Carbon flows in Baltic Sea food webs — a re-evaluation using a mass balance approach

The brackish Baltic Sea has been seen as particularly suitable for studies of food webs. Compared to fully marine ecosystems, it has low species diversity, which means fewer trophic linkages to analyse. The Baltic Sea is also one of the best-studied areas of the world, suggesting that most data requirements for food web models should be fulfilled. Nevertheless, the influence of physical and biological factors on trophic interactions and biogeochemical patterns varies spatially in the Baltic Sea, adding considerable complexity to food web studies. Food web structure and processes can be described and compared quantitatively between areas by estimating the flow of matter or energy through the organisms. Most such models have been based on carbon, though studies of complementary flows of other elements limiting production, such as nitrogen and phosphorus would be desirable. However, since ratios between carbon and other elements are used in calculating these flows, it is crucial, as a first step, to quantify the flows of carbon as accurately as possible.In this study, we used the EcopathII software (ver 3.1) to analyse models of carbon flow through the food webs in the three main areas of the Baltic Sea; the Baltic proper, Bothnian Sea and Bothnian Bay. A previously published study on carbon flow in the Baltic Sea [Elmgren, R. 1984. Trophic dynamics in the enclosed, brackish Baltic Sea. Rapp. P.-V. Reun. — Cons. Int. Explor. Mer. (183) 152–169.] was complemented with the data on respiration and flow to detritus [Wulff, F., Ulanowicz, R. 1989. A comparative anatomy of the Baltic Sea and Chesapeeake Bay ecosystems. In: F. Wulff, J.G. Field, K.H. Mann (Eds.), Flow Analysis of Marine Ecosystems: Theory and Practice. New York: Springer-Verlag.] in order to present complete mass balance models of carbon. The purpose of re-evaluating previous models with new analytic tools was to check how well their carbon flows balance, and to provide a basis for improved mass balance models using more recent data, including nutrients other than carbon.The resulting mass balance networks for the Baltic proper, Bothnian Sea and the Bothnian Bay were shown to deviate from steady state. There was an organic carbon surplus of 45, 25 and 18 g C m⁻² year⁻¹ in the pelagic zones of the Baltic proper, Bothnian Sea and Bothnian Bay, respectively. The Ecopath network analysis confirmed that the overall carbon flow was highest in the Baltic proper, somewhat lower in the Bothnian Sea and much lower in the Bothnian Bay. The only clear differences in food web structure between the basins was that the average trophic level was lower for demersal fish in the Bothnian Sea and higher for macrofauna in the Bothnian Bay, compared to the other basins. The analysis showed weakness in our current understanding in Baltic Sea food webs and highlighted areas where improvements could be made with more recent data.     

Description of a flexible and extendable physical–biogeochemical model system for the water column

A modelling system for coupled physical–biogeochemical simulations in the water column is presented here. The physical model component allows for a number of different statistical turbulence closure schemes, ranging from simple algebraic closures to two-equation turbulence models with algebraic second-moment closures. The biogeochemical module consists of models which are based on a number of state variables represented by their ensemble averaged concentrations. Specific biogeochemical models may range from simple NPZ (nutrient–phytoplankton–zooplankton) to complex ecosystem models. Recently developed modified Patankar solvers for ordinary differential equations allow for stable discretisations of the production and destruction terms guaranteeing conservative and non-negative solutions. The increased stability of these new solvers over explicit solvers is demonstrated for a plankton spring bloom simulation. The model system is applied to marine ecosystem dynamics the Northern North Sea and the Central Gotland Sea. Two different biogeochemical models are applied, a conservative nitrogen-based model to the North Sea, and a more complex model including an oxygen equation to the Baltic Sea, allowing for the reproduction of chemical processes under anoxic conditions. For both applications, earlier model results obtained with slightly different model setups could be basically reproduced. It became however clear that the choice for ecosystem model parameters such as maximum phytoplankton growth rates does strongly depend on the physical model parameters (such as turbulence closure models or external forcing).     

