Crabeater seals (Lobodon carcinophaga) in the Weddell Sea during DRE1998 campaign

The crabeater seal (Lobodon carcinophaga) is the most abundant Antarctic seal and inhabits the circumpolar pack ice zone of the Southern Ocean. Until now, information on important environmental factors affecting its distribution as well as on foraging behaviour is limited. In austral summer 1998, 12 crabeater seals of both sexes and different age classes were equipped with satellitelinked dive recorders at Drescher Inlet (72.85°S, 19.26°E), eastern Weddell Sea. To identify suitable habitat conditions within the Weddell Sea, a maximum entropy (Maxent) modelling approach was implemented. The model revealed that the eastern and southern Weddell Sea is especially suitable for crabeater seals. Distance to the continental shelf break and sea ice concentration were the two most important parameters in modelling species distribution throughout the study period. Model predictions demonstrated that crabeater seals showed a dynamic response to their seasonally changing environment emphasized by the favoured sea ice conditions. Crabeater seals utilized ice-free waters substantially, which is potentially explained by the comparatively low sea ice cover of the Weddell Sea during summer 1998. Diving behaviour was characterized by short (>90 % = 0-4 min) and shallow (>90 % = 0-51 m) dives. This pattern reflects the typical summer and autumn foraging behaviour of crabeater seals. Both the distribution and foraging behaviour corresponded well with the life history of the Antarctic krill (Euphausia superba), the preferred prey of crabeater seals. In general, predicted suitable habitat conditions were congruent with probable habitats of krill, which emphasizes the strong dependence on their primary prey.

Modelling calving front dynamics using a level-set method: application to Jakobshavn Isbræ, West Greenland, links model results in MATLAB format

Calving is a major mechanism of ice discharge of the Antarctic and Greenland ice sheets, and a change in calving front position affects the entire stress regime of marine terminating glaciers. The representation of calving front dynamics in a 2-D or 3-D ice sheet model remains non-trivial. Here, we present the theoretical and technical framework for a level-set method, an implicit boundary tracking scheme, which we implement into the Ice Sheet System Model (ISSM). This scheme allows us to study the dynamic response of a drainage basin to user-defined calving rates. We apply the method to Jakobshavn Isbræ, a major marine terminating outlet glacier of the West Greenland Ice Sheet. The model robustly reproduces the high sensitivity of the glacier to calving, and we find that enhanced calving triggers significant acceleration of the ice stream. Upstream acceleration is sustained through a combination of mechanisms. However, both lateral stress and ice influx stabilize the ice stream. This study provides new insights into the ongoing changes occurring at Jakobshavn Isbræ and emphasizes that the incorporation of moving boundaries and dynamic lateral effects, not captured in flow-line models, is key for realistic model projections of sea level rise on centennial timescales.

Surface velocity fields, digital elevation models and ice front positions derived from multi-mission SAR remote sensing data at Sjögren Inlet glaciers, Antarctic Peninsula

Substantial retreat or disintegration of numerous ice shelves have been observed on the Antarctic Peninsula. The ice shelf in the Prince Gustav Channel retreated gradually since the late 1980's and broke-up in 1995. Tributary glaciers reacted with speed-up, surface lowering and increased ice discharge, consequently contributing to sea level rise. We present a detailed long-term study (1993-2014) on the dynamic response of Sjögren Inlet glaciers to the disintegration of Prince Gustav Ice Shelf. We analyzed various remote sensing datasets to observe the reactions of the glaciers to the loss of the buttressing ice shelf. A strong increase in ice surface velocities was observed with maximum flow speeds reaching 2.82±0.48 m/d in 2007 and 1.50±0.32 m/d in 2004 at Sjögren and Boydell glaciers respectively. Subsequently, the flow velocities decelerated, however in late 2014, we still measured about two times the values of our first measurements in 1996. The tributary glaciers retreated 61.7±3.1 km² behind the former grounding line of the ice shelf. In regions below 1000 m a.s.l., a mean surface lowering of -68±10 m (-3.1 m/a) was observed in the period 1993-2014. The lowering rate decreased to -2.2 m/a in recent years. Based on the surface lowering rates, geodetic mass balances of the glaciers were derived for different time steps. High mass loss rate of -1.21±0.36 Gt/a was found in the earliest period (1993-2001). Due to the dynamic adjustments of the glaciers to the new boundary conditions the ice mass loss reduced to -0.59±0.11 Gt/a in the period 2012-2014, resulting in an average mass loss rate of -0.89±0.16 Gt/a (1993-2014). Including the retreat of the ice front and grounding line, a total mass change of -38.5±7.7 Gt and a contribution to sea level rise of 0.061±0.013 mm were computed. Analysis of the ice flux revealed that available bedrock elevation estimates at Sjögren Inlet are too shallow and are the major uncertainty in ice flux computations. This temporally dense time series analysis of Sjögren Inlet glaciers shows that the adjustments of tributary glaciers to ice shelf disintegration are still going on and provides detailed information of the changes in glacier dynamics.

