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.     

Long-term changes in the macrozoobenthos around the rocky island of Helgoland (German Bight,North Sea)

The paper briefly summarizes what is known about long-term changes (facts, causes, consequences) in the macrozoobenthos of intertidal and subtidal hard-bottom communities around the island of Helgoland (German Bight, North Sea). There is increasing observational evidence that these communities (spectrum and abundances of species) are changing on a long-term temporal scale. The reasons are diverse and mainly anthropogenic. A shift in North Sea climate towards more oceanic conditions may be among the most important factors driving the recent changes in species spectrum. Many of the species which have been recorded as new to the Helgoland area during the past decade are southern (oceanic) species which may be considered as indicators of a warming trend.Communicated by K. Wiltshire     

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.     

The epizoic diatom community on four bryozoan species from Helgoland (German Bight, North Sea)

The composition of the diatom community on the bryozoans Electra pilosa, Membranipora membranacea, Flustra foliacea, and Alcyonidium gelatinosum from the German Bight was studied by light and scanning electron microscopy. In total, members of 26 diatom genera were found, with Cocconeis, Tabularia, Licmophora, Amphora, and Navicula being the most abundant. The amount and the composition of the diatom covering seem to be typical for single bryozoan species. Electra pilosa and Membranipora membranacea showed a rather dense covering with 71–547 cells/mm² and 77–110 cells/mm², respectively. The most prominent genus on Electra pilosa was Cocconeis, reaching up to 58% of all diatoms in one sample, followed by Navicula, Tabularia and Amphora. The most abundant genera on Membranipora membranacea were Tabularia and Licmophora, making up almost 70% of all diatoms in one sample, followed by Navicula, Cocconeis and Amphora. The diatom composition was very stable on all Electra samples, but varied on Membranipora samples. With <1–27 cells/mm², diatoms were much less abundant on Alcyonidium gelatinosum. Members of the genera Tabularia and Navicula were the most frequently found benthic diatoms, whereas the planktonic forms Coscinodiscus, Cyclotella, and Thalassiosira made up 35% of the diatoms. On Flustra foliacea, diatoms were virtually absent, with fewer than 5 cells/mm². The low diatom numbers are probably due to toxic metabolites produced by the host. The same may be true for Alcyonidium gelatinosum, but here they might also be a consequence of the surface properties of the bryozoan. Electronic Publication     

A revised checklist of planktonic diatoms and dinoflagellates from Helgoland (North Sea,German Bight)

A checklist based on net samples taken twice weekly from 2001 until May 2003 is presented. Identification is based on observations under direct light microscopy and after taking some organisms in culture. The checklist includes 227 taxa observed at the Helgoland Reede sampling station. One hundred and thirty-two species of diatoms from 53 genera and 95 species of dinoflagellates from 35 genera have been recorded from net samples and cultures. Thirty-five diatom and 28 dinoflagellate taxa were documented in the Helgoland phytoplankton for the first time. The list does not claim to be complete, but provides an updated list of the taxa found at Helgoland and, for convenience, also includes data published for different adjacent areas.Communicated by K. Wiltshire     

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.     

Changes in the macrozoobenthos of the intertidal zone at Helgoland (German Bight, North Sea): a survey of 1984 repeated in 2002

Changes in the presence and absence of invertebrates as well as in species conspicuousness were documented in a rocky intertidal community based on surveys in 1984 and 2002. In 2002 six vertically and/or morphologically different stations of an intertidal platform were sampled. Five of these six habitats had already been surveyed in 1984. Replicating precisely the method of the first assessment, presence/absence changes as well as changes in species conspicuousness of 83 invertebrate species were documented, indicating that this intertidal community changed considerably during the 18-year interval. Compared with the study in 1984, 27 species newly appeared, whereas 32 species disappeared. Furthermore, 16 species increased in conspicuousness, whereas eight invertebrates decreased. The total number of species in 2002 was 154 versus 158 in 1984. Although algal species were not recorded as thoroughly as invertebrates, a massive decline in cover of Halidrys siliquosa was noted. Conversely, two invasive algal species became established after 1984, Sargassum muticum (since 1988), a cosmopolitan fucoid alga that prefers shallow subtidal areas for colonization, and Mastocarpus stellatus (introduction in the 1980s) that particularly colonized areas in the mid intertidal. In 1984 the mid intertidal zone was dominated by the brown alga Fucus serratus, whereas in 2002 the blue mussel Mytilus edulis and the periwinkle Littorina littorea were the most conspicuous organisms. Annual mean sea surface temperature (BAH measurements) warmed by 1.1°C over the past four decades. Range-related community shifts, introductions of non-indigenous species and the input of pollutants, are considered to explain long-term ecological changes in the invertebrate community at Helgoland.     

