The fate of an immigrant: Ensis directus in the eastern German Bight

We studied Ensis directus in the subtidal (7–16?m depth) of the eastern German Bight. The jack-knife clam that invaded in the German Bight in 1978 has all characteristics of a successful immigrant: Ensis directus has a high reproductive capacity (juveniles, July 2001: Amrumbank 1,914?m^?2, Eiderstedt/Vogelsand: 11,638?m^?2), short generation times and growths rapidly: maximum growth rates were higher than in former studies (mean: 3?mm?month^?1, 2nd year: up to 14?mm?month^?1). Ensis directus uses natural mechanisms for rapid dispersal, occurs gregariously and exhibits a wide environmental tolerance. However, optimal growth and population-structure annual gaps might be influenced by reduced salinity: at Vogelsand (transition area of Elbe river), maximum growth was lower (164?mm) than at the Eiderstedt site (outer range of Elbe river, L _∞?=?174?mm). Mass mortalities of the clams are probably caused by washout (video inspections), low winter temperature and strong storms. Ensis directus immigrated into the community finding its own habitat on mobile sands with strong tidal currents. Recent studies on E. directus found that the species neither suppresses native species nor takes over the position of an established one which backs up our study findings over rather short time scales. On the contrary, E. directus seems to favour the settlement of some deposit feeders. Dense clam mats might stabilise the sediment and function as a sediment-trap for organic matter. Ensis directus has neither become a nuisance to other species nor developed according to the ‘boom-and-bust’ theory. The fate of the immigrant E. directus rather is a story of a successful trans-ocean invasion which still holds on 23?years after the first findings in the outer elbe estuary off Vogelsand.     

Different zooplankton structures in the German Bight

In August 1982, a net of 48 stations with altogether 208 samples was investigated in the eastern German Bight with respect to temperature, salinity, as well as the amount and species composition of the mesozooplankton (>80 μm). The data were arranged into different structures by means of a cluster analysis. Four different clusters were found: (a) a “Wadden sea water” with few holoplankton organisms but a higher amount of spionid larvae; (b) a “German Bight water” with a maximum occurrence of turbellaria (Alaurina composita) and medium concentrations of copepods; (c) a mixing area between these two water masses with highest amounts ofOikopleura dioica, Temora longicornis, Acartia sp., mussel larvae and larvae of the spionid worms; (d) a “North Sea water” mass with highest concentrations ofPseudocalanus elongatus, Paracalanus parvus undOithona similis. The differences in the concentrations of the species mentioned between the four clusters were significant on the 0.1%-level.     

Wrecks as artificial lobster habitats in the German Bight

Once, the European lobster could be found in high abundances on rocky substrate around the island of Helgoland. Since the 1960s, the stock has been decreasing dramatically. Until now, it has been assumed that the lobster stock of Helgoland is the only one in the German Bight. Here, we provide first information about lobster distribution inside the German Bight off Helgoland. Diving in situ observations revealed that lobsters inhabit at least 15.6% of all 64 investigated wrecks. Considering the difficulties of detecting lobsters at wrecks, the true percentage is most likely much higher. Their locations are spatially homogenously distributed throughout the inspected area. The study indicates a broad distribution of the European lobster over the German Bight. The habitats provided by a considerable fraction of the more than one thousand wrecks outside the Wadden Sea are potential lobster refuges within the mud and sand dominated sea floor. Besides providing additional habitats, they represent stepping stones enhancing the connectivity of the North Sea lobster population.     

Zooplankton community and hydrographical properties of the Neretva Channel (eastern Adriatic Sea)