Air–sea exchange of mercury vapour over the Gulf of Gda sk and southern Baltic Sea

The total gaseous mercury (TGM) in air over the coastal station at Hel and over the southern Baltic Sea was measured during the summer and winter conditions. Recorded 30-min resolution TGM data showed both higher concentrations and variability during the summer compared to the winter conditions. The summer TGM data ranged from 1.1 to 7.5 ng m⁻³, while the winter data ranged from 0.8 to 4.4 ng m⁻³. The TGM content in air over the southern Baltic Sea indicated that, in general, during the summer conditions, the sea-to-air transport of gaseous mercury dominated, while during the winter season, a tendency of gaseous mercury to sink into the water has been found. The evidences of enhanced water-to-air transfer of mercury vapour were noted, in particular, over the shallow waters of the Gulf of Gda sk under the strong water-to-air temperature gradients. Obtained results indicate that under such conditions, the coastal waters could act as a significant source of mercury vapour that may contribute to the overall budget of atmospheric mercury over the Baltic proper.     

Inorganic carbon fluxes through the boundaries of the Greenland Sea Basin based on in situ observations and water transport estimates

A carbon budget for the exchange of total dissolved inorganic carbon C_T between the Greenland Sea and the surrounding seas has been constructed for winter and summer situations. An extensive data set of C_T collected over the years 1994–1997 within the European Sub-polar Ocean Programmes (ESOP1 and ESOP2) are used for the budget calculation. Based on these data, mean values of C_T in eight different boxes representing the inflow and outflow of water through the boundaries of the Greenland Sea Basin are estimated. The obtained values are then combined with simulated water transports taken from the ESOP2 version of the Miami Isopycnic Coordinate Ocean Model (MICOM). The fluxes of inorganic carbon are presented for three layers; a surface mixed layer, an intermediate layer and a deep layer, and the imbalance in the fluxes are attributed to air–sea exchange, biological fixation of inorganic carbon, and sedimentation. The main influx of carbon is found in the surface and the deep layers in the Fram Strait, and in the surface waters of direct Atlantic origin, whereas the main outflux is found in the surface layer over the Jan Mayen Fracture Zone and the Knipovich Ridge, transporting carbon into the Atlantic Ocean via the Denmark Strait and towards the Arctic Ocean via the Norwegian Sea, respectively. The flux calculation indicates that there is a net transport of carbon out of the Greenland Sea during wintertime. In the absence of biological activity, this imbalance is attributed to air sea exchange, and requires an oceanic uptake of CO₂ of 0.024±0.006 Gt C yr⁻¹. The flux calculations from the summer period are complicated by biological fixation of inorganic carbon, and show that data on organic carbon is required in order to estimate the air–sea exchange in the area.     

