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 degrees 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.

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.

Buoyancy and maximal diving depth in penguins: do they control inhaling air volume?

Using a newly developed data logger to measure acceleration, we demonstrate that free-ranging king and Adélie penguins only beat their flippers substantially during the first part of descent or when they were presumed to be chasing prey at the bottom of dives. Flipper beating stopped during the latter part of ascent: at 29+/-9 % (mean +/- S.D.) of dive depth (mean dive depth=136.8+/-145.1 m, N=425 dives) in king penguins, and at 52+/-20 % of dive depth (mean dive depth=72.9+/-70.5 m, N=664 dives) in Adélie penguins. Propulsive swim speeds of both species were approximately 2 m s(-1) during dives; however, a marked increase in speed, up to approximately 2.9 m s(-1), sometimes occurred in king penguins during the passive ascending periods. During the prolonged ascending, oblique ascent angle and slowdown near the surface may represent one way to avoid the potential risk of decompression sickness. Biomechanical calculations for data from free-ranging king and Adélie penguins indicate that the air volume of the birds (respiratory system and plumage) can provide enough buoyancy for the passive ascent. When comparing the passive ascents for shallow and deep dives, there is a positive correlation between air volume and the depth of the dive. This suggests that penguins regulate their air volume to optimize the costs and benefits of buoyancy.

Feeding behaviour of free-ranging penguins determined by oesophageal temperature.

Sea birds play a major role in marine food webs, and it is important to determine when and how much they feed at sea. A major advance has been made by using the drop in stomach temperature after ingestion of ectothermic prey. This method is less sensitive when birds eat small prey or when the stomach is full. Moreover, in diving birds, independently of food ingestion, there are fluctuations in the lower abdominal temperature during the dives. Using oesophageal temperature, we present here a new method for detecting the timing of prey ingestion in free-ranging sea birds, and, to our knowledge, report the first data obtained on king penguins (Aptenodytes patagonicus). In birds ashore, which were hand-fed 2-15 g pieces of fish, all meal ingestions were detected with a sensor in the upper oesophagus. Detection was poorer with sensors at increasing distances from the beak. At sea, slow temperature drops in the upper oesophagus and stomach characterized a diving effect per se. For the upper oesophagus only, abrupt temperature variations were superimposed, therefore indicating prey ingestions. We determined the depths at which these occurred. Combining the changes in oesophageal temperatures of marine predators with their diving pattern opens new perspectives for understanding their foraging strategy, and, after validation with concurrent applications of classical techniques of prey survey, for assessing the distribution of their prey.

Seabirds as monitors of upper-ocean thermal structure. King penguins at the Antarctic polar front, east of Kerguelen sector.

The main objective of this work was to assess the potential of diving birds to monitor the hydrographic features near the Antarctic polar front. We compared the temperature/depth profiles recorded by instrumented King penguins Aptenodytes patagonicus at Kerguelen Islands (South Indian Ocean) with the oceanographic and remote sensing (satellite) data available for the same area during the same season. The birds were equipped with time/depth/temperature recorders or Argos transmitters. In addition, two birds were instrumented (of which one successfully) both with a time/depth/temperature recorder and an Argos transmitter. King penguins foraged as far as 400 km from the coast, in water masses with a vertical temperature structure characteristic of the region just south of the polar front. The temperature/depth profiles recorded throughout the dives (up to 270 m) revealed a pronounced thermocline. A three-dimensional distribution of water temperature was reconstructed. Comparison with previous hydrographic data shows a high correlation. Instrumented predators may therefore usefully and cheaply complement the database provided by conventional hydrographic surveys and remote sensing, especially in distant and rough areas such as the Southern Ocean.

Foraging strategies of incubating and brooding king penguins Aptenodytes patagonicus.

For oceanic birds like king penguins, a major constraint is the separation of foraging areas from the breeding colony, largely because swimming increases foraging costs. However, the relationship between foraging strategy and breeding stage has been poorly investigated. Using time-depth recorders, we studied the diving behaviour of two groups of king penguins that were either incubating or brooding chicks at Crozet Islands (Southern Indian Ocean) at the same period of the year. Although birds with chicks had the highest predicted energy demand, they made foraging trips half as long as incubating birds (6 vs. 14 days) and modified their time and depth utilisation. Birds with chicks dived deeper during daylight (mean maximum depth of 280 m vs. 205 m for those incubating). At night, birds with chicks spent twice as much time diving as those incubating, but birds at both stages never dived beyond 30 m. Movements to greater depths by brooding birds are consistent with the vertical distribution of myctophid fish which are the main prey. As chick provisioning limits trip duration, it is suggested that it is more efficient for parents to change their diving patterns rather than to restrict their foraging range.

Energy and protein requirements for molt in the king penguin Aptenodytes patagonicus.

Adult king penguins annually fast ashore for 1 mo for molting. By the end of molt, they have lost 44% of their prefasting body mass. About 18% of new feather synthesis occurs at sea, thus reducing both nutrient requirement and fasting duration. Plumage synthesis continues during the first 3 wk of fasting. Loss of old feathers occurs between day 12 and day 21 of the molt, and it is associated with a peak in daily body mass loss. The dry mass of epidermal structure synthesized during molt is 395 g. Body composition analysis indicates that fat oxidation accounts for 85% of total energy expenditure. The proportion for protein is 15%, a value twofold higher than during the breeding (nonmolting) fast. The mean energy expenditure is also 21% higher during the molting fast (3.04 W/kg). Compared with other birds, the energetic cost of feather synthesis is the lowest in king penguins (85 kJ/g) and consequently the energetic efficiency is the highest (25%). Changes in tissue composition during molt show that integument is the main lipid source (72% of the lipid loss) and thus the main source of energy (61% of the total energy expenditure). The integument and the pectoral muscles play a major role in molting protein metabolism, providing 20 and 57%, respectively, of the total protein needs for feather synthesis and/or energy expenditure. This result emphasizes the role of integument as a protein source, because the large premolting muscle hypertrophy is not sufficient to account for the totality of the protein cost of molt.