Tidal impact on planktonic primary and bacterial production in the German Wadden Sea

 Tidal variation of biological parameters was studied at three anchor stations in selected inlet channels of the northern German Wadden Sea in May and July 1994. Concentrations of bacteria, chlorophyll a and suspended matter as well as primary and bacterial production were assessed over a period of 25 h in the surface and in the bottom water. Diurnal variation in primary production was found both under in situ light conditions and under constant illumination. Tidal turbulence caused the introduction of detritus, bacteria and pigments from the sediment into the water column. The impact of sediment resuspension was most evident in the bottom water, leading to tidally oscillating bacterial production rates which were high during high stream velocity and low during the slack times. Estimations of the areal daily phytoplankton production and corresponding bacterial carbon demands were unbalanced. Primary production accounted for only 25–45% of the total bacterial carbon requirement. This discrepancy is due to the shallow euphotic depth in the Wadden Sea, allowing net primary production only in the upper 2–3 m of the water column, while the relatively high levels of bacterial activity do not show a vertical decline. Assuming that the specific biological activities in the water columns over the tidal flats are similar to those found in the inlet channels, it was found that production processes dominate in shallow areas whereas decomposition processes dominate in the deep channels. Moreover, the predominance of heterotrophic processes in the inlet channels means that additional organic carbon sources must contribute to the heterotrophic metabolism in the deep parts of the Wadden Sea, and that the horizontal flux of material is important in this turbid mesotidal ecosystem. Received: 18 April 1998 / Accepted: 12 November 1998     

Nitrogen removal in coastal sediments of the German Wadden Sea

Although sediments of the German Wadden Sea are suspected to eliminate a considerable share of nitrate delivered to the SE North Sea, their denitrification rates have not been systematically assessed. We determined N₂ production rates over seasonal cycles (February 2009–April 2010) at two locations with two sediments types each, the first site (Meldorf Bight) receiving nitrate during all seasons from the Elbe river plume, and a second site on the island of Sylt, where nitrate is depleted during summer months. In sediments from the Sylt site, N₂ production ranged from 15 to 32 μmol N₂ m^?2 h^?1 in the fine sand station and from 7 to 13 μmol N₂ m^?2 h^?1 in the coarse sand station; N₂ production was not detected when nitrate was depleted in May and July of 2009. N₂ production in the Meldorf Bight sediments were consistently detected at higher rates (58–130 μmol N₂ m^?2 h^?1 in the very fine sand station and between 14 and 30 μmol N₂ m^?2 h^?1 in the medium sand station). Analysis of ancillary parameters suggests that major factors controlling N₂ production in coastal sediments of the German Wadden Sea are the nitrate concentrations in the overlying water, the ambient temperature, and the organic matter content of the sediment. Extrapolating our spot measurements to the zone of nitrate availability and sediment types, we estimate an annual nitrogen removal rate around 16 kt N year^?1 for the entire northern sector of the German Wadden Sea area. This corresponds to 14% of the annual Elbe river nitrogen load.     

Distribution patterns of bacterioplankton and chlorophyll-a in the German Wadden Sea

 In a first synoptic evaluation, the temporal and spatial distribution of bacterioplankton and chlorophyll-a were determined in the German Wadden Sea. Three surveys were undertaken in winter, spring, and summer of 1994 using up to eight ships simultaneously between the river Ems and Sylt island. Despite intensive hydrodynamic mixing of the Wadden Sea water, spatial gradients were obvious. The abundance of bacterioplankton ranged from 0.4 to 26×10⁵ ml^–1 and chlorophyll-a varied between <0.1 and 79 μg l^–1. In winter, relatively homogeneous distribution patterns of both parameters with small gradients were found. Highest chlorophyll-a values connected with a highly patchy structure were observed in spring, while in summer both total chlorophyll-a values and the complexity of the distribution pattern had decreased. In contrast, bacterial numbers increased steadily from January to July with the highest bacterial densities and greatest patchiness observed in summer. Moreover, in some regions of the Wadden Sea, a trophic succession of algae as carbon producers and bacteria as consumers was evident. Correlation analysis verified the relationship between bacteria and chlorophyll a, indicating bottom-up control of bacterial abundance in the northern part of the German Wadden Sea. Since the observed regression slope is remarkably low (0.12–0.46) compared to literature values (0.5–0.8), we suggest that the link between phytoplankton and bacteria found here is a special characteristic of the Wadden Sea as a transition zone between the coastal region and the outer North Sea. Received: 26 May 1998 / Accepted:12 November 1998     

