The recent arrival of the oceanic isopodIdotea metallica Bosc off Helgoland (German Bight, North Sea): an indication of a warming trend in the North Sea?

In 1988 a long-term study was started of the isopod fauna associated with surface drift material off Helgoland (German Bight, North Sea). In the summer of 1994 specimens ofIdotea metallica Bosc were recorded for the first time. There is no evidence that this species has ever been present in the German Bight before. The samples contained males, both gravid and non-gravid females, and juveniles, indicating that the species reproduced successfully in the Helgoland region. Interbreeding of specimens from Helgoland and the western Mediterranean produced fertile off-spring. As a neustonic species,I. metallica shows a high natural capacity for dispersal. It thus seems unlikely that the arrival of the species in the North Sea resulted from an accidental introduction by man. We are probably witnessing an extension of the species’ geographical range by natural means of dispersal, as a response to recent changes in the ecological conditions of the German Bight. Temperature data measured by the Biologische Anstalt Helgoland since 1962 show that the last decade (except 1996) was characterized by unusually mild winters. Following the severe winter of 1996,I. metallica was again absent from the Helgoland region. After the subsequent mild winters (1997 and 1998), however, the species reappeared in the summer of 1998 with higher numbers than ever before. This suggests that the observed phenomena are closely connected with the recent temperature anomalies.I. metallica can be regarded as a potential immigrant to a warmer North Sea, and may be useful as a sensitive indicator of the predicted long-term warming trend.     

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)     

Ecological long-term research at Helgoland (German Bight,North Sea): retrospect and prospect—an introduction

This paper summarizes and evaluates ecological long-term observations at the island of Helgoland (German Bight, North Sea). It is an introduction to a series of seven contributions to an issue of Helgoland Marine Research (vol. 58, no. 4) dealing with observations on the physico-chemical environment and the local biota (pelagic bacteria, phytoplankton, zooplankton, macroalgae, macrozoobenthos). More than 150 years of research at Helgoland (at the Biologische Anstalt Helgoland, BAH, since 1892), and particularly the Helgoland Roads time-series which started in 1962, has provided a multitude of data which represents a valuable basis for analysing past changes, evaluating the present status and predicting future changes in ecosystem parameters. Predicting the consequences of the recent rapid ecological change on regional and global scales requires increased efforts of focus on long-term ecological observations. Future efforts in this field will rely on the application of innovative advanced technology and on the concerted activities of a large number of institutions which only collectively can establish the large-scale patterns of temporal and spatial change in ecosystem functions.     

Rapid increase of the grey seal (<Emphasis Type="Italic">Halichoerus grypus</Emphasis>) breeding stock at Helgoland

Over the past 25 years, the grey seal (Halichoerus grypus) population in the North Sea increased on average at 5% per year, while founding a number of new colonies. Based on published data, this note compares recent trends in pup production at nine breeding sites, with focus on the German island of Helgoland. Since 1997, numbers born at Helgoland increased at 35% per year (95% CI: 29–42), reaching an all-time high of 53 in 2007. Yet, this was less than 1% of all grey seal offspring in the North Sea area. Some other small stocks had similarly high growth rates of 20–50%, which must be explained by immigration from the larger colonies, as sustained autochthonous increase cannot exceed 13% per year in the species. Further, the Helgoland breeding stock increased significantly faster than the neighbouring colonies at Amrum (13%) and in the Dutch Wadden Sea (20%), indicating higher offspring survival and/or higher relative influx of migrants. This is attributed to the fact that at Helgoland newborn seals are safe from floodwater. In the other two sites, in contrast, pups are born on sandbanks and are thus often hit by winterly storm tides, which poses a risk to their survival. Because of the favourable breeding conditions at Helgoland, this new colony is expected to soon become an important stronghold of the grey seal in the Dutch-German North Sea area.     

