Pteropods in Southern Ocean ecosystems

To date, little research has been carried out on pelagic gastropod molluscs (pteropods) in Southern Ocean ecosystems. However, recent predictions are that, due to acidification resulting from a business as usual approach to CO₂ emissions (IS92a), Southern Ocean surface waters may begin to become uninhabitable for aragonite shelled thecosome pteropods by 2050. To gain insight into the potential impact that this would have on Southern Ocean ecosystems, we have here synthesized available data on pteropod distributions and densities, assessed current knowledge of pteropod ecology, and highlighted knowledge gaps and directions for future research on this zooplankton group.Six species of pteropod are typical of the Southern Ocean south of the Sub-Tropical Convergence, including the four Thecosomes Limacina helicina antarctica, Limacina retroversa australis, Clio pyramidata, and Clio piatkowskii, and two Gymnosomes Clione limacina antarctica and Spongiobranchaea australis. Limacina retroversa australis dominated pteropod densities north of the Polar Front (PF), averaging 60 ind m⁻³ (max = 800 ind m⁻³) and 11% of total zooplankton at the Prince Edward Islands. South of the PF L. helicina antarctica predominated, averaging 165 ind m⁻³ (max = 2681 ind m⁻³) and up to >35% of total zooplankton at South Georgia, and up to 1397 ind m⁻³ and 63% of total zooplankton in the Ross Sea. Combined pteropods contributed <5% to total zooplankton in the Lazarev Sea, but 15% (max = 93%) to macrozooplankton in the East Antarctic. In addition to regional density distributions we have synthesized data on vertical distributions, seasonal cycles, and inter-annual density variation.Trophically, gymnosome are specialist predators on thecosomes, while thecosomes are considered predominantly herbivorous, capturing food with a mucous web. The ingestion rates of L. retroversa australis are in the upper range for sub-Antarctic mesozooplankton (31.2-4196.9 ng pig ind⁻¹ d⁻¹), while those of L. helicina antarctica and C. pyramidata are in the upper range for all Southern Ocean zooplankton, in the latter species reaching 27,757 ng pig ind⁻¹ d⁻¹ and >40% of community grazing impact. Further research is required to quantify diet selectivity, the effect of phytoplankton composition on growth and reproductive success, and the role of carnivory in thecosomes.Life histories are a significant knowledge gap for Southern Ocean pteropods, a single study having been completed for L. retroversa australis, making population studies a priority for this group. Pteropods appear to be important in biogeochemical cycling, thecosome shells contributing >50% to carbonate flux in the deep ocean south of the PF. Pteropods may also contribute significantly to organic carbon flux through the production of fast sinking faecal pellets and mucous flocs, and rapid sinking of dead animals ballasted by their aragonite shells. Quantification of these contributions requires data on mucous web production rates, egestion rates, assimilation efficiencies, metabolic rates, and faecal pellet morphology for application to sediment trap studies.Based on the available data, pteropods are regionally significant components of the Southern Ocean pelagic ecosystem. However, there is an urgent need for focused research on this group in order to quantify how a decline in pteropod densities may impact on Southern Ocean ecosystems.     

Pristine or altered: low-Mg calcite shells survived from massive dolomitization? A case study from Miocene carbonates

