a key element to adaptive capacity and reduced social and ecological vulnerability. Using a cluster analysis of landings, we identify four modes from 1974 to 2012 that are dominated (i.e., a given species accounting for the plurality of landings) by squid, sardine, anchovy, or lack any dominance, and seven points of transition among these periods. This approach enables us to determine which drivers are associated with each mode and each transition. Overall, we show that market and climate drivers are predominantly attributed to dominance transitions. Model selection of external drivers indicates that governance phases, reflected as perceived abundance, dictate long-term outcomes. Our findings suggest that globally, small-scale fishery managers should consider enabling shifts in effort among fisheries and retaining existing flexibility, as adaptive capacity is a critical determinant for social and ecological resilience.

Within a heterogeneous environment, animals must efficiently locate and utilise foraging patches. One way animals can achieve this is by increasing residency times in areas where foraging success is highest (area-restricted search). For air-breathing diving predators, increased patch residency times can be achieved by altering both surface movements and diving patterns. The current study aimed to spatially identify the areas where female Australian fur seals allocated the most foraging effort, while simultaneously determining the behavioural changes that occur when they increase their foraging intensity. To achieve this, foraging behaviour was successfully recorded with a FastLoc GPS logger and dive behaviour recorder from 29 individual females provisioning pups. Females travelled an average of 118 ± 50 km from their colony during foraging trips that lasted 7.3 ± 3.4 days. Comparison of two methods for calculating foraging intensity (first-passage time and first-passage time modified to include diving behaviour) determined that, due to extended surface intervals where individuals did not travel, inclusion of diving behaviour into foraging analyses was important for this species. Foraging intensity 'hot spots' were found to exist in a mosaic of patches within the Bass Basin, primarily to the south-west of the colony. However, the composition of benthic habitat being targeted remains unclear. When increasing their foraging intensity, individuals tended to perform dives around 148 s or greater, with descent/ascent rates of approximately 1.9 m•s-1 or greater and reduced postdive durations. This suggests individuals were maximising their time within the benthic foraging zone. Furthermore, individuals increased tortuosity and decreased travel speeds while at the surface to maximise their time within a foraging location. These results suggest Australian fur seals will modify both surface movements and diving behaviour to maximise their time within a foraging patch.

Unlike their terrestrial ancestors, marine mammals routinely confront extreme physiological and physical challenges while breath-holding and pursuing prey at depth. To determine how cetaceans and pinnipeds accomplish deep-sea chases, we deployed animal-borne instruments that recorded high-resolution electrocardiograms, behaviour and flipper accelerations of bottlenose dolphins (Tursiops truncatus) and Weddell seals (Leptonychotes weddellii) diving from the surface to >200 m. Here we report that both exercise and depth alter the bradycardia associated with the dive response, with the greatest impacts at depths inducing lung collapse. Unexpectedly, cardiac arrhythmias occurred in >73% of deep, aerobic dives, which we attribute to the interplay between sympathetic and parasympathetic drivers for exercise and diving, respectively. Such marked cardiac variability alters the common view of a stereotypic 'dive reflex' in diving mammals. It also suggests the persistence of ancestral terrestrial traits in cardiac function that may help explain the unique sensitivity of some deep-diving marine mammals to anthropogenic disturbances.

Background: Phylogenetic and population genetic studies often deal with multiple sequence alignments that require manipulation or processing steps such as sequence concatenation, sequence renaming, sequence translation or consensus sequence generation. In recent years phylogenetic data sets have expanded from single genes to genome wide markers comprising hundreds to thousands of loci. Processing of these large phylogenomic data sets is impracticable without using automated process pipelines. Currently no stand-alone or pipeline compatible program exists that offers a broad range of manipulation and processing steps for multiple sequence alignments in a single process run., Results: Here we present FASconCAT-G, a system independent editor, which offers various processing options for multiple sequence alignments. The software provides a wide range of possibilities to edit and concatenate multiple nucleotide, amino acid, and structure sequence alignment files for phylogenetic and population genetic purposes. The main options include sequence renaming, file format conversion, sequence translation between nucleotide and amino acid states, consensus generation of specific sequence blocks, sequence concatenation, model selection of amino acid replacement with ProtTest, two types of RY coding as well as site exclusions and extraction of parsimony informative sites. Convieniently, most options can be invoked in combination and performed during a single process run. Additionally, FASconCAT-G prints useful information regarding alignment characteristics and editing processes such as base compositions of single in- and outfiles, sequence areas in a concatenated supermatrix, as well as paired stem and loop regions in secondary structure sequence strings., Conclusions: FASconCAT-G is a command-line driven Perl program that delivers computationally fast and user-friendly processing of multiple sequence alignments for phylogenetic and population genetic applications and is well suited for incorporation into analysis pipelines.