Parameterization of air–sea gas fluxes at extreme wind speeds

Air–sea flux measurements of O₂ and N₂ obtained during Hurricane Frances in September 2004 [D'Asaro, E. A. and McNeil, C. L., 2006. Measurements of air–sea gas exchange at extreme wind speeds. Journal Marine Systems, this edition.] using air-deployed neutrally buoyant floats reveal the first evidence of a new regime of air–sea gas transfer occurring at wind speeds in excess of 35 m s^− 1. In this regime, plumes of bubbles 1 mm and smaller in size are transported down from near the surface of the ocean to greater depths by vertical turbulent currents with speeds up to 20−30 cm s^− 1. These bubble plumes mostly dissolve before reaching a depth of approximately 20 m as a result of hydrostatic compression. Injection of air into the ocean by this mechanism results in the invasion of gases in proportion to their tropospheric molar gas ratios, and further supersaturation of less soluble gases. A new formulation for air–sea fluxes of weakly soluble gases as a function of wind speed is proposed to extend existing formulations [Woolf, D.K, 1997. Bubbles and their role in gas exchange. In: Liss, P.S., and Duce, R.A., (Eds.), The Sea Surface and Global Change. Cambridge University Press, Cambridge, UK, pp. 173–205.] to span the entire natural range of wind speeds over the open ocean, which includes hurricanes. The new formulation has separate contributions to air–sea gas flux from: 1) non-supersaturating near-surface equilibration processes, which include direct transfer associated with the air–sea interface and ventilation associated with surface wave breaking; 2) partial dissolution of bubbles smaller than 1 mm that mix into the ocean via turbulence; and 3) complete dissolution of bubbles of up to 1 mm in size via subduction of bubble plumes. The model can be simplified by combining “surface equilibration” terms that allow exchange of gases into and out of the ocean, and “gas injection” terms that only allow gas to enter the ocean. The model was tested against the Hurricane Frances data set. Although all the model parameters cannot be determined uniquely, some features are clear. The fluxes due to the surface equilibration terms, estimated both from data and from model inversions, increase rapidly at high wind speed but are still far below those predicted using the cubic parameterization of Wanninkhof and McGillis [Wannikhof, R. and McGillis, W.R., 1999. A cubic relationship between air–sea CO₂ exchange and wind speed. Geophysical Research Letters, 26:1889–1892.] at high wind speed. The fluxes due to gas injection terms increase with wind speed even more rapidly, causing bubble injection to dominate at the highest wind speeds.     

Study of the seasonal cycle of the biogeochemical processes in the Ligurian Sea using a 1D interdisciplinary model

A one-dimensional coupled physical–biogeochemical model has been built to study the pelagic food web of the Ligurian Sea (NW Mediterranean Sea). The physical model is the turbulent closure model (version 1D) developed at the GeoHydrodynamics and Environmental Laboratory (GHER) of the University of Liège. The ecosystem model contains 19 state variables describing the carbon and nitrogen cycles of the pelagic food web. Phytoplankton and zooplankton are both divided in three size-based compartments and the model includes an explicit representation of the microbial loop including bacteria, dissolved organic matter, nano-, and microzooplankton. The internal carbon/nitrogen ratio is assumed variable for phytoplankton and detritus, and constant for zooplankton and bacteria. Silicate is considered as a potential limiting nutrient of phytoplankton's growth. The aggregation model described by Kriest and Evans in (Proc. Ind. Acad. Sci., Earth Planet. Sci. 109 (4) (2000) 453) is used to evaluate the sinking rate of particulate detritus. The model is forced at the air–sea interface by meteorological data coming from the “Côte d'Azur” Meteorological Buoy. The dynamics of atmospheric fluxes in the Mediterranean Sea (DYFAMED) time-series data obtained during the year 2000 are used to calibrate and validate the biological model. The comparison of model results within in situ DYFAMED data shows that although some processes are not represented by the model, such as horizontal and vertical advections, model results are overall in agreement with observations and differences observed can be explained with environmental conditions.     

Evaluating transfer velocity-wind speed relationship using a long-term series of direct eddy correlation CO2 flux measurements

Long-term observations of the marine atmospheric boundary layer were performed by an eddy correlation system, which was set-up on a platform in the Baltic Sea. In this experiment the three-dimensional wind vector and the turbulent fluxes of momentum, sensible and latent heat and CO₂ were measured for one and a half years. Simultaneously the CO₂ partial pressure p_CO2 in surface water was measured by a submersible autonomous moored instrument for CO₂ at the platform in 7-m depth. The high-resolution eddy correlation measurements of the atmospheric CO₂ flux F_CO2, together with the measurements of the CO₂ partial pressure differences between air and sea Δp_CO2 led to a long-term data set which provided the possibility to investigate the parameterization of the CO₂ transfer velocity k as a function of 10-m wind speed u in a statistical manner. From half-hour mean CO₂ fluxes and CO₂ partial pressure differences, k was calculated using k = F_CO2 / (K₀Δp_CO2), with K₀ the CO₂ solubility. The half-hour mean data points, used for the determination of the k–u parameterization, show large scatter. However, assuming a linear, quadratic dependency the analysis yields: k₆₆₀ = 0.365u² + 0.46u (k at 20 °C and salinity 35 psu) with a correlation coefficient of r² = 0.81. The large scatter indicates that the kinetics of the air–sea CO₂ transfer velocity is not only a function of the wind speed alone, but might also be controlled by other environmental parameters and mechanisms, such as sea state and surface coverage with surfactants.     