Eddy covariance and environmental data of boreal bogs plant species

In boreal bogs plant species are low in number, but they differ greatly in their growth forms and photosynthetic properties. We assessed how ecosystem carbon (C) sink dynamics were affected by seasonal variations in photosynthetic rate and leaf area of different species. Photosynthetic properties (light-response parameters), leaf area development and areal cover (abundance) of the species were used to quantify species-specific net and gross photosynthesis rates (PN and PG, respectively), which were summed to express ecosystem-level PN and PG. The ecosystem-level PG was compared with a gross primary production (GPP) estimate derived from eddy covariance measurements (EC). Species areal cover rather than differences in photosynthetic properties determined the species with the highest PG of both vascular plants and Sphagna. Species-specific contributions to the ecosystem PG varied over the growing season, which in turn determined the seasonal variation in ecosystem PG. The upscaled growing-season PG estimate, 230 g C/m**2, agreed well with the GPP estimated by the EC, 243 g C/m**2. Sphagna were superior to vascular plants in ecosystem-level PG throughout the growing season but had a lower PN. PN results indicated that areal cover of the species together with their differences in photosynthetic parameters shape the ecosystem-level C balance. Species with low areal cover but high photosynthetic efficiency appear to be potentially important for the ecosystem C sink. Results imply that functional diversity may increase the stability of C sink of boreal bogs.

The benthic foraminiferal community in a naturally CO2-rich coastal habitat of the southwestern Baltic Sea

It is expected that the calcification of foraminifera will be negatively affected by the ongoing acidification of the oceans. Compared to the open oceans, these organisms are subjected to much more adverse carbonate system conditions in coastal and estuarine environments such as the southwestern Baltic Sea, where benthic foraminifera are abundant. This study documents the seasonal changes of carbonate chemistry and the ensuing response of the foraminiferal community with bi-monthly resolution in Flensburg Fjord. In comparison to the surface pCO2, which is close to equilibrium with the atmosphere, we observed large seasonal fluctuations of pCO2 in the bottom and sediment pore waters. The sediment pore water pCO2 was constantly high during the entire year ranging from 1244 to 3324 µatm. Nevertheless, in contrast to the bottom water, sediment pore water was slightly supersaturated with respect to calcite as a consequence of higher alkalinity (AT) for most of the year. Foraminiferal assemblages were dominated by two calcareous species, Ammonia aomoriensis and Elphidium incertum, and the agglutinated Ammotium cassis. The one-year cycle was characterised by seasonal community shifts. Our results revealed that there is no dynamic response of foraminiferal population density and diversity to elevated sediment pore water pCO2. Surprisingly, the fluctuations of sediment pore water undersaturation (Omega calc) co-vary with the population densities of living Ammonia aomoriensis. Further, we observed that most of the tests of living calcifying foraminifera were intact. Only Ammonia aomorienis showed dissolution and recalcification structures on the tests, especially at undersaturated conditions. Therefore, the benthic community is subjected to high pCO2 and tolerates elevated levels as long as sediment pore water remains supersaturated. Model calculations inferred that increasing atmospheric CO2 concentrations will finally lead to a perennial undersaturation in sediment pore waters. Whereas benthic foraminifera indeed may cope with a high sediment pore water pCO2, the steady undersaturation of sediment pore waters would likely cause a significant higher mortality of the dominating Ammonia aomoriensis. This shift may eventually lead to changes in the benthic foraminiferal communities in Flensburg Fjord, as well as in other regions experiencing naturally undersaturated Omega calc levels.