Temporal and spatial variability of micro and mesozooplankton was studied in 1998 and 1999 at four stations in the Neretva Channel area influenced by the Neretva river and the open waters of the south Adriatic Sea. The area is orthophosphate limited, but an excessive accumulation of land derived nitrogen is prevented by phytoplankton uptake and the general circulation pattern. Microzooplankton was dominated by ciliates, with average abundances comparable to other Adriatic channel areas (122–543 ind. l⁻¹). Non-loricate ciliates (NLC) generally peaked in the warmer periods, but a winter increase was evident towards the inner part of the channel. Tintinnid abundances generally increased in autumn. A significant relationship with temperature was not recorded for either protozoan group. An inverse relationship between NLC and salinity might be indirectly caused by their preference for the food abundant surface layer. Mesozooplankton was dominated by copepods, with distinct summer maxima throughout the area and pronounced winter maxima of >10,000 ind. m⁻³ at the inner stations. The community was predominantly neritic but the open sea waters were important in structuring the mesozooplankton assemblage at all stations during the autumn–winter period. Although temperature regulated the seasonal dynamics of most metazoans and the species succession in the copepod community, small omnivorous copepods (Oncaea media complex, Oithona nana and Euterpina acutifrons) dominated regardless of the season. A trophic link between copepods and ciliates was evident in winter during low phytoplankton biomass.     

A population study of the shrimp Crangon allmanni in the German Bight

The only reproducing population of Crangon allmanni within the German Bight is in the Helgoland Trench (HTR), a depression of more than 50 m depth south of Helgoland. In coastal and shallower offshore waters the shrimp is much rarer and is recorded in higher numbers only in early spring when the water is still cold. The life cycle of the HTR population lasts for about 1.5 years maximum. Recruitment takes place in summer, and these recruits form the reproducing population of the next year. The shrimps are sensitive to environmental stress and are therefore good indicators of environmental changes. Electronic Publication     

Isolation of Novel Pelagic Bacteria from the German Bight and Their Seasonal Contributions to Surface Picoplankton

We tested new strategies for the isolation of abundant bacteria from coastal North Sea surface waters, which included reducing by several orders of magnitude the concentrations of inorganic N and P compounds in a synthetic seawater medium. Agar plates were resampled over 37 days, and slowly growing colonies were allowed to develop by repeatedly removing all newly formed colonies. A fivefold increase of colonies was observed on plates with reduced nutrient levels, and the phylogenetic composition of the culture collection changed over time, towards members of the Roseobacter lineage and other alpha-proteobacteria. Novel gamma-proteobacteria from a previously uncultured but cosmopolitan lineage (NOR5) formed colonies only after 12 days of plate incubation. A time series of German Bight surface waters (January to December 1998) was screened by fluorescence in situ hybridization (FISH) with isolate-specific and general probes. During spring and early summer, a prominent fraction of FISH-detectable bacteria (mean, 51%) were affiliated with the Cytophaga-Flavobacterium group (CF) of the Bacteroidetes. One Cytophaga sp. lineage with cultured representatives formed almost 20% of the CF group. Members of the Roseobacter cluster constituted approximately 50% of alpha-proteobacteria, but none of the Roseobacter-related isolates formed populations of >1% in the environment. Thus, the readily culturable members of this clade are probably not representative of Roseobacter species that are common in the water column. In contrast, members of NOR5 were found at high abundances (>10⁵ cells ml⁻¹) in the summer plankton. Some abundant pelagic bacteria are apparently able to form colonies on solid media, but appropriate isolation techniques for different species need to be developed.     

A seafloor crater in the German Bight and its effects on the benthos

 In 1963 a deep crater was formed about 65 m below sea level in the western part of the German Bight, due to a gas eruption caused by drilling carried out from the platform ’Mr. Louie’. The study area is situated in a sandy to muddy bottom area inhabited by an Amphiura filiformis association (sensu Salzwedel et al. 1985). The crater, sometimes called ’Figge-Maar’, functions as a sediment trap, concentrating particles and organisms from the water column, thus leading to extreme sedimentation rates of about 50 cm, on average, per year. Crater stations, compared with stations situated in the vicinity, show enrichments of juveniles. Echinoderms, especially the subsurface-dwelling heart urchin Echinocardium cordatum and ophiuroids are responsive to enrichment. Other species that are typical of the Amphiura filiformis association are shown to be unable to cope with the special conditions in the crater. Received: 12 June 1998 / Accepted:16 September 1998     

Biological interactions and their role in community structure in the rocky intertidal of Helgoland (German Bight,North Sea)