Although wildlife conservation actions have increased globally in number and complexity, the lack of scalable, cost-effective monitoring methods limits adaptive management and the evaluation of conservation efficacy. Automated sensors and computer-aided analyses provide a scalable and increasingly cost-effective tool for conservation monitoring. A key assumption of automated acoustic monitoring of birds is that measures of acoustic activity at colony sites are correlated with the relative abundance of nesting birds. We tested this assumption for nesting Forster's terns (Sterna forsteri) in San Francisco Bay for 2 breeding seasons. Sensors recorded ambient sound at 7 colonies that had 15-111 nests in 2009 and 2010. Colonies were spaced at least 250 m apart and ranged from 36 to 2,571 m(2) . We used spectrogram cross-correlation to automate the detection of tern calls from recordings. We calculated mean seasonal call rate and compared it with mean active nest count at each colony. Acoustic activity explained 71% of the variation in nest abundance between breeding sites and 88% of the change in colony size between years. These results validate a primary assumption of acoustic indices; that is, for terns, acoustic activity is correlated to relative abundance, a fundamental step toward designing rigorous and scalable acoustic monitoring programs to measure the effectiveness of conservation actions for colonial birds and other acoustically active wildlife., (© 2014 Society for Conservation Biology.)

Sea otters (Enhydra lutris) have the highest mass-specific metabolic rate of any marine mammal, which is superimposed on the inherently high costs of reproduction and lactation in adult females. These combined energetic demands have been implicated in the poor body condition and increased mortality of female sea otters nearing the end of lactation along the central California coast. However, the cost of lactation is unknown and currently cannot be directly measured for this marine species in the wild. Here, we quantified the energetic demands of immature sea otters across five developmental stages as a means of assessing the underlying energetic challenges associated with pup rearing that may contribute to poor maternal condition. Activity-specific metabolic rates, daily activity budgets and field metabolic rates (FMR) were determined for each developmental stage. Mean FMR of pre-molt pups was 2.29 ± 0.81 MJ day(-1) and increased to 6.16 ± 2.46 and 7.41 ± 3.17 MJ day(-1) in post-molt pups and dependent immature animals, respectively. Consequently, daily energy demands of adult females increase 17% by 3 weeks postpartum and continue increasing to 96% above pre-pregnancy levels by the average age of weaning. Our results suggest that the energetics of pup rearing superimposed on small body size, marine living and limited on-board energetic reserves conspire to make female sea otters exceptionally vulnerable to energetic shortfalls. By controlling individual fitness, maternal behavior and pup provisioning strategies, this underlying metabolic challenge appears to be a major factor influencing current population trends in southern sea otters (Enhydra lutris nereis)., (© 2014. Published by The Company of Biologists Ltd.)

A hallmark of the dive response, bradycardia, promotes the conservation of onboard oxygen stores and enables marine mammals to submerge for prolonged periods. A paradox exists when marine mammals are foraging underwater because activity should promote an elevation in heart rate (f(H)) to support increased metabolic demands. To assess the effect of the interaction between the diving response and underwater activity on f(H), we integrated interbeat f(H) with behavioral observations of adult bottlenose dolphins diving and swimming along the coast of the Bahamas. As expected for the dive response, f(H) while resting during submergence (40±6 beats min(-1)) was significantly lower than f(H) while resting at the water surface (105±8 beats min(-1)). The maximum recorded f(H) (f(H,max)) was 128±7 beats min(-1), and occurred during post-dive surface intervals. During submergence, the level of bradycardia was modified by activity. Behaviors such as simple head bobbing at depth increased f(H) by 40% from submerged resting levels. Higher heart rates were observed for horizontal swimming at depth. Indeed, the dolphins operated at 37-58% of their f(H,max) while active at depth and approached 57-79% of their f(H,max) during anticipatory tachycardia as the animals glided to the surface. f(H) was significantly correlated with stroke frequency (range=0-2.5 strokes s(-1), r=0.88, N=25 dives) and calculated swim speed (range=0-5.4 m s(-1), r=0.88, N=25 dives). We find that rather than a static reflex, the dive response is modulated by behavior and exercise in a predictable manner.