Mean dynamic topography and oceanographic parameters estimated from an inverse model and satellite geodesy, with link to model result in one single NetCDF file (392 MB), including the inverse data error covariance

Geostrophic surface velocities can be derived from the gradients of the mean dynamic topography-the difference between the mean sea surface and the geoid. Therefore, independently observed mean dynamic topography data are valuable input parameters and constraints for ocean circulation models. For a successful fit to observational dynamic topography data, not only the mean dynamic topography on the particular ocean model grid is required, but also information about its inverse covariance matrix. The calculation of the mean dynamic topography from satellite-based gravity field models and altimetric sea surface height measurements, however, is not straightforward. For this purpose, we previously developed an integrated approach to combining these two different observation groups in a consistent way without using the common filter approaches (Becker et al. in J Geodyn 59(60):99-110, 2012, doi:10.1016/j.jog.2011.07.0069; Becker in Konsistente Kombination von Schwerefeld, Altimetrie und hydrographischen Daten zur Modellierung der dynamischen Ozeantopographie, 2012, http://nbn-resolving.de/nbn:de:hbz:5n-29199). Within this combination method, the full spectral range of the observations is considered. Further, it allows the direct determination of the normal equations (i.e., the inverse of the error covariance matrix) of the mean dynamic topography on arbitrary grids, which is one of the requirements for ocean data assimilation. In this paper, we report progress through selection and improved processing of altimetric data sets. We focus on the preprocessing steps of along-track altimetry data from Jason-1 and Envisat to obtain a mean sea surface profile. During this procedure, a rigorous variance propagation is accomplished, so that, for the first time, the full covariance matrix of the mean sea surface is available. The combination of the mean profile and a combined GRACE/GOCE gravity field model yields a mean dynamic topography model for the North Atlantic Ocean that is characterized by a defined set of assumptions. We show that including the geodetically derived mean dynamic topography with the full error structure in a 3D stationary inverse ocean model improves modeled oceanographic features over previous estimates.

(Table S1) Boron isotope and trace element data and reconstructed carbonate system parameters, ODP Holes 122-761B and 154-926A

The middle Miocene Climatic Optimum (17-15 Ma; MCO) is a period of global warmth and relatively high CO2 and is thought to be associated with a significant retreat of the Antarctic Ice Sheet (AIS). We present here a new planktic foraminiferal d11B record from 16.6 to 11.8 Ma from two deep ocean sites currently in equilibrium with the atmosphere with respect to CO2. These new data demonstrate that the evolution of global climate during the middle Miocene (as reflected by changes in the cyrosphere) was well correlated to variations in the concentration of atmospheric CO2. What is more, within our sampling resolution (~1 sample per 300 kyr) there is no evidence of hysteresis in the response of ice volume to CO2 forcing during the middle Miocene, contrary to what is understood about the Antarctic Ice Sheet from ice sheet modelling studies. In agreement with previous data, we show that absolute levels of CO2 during the MCO were relatively modest (350-400 ppm) and levels either side of the MCO are similar or lower than the pre-industrial (200-260 ppm). These new data imply the presence of either a very dynamic AIS at relatively low CO2 during the middle Miocene or the advance and retreat of significant northern hemisphere ice. Recent drilling on the Antarctic margin and shore based studies indicate significant retreat and advance beyond the modern limits of the AIS did occur during the middle Miocene, but the complete loss of the AIS was unlikely. Consequently, it seems that ice volume and climate variations during the middle Miocene probably involved a more dynamic AIS than the modern but also some component of land-based ice in the northern hemisphere.