Kreuzungsexperimente anLaminaria saccharina von Helgoland und von der Isle of Man

The brown algaLaminaria saccharina growing at Helgoland (North Sea) has smooth blades, whereas the blades of the same species occurring at the Isle of Man are covered with deep bullations. Gametophytes of both morphological types were grown in the laboratory and the resulting sporophytes of each type as well as hybrids were planted out in the sublittoral zone of Helgoland for four months. The Isle of Man type and the hybrids both formed bullations. The Isle of Man type differed further from the Helgoland type in that it had broader blades and did not tolerate the water temperatures above 14° C which prevail near Helgoland in summer. All experimental plants were removed from the sea before the blades became sporogenous.     

Meeresalgen von Helgoland

This report represents a photographic documentation of the more conspicuous species of the marine algal vegetation of the island Helgoland (North Sea), based on collections and observations made since 1959. The catalogue is largely restricted to macroscopic forms, as adequate knowledge of most of the microscopic ones is not yet available. Remarks on some rare species are added in a special chapter; the list includes some 150 species. With a few exceptions, the figures have been obtained from living algae; they illustrate aspects of reproduction and development and, occasionally, demonstrate different seasonal habits. The simplified key may be useful for providing a guide both for naturalists and students of marine phycology. In addition, this algal flora enhances our knowledge of the distribution of benthic species along the European coasts, as the rocky island of Helgoland intervenes between southern Norway and the shores of the English Channel.     

Influence of oceanic factors on Anguilla anguilla (L.) over the twentieth century in coastal habitats of the Skagerrak,southern Norway

The European eel (Anguilla anguilla L.) is distributed in coastal and inland habitats all over Europe, but spawns in the Sargasso Sea and is thus affected by both continental and oceanic factors. Since the 1980s a steady decline has been observed in the recruitment of glass eels to freshwater and in total eel landings. The eel is considered as critically endangered on the International Union for the Conservation of Nature and Natural Resources Red List of species. The Skagerrak beach seine survey from Norway constitutes the longest fishery-independent dataset on yellow/silver eels (starting in 1904). The Skagerrak coastal region receives larvae born in the Sargasso Sea spawning areas that have followed the Gulf Stream/North Atlantic Drift before they penetrate far into the North Sea. The Skagerrak coastal time series is therefore particularly valuable for exploring the impacts of oceanic factors on fluctuations in eel recruitment abundance. Analyses showed that Sargasso Sea surface temperature was negatively correlated with eel abundance, with a lag of 12 years revealing a cyclic and detrimental effect of high temperatures on the newly hatched larvae. The North Atlantic Oscillation index and inflow of North Atlantic water into the North Sea were negatively correlated with eel abundance, with a lag of 11 years. Increased currents towards the North Atlantic during high North Atlantic Oscillation years may send larvae into the subpolar gyre before they are ready to metamorphose and settle, resulting in low recruitment in the northern part of the distribution area for these years. The Skagerrak time series was compared with glass eel recruitment to freshwater in the Netherlands (Den Oever glass eel time series), and similar patterns were found revealing a cycle linked to changes in oceanic factors affecting glass eel recruitment. The recent decline of eels in the Skagerrak also coincided with previously documented shifts in environmental conditions of the North Sea ecosystem.     

Long-term variation in trematode (Trematoda, Digenea) component communities associated with intertidal gastropods is linked to abundance of final hosts

Trematodes are abundant parasites essential for maintaining stability of marine intertidal ecosystems. Despite the great ecological significance of trematodes, long-term dynamics of their communities remains practically unstudied. This study, based on 12-year-long seasonal monitoring of infection of Littorina and Hydrobia snails with 16 trematode species in the White Sea, aimed to reveal factors determining long-term variation in infection of intertidal snails by trematode parthenitae and larvae. Using the state-of-the-art method of singular spectrum analysis, we revealed trends in this variation and assessed their significance. Interestingly, these trends were not associated with oceanic and climatic parameters but were mostly determined by changes in abundance of the trematode final hosts. Moreover, the prevalence trends turned out to be connected with both large-scale events and local factors at the scale of the intertidal site. The main factors determining the long-term dynamics of the trematode component communities in the study area were the decreasing abundance of birds due to growing anthropogenic disturbance and the increasing abundance of the three-spined stickleback. The analysis of long-term trends of trematode prevalence in intertidal snails may be a sensitive indicator of the abundance dynamics of final hosts in the coastal areas of temperate and northern seas.     