Over 3 successive seasonal cycles (April 1986 to October 1988), field experiments were established within 3 intertidal levels in the sheltered rocky intertidal of Helgoland (North Sea, German Bight). Competitors for space (Mytilus edulis, macroalgae), herbivores (Littorina spp.) and predators (Carcinus maenas) were either excluded from areas (0.25 m²) covered by undisturbed communities or enclosed at natural densities on areas that were cleared before of animals and plants. All the experimental fields (each 0.25 m²) were covered by cages with 4 mm gauze at the sides and a plexiglas top. The results of the experiments in the upper intertidal (occupied byLittorina spp. andEnteromorpha) showed that a natural density of herbivores could not prevent algal settlement and had only little influence on algal growth. Instead abiotic factors (storms, algae washed ashore) decreased the stock of the green algae. Experiments in the mid intertidal, dominated byMytilus (50% cover),Fucus spp. (20%) and grazingL. littorea (100 ind. m^?2) showed that community structure was directly changed both by grazing periwinkles and by competition for space between mussels and macroalgae. WheneverLittorina was excluded, the canopy ofFucus spp. increased continuously and reached total cover within two years. In addition to the increase ofFucus spp., the rock surface and the mussel shells were overgrown byUlva pseudocurvata, which covered the experimental fields during parts of the summer in the absence of herbivores. As soon as perennial species (fucoids) covered most of the experimental areas, the seasonal growth ofUlva decreased drastically. Presence and growth of macroalgae were also controlled by serious competition for space with mussels. EstablishedMytilus prevented the growth of all perennial and ephemeral algae on the rocks. However, the shells of the mussels provided free space for a new settlement ofFucus andUlva. In the lower intertidal (dominated by total algal cover ofF. serratus, herbivores such asL. littorea andL. mariae, and increasing number of predators such asCarcinus), the feeding activity of herbivores can neither prevent the settlement of the fucoid sporelings nor reduce the growth of macroalgae.F. serratus achieved a total canopy on the rock within one year. Doubled density of herbivores prevented the settlement ofFucus and most of the undercover algae. Predation byCarcinus onLittorina spp. had little influence on the herbivore community patterns. However, the crabs supported the establishment of macroalgae by excluding the mussels from the lower intertidal. In summary, the community organization and maintenance in the mid and lower intertidal is influenced to a high degree by biological interactions. Whereas both the relatively important herbivory byL. littorea and competition for space between mussels and macroalgae dominate in the mid intertidal, predation reaches its highest relative degree of importance for community structure in the lower intertidal.     

A multiannual outbreak of the turbellanan Alaurina composita Mecznikow 1865 in the German Bight

The plankton time series of ‘Helgoland Roads’ hasshown a multiannual population change in the abundance of theturbellarian Alaurina composita Mecznikow 1865. From the 1970sto the mid-1980s, the summer population increased by two ordersof magnitude; since then, it has fallen by about one order ofmagnitude. As limnetic investigations in related species show,A.cornposita must be considered a key predator of the NorthSea pelagial. Knowledge regarding the biology and ecology ofthe species is only rudimentary.     

Temporal patterns of microbial community structure in the Mid-Atlantic Bight

Although open ocean time-series sites have been areas of microbial research for years, relatively little is known about the population dynamics of bacterioplankton communities in the coastal ocean on kilometer spatial and seasonal temporal scales. To gain a better understanding of microbial community variability, monthly samples of bacterial biomass were collected in 1995-1996 along a 34-km transect near the Long-Term Ecosystem Observatory (LEO-15) off the New Jersey coast. Surface and bottom sampling was performed at seven stations along a transect line with depths ranging from 1 to 35 m (n=178). Microbial populations were fingerprinted using ribosomal 16S rRNA genes and terminal restriction fragment length polymorphism analysis. Results from cluster analysis revealed distinct temporal patterns among the bacterioplankton communities in the Mid-Atlantic Bight rather than grouping by sample location or depth. Principal components analysis models supported the temporal patterns. In addition, partial least squares regression modeling could not discern a significant correlation from traditional oceanographic physical and phytoplankton nutrient parameters on overall bacterial community variability patterns at LEO-15. These results suggest factors not traditionally measured during oceanographic studies are structuring coastal microbial communities.     