Survival depends on an animal's ability to find and acquire prey. In diving vertebrates, this ability is directly related to their physiological capability (e.g. oxygen stores). We studied the seasonal variation in oxygen stores, body temperature and body condition in California sea lions (Zalophus californianus) (CSL) as a function of seasonal variation in temperature, primary productivity, diving behavior and reproductive stage. During summer, blood oxygen stores were significantly greater and muscle oxygen stores were significantly lower than in winter. Total oxygen stores, body condition and body temperature did not change between seasons but variations in body temperature were greater during summer. Changes in oxygen stores are partly attributed to diving behavior, temperature and pregnancy that could increase oxygen consumption. Blood and muscle oxygen stores appear to be influenced by reproductive state. Blood oxygen stores are more likely influenced by diving behavior and temperature than muscle oxygen stores., (Copyright © 2012 Elsevier Inc. All rights reserved.)

Odontocetes have an exceptional range in body mass spanning 10(3) kg across species. Because, size influences oxygen utilization and carbon dioxide production rates in mammals, this lineage likely displays an extraordinary variation in oxygen store management compared to other marine mammal groups. To examine this, we measured changes in the partial pressures of respiratory gases ([Formula: see text], [Formula: see text]), pH, and lactate in the blood during voluntary, quiescent, submerged breath holds in Pacific white-sided dolphins (Lagenorhynchus obliquidens), bottlenose dolphins (Tursiops truncatus), and a killer whale (Orcinus orca) representing a mass range of 96-3,850 kg. These measurements provided an empirical determination of the effect of body size on the variability in blood biochemistry during breath hold and experimentally determined aerobic dive limits (ADL) within one taxonomic group (odontocetes). For the species in this study, maximum voluntary breath-hold duration was positively correlated with body mass, ranging from 3.5 min in white-sided dolphins to 13.3 min for the killer whale. Variation in breath-hold duration was associated with differences in the rate of change for [Formula: see text] throughout breath hold; [Formula: see text] decreased twice as fast for the two smaller species (-0.6 mmHg O(2) min(-1)) compared to the largest species (-0.3 mmHg O(2) min(-1)). In contrast, the rate of increase in [Formula: see text] during breath hold was similar across species. These results demonstrate that large body size in odontocetes facilitates increased aerobic breath-hold capacity as mediated by decreased mass-specific metabolic rates (rates of change in [Formula: see text] served as a proxy for oxygen utilization). Indeed the experimentally determined 5 min ADL for bottlenose dolphins was surpassed by the 13.3 min maximum breath hold of the killer whale, which did not end in a rise in lactate. Rather, breath hold ended voluntarily as respiratory gases and pH fell within a narrow range for both large and small species, likely providing cues for ventilation.

Constraints on locomotion could be an important component of the cost of reproduction as carrying an increased load associated with eggs or developing fetuses may contribute to decreased locomotor performance for females across taxa and environments. Diminished performance could increase susceptibility to predation, yet the mechanism(s) by which gravidity and pregnancy affect locomotion remains largely unexplored. Here we demonstrate that morphology, hydrodynamics and kinematics were altered during pregnancy, providing a mechanism for diminished locomotor performance in two near-term pregnant (10 days pre-parturition) bottlenose dolphins (Tursiops truncatus). Near-term pregnancy resulted in a 56 ± 13% [corrected] increase in frontal surface area, coinciding with dramatic increases in drag forces while gliding. For example, pregnant females encountered 80 N of drag at 1.7 m s(-1) whereas that magnitude of drag was not encountered until speed doubled for females 18 months post-parturition. Indeed, drag coefficients based on frontal surface area were significantly greater during pregnancy (C(d,F)=0.22 ± 0.04) than at 18 months post-parturition (C(d,F)=0.09 ± 0.01). Pregnancy also induced a gait change as stroke amplitude and distance per stroke were reduced by 13 and 14%, respectively, compared with non-pregnant periods (1-24 months post-parturition). This was concomitant with a 62 and 44% reduction in mean and maximum swim speeds, respectively, during the pregnancy period. Interestingly, attack speeds of known predators of dolphins surpass maximum speeds for the pregnant dolphins in this study. Thus, pregnant dolphins may be more susceptible to predation. This study demonstrates unequivocally that changes in morphology, hydrodynamics and kinematics are associated with diminished performance during pregnancy in dolphins.