Prevailing sea surface temperatures inhibit summer reproduction of the kelp Laminaria digitata at Helgoland (North Sea)

The impact of abiotic factors on kelp sporophyte reproduction has rarely been investigated. Laminaria digitata (Hudson) J.V. Lamouroux is one of the few summer fertile Laminaria species worldwide and reproduction is subjected to relatively high water temperatures. We investigated the impact of prevailing summer temperatures (~18°C in August) on the induction of sporangia, meiospore release, and germination at the island of Helgoland (North Sea). At Helgoland, fertile sporophytes are found between April and December with a maximum in late summer. While released meiospore numbers were constant between June and October, germination rates decreased significantly in summer. Short‐term exposure of mature sori to 17°C–22°C induced a significantly higher meiospore release indicating enhancement of sporulation by elevated temperatures. Induction of sporangia on vegetative blade disks was not possible at 20°C, and fertility was only 20% at 18°C–19°C, but it was 100% in cool temperatures of 1°C–10°C. It was shown for the first time in a kelp species that “sporogenesis” is the life‐cycle process with the narrowest temperature window compared to growth or survival of the sporophyte or reproduction, growth, and survival of the gametophyte. We incorporated several parameters (induction time, fertile area, and relative fertility) into a “Reproductive efficiency index.” This indicates that sporogenesis of L. digitata is a cold‐adapted process with an optimum at (5)–10°C. The results show that the population at Helgoland is at its reproduction limit despite the existence of other geographically more southerly located populations.     

40-year long-term study of microbial parameters near Helgoland (German Bight,North Sea): historical view and future perspectives

Since 1873, the waters at Helgoland Roads (sampling station Kabeltonne) have been sampled daily to determine temperature and salinity. In 1962, microbiological parameters were determined for the first time to establish microbiological long-term studies on marine bacteria, starting with the colony-forming units (CFU). In the following years, several other microbiological parameters were integrated for different periods of time (e.g. activity parameters like ATP and ectoenzymatic activity, marine yeasts, oil-degrading bacteria, flagellates and molecular methods like PCR followed by denaturing gradient gel electrophoresis). To date, the total count of bacteria, flagellates and viruses have been acquired using fluorescent DNA dyes and epifluorescence microscopy. Here we present both a historical overview of the methods used and examples of results obtained over the past 40 years. Furthermore, we try to evaluate challenging new methods for marine microbial ecology, appropriate for long-term studies of marine bacteria.Communicated by K. Wiltshire     

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.     

Dinoflagellates and ciliates at Helgoland Roads, North Sea

A monitoring programme for microzooplankton was started at the long-term sampling station “Kabeltonne” at Helgoland Roads (54°11.3′N; 7°54.0′E) in January 2007 in order to provide more detailed knowledge on microzooplankton occurrence, composition and seasonality patterns at this site and to complement the existing plankton data series. Ciliate and dinoflagellate cell concentration and carbon biomass were recorded on a weekly basis. Heterotrophic dinoflagellates were considerably more important in terms of biomass than ciliates, especially during the summer months. However, in early spring, ciliates were the major group of microzooplankton grazers as they responded more quickly to phytoplankton food availability. Mixotrophic dinoflagellates played a secondary role in terms of biomass when compared to heterotrophic species; nevertheless, they made up an intense late summer bloom in 2007. The photosynthetic ciliate Myrionecta rubra bloomed at the end of the sampling period. Due to its high biomass when compared to crustacean plankton especially during the spring bloom, microzooplankton should be regarded as the more important phytoplankton grazer group at Helgoland Roads. Based on these results, analyses of biotic and abiotic factors driving microzooplankton composition and abundance are necessary for a full understanding of this important component of the plankton.     

The warming trend at Helgoland Roads,North Sea: phytoplankton response

We combine the temperature and phytoplankton data from one of the longest aquatic data sets in history, the Helgoland Roads (North Sea, 54°11.3N, 7°54.0E) timeseries to evaluate the effects of climate change on the base of marine food webs. The data shows that, despite an obvious warming of 1.1°C since 1962, the mean diatom day of the algal spring bloom is delayed and shifted to the end of the first quarter of the year. This is apparently related to a warming of the autumn (October–December) temperatures. It is the first indication of a warming related shift in phytoplankton succession, the consequences of which would range from lifecycle/food resource mismatches to regime shifts in the North Sea system.     

