Dynamics of submersed aquatic vegetation on shallow soft bottoms in the Baltic Sea

Abstract. Persistence and colonization of submersed aquatic plant species were studied in permanent plots (20 cm × 20 cm) at three shallow sites adjacent to Askö island, in the northern Baltic Sea. The study started in 1991 at two of the sites, in 1992 at a third site and continued until 1997. Two major weather‐induced disturbances occurred during the study: a long, stormy period during a mild winter and a cold winter with extreme low water levels. The stormy period caused a large loss of species from plots (95 ± 5% SE) at the most wave‐exposed site, resulting in a low species number per plot (0.8 ± 0.2) the following summer. During the three following years the mean species number increased to 3.6 ± 0.2. The cold winter caused high species turnover and increased species number per plot at the two most wave‐exposed sites. The species turnover at the sheltered site was highest in the two years with low water level in May, possibly due to increased waterfowl herbivory. Annuals, loosely anchored and highly reproductive species had significantly higher plot colonization rates and lower persistence than perennials, well‐anchored species and species with none or low reproduction. The extent of lateral growth had no significant effect on colonization or persistence. Although these disturbances have a large effect on the dynamics, species mobility was also high in other years. Relative to other, similar, studies in terrestrial vegetation mean persistence in plots was remarkably low and mean mobility and species turnover rates were very high.     

Pore water dynamics in the sediment of a shallow and hypertrophic lake

Seasonal variations in pore water with main stress on pH and phosphate were investigated in the sediment of the shallow and hypertrophic Lake Søbygaard, Denmark. The purpose was to evaluate factors affecting the internal phosphorus loading. Pore water was obtained by in situ incubation of ceramic cups, sampled anaerobicaly from a fixed position in the sediment. The method is evaluated. During summer, pH and phosphate concentrations increased in the upper 8–10 cm of the sediment. Increased pH was most pronounced in the upper 5 cm, where pH increased to between 9 and 10. This is believed to be caused by the photosynthetically elevated pH in the above lake water. Phosphate concentrations increased with depth, from 0–2 mg P 1^–1 in the upper 5 cm to 3–6 mg P 1^–1 in 6–10 cm depth. Average phosphate gradient in the upper 6–8 cm was 1.0 mg P 1^–1 cm ^–1 in the summer decreasing to 0.2 mg P 1^–1 cm ^t1 in the autumn/winter. In spite of low redox potential, Fe(II) was not present in the upper 20 cm. The seasonal variation in pore water phosphate is believed mainly to be due to the variations in pore water pH inducing a substitution of phosphate ions with hydroxyl ions on ironhydroxides during summer. A considerable sedimentation of organic bound phosphorus and decomposition in the sediment is also considered important. Phosphorus release from the sediment is facilitated by bio- and gas turbation and by the frequent occurrence of resuspension caused by windaction. Net release rate is highly variable during the season. The summer average is 40 mg P m^–2 d^–1.     

Chemical evolution of the Salton Sea, California: nutrient and selenium dynamics

The Salton Sea is a 1000-km² terminal lake located in the desert area of southeastern California. This saline (44000 mg l^–1 dissolved solids) lake started as fresh water in 1905–07 by accidental flooding of the Colorado River, and it is maintained by agricultural runoff of irrigation water diverted from the Colorado River. The Salton Sea and surrounding wetlands have recently acquired substantial ecological importance because of the death of large numbers of birds and fish, and the establishment of a program to restore the health of the Sea. In this report, we present new data on the salinity and concentration of selected chemicals in the Salton Sea water, porewater and sediments, emphasizing the constituents of concern: nutrients (N and P), Se and salinity. Chemical profiles from a Salton Sea core estimated to have a sedimentation rate of 2.3 mm yr^–1 show increasing concentrations of OC, N, and P in younger sediment that are believed to reflect increasing eutrophication of the lake. Porewater profiles from two locations in the Sea show that diffusion from bottom sediment is only a minor source of nutrients to the overlying water as compared to irrigation water inputs. Although loss of N and Se by microbial-mediated volatilization is possible, comparison of selected element concentrations in river inputs and water and sediments from the Salton Sea indicates that most of the N (from fertilizer) and virtually all of the Se (delivered in irrigation water from the Colorado River) discharged to the Sea still reside within its bottom sediment. Laboratory simulation on mixtures of sediment and water from the Salton Sea suggest that sediment is a potential source of N and Se to the water column under aerobic conditions. Hence, it is important that any engineered changes made to the Salton Sea for remediation or for transfer of water out of the basin do not result in remobilization of nutrients and Se from the bottom sediment into the overlying water.     