Microbial diversity of cultivatable bacteria associated with the North Sea bryozoan Flustra foliacea

The microbial diversity of cultivatable bacteria associated with the bryozoan species Flustra foliacea from the North Sea was investigated by a molecular approach. Amplified ribosomal RNA restriction analyses (ARDRA) and 16S rDNA partial sequence analysis revealed differences in the composition of cultivatable bacteria populations from single bryozoan colonies collected from two different sampling sites in the North Sea as well from one site taken at different points in time. Whereas gamma-Proteobacteria identified as Shewanella frigidimarina, Pseudoalteromonas ssp. and Psycbrobacter ssp. were predominant on samples of Flustra I (taken near the island of Helgoland), most bacteria isolated from Flustra II, originating from the Steingrund, could be affiliated to Gram-positive taxa. Survey of the bryozoan samples from the latter site in February 2000 led to the detection of a phylogenetically mixed bacterial population, consisting of gamma-, and alpha-Proteobacteria and Gram-positive bacteria with low and high GC-content (Flustra III). As these bacteria are among the most widely isolated organisms from the marine environment, it may be concluded that the bryozoan Flustra foliacea accepts colonization of surfaces by bacteria which are common inhabitants of the marine environment and which may have been transferred into this environment from terrestrial sites.     

Species-Specific Bacterial Communities in the Phycosphere of Microalgae?

Specific associations of bacteria with phytoplankton have recently been reported in the literature. In our study, we analyzed bacterial communities of microalgal cultures related to algal growth phases. Seven freshly isolated key diatom and dinoflagellate species from Helgoland Roads, North Sea, were investigated. The community composition of associated bacteria as well as the cell numbers, the photosynthetic efficiency of the algae, and the depletion of inorganic nutrients in the medium were recorded over a period of 8 weeks in batch cultures. Diversity and succession of bacterial communities was analyzed by ribosomal intergenic spacer analysis. Phylogenetic analysis of bacterial populations was performed by denaturing gradient gel electrophoresis of 16S rRNA genes followed by DNA sequence analysis. Members of Alphaproteobacteria and Gammaproteobacteria and the Flavobacteria–Sphingobacteria group within the Bacteroidetes phylum predominated in the cultures. Differences in free-living and attached bacterial populations were observed between the phylogenetic groups. Shifts in the bacterial communities could not be correlated to changes of nutrient levels or algal growth phases. Regarding our results, it should not be generalized that the compositions of the bacterial communities are strictly species specific for microalgae. The importance of factors like the composition of exudates is apparent.     

Timing of spring migration in birds: long-term trends, North Atlantic Oscillation and the significance of different migration routes

We studied long-term trends and the yearly variation in mean spring passage time in 36 passerine bird species trapped at Ottenby Bird Observatory in south-eastern Sweden. Between the years 1952–2002, data were available for 22–45 years depending on species. Most long-distance migrant species passed progressively earlier over the study period (range: 2.5 days earlier to 0.7 days later per 10 years, with an average of 0.9 days earlier per 10 years). The annual variation in timing of migration in most species, regardless of migration distance, correlated negatively with the winter index of the North Atlantic Oscillation (NAO), a large-scale climate phenomenon influencing the climate in the North Atlantic region. Birds passed earlier after mild and humid winters, corresponding to the high phase of the NAO. This corroborates the pattern found at a nearby migration site with a comparable dataset (Helgoland, 600 km WSW of Ottenby). However, short/medium-distance migrant species at Ottenby, in contrast to the situation at Helgoland, have shown no general trend of earlier passage in recent years. This was probably a consequence of the shorter study period at Ottenby, which included only the last 22–32 years (41 years at Helgoland), when the NAO showed no significant trend. At the species-specific level, the long-term trends in passage time were similar at the two sites, and there was some congruence to the extent that a given species was affected by NAO. Long-distance migrants wintering south and south-east of the breeding grounds showed some of the strongest changes in long-term trends (passing progressively earlier) at Ottenby, and for some of these species passage time varied negatively with NAO. Obviously, and contrary to previous suggestions, variations in NAO also influence birds migrating through eastern Europe, although the direct or indirect mechanisms through which this is achieved are unknown.     

