Tracking of marine predators to protect Southern Ocean ecosystems.

Southern Ocean ecosystems are under pressure from resource exploitation and climate change1,2. 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.

Sink fast and swim harder! Round-trip cost-of-transport for buoyant divers.

Efficient locomotion between prey resources at depth and oxygen at the surface is crucial for breath-hold divers to maximize time spent in the foraging layer, and thereby net energy intake rates. The body density of divers, which changes with body condition, determines the apparent weight (buoyancy) of divers, which may affect round-trip cost-of-transport (COT) between the surface and depth. We evaluated alternative predictions from external-work and actuator-disc theory of how non-neutral buoyancy affects round-trip COT to depth, and the minimum COT speed for steady-state vertical transit. Not surprisingly, the models predict that one-way COT decreases (increases) when buoyancy aids (hinders) one-way transit. At extreme deviations from neutral buoyancy, gliding at terminal velocity is the minimum COT strategy in the direction aided by buoyancy. In the transit direction hindered by buoyancy, the external-work model predicted that minimum COT speeds would not change at greater deviations from neutral buoyancy, but minimum COT speeds were predicted to increase under the actuator disc model. As previously documented for grey seals, we found that vertical transit rates of 36 elephant seals increased in both directions as body density deviated from neutral buoyancy, indicating that actuator disc theory may more closely predict the power requirements of divers affected by gravity than an external work model. For both models, minor deviations from neutral buoyancy did not affect minimum COT speed or round-trip COT itself. However, at body-density extremes, both models predict that savings in the aided direction do not fully offset the increased COT imposed by the greater thrusting required in the hindered direction.

Effects of handling regime and sex on changes in cortisol, thyroid hormones and body mass in fasting grey seal pups.

Survival of seal pups may be affected by their ability to respond appropriately to stress. Chronic stress can adversely affect secretion of cortisol and thyroid hormones, which contribute to the control of fuel utilisation. Repeated handling could disrupt the endocrine response to stress and/or negatively impact upon mass changes during fasting. Here we investigated the effects of handling regime on cortisol and thyroid hormone levels, and body mass changes, in fasting male and female grey seal pups (Halichoerus grypus). Females had higher thyroid hormone levels than males throughout fasting and showed a reduction in cortisol midway through the fast that was not seen in males. This may reflect sex-specific fuel allocation or development. Neither handling frequency nor cumulative contact time affected plasma cortisol or thyroid hormone levels, the rate of increase in cortisol over the first five minutes of physical contact or the pattern of mass loss during fasting in either sex. The endocrine response to stress and the control of energy balance in grey seal pups appear to be robust to repeated, short periods of handling. Our results suggest that routine handling should have no additional impact on these animals than general disturbance caused by researchers moving around the colony.

Northern elephant seals adjust gliding and stroking patterns with changes in buoyancy: validation of at-sea metrics of body density.

Many diving animals undergo substantial changes in their body density that are the result of changes in lipid content over their annual fasting cycle. Because the size of the lipid stores reflects an integration of foraging effort (energy expenditure) and foraging success (energy assimilation), measuring body density is a good way to track net resource acquisition of free-ranging animals while at sea. Here, we experimentally altered the body density and mass of three free-ranging elephant seals by remotely detaching weights and floats while monitoring their swimming speed, depth and three-axis acceleration with a high-resolution data logger. Cross-validation of three methods for estimating body density from hydrodynamic gliding performance of freely diving animals showed strong positive correlation with body density estimates obtained from isotope dilution body composition analysis over density ranges of 1015 to 1060 kg m(-3). All three hydrodynamic models were within 1% of, but slightly greater than, body density measurements determined by isotope dilution, and therefore have the potential to track changes in body condition of a wide range of freely diving animals. Gliding during ascent and descent clearly increased and stroke rate decreased when buoyancy manipulations aided the direction of vertical transit, but ascent and descent speed were largely unchanged. The seals adjusted stroking intensity to maintain swim speed within a narrow range, despite changes in buoyancy. During active swimming, all three seals increased the amplitude of lateral body accelerations and two of the seals altered stroke frequency in response to the need to produce thrust required to overcome combined drag and buoyancy forces.