Long-term population dynamics of breeding bird species in the German Wadden Sea area

For no other group of organisms in coastal areas are there so exact and long-term data available as there are for seabirds. Since the beginning of the 20th century, documentation of population size, especially for species breeding in colonies from the groups gulls, terns and auks, is almost complete. These species act as bio-indicators, and data on fluctuations in their population size are useful as they reflect changes in the state of the marine ecosystem. The population development of some of these seabird species (Herring Gull, Guillemot, Common, Arctic and Sandwich Tern) from the German North Sea coast, which primarily feed on fish, is given. Common to all these species is an exponential increase in numbers in recent years (1970–1985). Possible causes for this development, e.g. pressure from enemies or competitors, availability of breeding places, anthropogenic stress and mortality factors, as well as the direct and indirect anthropogenic-influenced changes in the trophic system due to the increasing eutrophication of coastal waters, are evaluated. Signs of a collapse in the stocks of seabrids resulting from environmental pollution are discussed. Consequences resulting from the ecosystem changes, such as reduction of nutrient discharge into the North Sea and the expansion of biological monitoring, are described. Presented at the VI International Wadden Sea Symposium (Biologische Anstalt Helgoland, Wattenmeerstation Sylt, D-2282 List, FRG, 1–4 November 1988)     

Habitat selection of wintering passerines in salt marshes of the German Wadden Sea

The salt marshes of the Wadden Sea are important wintering areas for some species of granivorous passerines, which have declined considerably since the 1960s. We investigated the habitat choice of all wintering passerines in eight study areas in German salt marshes with special consideration of human impact on these habitats. Granivorous species that almost exclusively winter in salt marshes, Shorelark (Eremophila alpestris), Snow Bunting (Plectrophenax nivalis) and Twite (Carduelis flavirostris) were concentrated in the lower salt marsh vegetation and in the driftlines, while all other species preferred the high upper salt marsh communities, although Rock Pipits (Anthus petrosus littoralis) fed in muddy areas along ditches. Shorelarks switched habitat in conditions where seeds were scarce to feed instead on arthropods in upper salt marshes. Intensively sheep-grazed upper salt marshes resemble lower salt marshes in their vegetation and were therefore mainly visited by Shorelarks, Snow Buntings and Twites. In winter, the driftline is preferred by the two former species, while in autumn and spring more birds foraged in the salt marshes. Twites prefer to feed mainly on seeds of Salicornia. Areas with S. europaea are visited mainly in late autumn and early winter, while areas with S. stricta are used throughout the winter because of a steady supply of seeds. Several years after embankment, polders are hardly used any more by the lower salt marsh species as the habitat changes into freshwater marshes. Large embankment projects since the early 1960s have included salt marshes and intertidal flats, and the resultant loss of habitat is responsible for the decline of lower salt marsh species. For other passerine species the effects of reclamation are unknown. The effects of intensified grazing on the wintering populations of Shorelark, Snow Bunting and Twite are still unresolved. Although grazing supports lower salt marsh vegetation, the seed production per plant is much lower there and some important seed producers hardly occur. Since grazing was reduced and embankment projects have been stopped, the salt marsh areas (especially lower salt marshes) have increased and so have the wintering populations of Shorelark, Snow Bunting and Twite. For the other species, the consequences of habitat changes are unknown, although it is suggested that reduced grazing will support them. Reducing the human impact on salt marshes will, in the long run, probably lead to a natural salt marsh with much variety in elevation and in its corresponding vegetation and bird communities. Meanwhile, management by grazing might be required in parts of the salt marshes.     

