Martin Biuw

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.

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.

Phylogenetic analyses of sexual selection and sexual size dimorphism in pinnipeds

Abstract. Pinnipedia contains some of the most spectacular examples of sexual size dimorphism, examples that are therefore frequently used to illustrate the theory of sexual selection. This paper addresses the question of whether a significant relationship between sexual selection and size dimorphism exists in a comparative context. Thus, harem size and body size data gathered from the literature were analysed with independent contrasts analyses. These investigations showed that sexual size dimorphism is not a consequence of an allometric relationship between male and female size. Instead, there is a clear relationship between harem size and sexual size dimorphism. Further analyses also revealed a significant relationship between harem size and male size whereas no such relationship existed for females. These results support the hypothesis that sexual size dimorphism in pinnipeds is the product of an exclusively male response to sexual selection.

Successful foraging zones of southern elephant seals from the Kerguelen Islands in relation to oceanographic conditions

Southern elephant seals, Mirounga leonina, undertake large-scale oceanic movements to access favourable foraging areas. Successful foraging areas of elephant seals from the Kerguelen Islands are investigated here in relation to oceanographic parameters. Movements and diving activity of the seals as well as oceanographic data were collected through a new generation of satellite relayed devices measuring and transmitting locations, pressure, temperature and salinity. For the first time, we have associated foraging behaviour, determined by high increased sinuosity in tracks, and dive density (i.e. number of dives performed per kilometre covered), and changes in body condition, determined by variations in drift rate obtained from drift dives, to identify the oceanographic conditions of successful foraging zones for this species. Two main sectors, one close to the Antarctic continent and the other along the Polar Front (PF), where both foraging activity and body condition increase, seem to be of particular interest for the seals. Within these regions, some seals tended to focus their foraging activity on zones with particular temperature signatures. Along the Antarctic continent, some seals targeted colder waters on the sea bottom during benthic dives, while at the PF the favourable zones tended to be warmer. The possible negative effect of colder waters in Antarctic on the swimming performances of potential fish or squid prey could explain the behaviour of elephant seals in these zones, while warmer waters within the PF could correspond to the optimal conditions for potential myctophid prey of elephant seals.

Estimates of the Southern Ocean general circulation improved by animal‐borne instruments

Over the last decade, several hundred seals have been equipped with conductivity-temperature-depth sensors in the Southern Ocean for both biological and physical oceanographic studies. A calibrated collection of seal-derived hydrographic data is now available, consisting of more than 165,000 profiles. The value of these hydrographic data within the existing Southern Ocean observing system is demonstrated herein by conducting two state estimation experiments, differing only in the use or not of seal data to constrain the system. Including seal-derived data substantially modifies the estimated surface mixed-layer properties and circulation patterns within and south of the Antarctic Circumpolar Current. Agreement with independent satellite observations of sea ice concentration is improved, especially along the East Antarctic shelf. Instrumented animals efficiently reduce a critical observational gap, and their contribution to monitoring polar climate variability will continue to grow as data accuracy and spatial coverage increase.

Eddy overturning of the Antarctic Slope Front controls glacial melting in the Eastern Weddell Sea

[1] The Eastern Weddell Sea is characterized by narrow continental shelves and Warm Deep Water (WDW) is located in close proximity to the ice shelves in this region. The exchange of WDW across the Antarctic Slope Front (ASF) determines the rate of basal ice shelf melting. Here, we present a unique data set consisting of 2351 vertical profiles of temperature and salinity collected by southern elephant seals (Mirounga leonina) and a profile beneath the Fimbul Ice Shelf (FIS), obtained via drilling through 395 m of ice. This data set reveals variations in salinity and temperature through winter, and using a conceptual model of the coastal salt budget we quantify the main exchange processes. Our data show that modified WDW, with temperatures below −1.5°C, is advected onto the shelf and into the ice shelf cavities by an eddy overturning of the ASF. The onshore Ekman flux of surface waters during summer is the main source of freshwater that leads to the formation of low salinity shelf waters in the region. The modified WDW that reaches beneath the ice shelves is too cold for basal ice shelf melting to create such low salinity water. A high‐resolution model of an idealized ASF–continental shelf–ice shelf system supports the conclusions from the data analysis. The inflow of WDW onto the continental shelf and into the ice shelf cavity occurs within a bottom boundary layer where the eddy advection in the model is particularly strong, in close agreement with the observed vertical profile of temperature beneath the FIS. Citation: Nost, O. A., M. Biuw, V. Tverberg, C. Lydersen, T. Hattermann, Q. Zhou, L. H. Smedsrud, and K. M. Kovacs (2011), Eddy overturning of the Antarctic Slope Front controls glacial melting in the Eastern Weddell Sea, J. Geophys. Res., 116, C11014, doi:10.1029/2011JC006965.