Using GPS data to evaluate the accuracy of state-space methods for correction of Argos satellite telemetry error.

Recent studies have applied state-space models to satellite telemetry data in order to remove noise from raw location estimates and infer the true tracks of animals. However, while the resulting tracks may appear plausible, it is difficult to determine the accuracy of the estimated positions, especially for position estimates interpolated to times between satellite locations. In this study, we use data from two gray seals (Halichoerus grypus) carrying tags that transmitted Fastloc GPS positions via Argos satellites. This combination of Service Argos data and highly accurate GPS data allowed examination of the accuracy of state-space position estimates and their uncertainty derived from satellite telemetry data. After applying a speed filter to remove aberrant satellite telemetry locations, we fit a continuous-time Kalman filter to estimate the parameters of a random walk, used Kalman smoothing to infer positions at the times of the GPS measurements, and then compared the filtered telemetry estimates with the actual GPS measurements. We investigated the effect of varying maximum speed thresholds in the speed-filtering algorithm on the root mean-square error (RMSE) estimates and used minimum RMSE as a criterion to guide the final choice of speed threshold. The optimal speed thresholds differed between the two animals (1.1 m/s and 2.5 m/s) and retained 50% and 65% of the data for each seal. However, using a speed filter of 1.1 m/s resulted in very similar RMSE for both animals. For the two seals, the RMSE of the Kalman-filtered estimates of location were 5.9 and 12.76 km, respectively, and 75% of the modeled positions had errors less than 6.25 km and 11.7 km for each seal. Confidence interval coverage was close to correct at typical levels (80-95%), although it tended to be overly generous at smaller sizes. The reliability of uncertainty estimates was also affected by the chosen speed threshold. The combination of speed and Kalman filtering allows for effective calculation of location and also indicates the limits of accuracy when correcting service Argos locations and linking satellite telemetry data to spatial covariate and habitat data.

The retrospective analysis of Antarctic tracking data project.

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.

Aquatic behaviour of polar bears (Ursus maritimus) in an increasingly ice-free Arctic.

Polar bears are ice-associated marine mammals that are known to swim and dive, yet their aquatic behaviour is poorly documented. Reductions in Arctic sea ice are clearly a major threat to this species, but understanding polar bears' potential behavioural plasticity with respect to the ongoing changes requires knowledge of their swimming and diving skills. This study quantified time spent in water by adult female polar bears (n = 57) via deployment of various instruments bearing saltwater switches, and in some case pressure sensors (79 deployments, 64.8 bear-years of data). There were marked seasonal patterns in aquatic behaviour, with more time spent in the water during summer, when 75% of the polar bears swam daily (May-July). Females with cubs-of-the-year spent less time in the water than other females from den emergence (April) until mid-summer, consistent with small cubs being vulnerable to hypothermia and drowning. Some bears undertook notable long-distance-swims. Dive depths up to 13.9 m were recorded, with dives ≥5 m being common. The considerable swimming and diving capacities of polar bears might provide them with tools to exploit aquatic environments previously not utilized. This is likely to be increasingly important to the species' survival in an Arctic with little or no persistent sea ice.

Hooded seal Cystophora cristata foraging areas in the Northeast Atlantic Ocean-Investigated using three complementary methods.