Biotic and abiotic parameters and cynobacteria abundance during an experimental set-up with varying pCO2 conditions

Heterocystous cyanobacteria of the genus Nodularia form extensive blooms in the Baltic Sea and contribute substantially to the total annual primary production. Moreover, they dispense a large fraction of new nitrogen to the ecosystem when inorganic nitrogen concentration in summer is low. Thus, it is of ecological importance to know how Nodularia will react to future environmental changes, in particular to increasing carbon dioxide (CO2) concentrations and what consequences there might arise for cycling of organic matter in the Baltic Sea. Here, we determined carbon (C) and dinitrogen (N2) fixation rates, growth, elemental stoichiometry of particulate organic matter and nitrogen turnover in batch cultures of the heterocystous cyanobacterium Nodularia spumigena under low (median 315 µatm), mid (median 353 µatm), and high (median 548 µatm) CO2 concentrations. Our results demonstrate an overall stimulating effect of rising pCO2 on C and N2 fixation, as well as on cell growth. An increase in pCO2 during incubation days 0 to 9 resulted in an elevation in growth rate by 84 ± 38% (low vs. high pCO2) and 40 ± 25% (mid vs. high pCO2), as well as in N2 fixation by 93 ± 35% and 38 ± 1%, respectively. C uptake rates showed high standard deviations within treatments and in between sampling days. Nevertheless, C fixation in the high pCO2 treatment was elevated compared to the other two treatments by 97% (high vs. low) and 44% (high vs. mid) at day 0 and day 3, but this effect diminished afterwards. Additionally, elevation in carbon to nitrogen and nitrogen to phosphorus ratios of the particulate biomass formed (POC : POP and PON : POP) was observed at high pCO2. Our findings suggest that rising pCO2 stimulates the growth of heterocystous diazotrophic cyanobacteria, in a similar way as reported for the non-heterocystous diazotroph Trichodesmium. Implications for biogeochemical cycling and food web dynamics, as well as ecological and socio-economical aspects in the Baltic Sea are discussed.

(Table 1) Stage of the lemming cycle and nesting parameters of the greater snow geese between 1996-2005, Bylot Island

Resource pulses are common in various ecosystems and often have large impacts on ecosystem functioning. Many animals hoard food during resource pulses, yet how this behaviour affects pulse diffusion through trophic levels is poorly known because of a lack of individual-based studies. Our objective was to examine how the hoarding behaviour of arctic foxes (Alopex lagopus) preying on a seasonal pulsed resource (goose eggs) was affected by annual and seasonal changes in resource availability. We monitored foraging behaviour of foxes in a greater snow goose (Chen caerulescens atlanticus) colony during 8 nesting seasons that covered 2 lemming cycles. The number of goose eggs taken and cached per hour by foxes declined 6-fold from laying to hatching, while the proportion of eggs cached remained constant. In contrast, the proportion of eggs cached by foxes fluctuated in response to the annual lemming cycle independently of the seasonal pulse of goose eggs. Foxes cached the majority of eggs taken (> 90%) when lemming abundance was high or moderate but only 40% during the low phase of the cycle. This likely occurred because foxes consumed a greater proportion of goose eggs to fulfill their energy requirement at low lemming abundance. Our study clearly illustrates a behavioural mechanism that extends the energetic benefits of a resource pulse. The hoarding behaviour of the main predator enhances the allochthonous nutrients input brought by migrating birds from the south into the arctic terrestrial ecosystem. This could increase average predator density and promote indirect interactions among prey.

Shape-invariant multibeam backscatter data of the eastern slope of the Porcupine Seabight (50 m grid size)

In this study multibeam angular backscatter data acquired in the eastern slope of the Porcupine Seabight are analysed. Processing of the angular backscatter data using the 'NRGCOR' software was made for 29 locations comprising different geological provinces like: carbonate mounds, buried mounds, seafloor channels, and inter-channel areas. A detailed methodology is developed to produce a map of angle-invariant (normalized) backscatter data by correcting the local angular backscatter values. The present paper involves detailed processing steps and related technical aspects of the normalization approach. The presented angle-invariant backscatter map possesses 12 dB dynamic range in terms of grey scale. A clear distinction is seen between the mound dominated northern area (Belgica province) and the Gollum channel seafloor at the southern end of the site. Qualitative analyses of the calculated mean backscatter values i.e., grey scale levels, utilizing angle-invariant backscatter data generally indicate backscatter values are highest (lighter grey scale) in the mound areas followed by buried mounds. The backscatter values are lowest in the inter-channel areas (lowest grey scale level). Moderate backscatter values (medium grey level) are observed from the Gollum and Kings channel data, and significant variability within the channel seafloor provinces. The segmentation of the channel seafloor provinces are made based on the computed grey scale levels for further analyses based on the angular backscatter strength. Three major parameters are utilized to classify four different seafloor provinces of the Porcupine Seabight by employing a semi-empirical method to analyse multibeam angular backscatter data. The predicted backscatter response which has been computed at 20° is the highest for the mound areas. The coefficient of variation (CV) of the mean backscatter response is also the highest for the mound areas. Interestingly, the slope value of the buried mound areas are found to be the highest. However, the channel seafloor of moderate backscatter response presents the lowest slope and CV values. A critical examination of the inter-channel areas indicates less variability within the estimated three parameters.