Antibiotic production by a Roseobacter clade-affiliated species from the German Wadden Sea and its antagonistic effects on indigenous isolates

A strain affiliated with the Roseobacter clade and producing a new antibiotic named tropodithietic acid (L. Liang, Ph.D. thesis, University of G?ttingen, G?ttingen, Germany, 2003) was isolated from the German Wadden Sea. The compound showed strong inhibiting properties with respect to marine bacteria of various taxa and marine algae. Antibiotic production was found to occur during the complete growth phase. Strain mutants without antagonistic properties appeared several times spontaneously.     

Management of the Wadden Sea

The Wadden Sea situated along the North Sea coasts of Denmark, the Federal Republic of Germany and The Netherlands represents one of the world's largest bar-built type of estuaries. The area is a typical sedimentation and mineralization basin, with a large influx of organic matter from the adjoining North Sea, consequently a delicate oxygen balance and a rich benthic macrofauna, poor in species, which serves as food for juveniles of some commercially important North Sea fishes and for large numbers of migrating and wintering waders and waterfowl. Past and present activities of the human society in the area include fisheries (mainly for shrimp and mussels, semi-culture), shipping, land reclamation, recreation, dredging for sand and shells, and waste discharge from industries and human communities. Until the present these activities, although sometimes conflicting, did not fundamentally affect the area and its biota (pollution excluded), but future claims, including the construction of large deep-sea harbours, drilling for natural gas and oil, large-scale land reclamation and increased industrialization etc., might gradually induce degradation. For instance, area reduction by continued land reclamation could lead to irreversible losses of specific biotopes (e. g. salt-marshes, mud-flats), which could affect the size of bird and fish populations in a much wider region. Increased pollution, which has already inflicted damage on bird and seal populations, could reduce the fauna and hence the value of the area as a natural sanctuary. In the event of a proposal for a new human activity in the area, the present standing practice in the countries concerned requires an evaluation of its safety and economic aspects and its environmental impact. However, the various plans are considered separately and there is a general need for integrated management of the area.     

Assessment of primary production in the White Sea

The literature and original data on the primary production of phytoplankton in the White Sea are analyzed. By this parameter, the White Sea is significantly inferior only to the Chukchi Sea; it is similar to the Barents Sea, and exceeds other Russian Arctic seas by two to three times (the Kara Sea, Laptev Sea, and East Siberian Sea).     

A review of data on production and import of organic carbon in the western Wadden Sea

Summary Annual phytoplankton primary production in the tidal channels of the western Wadden Sea cannot be estimated more precisely than 150±50 g C.m^–2a^–1, due to large spatial and short-term temporal variations. This implies that year-to-year variations and eventually long-term trends have to be very pronounced to be measurable even with a weekly sampling program.Short-term temporal variation in primary production of microphytobenthos living on the tidal flats is less pronounced, but spatial variation is large. Primary production on high tidal flats is larger than on flats lower in the tidal zone. Year-to-year variation on a tidal flat station occupied now for 12 years was large. This variation cannot be explained by year-to-year variations in nutrients, light or temperature, but probably by year-to-year variations in grazing. Macrophytobenthic primary production plays a subordinate role in the western Wadden Sea after the disappearance ofZostera fields. It is difficult to give one figure for the annual primary production of an average tidal flat due to the variations mentioned. Tentatively, microphytobenthic plus macrophytobenthic plus phytoplankton primary production on and above an average tidal flat is estimated at 150±50 g C.m^–2a^–1,i.e. the same as estimated for phytoplankton primary production in the tidal channels.The western Wadden Sea receives a considerable amount of particulate organic carbon from outside the area, estimated at 240 g C.m^–2a^–1. Formerly the North Sea was thought to be the only source. New data indicate also an import from the Ysselmeer. The import of total organic carbon, particulate plus dissolved, from the Ysselmeer exceedsin situ primary production. From this import Ysselmeer DOC is probably not used very much in the Wadden Sea. However, Ysselmeer POC, consisting for 50% on average of living phytoplankton cells, may form a suitable food source for Wadden Sea invertebrates. Cell counts ofScenedesmus sp., a freshwater alga used as a natural tracer for Ysselmeer POC in the western Wadden Sea, indicate that a large part of the Ysselmeer POC is retained here.     