Impacts of nutrient enrichment and sediment on phytoplankton community structure in the northern Baltic Sea

A three-week mesocosm experiment was conducted in order to study the effects of bottom sediment and nutrient enrichment on phytoplankton and zooplankton community structure in the Archipelago Sea, northern Baltic Sea. The transparent polyethylene enclosures included the whole water column and varied in volume from 30 to 40 m³. There were two types of enclosures: some with natural sediment as a bottom and others with a plastic bottom. The experiment was a 2 × 2 factorial design with presence of sediment and nutrient enrichment as treatment factors. Both the sediment presence and nutrient enrichment significantly increased water nutrient concentrations and the rate of primary production. However, external nutrient enrichment and the presence of sediment stimulated the growth of different phytoplankton groups, indicating that the effect of sediment was not related to nutrient fluxes alone, but involved more complex interactions. External nutrient enrichment was primarily channelled to picoplanktonic cyanobacteria, the biomass of which increased four- to fivefold due to enrichment. The presence of sediment increased the biomass of cryptophytes, chrysophytes and prasinophytes, but decreased the biomass of N₂-fixing cyanobacteria. Zooplankton biomass increased during the experiment, but was not affected by the treatments. The study shows that sediment plays a significant role in phytoplankton dynamics, underlining the importance of including sediment in shallow-water mesocosm experiments. Handling editor: J. Padisak     

Spatially differing bacterial communities in water columns of the northern Baltic Sea

The Baltic Sea is a large, shallow, and strongly stratified brackish water basin. It suffers from eutrophication, toxic cyanobacterial blooms, and oxygen depletion, all of which pose a threat to local marine communities. In this study, the diversity and community structure of the northern Baltic Sea bacterial communities in the water column were, for the first time, thoroughly studied by 454 sequencing. The spring and autumn bacterial communities were one order of magnitude less diverse than those in recently studied oceanic habitats. Patchiness and strong stratification were clearly detectable; <1% of operational taxonomic units were shared among 11 samples. The community composition was more uniform horizontally (at a fixed depth) between different sites than vertically within one sampling site, implying that the community structure was affected by prevailing physical and hydrochemical conditions. Taxonomic affiliations revealed a total of 23 bacterial classes and 169 genera, while 5% of the sequences remained unclassified. The cyanobacteria accounted for <2% of the sequences, and potentially toxic cyanobacterial genera were essentially absent during the sampling seasons.     

Microphytobenthos and chironomid larvae attenuate nutrient recycling in shallow‐water sediments

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Mesozooplankton dynamics in the northern Baltic Sea: implications of variations in hydrography and climate

An 18-year mesozooplankton time series from 1967 to 1984 fromthe SW coast of Finland, northern Baltic Sea, was studied inrelation to climatic parameters (air temperature, air pressure,precipitation, and wind speed and direction), freshwater dischargeand hydrographical factors (water temperature, salinity andwater column stability). Climatic factors influenced mesozooplanktoncommunity through their effects on hydrography. The averagewater salinity responded to fluctuations in river dischargewith a 1–2 year lag. River discharge was in turn regulatedby variations in atmospheric pressure and precipitation, whichoscillated on 3–5 year cycles. Shorter-term variationsin salinity and temperature were related to changes in the dominatmgwind directions. The seasonally adjusted residuals of totalmesozooplankton biomass, cladoceran biomass and those of manyof the dominant mesozooplankton species were significantly negativelycorrelated with salinity. The annual peaks of the most abundaiitcladoceran species and one of the most abundant calanoid speciesco-varied with the seasonal fluctuation of surface water temperature.Canonical correspondence analysis identified the mesozooplanktonspecies by their relationship to water stability: rotifers andcladocerans had their optima in warm and stabile conditions,while the nentic calanoid copepods were more abundant in relativelysaline and mixed conditions.     