Impacts of passive warming chambers on soil parameters in Antarctica

Passive chambers are used to examine the impacts of summer warming in Antarctica but, so far, impacts occurring outside the growing season, or related to extreme temperatures, have not been reported, despite their potentially large biological significance. In this review, we synthesise and discuss the microclimate impacts of passive warming chambers (closed, ventilated and Open Top Chamber-OTC) commonly used in Antarctic terrestrial habitats, paying special attention to seasonal warming, during the growing season and outside, extreme temperatures and freeze-thaw events. Both temperature increases and decreases were recorded throughout the year. Closed chambers caused earlier spring soil thaw (8-28 days) while OTCs delayed soil thaw (3-13 days). Smaller closed chamber types recorded the largest temperature extremes (up to 20°C higher than ambient) and longest periods (up to 11 h) of above ambient extreme temperatures, and even OTCs had above ambient temperature extremes over up to 5 consecutive hours. The frequency of freeze-thaw events was reduced by ~25%. All chamber types experienced extreme temperature ranges that could negatively affect biological responses, while warming during winter could result in depletion of limited metabolic resources. The effects outside the growing season could be as important in driving biological responses as the mean summer warming. We make suggestions for improving season-specific warming simulations and propose that seasonal and changed temperature patterns achieved under climate manipulations should be recognised explicitly in descriptions of treatment effects.

Ocean acidification decreases the light-use efficiency in an Antarctic diatom under dynamic but not constant light, link to supplementary data

There is increasing evidence that different light intensities strongly modulate the effects of ocean acidification (OA) on marine phytoplankton. The aim of the present study was to investigate interactive effects of OA and dynamic light, mimicking natural mixing regimes. The Antarctic diatom Chaetoceros debilis was grown under two pCO2 (390 and 1000 latm) and light conditions (constant and dynamic), the latter yielding the same integrated irradiance over the day. To characterize interactive effects between treatments, growth, elemental composition, primary production and photophysiology were investigated. Dynamic light reduced growth and strongly altered the effects of OA on primary production, being unaffected by elevated pCO2 under constant light, yet significantly reduced under dynamic light. Interactive effects between OA and light were also observed for Chl production and particulate organic carbon (POC) quotas. Response patterns can be explained by changes in the cellular energetic balance. While the energy transfer efficiency from photochemistry to biomass production (Phi_e,C) was not affected by OA under constant light, it was drastically reduced under dynamic light. Contrasting responses under different light conditions need to be considered when making predictions regarding a more stratified and acidified future ocean.

Global Ocean Particulate Organic Carbon flux merged with satellite parameters

The efficiency of the biological pump of carbon to the deep ocean depends largely on the biologically mediated export of carbon from the surface ocean and its remineralization with depth. Global satellite studies have primarily focused on chlorophyll concentration and net primary production (NPP) to understand the role of phytoplankton in these processes. Recent satellite retrievals of phytoplankton composition now allow for the size of phytoplankton cells to be considered. Here, we improve understanding of phytoplankton size structure impacts on particle export, remineralization and transfer. Particulate organic carbon (POC) flux observations from sediment traps and 234Th are compiled across the global ocean. Annual climatologies of NPP, percent microplankton, and POC flux at four time series locations and within biogeochemical provinces are constructed, and sinking velocities are calculated to align surface variables with POC flux at depth. Parameters that characterize POC flux vs. depth (export flux ratio, labile fraction, remineralization length scale) are then fit to the aligned dataset. Times of the year dominated by different size compositions are identified and fit separately in regions of the ocean where phytoplankton cell size showed enough dynamic range over the annual cycle. Considering all data together, our findings support the paradigm of high export flux but low transfer efficiency in more productive regions and vice versa for oligotrophic regions. However, when parsing by dominant size class, we find periods dominated by small cells to have both greater export flux and lower transfer efficiency than periods when large cells comprise a greater proportion of the phytoplankton community.