Tropical cyclone genesis over the south China sea

The South China Sea (SCS) is among areas in the Northwest Pacific most frequented by tropical cyclones (TCs) with intensity reaching a tropical storm or stronger. It is also an area of significant TC genesis. In this study, TC genesis in SCS and its monsoonal variability for 1948–2003 are analyzed. Altogether, in May–September (southwest monsoon period) 157 TC geneses have occurred north of 12°N in SCS, while in October–December (northeast monsoon period) 64 out of 65 TC geneses have happened south of 18°N. It is found that the monsoonal characteristics of the SCS basically determine the region of TC genesis in each monsoon season. Winter TC genesis in the SCS happens over the region where the marine environment satisfies the four criterions on, respectively, the sea surface temperature (SST), mid-troposphere relative humidity, vertical shear of the horizontal winds and low-level atmospheric vorticity. During the summer, as the two criterions on SST and the mid-troposphere relative humidity are satisfied for the whole SCS, TC genesis occurs in the region where both the low-level vorticity and the vertical shear satisfy the criterion. In addition, there is likely more TC genesis in the winter during the onset of La Nina, and more TC genesis in the summer following the onset of El Nino.     

Future aspects in marine ecosystem modelling

Existing ecosystem models are briefly presented and summarised. The problem of coupling physical and biological models as well as aspects of prediction and predictability are discussed. The general perception that marine ecosystems are inherently unpredictable due to non-linearity becomes questionable if the response of climate variability in marine ecosystems is analysed. Many authors have shown correlations between climate variability and the variability of abundance or biomass of marine organisms such as phytoplankton, zooplankton, benthos or fish recruitment in different parts of the world ocean. In the northern hemisphere, certain species show a linear response to climate variability mainly during winter and spring. However, the underlying mechanisms are not well understood. Often, a phase lag can be observed between climate variability and the reaction of organisms. The identification of a plausible mediator between climate and biology is difficult, since all possible physicochemical mechanisms having a direct or indirect influence on the variability of abundance or biomass of marine organisms have to be considered as mediator.The understanding of the reason of the phase lag, which possibly implies a “biological memory”, and the identification of all possible mediators are necessary to predict the response of marine organisms to climate variability. The identification of mediators will result in an improvement of coupled models, a deeper understanding of physical–biological interaction and the improvement of predictive capability of marine ecosystem models.     

Use of SeaWiFS data for light availability and parameter estimation of a phytoplankton production model of the Bay of Biscay

Processing SeaWiFS (Sea-viewing Wide Field-of-view Sensor) data provides useful information for the observation and modelling of the phytoplankton production of the Bay of Biscay. Empirical algorithms allow the retrieval of chlorophyll a and non-living Suspended Particulate Matter (SPM) concentrations. These data are used to constrain a coupled 3D physical–biogeochemical model of the Bay of Biscay continental shelf. Two issues are investigated, depending on the variable used, to constrain the winter to spring phytoplankton production for the year 2001. First, SPM data is used as forcing data to correct the corresponding state variable of our model. This allows the realistic simulation of the light limited bloom at the end of February 2001, as observed with SeaWiFS chlorophyll a images and from the NUTRIGAS field cruise. Second, chlorophyll a data is used for parameter estimation of the biogeochemical model. The ability of assimilating these data is tested to improve the simulation of strong blooms observed in late May 2001 in the Loire and Gironde plumes. A global optimization method (Evolutive Strategies) is adapted to the complete 3-D coupled model, in order to find the best set of parameters. The hydrological conditions during the bloom can be validated with data from the PEL01 field cruise. After selection of the most sensitive parameters, the method is tested with twin experiments. Then, the use of real SeaWiFS data reduces the model/data misfit by a factor of two, improving the simulation of bloom intensities and extensions. The sets of parameters retrieved in each plume are discussed.     