Chemical limnology of soft water lakes in the Upper Midwest

Water samples from 36 lakes in northern Minnesota, Wisconsin, and Michigan were collected and analyzed during 1983–1984. All study lakes were dilute and had total alkalinities of less than 150 eq · L^–1. Minnesota lakes have hydrologic inputs from the watershed and inputs of base cations derived from the watershed. Study lakes in Minnesota had higher total alkalinities, dissolved organic carbon, and noncarbonate alkalinity as a result of watershed inputs. Lakes in Michigan and Wisconsin were precipitation-dominated seepage lakes that have lower concentrations of base cations than lakes in Minnesota. All of the study lakes have lower sulfate concentrations than expected, based on atmospheric wet deposition and evapotranspiration.Pore water samples collected from one of the study lakes—Little Rock Lake—in Wisconsin were used to calculate diffusive fluxes between the sediment and water column. According to these calculations, the sediments were a source of total alkalinity and Ca²⁺ and a sink for SO₄ ^2–. The sediment-water exchange of total alkalinity, Ca²⁺, and SO₄ ^2– appears to be important in the whole-lake budgets of these ions for Little Rock Lake.     

Marine Benthos in Relation to Pore Water Chemistry and Sediment Geochemistry of Simulated Dredged Material

During the course of study on vertical migration and mortality of benthos in simulated dredged material, changes in pore water chemistry and sediment geochemistry were also examined. Dissolved oxygen, ammonia, and sulfide showed significant changes between overlying water and pore water over the two week experimental period. Chemical effects in dredging operations can be important on infaunal organisms.     

Effect of nutrient enrichment on bacterioplankton biomass and community composition in mesocosms in the Archipelago Sea, northern Baltic

Effects of nutrient enrichment on the biomass and communitycomposition of heterotrophic bacteria and picocyanobacteriawere studied in large (42 m³) mesocosms in the brackish-waterArchipelago Sea (Baltic Sea) in late summer 2000 using cellcounts and denaturing gradient gel electrophoresis (DGGE) ofpolymerase chain reaction (PCR)-amplified 16S rRNA gene fragments.The identity of the major DNA bands was determined by sequencing.The obtained sequences were related to - and -proteobacteria,actinobacteria, verrucomicrobia and cyanobacteria. Nitrogenand phosphorus additions increased the biomasses of heterotrophicbacteria and picocyanobacteria and caused significant changesin their community composition judging from the DGGE bandingpatterns. Most verrucomicrobial bands had their highest relativeintensity in the control treatment and their lowest in the highernutrient addition treatment, whereas most Synechococcus-relatedbands had their lowest relative intensity in the lower nutrientaddition treatment. The responses of proteobacteria and actinobacteriawere more variable. The presence of both freshwater and marinesequences among the closest relatives to our sequences highlightsthe intermediate character of the Archipelago Sea between afreshwater and truly marine environment.     

The cycle of mussels: long-term dynamics of mussel beds on intertidal soft bottoms at the White Sea

Dense blue mussel assemblages are unstable, their structure changing from year to year. Three types of models can be used to describe this instability: (1) “exogenous” model based on regional temperature fluctuations, (2) “endogenous” deterministic model associated with negative impact of adult mussels on juveniles and (3) “density-linked stochasticity” model based on positive feedbacks resulting in overcrowding and destabilizing the settlement. We compared predictions deduced from these models with a time series based on the results of long-term (18 years) monitoring of abundance and demographic structure of three mussel beds at the White Sea. Most of our findings agreed well with the predictions deduced from the endogenous model. In particular, (1) long-term changes in mussel abundance and demographic structure were strictly cyclic, with non-matching periods (5–9 years) at different sites; (2) stages with the dominance of old mussels alternated with those where juveniles dominated and (3) some signals of delayed density dependence were revealed. However, the time series also contained elements of long-term trends, which may testify to the involvement of some exogenous factors (probably long-term climate changes).     

