Michael E. Goebel

Variations in behavior and condition of a Southern Ocean top predator in relation to in situ oceanographic conditions

Responses by marine top predators to environmental variability have previously been almost impossible to observe directly. By using animal-mounted instruments simultaneously recording movements, diving behavior, and in situ oceanographic properties, we studied the behavioral and physiological responses of southern elephant seals to spatial environmental variability throughout their circumpolar range. Improved body condition of seals in the Atlantic sector was associated with Circumpolar Deep Water upwelling regions within the Antarctic Circumpolar Current, whereas High-Salinity Shelf Waters or temperature/salinity gradients under winter pack ice were important in the Indian and Pacific sectors. Energetic consequences of these variations could help explain recently observed population trends, showing the usefulness of this approach in examining the sensitivity of top predators to global and regional-scale climate variability.

Interpolation of animal tracking data in a fluid environment

SUMMARY 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.

Southern Ocean frontal structure and sea-ice formation rates revealed by elephant seals

Polar regions are particularly sensitive to climate change, with the potential for significant feedbacks between ocean circulation, sea ice, and the ocean carbon cycle. However, the difficulty in obtaining in situ data means that our ability to detect and interpret change is very limited, especially in the Southern Ocean, where the ocean beneath the sea ice remains almost entirely unobserved and the rate of sea-ice formation is poorly known. Here, we show that southern elephant seals (Mirounga leonina) equipped with oceanographic sensors can measure ocean structure and water mass changes in regions and seasons rarely observed with traditional oceanographic platforms. In particular, seals provided a 30-fold increase in hydrographic profiles from the sea-ice zone, allowing the major fronts to be mapped south of 60°S and sea-ice formation rates to be inferred from changes in upper ocean salinity. Sea-ice production rates peaked in early winter (April–May) during the rapid northward expansion of the pack ice and declined by a factor of 2 to 3 between May and August, in agreement with a three-dimensional coupled ocean–sea-ice model. By measuring the high-latitude ocean during winter, elephant seals fill a “blind spot” in our sampling coverage, enabling the establishment of a truly global ocean-observing system.

Aerobic dive limit: how often does it occur in nature? ☆

Diving animals offer a unique opportunity to study the importance of physiological constraint in their everyday behaviors. An important component of the physiological capability of any diving animal is its aerobic dive limit (ADL). The ADL has only been measured in a few species. The goal of this study was to estimate the aerobic dive limit from measurements of body oxygen stores and at sea metabolism. This calculated ADL (cADL) was then compared to measurements of diving behavior of individual animals of three species of otariids, the Antarctic fur seal, Arctocephalus gazella, the Australian sea lion, Neophoca cinerea, and the New Zealand sea lion, Phocarctos hookeri. Antarctic fur seals dove well within the cADL. In contrast, many individuals of both sea lion species exceeded the cADL, some by significant amounts. Australian sea lions typically dove 1.4 times longer than the cADL, while New Zealand sea lions on average dove 1.5 times longer than the cADL. The tendency to exceed the cADL was correlated with the dive pattern of individual animals. In both Antarctic Fur Seals and Australian sea lions, deeper diving females made longer dives that approached or exceeded the cADL (P<0.01, r(2)=0.54). Australian and New Zealand sea lions with longer bottom times also exceeded the cADL to a greater degree. The two sea lions forage on the benthos while the fur seals feed shallow in the water column. It appears that benthic foraging requires these animals to reach or exceed their aerobic dive limit.

Diving behaviour of chinstrap penguins at Seal Island

Diving behaviour of chinstrap penguins (Pygoscelis anturctica) was studied in four adults brooding chicks on Seal Island, South Shetland Islands, Antarctica. During foraging trips to sea, chinstrap penguins made shallow, short duration dives almost continuously, for the most part within 50 m of the surface. Diving effort was concentrated during the daylight hours (10h00-15h00), although a second peak in effort was seen around midnight (22h00-02hOO). These peaks were possibly due to the constraints of visual location of prey, chick provisioning, or the need to take advantage of diurnal changes in krill swarm densities or behaviour. It was estimated that most effort was concentrated 3-20 km from shore. Dive depth and duration averaged 31.0 m (2 26.3m)and72s(+36 s),respectively. Maximumdivedepthanddurationwere 121mand180s, respectively. Received 23 April 1991, accepted 20 August 1992

Oceanic controls on the mass balance of Wilkins Ice Shelf, Antarctica

� 0.8 m a � 1 , driven by a mean basal melt rate of 〈wb〉 = 1.3 � 0.4 m a � 1 . Interannual variability was large, associated with changes in both surface mass accumulation and 〈wb〉. Basal melt rate declined significantly around 2000 from 1.8 � 0.4 m a � 1 for 1992–2000 to � 0.75 � 0.55 m a � 1 for 2001–2008; the latter value corresponding to approximately steady-state ice-shelf mass. Observations of ocean temperature T obtained during 2007–2009 by instrumented seals reveal a cold, deep halo of Winter Water (WW; T ≈ � 1.6°C) surrounding WIS. The base of the WW in the halo is � 170 m, approximately the mean ice draft for WIS. We hypothesize that the transition in 〈wb〉 in 2000 was caused by a small perturbation (� 10–20 m) in the relative depths of the ice base and the bottom of the WW layer in the halo. We conclude that basal melting of thin ice shelves like WIS is very sensitive to upper-ocean and coastal processes that act on shorter time and space scales than those affecting basal melting of thicker West Antarctic ice shelves such as George VI and Pine Island Glacier.