Low-Mg calcite shells have been widely used to reconstruct the chemistry of ancient seawater. There is always a question: are the shells chemically pristine? This paper presents the isotope and elemental geochemistry of low-Mg calcite bivalve shells in late Miocene platform carbonates, SE Spain. The platform carbonates were extensively dolomitized, and limestone is restricted to older stratal units, and to units mainly in topographically higher and more landward strata. Low-Mg calcite oyster shells were completely dissolved out in the basinward dolomite, but are well preserved in the limestone. These shells appear to retain the original growth microstructures, based on hand samples. Under the microscope, however, dissolution and recrystallization, as well as pristine growth lines are all present. Sr isotopes in these shells range from that of normal Miocene seawater to radiogenic values. δ¹⁸O and δ¹³C values, and Mg, Sr, and Na concentrations in these shells are rather variable. The high end members are consistent with the typical values of modern/late Miocene normal-seawater low-Mg calcite shells, whereas the low end members are close to those of diagenetic calcite cements, which have low δ¹⁸O, δ¹³C, Sr and Na values, and radiogenic Sr. The Nijar shells were altered physically and chemically to different degrees by diagenesis, although these shells are consistent with some “criteria” of unalteration. The isotopic and trace-element data collected in altered and pristine (or less altered) portions coexisting in the same shells are clearly differentiable. Quantitative simulation of covariations of geochemical pairs indicates that solid mixing of unaltered and altered portions by sampling is consistent with the variations in isotopic and elemental data recorded in the Nijar shells. The geological significance of this study is that ancient fabric-retentive calcite shells may have been altered geochemically although they may appear pristine. Calcite shells that underwent intensive diagenesis should be examined rigorously under the microscope, coupled with investigations of multiple geochemical proxies to assess chemical alteration. Only the data of unaltered shells can be used to reconstruct the chemistry of ancient seawater.     

Gradients in coral reef communities exposed to muddy river discharge in Pohnpei,Micronesia

This study analyzed how coral communities change along a gradient of increasing exposure to a mud-discharging river in the Enipein Catchment, Pohnpei, Micronesia. Using video transects, we quantified benthic communities at five sites along a gradient moving away from the river mouth towards the barrier reef. The most river-impacted site was characterized by a high accumulation of mud, low coral cover and low coral diversity. Although coral cover leveled off at ∼400 m from the river mouth to values found at the outer-most sites, coral diversity continued to increase with increasing distance, suggesting that the most distant site was still impacted by the river discharges. Fungiidae, Pavona, Acropora, Pachyseris and Porites rus all significantly increased in cover with distance from the river, while Turbinaria decreased. The combined presence and abundance of these six species groups, together with coral species richness, may help to indicate the effects of terrestrial runoff in similar runoff-exposed settings around Micronesia, whereas coral cover is not a sensitive indicator for river impact. Coral reefs are important resources for the people of Pohnpei. To prevent further degradation of this important resource, an integrated watershed approach is needed to control terrestrial activities.     

The effect of a topographic high on the morphological stability of a two-inlet bay system

The cross-sectional stability of two tidal inlets connecting the same back-barrier lagoon to the ocean is investigated. The condition for equilibrium is the cross-sectional area tidal prism relationship. In an earlier study [Van de Kreeke, J., 1990. Can multiple inlets be stable? Estuarine, Coastal and Shelf Science 30: 261–273.], using the same equilibrium condition, it was concluded that where two inlets connect the same basin to the ocean ultimately one inlet will close. One of the major assumptions in that study was that the water level in the basin fluctuated uniformly. In hindsight this assumption might be too restrictive. For example, in the Wadden Sea the back barrier lagoon consists of a series of basins, rather than one single basin, separated by topographic highs. These topographic highs limit but do not exclude the exchange of water between the sub-basins. For this reason in the present study, a topographic high in the form of a weir was added, separating the back-barrier lagoon in two sub-basins. The water level in the sub-basins, rather than in the back-barrier as a whole, is assumed to fluctuate uniformly. For this schematization the hydrodynamic equations are solved using a finite difference method. The results, together with the equilibrium condition, yield the equilibrium flow curve for each of the inlets. The intersections of the two equilibrium flow curves represent combinations of cross-sectional areas for which both inlets are in equilibrium. The stability of the equilibriums was investigated using a non-linear stability analysis resulting in a flow diagram. Calculations were carried out for different inlet and weir characteristics. Sinussoidal tides were the same for both inlets. The results show that for relatively large wetted cross-sectional areas over the topographic high, approaching the situation of a single basin, there are no combinations of inlet cross-sectional areas for which both inlets are in a stable equilibrium. This supports the conclusion in the earlier study. For relatively small wetted cross-sectional areas over the topographic high there is one set of stable equilibriums. In that case the two-inlet bay system approaches that of two single-inlet bay systems.     