Shotgun proteomics reveals physiological response to ocean acidification in Crassostrea gigas

Background. Ocean acidification as a result of increased anthropogenic CO2 emissions is occurring in marine and estuarine environments worldwide. The coastal ocean experiences additional daily and seasonal fluctuations in pH that can be lower than projected end of century open ocean pH reductions. Projected and current ocean acidification have wide-ranging effects on many aquatic organisms, however the exact mechanisms of the impacts of ocean acidification on many of these animals remains to be characterized. Methods. In order to assess the impact of ocean acidification on marine invertebrates, Pacific oysters (Crassostrea gigas) were exposed to one of four different pCO2 levels for four weeks: 400 µatm (pH 8.0), 800 µatm (pH 7.7), 1000 µatm (pH 7.6), or 2800 µatm (pH 7.3). At the end of 4 weeks a variety of physiological parameters were measured to assess the impacts of ocean acidification: tissue glycogen content and fatty acid profile, shell micromechanical properties, and response to acute heat shock. To determine the effects of ocean acidification on the underlying molecular physiology of oysters and their stress response, some of the oysters from 400 µatm and 2800 µatm were exposed to an additional mechanical stress and shotgun proteomics were done on oysters from high and low pCO2 and from with and without mechanical stress. Results. At the end of the four week exposure period, oysters in all four pCO2 environments deposited new shell, but growth rate was not different among the treatments. However, micromechanical properties of the new shell were compromised by elevated pCO2. Elevated pCO2 affected neither whole body fatty acid composition, nor glycogen content, nor mortality rate associated with acute heat shock. Shotgun proteomics revealed that several physiological pathways were significantly affected by ocean acidification, including antioxidant response, carbohydrate metabolism, and transcription and translation. Additionally, the proteomic response to a second stress differed with pCO2, with numerous processes significantly affected by mechanical stimulation at high versus low pCO2 (all proteomics data are available in the ProteomeXchange under the identifier PXD000835). Discussion. Oyster physiology is significantly altered by exposure to elevated pCO2, indicating changes in energy resource use. This is especially apparent in the assessment of the effects of pCO2 on the proteomic response to a second stress. The altered stress response illustrates that ocean acidification may impact how oysters respond to other changes in their environment. These data contribute to an integrative view of the effects of ocean acidification on oysters as well as physiological trade-offs during environmental stress.

Regulation of intracellular pH in cnidarians: response to acidosis in Anemonia viridis

The regulation of intracellular pH (pHi) is a fundamental aspect of cell physiology that has received little attention in studies of the phylum Cnidaria, which includes ecologically important sea anemones and reef-building corals. Like all organisms, cnidarians must maintain pH homeostasis to counterbalance reductions in pHi, which can arise because of changes in either intrinsic or extrinsic parameters. Corals and sea anemones face natural daily changes in internal fluids, where the extracellular pH can range from 8.9 during the day to 7.4 at night. Furthermore, cnidarians are likely to experience future CO2-driven declines in seawater pH, a process known as ocean acidification. Here, we carried out the first mechanistic investigation to determine how cnidarian pHi regulation responds to decreases in extracellular and intracellular pH. Using the anemone Anemonia viridis, we employed confocal live cell imaging and a pH-sensitive dye to track the dynamics of pHi after intracellular acidosis induced by acute exposure to decreases in seawater pH and NH4Cl prepulses. The investigation was conducted on cells that contained intracellular symbiotic algae (Symbiodinium sp.) and on symbiont-free endoderm cells. Experiments using inhibitors and Na-free seawater indicate a potential role of Na/H plasma membrane exchangers (NHEs) in mediating pHi recovery following intracellular acidosis in both cell types. We also measured the buffering capacity of cells, and obtained values between 20.8 and 43.8 mM per pH unit, which are comparable to those in other invertebrates. Our findings provide the first steps towards a better understanding of acid-base regulation in these basal metazoans, for which information on cell physiology is extremely limited.