Electronic tracking tag programming is critical to data collection for behavioral time-series analysis

Electronic tracking tags are major tools of ecological research and management, but programming sophisticated tags can be challenging. We discovered that a common programming scheme can negatively affect the quality of tracks collected by Argos tags. Here we describe the problem and how it occurred. We then simulated a series of tracks with different data collection schemes to investigate how spatial precision and temporal frequency affect the overall quality of tracking data. Tracks were simulated using a two-state composite correlated random walk (CCRW). Tracks were simulated with two spatial scales, using parameters estimated from northern elephant seal (large scale) and California sea lion (small scale) tracking data. Onto each simulated track, observations of varying precision, frequency, and censoring were imposed. We then fit the CCRW in a state-space model (SSM) to the simulated observations in order to assess how data quality and frequency affected recovery of known behavioral state and location. We show that when movement scales are small, regular observations were critical to recover behavior and location. In addition, tracks with frequent regular locations (increasing N) overcame low spatial accuracy (e.g., Argos) to detect small-scale movement patterns, suggesting frequently collected Argos locations may be as good as infrequently collected GPS in some circumstances. From these results and our experience tracking animals generally, we produce a set of guidelines for those manufacturing, programming, and deploying electronic tracking tags to maximize the utility of the data they produce.

Seals map bathymetry of the Antarctic continental shelf

We demonstrate the first use of marine mammal dive-depth data to improve maps of bathymetry in poorly sampled regions of the continental shelf. A group of 57 instrumented elephant seals made on the order of 2 x 10(5) dives over and near the continental shelf on the western side of the Antarctic Peninsula during five seasons, 2005-2009. Maximum dive depth exceeded 2000 m. For dives made near existing ship tracks with measured water depths H<700 m, similar to 30% of dive depths were to the seabed, consistent with expected benthic foraging behavior. By identifying the deepest of multiple dives within small areas as a dive to the seabed, we have developed a map of seal-derived bathymetry. Our map fills in several regions for which trackline data are sparse, significantly improving delineation of troughs crossing the continental shelf of the southern Bellingshausen Sea.

Ecological implications of body composition and thermal capabilities in young antarctic fur seals (Arctocephalus gazella).

In comparison with other homeotherms, young recently weaned marine mammals in high latitudes face exceptional energetic demands when foraging and thermoregulating. Lipids are an important source of energy and a major component of insulation that allows them to meet these demands. To examine the role of lipid stores in a high‐latitude pinniped, we measured the body composition and thermoregulatory capabilities of Antarctic fur seal (Arctocephalus gazella) pups and yearlings by using flow‐through respirometry and hydrogen isotope dilution. From these data, we constructed a model to examine the importance of postweaning fasting capability in free‐ranging young fur seals. Resting metabolic rates were different for pups and yearlings measured in 0.6°C water, 10.3°C water, and ambient air; however, mass and percent lipid as covariates accounted for the different metabolic responses in pups and yearlings for all treatments. The estimated lower critical temperature for combined pups and yearlings was 14.4°C, 10°–15°C above water temperatures normally experienced by Antarctic fur seals. Modeling predicted that a weaned fur seal pup would survive at sea from 9.8 to 36.2 d before succumbing to starvation. The most likely maximum travel distance within this time constraint suggests that food resources close to the natal rookery are important to first‐year survival for this species.

Top-down and bottom-up influences on demographic rates of Antarctic fur seals Arctocephalus gazella

Two major drivers in population dynamics are bottom-up processes, such as environmental factors that affect foraging success, and the top-down impacts of predation. Many populations of marine mammal and seabird species appear to be declining in response to reductions in prey associated with the bottom-up effects of climate change. However, predation, which usually occurs at sea and is difficult to observe, may also play a key role. We analysed drivers of population dynamics of Antarctic fur seals Arctocephalus gazella at Cape Shirreff from 1997 to 2009, including a predator that targets pre-weaned pups and bottom-up environmental effects in an ecosystem particularly sensitive to small changes in temperature. We use Bayesian mark-recapture analysis to demonstrate that although large-scale environmental variability affects annual adult survival and reproduction, first year survival appears to be driving the current decline in this population (as defined by a decline in the annual number of pups born). Although the number of pups increased during the first third of the study, first year survival and recruitment of those pups in later years was very low. Such low survival may be driven by leopard seal Hydrurga leptonyx predation, particularly prior to weaning. Our results suggest that without leopard seal predation, this population would most likely increase in size, despite the observed bottom-up effects of climate changes on adult vital rates. More broadly, our results show how age-targeted predation could be a major factor in population decline of K-selected colonial breeders.