Sampling the vertical particle flux in the upper water column using a large diameter free-drifting NetTrap adapted to an Indented Rotating Sphere sediment trap

Further development of the large, surface-tethered sediment trap (NetTrap) employed as part of the MedFlux program is described whereby the large collection capacity of the NetTrap is combined with an Indented Rotating Sphere/Sample Carousel (IRSC) sediment trap (IRSC–NT). This trap is capable of collecting particle flux either in a time series or settling velocity mode; settling velocity mode allows the collection of particles that fall within discrete settling velocity intervals. During short field deployments in the Mediterranean Sea the IRSC–NT configured in the settling velocity mode successfully collected unpoisoned samples for chemical and microbiological experiments. In addition to the development of the IRSC–NT, particle-settling behavior above and below the swimmer-excluding IRS valve was tested during on-deck experiments using a specially constructed water-tight trap. Chemical analyses of settling materials (published elsewhere) suggested that separation of particles by settling velocity was achieved. However, due to the motion of the ship, it was not possible to directly measure particle-settling velocities within the trap. Particle release from the IRS did not bias the apparent settling velocity spectrum. Rotation of the IRS did not engender turbulence at the surface of the sphere or within the skewed funnel below. Tests of different ball designs over the course of the MedFlux program showed that a “ridge and saddle” pattern was optimal for efficiently transferring particles under the IRS seal while still reducing swimmer entrance to the collection funnel. The large size of the IRSC–NT did not prevent it from drifting effectively with the current. Several modifications of the present design are proposed that should improve the accuracy of the settling velocity measurements.     

Composition of ferromanganese nodules from Riga Bay (Baltic Sea)

An average sample of ferromanganese nodules from Riga Bay was investigated by analytical electron microscopy coupled with atomic absorption, neutron activation, and ICP-MS methods. It is established that the nodules consist of colloform and poorly crystallized Fe and Mn hydroxides with an admixture of dispersed inclusions of better crystallized particles allowing revealing their mineralogical nature by the microdiffraction method. The major mineral phases are ferroxigite and Fe-vernadite with subordinate magnetite, asbolane, and todorokite. The chemical composition of the nodules is dominated by iron followed by cobalt and lead, while nickel, molybdenum, and tungsten associate with the manganese phase. The concentration of several minor elements in the Riga Bay nodules is slightly lower as compared with their average values for the entire Baltic Sea; some elements such as silver, bismuth, cadmium, niobium, tin, and thallium were never analyzed before.     

Fundamental study for morphodynamic modelling: Sand mounds in oscillatory flows

Experiments on sand mounds in oscillatory flow, undertaken in controlled, large-scale laboratory conditions, have produced well-defined data sets for model comparison. Three bathymetries with different levels of submergence, including a surface-piercing case, were tested. The maximum slope was about 1:5.5. Sediment transport is due to bed load with ripple formation. The principal time-dependent bulk parameters are the vertical distance of the centre of gravity above the base and the volume of the mound. A semi-implicit finite-volume depth-averaged hydrodynamic model is used to drive morphodynamics, using van Rijn's sediment flux model generalized to take account of bed slope, and some justification is given for depth-averaged modeling in these conditions. Starting the model runs with the conditions at the end of the first cycle avoided initial atypical physical behaviour. In general good predictions were obtained with an angle of repose reduced from the standard value of about 30° for stationary beds to 15°. For these situations, morphodynamics was largely unaffected by a hydrodynamic roughness height in the range 2.5D₅₀ to 51D₅₀, with larger values accounting for ripple roughness. The reduced angle of repose may be physically expected with mobile beds but this specific value is only expected to be suited to this form of bed motion. In one case an exaggerated ripple formed near the top of the mound reducing agreement with experiment. For the submerged case with normal ripple structure excellent predictions were obtained. For the initially surface-piercing mound, the time of submergence was better predicted with a 30° angle of repose, presumably due to the prominent influence of the near stationary bed near the wet/dry interface, although long term predictions were better predicted with 15°. The occurrence of vortex shedding in the first cycle modeled was in agreement with experimental observation.     