Mesoscale subduction at the Almeria–Oran front: Part 1: Ageostrophic flow

This paper presents a detailed diagnostic analysis of hydrographic and current meter data from three, rapidly repeated, fine-scale surveys of the Almeria–Oran front. Instability of the frontal boundary, between surface waters of Atlantic and Mediterranean origin, is shown to provide a mechanism for significant heat transfer from the surface layers to the deep ocean in winter. The data were collected during the second observational phase of the EU funded OMEGA project on RRS Discovery cruise 224 during December 1996. High resolution hydrographic measurements using the towed undulating CTD vehicle, SeaSoar, traced the subduction of Mediterranean Surface Water across the Almeria–Oran front. This subduction is shown to result from a significant baroclinic component to the instability of the frontal jet. The Q-vector formulation of the omega equation is combined with a scale analysis to quantitatively diagnose vertical transport resulting from mesoscale ageostrophic circulation. The analyses are presented and discussed in the presence of satellite and airborne remotely sensed data; which provide the basis for a thorough and novel approach to the determination of observational error.     

Changes in the shelf macrobenthic community over large temporal and spatial scales in the Bohai Sea, China

Over the past 20 years, the Bohai Sea has been subjected to a considerable human impact through over-fishing and pollution. Together with the influence of the Yellow River cut-off, the ecosystem experienced a dramatic change. In order to integrate available information to detect any change in macrobenthic community structure and diversity over space and time, data collected during the 1980s and the 1990s from 3 regions of the Bohai Sea (Laizhou Bay, 16 stations, 37–38°N, 119–120.5°E; central Bohai Sea, 25 stations, 38–39°N, 119–121°E; eastern Bohai Bay, 12 stations, 38–39°N, 118.5–119°E) were reanalyzed in a comparative way by means of a variety of statistical techniques. A considerable change in community structure between the 1980s and the 1990s and over the geographical regions at both the species and family level were revealed. After 10 years, there was a considerable increase in abundance of small polychaetes, bivalves and crustaceans but decreased number of echinoderms. Once abundant in Laizhou Bay in the 1980s, a large echinoderm Echinocardium cordatum and a small mussel Musculista senhousia almost disappeared from the surveying area in the 1990s. Coupled with the increased abundance was the increased species richness in general whereas evenness was getting lower in central Bohai Sea and Bohai Bay but increased in Laizhou Bay. K-dominance plot showed the same trend as evenness J′. After 10 years, the macrobenthic diversity in the Bohai Sea as a whole was slightly reduced and a diversity ranking of central Bohai Sea > Laizhou Bay > eastern Bohai Bay over space was also suggested. Sediment granulometry and organic content were the two major agents behind the observed changes.     