Phytoplankton community responses to nutrient and iron enrichment under different nitrogen to phosphorus ratios in the northern Baltic Sea

The impact of nutrient enrichment on the phytoplankton community structure, and particularly cyanobacteria, was studied in a 3-week mesocosm experiment conducted in August 2001 in the Archipelago Sea, a part of the northern Baltic Sea. The factorial design experiment included daily additions of nitrogen (N) and phosphorus (P) at two mass ratios, 1N:1P and 7N:1P, respectively, additions of iron (Fe) and a synthetic chelator, ethylenediaminetetraacetic acid (EDTA). The floating enclosures (400 l) were sampled for analyses of phytoplankton biomass and community structure, phytoplankton primary production, chlorophyll a, nutrients, and hepatotoxins. Chlorophyll a concentration, phytoplankton biomass and primary production increased most in the 7N:1P treatment. The increase was mainly due to an abundant growth of chlorophytes (Dictyosphaerium subsolitarium, Kirchneriella spp., Monoraphidium contortum, and Oocystis spp.), pennate diatoms (especially Nitzschia spp.), dinophytes and the chroococcalean cyanobacterium Synechococcus sp. The nutrient enrichments had no effect on the total biomass of N₂-fixing cyanobacteria. Nevertheless, the biomass of Anabaena spp. was highest in the enrichments with a low N/P ratio. Chlorophyll a concentration and total phytoplankton biomass were not affected by Fe or EDTA, but Fe alone had a positive effect on the chlorophyte Kirchneriella sp. The N₂-fixing cyanobacteria Aphanizomenon sp. responded positively to Fe alone and to both Fe and EDTA added together. The hepatotoxin concentration increased during the experiment, but no clear responses to nutrient enrichments were found. Our study showed species-specific responses to nutrient enrichments among the N₂-fixing cyanobacteria. Although the total phytoplankton production was not Fe-limited; the availability of Fe clearly affected the phytoplankton community structure.     

Diffuse nutrient reduction in the German Baltic Sea catchment: Cost-effectiveness analysis of water protection measures

Water quality of groundwater, rivers, and seas is still negatively influenced by high diffuse nitrogen and phosphorus discharge. Increasing importance is attached to cost-effectiveness to indicate sufficient measures to reduce nutrient input of land use. The aim of this paper is to find cost-effective solutions for diffuse nutrient load reductions for the German Baltic Sea region (by formulating scenarios for a 25% and 50% reduction of nitrogen and phosphorus). I combined nutrient emission and land use modelling (literature analyses, MONERIS-data) with economic land use analysis (e.g. opportunity costs, costs of the production methods) in a cost-effectiveness analysis.The results show that total reduction costs need not be as expensive as often anticipated (e.g. 25% simultaneous reduction of nitrogen and phosphorus costs about 8–20 € (ha y)⁻¹).Advisory service turned out as the most cost-effective measure and the most important indicator for the cost-saving potential. The measure included guidelines to the farmers on land use and water quality optimization. For example, if a 100% advisory service is successful, total costs for a 25% simultaneous reduction of nitrogen and phosphorus could drop by 65%, corresponding to 23 million € y⁻¹ compared to a 50% advisory service. Due to a lack of data, I could not consider the additional nutrient retention capacity of wetlands and buffer strips in the analysis.The policy recommendations derived from this study deliver insights for the strategic orientation of water protection policy and useful indicators to measure the performance. The results are of interest not only in the Baltic Sea context, but also for supporting and facilitating the application of the cost-effectiveness analysis postulated in the Water Framework Directive. Future research should concentrate on improving data availability (e.g. livestock husbandry) and models and on implementing further transaction costs and additional indicators like nutrient retention of wetlands.     