Historical changes in the benthos of the Wadden Sea around the island of Sylt in the North Sea

The situation regarding the distribution and abundance of seagrass, macroalgae and benthic fauna near the island of Sylt in the south-eastern North Sea during the period 1923 to 1940 is compared with that of the 1980s. Evidence of organic enrichment in recent times is provided by (1) massive growth of green algal mats on sheltered tidal flats, (2) a decline of red algae in the subtidal zone, (3) an expansion of mussel beds along low water line and down to 20 m depth, (4) increased abundance of polychaetes inhabiting intertidal and subtidal sandy bottoms. Seagrass beds have undergone complex changes which remain unexplained. Intensified erosion has contributed to the loss of habitats in the intertidal zone, and probably affected sessile epifauna in the deep channels. Here, direct removal and disturbance by the bottom-trawling fishery may also have contributed to the observed species impoverishment. Presented at the VI International Wadden Sea Symposium (Biologische Anstalt Helgoland, Wattenmeerstation Sylt, D-2282 List, FRG, 1–4 November 1988)     

THE TYPE SPECIES OF CERAMIUM (RHODOPHYTA), CERAMIUM VIRGATUM ROTH: TYPIFICATION AND PHYLOGENY

The type species of Ceramium is Ceramium virgatum Roth (1797, Cat. Bot I: 148, pl. VIII, fig. 1). This has been treated as a probable synonym of Ceramium nodulosum (Lightfoot) Ducluzeau, one of the species formerly confused under the illegitimate name Ceramium rubrum (Hudson) C. Agardh. However, in 1996 Silva showed that C. nodulosum (Lightfoot) Ducluzeau is a later homonym of C. nodulosum de Candolle 1805, and advocated conserving C. rubrum with a neotype. As an alternative to this, we investigated whether the name C. virgatum might be available for this species. The lectotype of C. virgatum is Roth's illustration of a much-branched, fully corticate, Ceramium specimen from Eckwarden, North Sea, which could represent any member of the ' C. rubrum' group. We obtained material resembling Roth's, from the North Sea island of Helgoland, as a potential epitype. Its rbc L sequence was aligned with sequences of all C. rubrum -like species in the British Isles, i.e. C. secundatum , C. botryocarpum , C. pallidum and ' C. nodulosum '. The Helgoland material was clearly conspecific with ' C. nodulosum ' from Ireland (0.08% divergence). We therefore suggest that the name C. virgatum Roth should be employed for this species. Phylogenetic analyses of these sequences aligned with various representatives of the tribe Ceramieae, placed the North Atlantic C. rubrum -like species in two well-supported clades. In one of these, C. virgatum is basal to C. secundatum and C. botryocarpum. The other clade consists of C. pallidum and a partly ecorticate species resembling C. diaphanum.     