Parameterisation of clastic sediments including benthic structures

The sediment transport processes in the south-western Baltic Sea are predicted by means of a numerical model in the project DYNAS. There are two sediment parameters that influence the results of modelling remarkably: critical shear stress velocity and bottom roughness. This paper presents the way how to parameterise these factors and extrapolate them into the investigation area. The critical shear stress velocity is parameterised basing on grain size data, combining approximations after Hjulström [Hjulström, F., 1935: Studies in the morphological activity of rivers as illustrated by the river Fyris. Geological Institution of University of Uppsala: Bulletin (25): 221–528.], Shields [Shields, A., 1936: Anwendung der Ähnlichkeits-Mechanik und der Turbulenzforschung auf die Geschiebebewegung. Mitteilungen der Preussischen Versuchsanstalt für Wasserbau und Schiffahrt (26): 26 pp.] and Bohling [Bohling, B., 2003: Untersuchungen zur Mobilität natürlicher und anthropogener Sedimente in der Mecklenburger Bucht. unpublished doctoral thesis, Mathematisch-Naturwissenschaftliche Fakultät, Ernst-Moritz-Arndt-Universität Greifswald/Germany, 156 pp.]. The roughness length, in the case of absence of macro zoo-benthos and their structures, is parameterised basing on grain size too employing Soulsby [Soulsby, R.L., 1997: Dynamics of Marine Sands: a Manual for Practical Applications. London, Thomas Telford Publications. 249 pp.], Nielsen [Nielsen, P., 1983: Analytical determination of nearshore wave height variation due to refraction shoaling and friction. Coastal Engineering 7, 233–251.] and Yalin [Yalin, M.S., 1977: Mechanics of Sediment Transport. Pergamon Press, New York. 298 pp.]. No equivalent simple parameterisations for biologically caused bed roughness exist. Here, findings of Friedrichs [Friedrichs, M., 2004: Flow-induced effects of macro zoo-benthic structures on the near-bed sediment transport. Dissertation, Universität Rostock, 80 S.] and estimations by the DYNAS biologists group were combined in order to derive roughness lengths from abundance measurements of four previously selected key species which represent the originators of the dominating benthic structures at the sea floor in the south-western Baltic Sea. Critical shear stress velocity and bed roughness are known at few sample sites only. They were extrapolated into the larger investigation area using a proxy-target concept. The mean near bottom milieu (bathymetry, median grain size, salinity, oxygen) which was derived using results from numerical modelling serves as the proxy. Since the milieu parameters are measured at the sampling sites for which the target parameters have been determined, a combined hierarchical and supervised classification was employed to transfer the local knowledge into the unknown investigation area.     

Surface chlorophyll in the Black Sea over 1978–1986 derived from satellite and in situ data

Absolute values of chlorophyll a concentration and its spatial and seasonal variations in the Black Sea were assessed by using satellite CZCS and in situ data. Since the satellite CZCS had operated for the 1978–1986 period, CZCS data was used for assessing the past state of the Black Sea just before the onset of drastic changes observed in late 1980s. The approach used for the calculation of the absolute values of chlorophyll a concentration from CZCS data was based on the direct comparison of in situ chlorophyll a data and those of CZCS and by applying the algorithm developed for the transformation of CZCS data into chlorophyll a values. CZCS Level 2 data related with pigment concentration having a spatial resolution of 1 km at nadir were used. The daily Level 3 files were derived by binning Level 2 values into 4-km grid cells and the monthly and seasonal Level 3 files were created by averaging the daily Level 3 files over the corresponding period. In situ chlorophyll a data were obtained by spectrophotometric and fluorometric methods in 15 scientific cruises over the 1978–1986 period. Total number of ship-measured data used for the comparison with those CZCS values was 590.Chlorophyll a concentration (Chl) was derived from CZCS values (C) with regression equations Chl=kC; the coefficient of transformation k was calculated from six different data sets by taking into account distinctions between subregions and seasons. The reasons for difference in the k values have been analyzed.Statistical comparison of the chlorophyll a values measured in situ and those derived from CZCS data was based on log-transformed data and gave the following results: regression SLOPE=0.842, regression INTERCEPT=−0.081, coefficient of determination (R²)=0.806, root–mean–square ERROR=0.195. The mean monthly chlorophyll a distributions derived from CZCS data over 1978–1986 have been constructed and the mean seasonal chlorophyll a values in different regions have been calculated and analyzed. The significant difference in chlorophyll concentration between the western shelf regions and the open part of the Black Sea has been demonstrated, especially in warm season. At almost all seasons, the highest chlorophyll concentration is observed in the western interior shelf region which is under strong influence of Danube. The summer mean chlorophyll concentration in this region is 18 times higher than that in the open parts and about nine times higher than in the eastern shelf region. The greatest seasonal variations are observed in the open part of the Black Sea: chlorophyll concentration in cold season is four to six times higher than in summer and three to five times higher than in April and October. To the contrary, in the western interior shelf regions, the concentration is higher in May–October (about twice than that in November–March). Seasonal variations in the western outer shelf regions are smoothed out as compared with both the western interior shelf and the open regions.