Large-Scale Spatial Dynamics of Intertidal Mussel (Mytilus edulis L.) Bed Coverage in the German and Dutch Wadden Sea

Intertidal blue mussel beds are important for the functioning and community composition of coastal ecosystems. Modeling spatial dynamics of intertidal mussel beds is complicated because suitable habitat is spatially heterogeneously distributed and recruitment and loss are hard to predict. To get insight into the main determinants of dispersion, growth and loss of intertidal mussel beds, we analyzed spatial distributions and growth patterns in the German and Dutch Wadden Sea. We considered yearly distributions of adult intertidal mussel beds from 36 connected tidal basins between 1999 and 2010 and for the period 1968–1976. We found that in both periods the highest coverage of tidal flats by mussel beds occurs in the sheltered basins in the southern Wadden Sea. We used a stochastic growth model to investigate the effects of density dependence, winter temperature and storminess on changes in mussel bed coverage between 1999 and 2010. In contrast to expectation, we found no evidence that cold winters consistently induced events of synchronous population growth, nor did we find strong evidence for increased removal of adult mussel beds after stormy winter seasons. However, we did find synchronic growth within groups of proximate tidal basins and that synchrony between distant groups is mainly low or negative. Because the boundaries between synchronic groups are located near river mouths and in areas lacking suitable mussel bed habitat, we suggest that the metapopulation is under the control of larval dispersal conditions. Our study demonstrates the importance of moving from simple habitat suitability models to models that incorporate metapopulation processes to understand spatial dynamics of mussel beds. The spatio-dynamic structure revealed in this paper will be instrumental for that purpose.     

Red List of marine macroalgae of the Wadden Sea

This list forms part of the Report on the RED LISTS OF BIOTOPES, FLORA AND FAUNA OF THE TRILATERAL WADDEN SEA AREA. For basic information concerning, for example, function of these lists, species taken into account, structure of the lists and abbreviations used, see also the general introduction to the Red Lists.     

Planktonic primary production in a tidally influenced mangrove forest on the Pacific coast of Costa Rica

The seasonal variation of planktonic primary productivity was measured during one year in the main channel in the interior part of the mangrove forest of the Estero de Morales (Estero de Punta Morales), a mangrove system located in the Golfo de Nicoya at the Pacific coast of Costa Rica. Samples were incubated at the surface, 0.5 m and 1.0 m depth and the "light and dark bottle technique" was employed. The annual gross primary productivity (PPg) was 457 and the net primary productivity (PPn) was 278 g C m(-2) a(-1). Daily PPg ranged from 0.29 to 3.88 and PPn from 0.12 to 2.76 g C m(-2) d(-1). The highest rates observed in May and September were due to red tide blooms. The seasonal variation of primary productivity inside the mangrove forest depends closely on the PP in the adjacent area of the upper Golfo de Nicoya. Obviously the PP was light-limited since the compensation depth in the ebb current was found at only 1 m depth. In the flood current it was somewhat deeper. The planktonic primary productivity inside the mangrove forest was completely restricted to the open channels. A simultaneous measurement demonstrated that PPn of the phytoplankton could not take place under the canopy of the mangroves. Additional studies on the time course of the oxygen concentration in the mouth of the main channel over 24 hrs demonstrated a relation between the O2 and the tidal curves. The ebb current had always lower O2 concentrations than the flood current, regardless of the time of the day. The difference to the foregoing high tide, however, was much smaller when the low tide occurred during the day. This indicates that under the canopy the net primary production and hence O2 liberation of the attached macro- and microalgae, together with the high PPn of the phytoplankton in the channels, helped the oxygen concentration not to decrease as far as during the night. Nevertheless it shows that the consumtion of organic material in the submersed part of the mangrove forest exceeds always its production.     

Concentrations of particulate matter in the North Frisian Wadden Sea

The amount of particulate matter for different size classes was determined in the North Frisian Wadden Sea over a period of 10 months from November 1973 to August 1974. Particle volume, particulate organic carbon and phaeopigment concentrations decrease with advancing season, reaching minima in June. It is concluded that concentrations of particulate matter including chlorophyll were sufficiently high throughout the study period to support the growth and reproduction of appendicularia which abundantly occur in the Wadden Sea.