Eine “red tide” in der südlichen Nordsee und ihre Beziehungen zur Umwelt

Phytoplankton blooms have attracted attention since times of old, especially the red tides caused by peridinians. Observed in all regions of the world, red tides occur predominantly in the warmer seas (e. g. West Coast of Florida). An opportunity to study a bloom, caused by the non-toxic dinoflagellateCeratium furca, was provided in North Sea coastal waters of an area ranging from Belgium to Sweden. Its development could be followed from July to October 1981 with the aid of samples taken 5 days a week near the island of Helgoland. A decline in the bloom coincided with a drop in water temperature (increasing turbulence). This condition and the presence, right up to the end, of relatively high amounts of nutrients indicate that nutrient deficiency did not terminate the bloom. Similarly, there was no evidence for a dependence on micronutrients by freshwater input or by the preceding diatom bloom. Longterm investigations indicate that climatic changes coincide with an increase in the phytoplankton stock during all seasons (especially noticeable from July to September) and with a decrease in salinity (there was a minimum in 1981). The stratification of the sea water causes the peridinians to move to depths optimal for their growth which may bring about an aggregation at the surface. There are numerous indications in literature of similar changes of environmental conditions in the North Sea. One example from the Western Baltic Sea demonstrates that the presence of a high vertical density gradient in the sea leads to oxygen deficiency in deeper zones, which, however, is not directly correlated to the coincident phytoplanktonic bloom in the upper layers. It is probable that, under appropriate hydrographical conditions, “red tides” can occur repeatedly in the southern North Sea, produced by non-toxic as well as toxic species.     

Unchanging diet in a stable colony: contemporary and past diet composition of black-legged kittiwakes <Emphasis Type="Italic">Rissa tridactyla</Emphasis> at Helgoland,south-eastern North Sea

In contrast to the situation at the west coast of the North Sea, the breeding colony of black-legged kittiwakes Rissa tridactyla at Helgoland in the south-eastern North Sea did not exhibit severe declines since 1990 but instead numbers increased and only lately stabilised. Declines at the west coast of the North Sea were attributed to a lower abundance and lower quality of the key prey, sandeels. We hypothesised that kittiwakes at Helgoland do not rely as heavily on sandeels as their conspecifics. We analysed stomach contents of nestlings and adults of 2001, 2002, 2004 and 2006. In concordance with earlier studies of the 1980s and 1990s, young whiting Merlangius merlangus was the most important prey species in 2001, 2002 and 2004. Clupeids and sandeels were consumed in lower proportions. While earlier studies suggested whiting to originate from fisheries discards, evidence now supports that kittiwakes prey upon whiting in areas of hydrographic fronts. No whiting was recovered in samples of 2006 and the proportion of fish prey was low. Main prey items were polychaete worms (Nereidae), which were presumably consumed as swarming Heteronereis stages. An observed strong rise in water temperature in summer 2006 might have influenced food availability of kittiwakes by inducing swarming of Nereidae. Overall, kittiwakes breeding on Helgoland showed a positive population trend for several decades while mainly feeding on whiting.     

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.     

Ostreobium quekettii (Codiales,Chlorophyta)

The shell-inhabiting, marine algaOstreobium quekettii Bornet & Flahault is a distinct species reproducing by zoospores with four flagella. Sporangium formation and the development of zoospores are described. The material obtained from Helgoland (North Sea) is assumed to be identical with the type specimen from the French Atlantic coast.O. quekettii is said to be of worldwide distribution; in the absence of further information, it must be doubted whether other finds reported to belong to this species are identical with the type species.     

New complete mitochondrial DNA sequence of the lancelet Branchiostoma lanceolatum (Cephalochordata) and the identity of this species' sequences

Three mitochondrial (mt) genes were sequenced for two Atlantic lancelet species, Branchiostoma lanceolatum and B. floridae, to examine a serious discrepancy among previously published results of molecular studies: substantial sequence difference in a nuclear gene vs. virtual identity in the mt genome sequence. The results revealed that three mt genes of B. lanceolatum, collected from Helgoland in the North Sea and Naples in the Mediterranean, were quite diverged from those of B. floridae, collected from Tampa Bay, Florida. Therefore, the previously recognized identity in the mt genome between the two species is attributable to misidentification of materials used. To correct this misleading information, the complete mtDNA sequence of B. lanceolatum was determined for an individual from Helgoland.