Identifying environmental characteristics that define the ecological niche of a species is essential to understanding how changes in physical conditions might affect its distribution and other aspects of its ecology. The present study used satellite relay data loggers (SRDLs) to study habitat use by Northeast Atlantic hooded seals (N = 20; 9 adult females, 3 adult males, and 8 juveniles). Three different methods were used in combination to achieve maximum insight regarding key foraging areas for hooded seals in this region, which have decline by 85% in recent decades: 1) first passage time (FPT); 2) vertical transit rate and; 3) change in dive drift rate. Generalized additive mixed models (GAMM) were applied to each method to determine whether specific habitat characteristics were associated with foraging. Separate models were run for the post-molting and the post-breeding seasons; sex and age classes were included in the GAMMs. All three methods highlighted a few common geographic areas as being important foraging zones; however, there were also some different areas identified by the different methods, which highlights the importance of using multiple indexes when analyzing tracking and diving data to study foraging behavior. Foraging occurred most commonly in relatively shallow areas with high Sea Surface Temperatures (SST), corresponding to continental shelf areas with Atlantic Water masses. All age and sex classes overlapped spatially to some extent, but the different age and sex groups showed differences in the bathymetry of their foraging areas as well as in their vertical use of the water column. When foraging, pups dove in the upper part of the water column in relatively deep areas. Adult females foraged relatively shallowly in deep water areas too, though in shallower areas than pups. Adult males foraged close to the bottom in shallower areas.

Bimodal Winter Haul-Out Patterns of Adult Weddell Seals (Leptonychotes weddellii) in the Southern Weddell Sea.

Hauling out is an essential component of pinniped life-history. Haul-out behaviour may be affected by biological (e.g. sex, age and condition) and physical factors (e.g. food availability and environmental conditions), and identifying these factors may help explain the spatio-temporal distribution and habitat use of pinnipeds. The aim of this study is to describe observed winter haul-out patterns of adult Weddell seals in the Weddell Sea and investigate the role of potential predictors to gain insight into the way these animals interact with the physical environment in this region. We examined the haul-out behaviour in relation to available biological (i.e., diving effort, sex) and physical information (i.e., sun angle). Thirty-three satellite telemetry tags were deployed on adult Weddell seals in the southern Weddell Sea during February 2007, 2009 and 2011, following their annual moult recording information on the behavioural mode of the animal: at surface, hauled out or diving. At the end of the austral summer Weddell seals spent, on average, more than 40% of their time hauled out on the ice. Under constant light conditions, it appears that physiological factors drive sex differences in the timing and duration of haul-out behaviour, with females spending on average more time hauled out than males during daylight hours. This time spent hauled-out declined to around 15% in both sexes by the beginning of autumn and remained at this level with a clear nocturnal haul-out pattern during the winter. The time spent diving increased during this period, indicating an increase in foraging effort during the winter months, and led to a common haul-out pattern in both sexes over winter. We found a positive relationship between haul-out duration and the percentage of time spent diving prior to a haul-out in both sexes, with the exception of female daytime haul-outs early in the year.

The Argos-CLS Kalman Filter: Error Structures and State-Space Modelling Relative to Fastloc GPS Data.

Understanding how an animal utilises its surroundings requires its movements through space to be described accurately. Satellite telemetry is the only means of acquiring movement data for many species however data are prone to varying amounts of spatial error; the recent application of state-space models (SSMs) to the location estimation problem have provided a means to incorporate spatial errors when characterising animal movements. The predominant platform for collecting satellite telemetry data on free-ranging animals, Service Argos, recently provided an alternative Doppler location estimation algorithm that is purported to be more accurate and generate a greater number of locations that its predecessor. We provide a comprehensive assessment of this new estimation process performance on data from free-ranging animals relative to concurrently collected Fastloc GPS data. Additionally, we test the efficacy of three readily-available SSM in predicting the movement of two focal animals. Raw Argos location estimates generated by the new algorithm were greatly improved compared to the old system. Approximately twice as many Argos locations were derived compared to GPS on the devices used. Root Mean Square Errors (RMSE) for each optimal SSM were less than 4.25 km with some producing RMSE of less than 2.50 km. Differences in the biological plausibility of the tracks between the two focal animals used to investigate the utility of SSM highlights the importance of considering animal behaviour in movement studies. The ability to reprocess Argos data collected since 2008 with the new algorithm should permit questions of animal movement to be revisited at a finer resolution.