Pore water phosphorus and iron concentrations in a shallow,eutrophic lake — indications of bacterial regulation

Peak pore water SRP and iron(II) concentrations were found during summer in surface sediments in the shallow and eutrophic L. Finjasjön, Sweden, and the concentrations generally increased with water depth. The SRP variation in surface sediments (0–2 cm) was correlated with temperature (R² = 0.82–0.95) and iron(II) showed a correlation with sedimentary carbon on all sites (R² = 0.42–0.96). In addition, sedimentary Chla, bacterial abundances and production rates in surface sediments (0–2 cm) varied seasonally, with peaks during spring and fall sedimentation. Bacterial production rates were correlated with phosphorus and carbon in the sediment (R² = 0.90–0.95 and R² = 0.31–0.95, respectively), indicating a coupling with algal sedimentation. A general increase in sediment Chla and bacterial abundances towards sediments at greater water depth was found. Further, data from 1988–90 reveal that TP and TFe concentrations in the lake were significantly correlated during summer (R² = 0.81 and 0.76, in the hypolimnion and epilimnion, respectively). The results indicate that the increase in pore water SRP and Fe(II) in surface sediments during summer is regulated by bacterial activity and the input of organic matter. In addition, spatial and temporal variations in pore water composition are mainly influenced by temperature and water depth and the significant correlation between TP and TFe in the water suggests a coupled release from the sediment. These findings support the theory of anoxic microlayer formation at the sediment-water interface.     

Porewater nutrient profiles and nutrient sediment–water exchange in a tropical mangrove waterway, Mapopwe Creek, Chwaka Bay, Zanzibar

Sediments were examined in the Mapopwe Creek, a tidally dominated mangrove waterway in the Chwaka Bay mangrove forest, Zanzibar, to assess their significance in the nutrient dynamics of the mangrove forest and the adjacent bay. Porewater concentrations of dissolved ammonium and that of soluble reactive phosphorus (SRP) were generally higher during the dry season than during the wet season. NO₃^? plus NO₂^? concentration averaged 1 µm and did not vary much between the two periods. Fluxes of ammonium ranged from ?575 to 523 µm m^?2 h^?1 and those of SRP from ?55.7 to 69.5 µm m^?2 h^?1. Measurements of NO_x did not show any consistent fluxes of this dissolved nitrogen species. Variations of flux rates between the two seasons were not significant even though there were small variations in the flux direction in both nutrients. Results imply that Mapopwe sediments are a source of NH₄⁺ but act as a sink for SRP.     

Spatio-temporal dynamics of chlorophyll in the open Baltic Sea

Five charts of the chlorophyll and hydrographic fields by verticalprofiles of in situ fluorescence and CTD were made on a stationarygrid of 20 ? 25 nautical miles with a 5-mile spacing in theopen Baltic Sea. Both chlorophyll levels and variability weremaximal close to the spring bloom. High chlorophyll levels insummer are sustained by recurrent nutrient injections from thedeep saline layer. Two of the surveys showed close couplingbetween the coarse-scale (10 km) chlorophyll distribution andthe hydrographic structure determining the intensity of nutrienttransfer. Vigorous advection, stirring and current shear, associatedwith a strong mesoscale eddy, probably dominated the chlorophyllpattern on three surveys. The upward velocities in the cycloniceddy resulted in accumulation of phytoplankton in the aphoticzone. Intensive heat input from the surface caused a suddensinking of the phytoplankton and the formation of a pronouncedsub-surface chlorophyll maximum.     