Recovery criteria for depleted species or populations normally are based on demographic measures, the goal being to maintain enough individuals over a sufficiently large area to assure a socially tolerable risk of future extinction. Such demographically based recovery criteria may be insufficient to restore the functional roles of strongly interacting species. We explored the idea of developing a recovery criterion for sea otters (Enhydra lutris) in the Aleutian archipelago on the basis of their keystone role in kelp forest ecosystems. We surveyed sea otters and rocky reef habitats at 34 island-time combinations. The system nearly always existed in either a kelp-dominated or deforested phase state, which was predictable from sea otter density. We used a resampling analysis of these data to show that the phase state at any particular island can be determined at 95% probability of correct classification with information from as few as six sites. When sea otter population status (and thus the phase state of the kelp forest) was allowed to vary randomly among islands, just 15 islands had to be sampled to estimate the true proportion that were kelp dominated (within 10%) with 90% confidence. We conclude that kelp forest phase state is a more appropriate, sensitive, and cost-effective measure of sea otter recovery than the more traditional demographically based metrics, and we suggest that similar approaches have broad potential utility in establishing recovery criteria for depleted populations of other functionally important species.

Body size is an important determinant of the diving and foraging ability in air-breathing marine vertebrate predators. Satellite-linked dive recorders were used during 2003-2004 to investigate the foraging behavior of 22 male California sea lions (Zalophus californianus, a large, sexually dimorphic otariid) and to evaluate the extent to which body size explained variation among individuals and foraging strategies. Multivariate analyses were used to reduce the number of behavioral variables used to characterize foraging strategies (principal component analysis, PCA), to identify individually based foraging strategies in multidimensional space (hierarchical cluster analysis), and to classify each individual into a cluster or foraging strategy (discriminant analysis). Approximately 81.1% of the variation in diving behavior among individuals was explained by three factors: diving patterns (PC1), foraging effort (PC2), and behavior at the surface (PC3). Individuals were classified into three distinct groups based on their diving behavior (shallow, mixed depth, and deeper divers), and jackknife resampling of the data resulted in correct group assignment 86% of the time. Body size as an independent variable was positively related to dive duration and time spent ashore and negatively related to time at sea, and it was a key parameter in PC2 used to classify the three distinct clusters. Differences among individual-based foraging strategies probably were driven by differences in body size, which enabled larger animals to dive deeper and forage more efficiently by targeting different and perhaps larger prey items. The occurrence of foraging specializations within a species and age class has implications for quantitative modeling of population-level predator-prey interactions and ecosystem structure.

Coastal marine habitats at the interface of land and sea are subject to threats from human activities in both realms. Researchers have attempted to quantify how these various threats impact different coastal ecosystems, and more recently have focused on understanding the cumulative impact from multiple threats. Here, the top threats to coastal marine ecosystems and recent efforts to understand their relative importance, ecosystem-level impacts, cumulative effects, and how they can best be managed and mitigated, are briefly reviewed. Results of threat analysis and rankings will differ depending on the conservation target (e.g., vulnerable species, pristine ecosystems, mitigatable threats), scale of interest (local, regional, or global), whether externalities are considered, and the types of management tools available (e.g., marine-protected areas versus ecosystem-based management). Considering the cumulative effect of multiple threats has only just begun and depends on spatial analysis to predict overlapping threats and a better understanding of multiple-stressor effects and interactions. Emerging conservation practices that hold substantial promise for protecting coastal marine systems include multisector approaches, such as ecosystem-based management (EBM), that account for ecosystem service valuation; comprehensive spatial management, such as ocean zoning; and regulatory mechanisms that encourage or require cross-sector goal setting and evaluation. In all cases, these efforts require a combination of public and private initiatives for success. The state of our ecological understanding, public awareness, and policy initiatives make the time ripe for advancing coastal marine management and improving our stewardship of coastal and marine ecosystems.

Humans impact natural systems in a multitude of ways, yet the cumulative effect of multiple stressors on ecological communities remains largely unknown. Here we synthesized 171 studies that manipulated two or more stressors in marine and coastal systems and found that cumulative effects in individual studies were additive (26%), synergistic (36%), and antagonistic (38%). The overall interaction effect across all studies was synergistic, but interaction type varied by response level (community: antagonistic, population: synergistic), trophic level (autotrophs: antagonistic, heterotrophs: synergistic), and specific stressor pair (seven pairs additive, three pairs each synergistic and antagonistic). Addition of a third stressor changed interaction effects significantly in two-thirds of all cases and doubled the number of synergistic interactions. Given that most studies were performed in laboratories where stressor effects can be carefully isolated, these three-stressor results suggest that synergies may be quite common in nature where more than two stressors almost always coexist. While significant gaps exist in multiple stressor research, our results suggest an immediate need to account for stressor interactions in ecological studies and conservation planning.