Effects of Hydrographic Variability on the Spatial, Seasonal and Diel Diving Patterns of Southern Elephant Seals in the Eastern Weddell Sea

Weddell Sea hydrography and circulation is driven by influx of Circumpolar Deep Water (CDW) from the Antarctic Circumpolar Current (ACC) at its eastern margin. Entrainment and upwelling of this high-nutrient, oxygen-depleted water mass within the Weddell Gyre also supports the mesopelagic ecosystem within the gyre and the rich benthic community along the Antarctic shelf. We used Conductivity-Temperature-Depth Satellite Relay Data Loggers (CTD-SRDLs) to examine the importance of hydrographic variability, ice cover and season on the movements and diving behavior of southern elephant seals in the eastern Weddell Sea region during their overwinter feeding trips from Bouvetøya. We developed a model describing diving depth as a function of local time of day to account for diel variation in diving behavior. Seals feeding in pelagic ice-free waters during the summer months displayed clear diel variation, with daytime dives reaching 500-1500 m and night-time targeting of the subsurface temperature and salinity maxima characteristic of CDW around 150–300 meters. This pattern was especially clear in the Weddell Cold and Warm Regimes within the gyre, occurred in the ACC, but was absent at the Dronning Maud Land shelf region where seals fed benthically. Diel variation was almost absent in pelagic feeding areas covered by winter sea ice, where seals targeted deep layers around 500–700 meters. Thus, elephant seals appear to switch between feeding strategies when moving between oceanic regimes or in response to seasonal environmental conditions. While they are on the shelf, they exploit the locally-rich benthic ecosystem, while diel patterns in pelagic waters in summer are probably a response to strong vertical migration patterns within the copepod-based pelagic food web. Behavioral flexibility that permits such switching between different feeding strategies may have important consequences regarding the potential for southern elephant seals to adapt to variability or systematic changes in their environment resulting from climate change.

Ecosystem Impacts of a Range Expanding Forest Defoliator at the Forest-Tundra Ecotone

Insect outbreaks in northern-boreal forests are expected to intensify owing to climate warming, but our understanding of direct and cascading impacts of insect outbreaks on forest ecosystem functioning is deficient. The duration and severity of outbreaks by geometrid moths in northern Fennoscandian mountain birch forests have been shown to be accentuated by a recent climate-mediated range expansion, in particular of winter moth (Operophtera brumata). Here, we assess the effect of moth outbreak severity, quantified from satellite-based defoliation maps, on the state of understory vegetation and the abundance of key vertebrate herbivores in mountain birch forest in northern Norway. We show that the most recent moth outbreak caused a regional-scale state change to the understory vegetation, mainly due to a shift in dominance from the allelopathic and unpalatable dwarf-shrub Empetrum nigrum to the productive and palatable grass Avenella flexuosa. Both these central understory plant species responded significantly and nonlinearly to increasing outbreak severity. We further provide evidence that the effects of the outbreak on understory vegetation cascaded to cause strong but opposite impacts on the abundance of the two most common herbivore groups. Rodents increased with defoliation, largely mirroring the increase in A. flexuosa, whereas ungulate abundance instead showed a decreasing trend. Our analyses also suggest that the response of understory vegetation to defoliation may depend on the initial state of the forest, with poorer forest types potentially allowing stronger responses to defoliation.

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

SUMMARY 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.

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

SUMMARY 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.