A Southern Indian Ocean database of hydrographic profiles obtained with instrumented elephant seals.

The instrumentation of southern elephant seals with satellite-linked CTD tags has offered unique temporal and spatial coverage of the Southern Indian Ocean since 2004. This includes extensive data from the Antarctic continental slope and shelf regions during the winter months, which is outside the conventional areas of Argo autonomous floats and ship-based studies. This landmark dataset of around 75,000 temperature and salinity profiles from 20-140 °E, concentrated on the sector between the Kerguelen Islands and Prydz Bay, continues to grow through the coordinated efforts of French and Australian marine research teams. The seal data are quality controlled and calibrated using delayed-mode techniques involving comparisons with other existing profiles as well as cross-comparisons similar to established protocols within the Argo community, with a resulting accuracy of ±0.03 °C in temperature and ±0.05 in salinity or better. The data offer invaluable new insights into the water masses, oceanographic processes and provides a vital tool for oceanographers seeking to advance our understanding of this key component of the global ocean climate.

How many seals were there? The global shelf loss during the last glacial maximum and its effect on the size and distribution of grey seal populations.

Predicting how marine mammal populations respond to habitat changes will be essential for developing conservation management strategies in the 21st century. Responses to previous environmental change may be informative in the development of predictive models. Here we describe the likely effects of the last ice age on grey seal population size and distribution. We use satellite telemetry data to define grey seal foraging habitat in terms of the temperature and depth ranges exploited by the contemporary populations. We estimate the available extent of such habitat in the North Atlantic at present (between 1.42 · 10(6) km(2) and 2.07 · 10(6) km(2)) and at the last glacial maximum (between 4.74 · 10(4) km(2) and 2.11 · 10(5) km(2)); taking account of glacial and seasonal sea-ice coverage, estimated reductions of sea-level (123 m) and sea surface temperature hind-casts. Most of the extensive continental shelf waters (North Sea, Baltic Sea and Scotian Shelf), currently supporting >95% of grey seals, were unavailable during the last glacial maximum. A combination of lower sea-level and extensive ice-sheets, massively increased seasonal sea-ice coverage and southerly extent of cold water would have pushed grey seals into areas with no significant shelf waters. The habitat during the last glacial maximum might have been as small as 3% of today's extent and grey seal populations may have fallen to similarly low numbers. An alternative scenario involving a major change to a pelagic or bathy-pelagic foraging niche cannot be discounted. However, hooded seals currently dominate that niche and may have excluded grey seals from such habitat. If as seems likely, the grey seal population fell to very low levels it would have remained low for several thousand years before expanding into current habitats over the past 12,000 years or so.

Eat now, pay later? Evidence of deferred food-processing costs in diving seals.

Seals may delay costly physiological processes (e.g. digestion) that are incompatible with the physiological adjustments to diving until after periods of active foraging. We present unusual profiles of metabolic rate (MR) in grey seals measured during long-term simulation of foraging trips (4-5 days) that provide evidence for this. We measured extremely high MRs (up to almost seven times the baseline levels) and high heart rates during extended surface intervals, where the seals were motionless at the surface. These occurred most often during the night and occurred frequently many hours after the end of feeding bouts. The duration and amount of oxygen consumed above baseline levels during these events was correlated with the amount of food eaten, confirming that these metabolic peaks were related to the processing of food eaten during foraging periods earlier in the day. We suggest that these periods of high MR represent a payback of costs deferred during foraging.

Effects of mass and body composition on fasting fuel utilisation in grey seal pups (Halichoerus grypus Fabricius): an experimental study using supplementary feeding.