Mammals balance heat dissipation with heat production to maintain core body temperatures independent of their environment. Thermal balance is undoubtedly most challenging for mammals born in polar regions because small body size theoretically results in high surface-area-to-volume ratios (SA:V), which facilitate heat loss (HL). Thus, we examined the ontogeny of thermoregulatory characteristics of an ice-breeding seal (Weddell seal Leptonychotes weddelli). Morphology, blubber thickness, rectal temperature (T(r)), muscle temperature (T(m)), and skin temperatures on the trunk (T(s)) and flipper (T(f)) in 3-5-wk-old pups, yearlings, and adults were measured. Adults maintained the thickest blubber layers, while yearlings had the thinnest; T(r) and T(m) fell within a narrow range, yet T(r) and T(m) decreased significantly with body length. All seals maintained skin temperatures lower than T(r), our index of core body temperature. The T(s)'s were positively correlated with environmental temperatures; conversely, T(f)'s were not. Although pups had the greatest proportion of blubber, their greater SA:V and limited ability to minimize body-to-environment temperature gradients led to the greatest calculated mass-specific HL. This implies that pups relied on elevated metabolic heat production to counter HL. Heat production in pups and yearlings may have been aided by nonshivering thermogenesis in the skeletal muscle via the enhanced muscle mitochondrial densities that have been observed in these segments of this population.

The vulnerability of the human brain to injury following just a few minutes of oxygen deprivation with submergence contrasts markedly with diving mammals, such as Weddell seals (Leptonychotes weddellii), which can remain underwater for more than 90 min while exhibiting no neurological or behavioural impairment. This response occurs despite exposure to blood oxygen levels concomitant with human unconsciousness. To determine whether such aquatic lifestyles result in unique adaptations for avoiding ischaemic-hypoxic neural damage, we measured the presence of circulating (haemoglobin) and resident (neuroglobin and cytoglobin) oxygen-carrying globins in the cerebral cortex of 16 mammalian species considered terrestrial, swimming or diving specialists. Here we report a striking difference in globin levels depending on activity lifestyle. A nearly 9.5-fold range in haemoglobin concentration (0.17-1.62 g Hb 100 g brain wet wt(-1)) occurred between terrestrial and deep-diving mammals; a threefold range in resident globins was evident between terrestrial and swimming specialists. Together, these two globin groups provide complementary mechanisms for facilitating oxygen transfer into neural tissues and the potential for protection against reactive oxygen and nitrogen groups. This enables marine mammals to maintain sensory and locomotor neural functions during prolonged submergence, and suggests new avenues for averting oxygen-mediated neural injury in the mammalian brain.

Dietary diversity often varies inversely with prey resource abundance. This pattern, although typically measured at the population level, is usually assumed to also characterize the behavior of individual animals within the population. However, the pattern might also be produced by changes in the degree of variation among individuals. Here we report on dietary and associated behavioral changes that occurred with the experimental translocation of sea otters from a food-poor to a food-rich environment. Although the diets of all individuals were broadly similar in the food-rich environment, a behaviorally based dietary polymorphism existed in the food-poor environment. Higher dietary diversity under low resource abundance was largely driven by greater variation among individuals. We further show that the dietary polymorphism in the food-poor environment included a broad suite of correlated behavioral variables and that the individuals that comprised specific behavioral clusters benefited from improved foraging efficiency on their individually preferred prey. Our findings add to the growing list of examples of extreme individuality in behavior and prey choice within populations and suggest that this phenomenon can emerge as a behavioral manifestation of increased population density. Individuality in foraging behavior adds complexity to both the fitness consequences of prey selection and food web dynamics, and it may figure prominently as a diversifying process over evolutionary timescales.