Contribution to the structure of respiration control]

A structure is reported which describes the relatively free availability of the motor respiratory apparatus to non-respiratory functions due to the presence of a respiratory control both on homeostatic and motor levels. It is assumed that, at the motor level, a competition develops of two or more regulators for the same parameter (e.g. the activity of neurons that can be claimed both by respiratory and by a second skeletal motor process). The compromise being reached is dependent on the amplification in the competing circuits. The correlation between the metabolic value and that of the respiration-time volume may be accomplished by the fact that amplification in the respiratory-motor circuit is rendered variable as a result of its interconnection with the chemical respiratory control. The high time constant of the latter allows for the instantaneous free availability of the motor respiratory apparatus to non-respiratory functions. The postulated structure offers better interpretations for the actual respiration behaviour than those considering the respiratory function alone.     

Contribution of vines to the evapotranspiration of a secondary forest in eastern Amazonia

The contribution of vines to the evapotranspiration (ET) of a secondary forest in eastern Amazonia was estimated based on field measurements of vine and tree transpiration, and seasonal changes in soil water content to 12 meters depth. Transpiration of vines and trees was measured with sapflow gauges placed around stems or branches. Total ET of the secondary forest was estimated as the sum of rainfall and reductions in soil moisture measured using Time Domain Reflectometry sensors installed in the walls of soil shafts. Our results suggest that vines transpire more than trees with stems of similar diameter, and with similar leaf crown exposure to sunlight. Trees experienced a smaller reduction in transpiration from the wet to the dry season than did vines. During the dry season, vines represented 8% (0.4 mm d^–1) of total secondary forest ET (5.4 mm d^–1), but they represented only 5.5% (0.5 m² ha^–1) of total secondary forest basal area (9.6 m² ha^–1). Considering that transpiration corresponds to 66–90% of forest ET, vines may contribute 9–12% to the transpiration of the forest. Hence, vine cutting, which is a commonly recommended management practice to favor the growth of tropical timber trees, may result in a proportionally larger reduction in evapotranspiration than in forest basal area.     

Importance and abundance of the recently established speciesCoscinodiscus wailesii Gran & Angst in the German Bight

Grids of 17 to 50 stations in the German Bight were sampled 18 times within the framework of the multidisciplinary programmes ZISCH and PRISMA in winter and spring of 1988/89 and from April 1991–April 1992. The frequent abundance ofCoscinodiscus wailesii Gran & Angst, a recently established large diatom, was noteworthy, as it dominated the phytoplankton biomass over long periods (e.g. 12/88–3/89 and 8/91–11/91). The bulk of the phytoplankton carbon during these periods (up to 90%) could be attributed to this species. Blooms ofCoscinodiscus wailesii producing up to 1400 μg carbon 1⁻¹, were recorded in early spring of 1989 and autumn of 1991. The potential consequences for the whole ecosystem refer to the huge size of this organism, which may cause reduced exploitation of its primary production by native consumers. Furthermore, sedimentation and remineralisation processes may be affected. ForC. wailesii, a doubling of biomass in 70 h could be estimated on the basis of data from four successive surveys in 1991. Results from laboratory cultures under comparable conditions confirm this rate. In the survey area, no increase in biomass was recorded for the relatedCoscinodiscus granii Gough, although there was sufficient silicate supply for growth. Field data showed 8–10 times lower copper and 10–20 times lower cadmium and zinc accumulation inC. wailesii, compared to concentrations found in native phytoplankton species. Subsequent laboratory tests suggested that one reason for the remarkable success ofC. wailesii, mainly in inshore regions, may be derived from its tolerance of higher doses of heavy metals due to very low sorption.     

The macrofauna of a stony sand area in the German Bight (North Sea)

The sublittoral macrofauna of the Steingrund, a stony area east-northeast of Helgoland, was investigated from May till October 1991 using a van Veen grab and a small dredge. The diverse endo- and epifauna of the sandy bottoms, pebbles and boulders of this Saalian end moraine comprised 289 taxa, whereby the polychaetesMagelona papillicornis, Lanice conchilega andSpiophanes bombyx dominated in terms of abundance. Species rare in the German Bight, such as the spongeLeucandra fistulosa, the sea urchinEchinus esculentus, and the sea anemoneHaliplanella lineata, were also found. The two sample sets were processed separately with multivariate techniques, and differentiated on the basis of occurrence and numbers of the abundant species. The analysis of the grab samples revealed two types of theTellina-fabula-community. These associations were differentiated by the presence of species of the coarse-sand-inhabitingGoniadella-Spisula-community and were related to the distribution of the grain size of the sediment. Likewise, two epifaunal assemblages were distinguished. Sandy bottoms were characterized byOphiura albida, Liocarcinus holsatus andPagurus bernhardus, while boulders and pebbles were covered by a varied sessile and mobile epifauna dominated by the sessile bryozoanFlustra foliacea and the mobile pantopodeAechelia echinata. Numerical density, biomass and annual production estimates are in the range of values determined for the macrobenthos of the German Bight, while annual P/B ratios mounted up to 5.     