Experimental investigation of water impact on axisymmetric bodies

The results of an elaborate experimental investigation on bottom slamming of axisymmetric objects are presented. Drop tests have been performed on a hemisphere and two conical shapes with different deadrise angles. The test setup is designed so as to prevent small rotations of the test objects which cause scatter in the measurement data. The pressure distribution and evolution as well as the body motion parameters are measured during impact. By means of a high speed camera the water uprise is visualized and the wetting factor is determined for the cones. The results are compared with a three-dimensional asymptotic theory for axisymmetric rigid bodies with constant entry velocity. The ratio between the registered peak pressures and the asymptotic theory are in accordance with comparable experiments in the literature. The asymptotic theory, however, is found to be quite conservative, since the measured peak pressure levels appear to be approximately 50% to 75% of the theoretical levels.     

Reevaluation of historical ocean heat content variations with time-varying XBT and MBT depth bias corrections

As reported in former studies, temperature observations obtained by expendable bathythermographs (XBTs) and mechanical bathythermographs (MBTs) appear to have positive biases as much as they affect major climate signals. These biases have not been fully taken into account in previous ocean temperature analyses, which have been widely used to detect global warming signals in the oceans. This report proposes a methodology for directly eliminating the biases from the XBT and MBT observations. In the case of XBT observation, assuming that the positive temperature biases mainly originate from greater depths given by conventional XBT fall-rate equations than the truth, a depth bias equation is constructed by fitting depth differences between XBT data and more accurate oceanographic observations to a linear equation of elapsed time. Such depth bias equations are introduced separately for each year and for each probe type. Uncertainty in the gradient of the linear equation is evaluated using a non-parametric test. The typical depth bias is +10 m at 700 m depth on average, which is probably caused by various indeterminable sources of error in the XBT observations as well as a lack of representativeness in the fall-rate equations adopted so far. Depth biases in MBT are fitted to quadratic equations of depth in a similar manner to the XBT method. Correcting the historical XBT and MBT depth biases by these equations allows a historical ocean temperature analysis to be conducted. In comparison with the previous temperature analysis, large differences are found in the present analysis as follows: the duration of large ocean heat content in the 1970s shortens dramatically, and recent ocean cooling becomes insignificant. The result is also in better agreement with tide gauge observations. On leave from the Meteorological Research Institute of the Japan Meteorological Agency.     

Large bio-geographical shifts in the north-eastern Atlantic Ocean: From the subpolar gyre, via plankton, to blue whiting and pilot whales

Pronounced changes in fauna, extending from the English Channel in the south to the Barents Sea in the north-east and off Greenland in the north-west, have occurred in the late 1920s, the late 1960s and again in the late 1990s. We attribute these events to exchanges of subarctic and subtropical water masses in the north-eastern North Atlantic Ocean, associated with changes in the strength and extent of the subpolar gyre. These exchanges lead to variations in the influence exerted by the subarctic or Lusitanian biomes on the intermediate faunistic zone in the north-eastern Atlantic. This strong and persistent bottom-up bio-physical link is demonstrated using a numerical ocean general circulation model and data on four trophically connected levels in the food chain – phytoplankton, zooplankton, blue whiting, and pilot whales. The plankton data give a unique basin-scale depiction of these changes, and a long pilot whale record from the Faroe Islands offers an exceptional temporal perspective over three centuries. Recent advances in simulating the dynamics of the subpolar gyre suggests a potential for predicting the distribution of the main faunistic zones in the north-eastern Atlantic a few years into the future, which might facilitate a more rational management of the commercially important fisheries in this region.