Long-term dynamics of zooplankton in the southeastern Baltic Sea

Long-term research in the Baltic Sea revealed the basic trends of zooplankton community variations depending on oceanographic processes. Alternation of the periods of increase and decrease in salinity of the Baltic Sea against the background of climate changes (temperature increase) and eutrophication affect the state of the entire Baltic ecosystem, including zooplankton. For these periods, the dynamics of zooplankton in the Baltic Sea were analyzed based on literature data and results of regular research in the southeastern Baltic Sea during 1998–2007. The changes in the hydrological situation were accompanied by significant changes in the zooplankton community. In the 1990s–2000s, the abundance and biomass of brackish-water and thermophilous species primarily of Cladocera and Copepoda increased markedly. The role of the previously dominant marine copepod Pseudocalanus elongatus decreased due to salinity reduction in the deep-water part of the Baltic Sea. Maximum development of zooplankton occurred in years of the greatest warming-up of the water (2001, 2005–2007) against the background of a general positive trend of zooplankton abundance in the last decade.     

Reproductive characteristics of the male herring in the northern Baltic Sea

The gonad weight and gonadosomatic index of the male Baltic herring Clupea harengus membras L., were highest at the beginning of the reproductive season (April/May), the values decreasing towards the end of it (July/August) during 1988–1991. The decline could not be explained by fish size but may have been due to fish condition. A high individual variation was typical for both gonad weights and gonadosomatic indices in fish of the same size and maturity stage. The mean density of sperm cells was significantly higher in June (34·9 × 10⁹ ml⁻¹) than in July (19·2 × 10⁹ ml⁻¹, Mann-Whitney U= 17; P<0·05), the variation among the males being high in both groups. Electron microscope analysis showed a severe disruption of the mitochondrial elements in males spawning at 22°C.     

Long-term modeling of large-scale nutrient cycles in the entire Baltic Sea

Management of eutrophication in marine ecosystems requires a good understanding of nutrient cycles at the appropriate spatial and temporal scales. Here, it is shown that the biogeochemical processes controlling large-scale eutrophication of the Baltic Sea can be described with a fairly aggregated model: simple as necessary Baltic long-term large scale (SANBALTS). This model simulates the dynamics of nitrogen, phosphorus, and silica driven by the external inputs, the major physical transports, and the internal biogeochemical fluxes within the seven major sub-basins. In a long-term hindcast (1970–2003), the model outputs reasonably matched observed concentrations and fluxes. The model is also tested in a scenario where nutrient inputs are reduced to levels that existed over 100 years ago. The simulated response of the Baltic Sea trophic state to this very large reduction is verified by a similar simulation made with a much more complex process-oriented model. Both models indicate that after initial, rather rapid changes the system goes into much slower evolution, and nutrient cycles would not become balanced even after 130 years. Guest editors: J. H. Andersen & D. J. Conley Eutrophication in Coastal Ecosystems: Selected papers from the Second International Symposium on Research and Management of Eutrophication in Coastal Ecosystems, 20–23 June 2006, Nyborg, Denmark     

Denitrification,Nitrificastion and Nitrogen Fixation in Sublittoral Sediments of the South China Sea

Measurements of denitrification, nitrification and nitrogen fixation rates were made alongside with measuring of chemical and physical properties in sublittoral sediments of the South China Sea near the coast of Vietnam. Studied sediments were suboxic (Eh was positive as a rule), had 0.18–1.5 % of organic carbon, 0.004–0.135 % of total nitrogen and 3-12 % of total iron. The numbers of denitrifying and nitrate-reducing bacteria were as high as millions and hundreds of millions cells per gram wet weight of sediment matter, respectively. The processes of nitrification and denitrification were not spearated spacely. The nitrification was measured in both superficial layer and in a 10-cm sediment column. There were indirect evidences suggesting possibility of anaerobic ammonium oxidation. Denitrification was detectable in the sediments from two sites of sampling; maximal value was 86.2 μmoles N m⁻²h⁻¹. The denitrification potential determined at 1 mM nitrate decreased regularly from the upper to lower layers. Its values in the different sediments ranged from 134 to 532 μmoles N m⁻²h⁻¹. Nitrogen fixation (from 4.8 to 86μmoles N m⁻²h⁻¹) was close to that found in similar sediments in temperate waters in summer, and was not a significant source of nitrogen. It was comparable with diffusion of ammonium from sedimnts.