Seasonal occurrence and abundance of caridean shrimp larvae at Helgoland,German Bight

Plankton samples were collected from January 1985 to January 1986 three times per week at Helgoland to study seasonal occurrence and abundance of caridean shrimp larvae. A total of eleven species were obtained. Ninety-one % of all larvae collected during the sample period belonged toCrangon crangon L. andCrangon allmanni Kinahan, 6% toPhilocheras trispinosus Hailstone and 3% to the remaining eight species. Collections were generally dominated byC. crangon larvae. However,C. allmanni larvae were most abundant in June coinciding with hatching activities of the population near Helgoland.C. allmanni was observed to have the highest density of all species with approximately 8 larvae per m³. Larvae ofEualus occultus (Lebour),Eualus pusiolus (Kroyer),Hippolyte varians Leach andAthanas nitescens Leach were most likely released by populations inhabiting the rocky intertidal zone around Helgoland. The presence ofProcessa modica Williamson & Rochanaburanon andProcessa nouveli holthuisi Al-Adhub & Williamson in the German Bight was verified by observations of a series of different developmental stages. Larvae of the rare speciesCaridion steveni Lebour were also recorded. The observed shrimp species were placed into three different groups with respect to their seasonal occurrence. Possible advantages of the timing of larval dispersal relative to predation and food availability are given. The results on seasonal occurrence and relative abundance are discussed in relation to environmental factors (temperature, salinity) as well as to the geographical distribution of the species.     

Community composition of the rocky intertidal at Helgoland (German Bight,North Sea)

At the rocky island of Helgoland (North Sea), the distribution and abundances of intertidal communities were assessed and the effects of wave exposure and tidal height on the spatial distribution patterns of the communities were evaluated. Macroalgae and invertebrates were sampled quantitatively along line transects in three intertidal locations, a semi-exposed, an exposed and a sheltered one. The semi-exposed location was characterised by (1) Ulva spp. at the high intertidal (Ulva-community), (2) mussels and periwinkles at the mid intertidal (Mytilus-community) and (3) Corallina officinalis and mainly the large brown alga Fucus serratus at the low intertidal (Fucus-community). The exposed location encompassed the mid and low intertidal; at both zones the Fucus-community occurred. The sheltered location was characterised by (1) barnacles (Balanus-community) and (2) bryozoans, hydrozoans and mainly the large brown alga Ascophyllum nodosum (Ascophyllum-community). At the semi-exposed, but not at the exposed location the communities changed with the intertidal position. A relationship between wave exposure and the occurrence of specific communities was shown for the sheltered location; in contrast, communities of the semi-exposed and the exposed location appear to be little influenced by wave exposure directly. The community concept and the potential causes of distribution patterns of the defined communities are discussed and suggestions for a future monitoring are given. Variations in the communities at different spatial scales speak in favour of a multiple scale sampling design to monitor changes in the intertidal communities at Helgoland. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Comparison of trace heavy-metal levels from monitoring in the German Bight and the southwestern Baltic Sea

Since 1973, monitoring for selected trace heavy metals has been performed in the German Bight (North Sea) and the western Baltic Sea. This paper deals with a set of filtered and unfiltered samples from a network of 22 stations covering the whole German Bight based on a cruise with RV “Gauss” in May 1974, and a similar series of samples taken at different depths at 18 stations in the Baltic Sea in October 1974. The metals Cd, Cu, Fe, Mn, and Ni were determined by flameless atomic absorption spectrometry. Data comparison is facilitated since sampling and analytical methods were identical. Distributions of metal-content data have been calculated from unfiltered and filtered samples. Qualitative and semi-quantitative aspects of the major differences or similarities between the histograms obtained from both sea areas are discussed.