Otariids, like other wild mammals, contend with a wide variety of energetic demands across seasons. However, due to the cryptic behaviors of this marine group, few studies have been able to examine longitudinal energetic costs or the potential impact of these costs on seasonal or annual prey requirements. Here we evaluated the changes in energy demand and intake of female California sea lions (Zalophus californianus) during reproductive (n=2 sea lions) and nonreproductive (n=3) periods. Monthly measurements included resting metabolic rate, blood hormone levels, body condition (blubber thickness and body mass), and caloric intake for adult sea lions throughout molting, late pregnancy, lactation, and postweaning. We found that maintenance energy demands decreased from 32.0 to 23.1 MJ d(-1) before pupping, remaining stable at 19.4+/-0.6 MJ d(-1) during lactation and postweaning. Energy intake rates to meet these demands showed marked changes with activity level and the reproductive cycle, reaching a peak intake of 3.6 times baseline levels during lactation. Translating this into prey demands, we find that 20,000 reproductively active females on San Nicolas Island rookeries would maximally require 4,950 metric tons of Pacific whiting during a month of the breeding season. This localized impact is reduced significantly with postbreeding dispersal and demonstrates the importance of considering spatial and temporal factors driving the energetic requirements of predators when designing marine protected areas.

Quantifying spatial and temporal patterns of prey searching is of primary importance for understanding animals' critical habitat and foraging specialization. In patchy environments, animals forage by exhibiting movement patterns consisting of area-restricted searching (ARS) at various scales. Here, we present a new method, the fractal landscape method, which describes the peaks and valleys of fractal dimension along the animal path. We describe and test the method on simulated tracks, and quantify the effect of track inaccuracies. We show that the ARS zones correspond to the peaks from this fractal landscape and that the method is near error-free when analyzing high-resolution tracks, such as those obtained using the Global Positioning System (GPS). When we used tracks of lower resolution, such as those obtained with the Argos system, 9.6-16.3% of ARS were not identified, and 1-25% of the ARS were found erroneously. The later type of error can be partially flagged and corrected. In addition, track inaccuracies erroneously increased the measured ARS size by a factor of 1.2 to 2.2. Regardless, the majority of the times and locations were correctly flagged as being in or out of ARS (from 83.8 to 89.5% depending on track quality). The method provides a significant new tool for studies of animals' foraging behavior and habitat selection, because it provides a method to precisely quantify each ARS separately, which is not possible with existing methods.

A defining physiological capability for air-breathing marine vertebrates is the amount of oxygen that can be stored in tissues and made available during dives. To evaluate the influence of oxygen storage capacity on aerobic diving capacity, we examined the extent to which blood and muscle oxygen stores varied as a function of age, body size and sex in the sexually dimorphic California sea lion, Zalophus californianus. We measured total body oxygen stores, including hematocrit, hemoglobin, MCHC, plasma volume, blood volume and muscle myoglobin in pups through adults of both sexes. Blood and muscle oxygen storage capacity was not fully developed by the end of the dependency period, with blood stores not fully developed until animals were larger juveniles (70 kg; 1.5-2.5 years) and muscle stores not until animals were sub-adult size (125 kg; 4-6 years). Differences in aerobic diving capacity among size classes were reflective of these major milestones in the development of oxygen stores. Male sea lions had greater absolute blood volume than females and reflected the larger mass of males, which became apparent when animals were large juveniles. Adult female sea lions had greater muscle myoglobin concentrations compared to males, resulting in greater mass-specific muscle and total oxygen stores. Delayed development of oxygen stores is consistent with the shallow epi-mesopelagic foraging behavior in this species. We hypothesize that the greater mass-specific oxygen stores of female sea lions compared to males is related to differences in foraging behavior between the sexes.

Reliable information on historical and current population dynamics is central to understanding patterns of growth and decline in animal populations. We developed a maximum likelihood-based analysis to estimate spatial and temporal trends in age/sex-specific survival rates for the threatened southern sea otter (Enhydra lutris nereis), using annual population censuses and the age structure of salvaged carcass collections. We evaluated a wide range of possible spatial and temporal effects and used model averaging to incorporate model uncertainty into the resulting estimates of key vital rates and their variances. We compared these results to current demographic parameters estimated in a telemetry-based study conducted between 2001 and 2004. These results show that survival has decreased substantially from the early 1990s to the present and is generally lowest in the north-central portion of the population's range. The greatest temporal decrease in survival was for adult females, and variation in the survival of this age/sex class is primarily responsible for regulating population growth and driving population trends. Our results can be used to focus future research on southern sea otters by highlighting the life history stages and mortality factors most relevant to conservation. More broadly, we have illustrated how the powerful and relatively straightforward tools of information-theoretic-based model fitting can be used to sort through and parameterize quite complex demographic modeling frameworks.