This study used supplementary feeding to test the hypothesis that fuel partitioning during the postweaning fast in grey seal pups is affected by size and composition of energy reserves at weaning, and by extra provisioning. Mass and body composition changes were measured during suckling and fasting to investigate the effect of natural differences in energy reserves at weaning on subsequent allocation of fat and protein to energy use. We fed seven pups for 5 days after weaning, to investigate the effect of increased fuel availability, and particularly protein, on fuel utilisation. After correcting for protein used during the moult, the proportional contribution of fat was 86-99% of total energy use. Pups with greater energy reserves, i.e. those that were heavier and fatter at weaning, had higher rates of fat and energy use. There was no significant relationship between adiposity at weaning and proportional contribution of fat to energy use, perhaps due to a limited sample size or range of body masses and adiposity. Supplemented individuals used energy, specifically fat, much faster and utilised proportionally less of their endogenous protein by departure than non-supplemented individuals. Fat metabolism contributed a similar percentage to daily energy use in both groups. These findings show that pups spare protein, even when energy use is dramatically increased. Pups that receive greater maternal provisioning and lay down more protein may have increased survival chances at sea. This study highlights the importance of protein reserves in first year survival of grey seal pups.

Overcoming the constraints of long range radio telemetry from animals: getting more useful data from smaller packages.

Many species carry out their most interesting activities where they cannot readily be observed or monitored. Marine mammals are extreme among this group, accomplishing their most astounding activities both distant from land and deep in the sea. Collection, storage and transmission of data about these activities are constrained by the energy requirements and size of the recording loggers and transmitters. The more bits of information collected, stored and transmitted, the more battery is required and the larger the tag must be. We therefore need to be selective about the information we collect, while maintaining detail and fidelity. To accomplish this in the study of marine mammals, we have designed "intelligent" data logger/transmitters that provide context-driven data compression, data relay, and automated data base storage. We later combine these data with remotely sensed environmental information and other oceanographic data sets to recreate the environmental context for the animal's activity, and we display the combined data using computer animation techniques. In this way, the system can provide near real time "observation" of animal behavior and physiology from the remotest parts of the globe.

Blubber and buoyancy: monitoring the body condition of free-ranging seals using simple dive characteristics.

Elephant seals regularly perform dives during which they spend a large proportion of time drifting passively through the water column. The rate of vertical change in depth during these "drift" dives is largely a result of the proportion of lipid tissue in the body, with fatter seals having higher (more positive or less negative) drift rates compared with leaner seals. We examined the temporal changes in drift rates of 24 newly weaned southern elephant seal (Mirounga leonina) pups during their first trip to sea to determine if this easily recorded dive characteristic can be used to continuously monitor changes in body composition of seals throughout their foraging trips. All seals demonstrated a similar trend over time: drift rates were initially positive but decreased steadily over the first 30-50 days after departure (Phase 1), corresponding to seals becoming gradually less buoyant. Over the following approximately 100 days (Phase 2), drift rates again increased gradually, while during the last approximately 20-45 days (Phase 3) drift rates either remained constant or decreased slightly. The daily rate of change in drift rate was negatively related to the daily rate of horizontal displacement (daily travel rate), and daily travel rates of more than approximately 80 km were almost exclusively associated with negative changes in drift rate. We developed a mechanistic model based on body compositions and morphometrics measured in the field, published values for the density of seawater and various body components, and values of drag coefficients for objects of different shapes. We used this model to examine the theoretical relationships between drift rate and body composition and carried out a sensitivity analysis to quantify errors and biases caused by varying model parameters. While variations in seawater density and uncertainties in estimated body surface area and volume are unlikely to result in errors in estimated lipid content of more than +/-2.5%, variations in drag coefficient can lead to errors of >or =10%. Finally, we compared the lipid contents predicted by our model with the lipid contents measured using isotopically labelled water and found a strong positive correlation. The best-fitting model suggests that the drag coefficient of seals while drifting passively is between approximately 0.49 (roughly corresponding to a sphere-shaped object) and 0.69 (a prolate spheroid), and we were able to estimate relative lipid content to within approximately +/-2% lipid. Our results suggest that this simple method can be used to estimate the changes in lipid content of free-ranging seals while at sea and may help improve our understanding of the foraging strategies of these important marine predators.