Gymnodinium blooms in the Helgoland bight (North Sea) during August, 1968

Summary 1. As in other parts of the North Sea, dinoflagellate red tides occurred in Helgoland waters in August, 1968. Measurements of plankton, and physical and chemical water properties at the permanent station Helgoland Roads were analyzed to describe the blooms. In addition, planktological and hydrographical investigations at three areas south, southwest and northwest of Helgoland on 27 and 28 August, as well as at two drifting stations off the mouths of the Elbe and Eider rivers on 6 and 8 August, were used for this work.2.Gymnodinium sp. was abundant at all these localities, forming blooms near Helgoland from 14 to 30 August, with a maximum of 3 to 3.25 × 10⁶ cells/l (and 18 to 19µg chlorophylla/l) on 28 and 30 August at Helgoland Roads. The primary production was as high as 0.98 mg C/l in 6 hours in a suspension of 3.7 × 10⁶ Gymnodinium/l, where diatoms had been removed. This means that one millionGymnodinium produced 0.265 mg C in 6 hours. At Helgoland Roads nitrate and nitrite were depleted at times, but not phosphate.3. About 3 × 10⁶ Gymnodinium sp./l were found in the upper 16.5 m of water investigated around Helgoland on 27 and 28 August (maximal 7.8 × 10⁶ cells/l at 3 m). There was a marked vertical stratification ofGymnodinium with a concentration towards the surface during the day. This was particularly the case in the turbid water off the Elbe estuary, where numbers up to 0.3 × 10⁶/l were counted.Gymnodinium sp. formed 96 to 99 % of the phytoplankton biomass during maximal development. The chlorophylla content of one millionGymnodinium was only 3.5µg. An extinktion of E=0.083/1 m was measured in a suspension of 10⁶ Gymnodinium/l in samples with minimal other plankton and detritus.4. The water masses containingGymnodinium blooms, which reached Helgoland after a change of wind direction, were characterized by lower salinity and higher temperature. This indicates that blooms developed in the coastally influenced water masses east of Helgoland. The red tides occurred during a period of minimal discharge of Elbe river water and of relatively high salinity of the coastal water in the Inner German Bight. They developed after a long period of calm winds. There were no records of reported fish or shellfish poisoning.

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Gymnodinium-Wucherungen in der Helgoländer Bucht (Nordsee) im August 1968

Kurzfassung Wie in anderen Teilen der Nordsee traten auch in der Helgoländer Bucht im August 1968 starke Wucherungen von Dinoflagellaten auf. Untersuchungen des Planktons und der Wassereigenschaften auf der Helgoland-Reede sowie auf weiteren Stationen in der Helgoländer Bucht wurden für die Erklärung dieser Erscheinung ausgewertet. In allen untersuchten Gebieten (Fig. 1) fand sichGymnodinium sp. als dominante Planktonform. Gymnodinien-Wucherungen traten auf der Helgoland-Reede in der Zeit vom 14.–30. August auf und erreichten Zellzahlen von über 3 × 10⁶/l am 28. und 30. August, bei nur 18–19µg Chlorophylla/l. Eine Million Gymnodinien produzierten 0,265 mg C in 6 Stunden im Inkubator. Etwa 3 × 10⁶ Gymnodinium sp. wurden in den obersten 16,5 m der Wassersäule in der Nähe Helgolands während des Blüte-Maximums gemessen (maximal 7,8 × 10⁶/l in 3 m). Die Gymnodinien waren tagsüber in den obersten Metern konzentriert. Eine solche Vertikalschichtung war besonders im Küstenwasser vor der Elbmündung ausgeprägt, wo bis zu 0,3 × 10⁶ Zellen/l gefunden wurden.Gymnodinium sp. bildete 96–99 % der Biomasse während der Blüte bei Helgoland. Es wurden ein Chlorophyll-a-Gehalt von nur 3,5µg für 10⁶ Zellen ermittelt und eine Extinktion von E=0,083/1 m für eine Suspension von 10⁶ Gymnodinien/l in Proben mit minimalem Gehalt an übrigem Plankton und Detritus gemessen. Der Nitrat- und Nitritgehalt im Wasser war zeitweise erschöpft, nicht aber das Phosphat. Wenn Wassermassen mitGymnodinium-Blüten nach Helgoland gedriftet wurden, fiel der Salzgehalt, und die Temperatur stieg an, was auf eine Entstehung der Wucherungen im brackwasser-beeinflußten Gebiet östlich Helgolands hindeutet. DieGymnodinium-Blüten entwickelten sich nach einer außergewöhnlich langen windarmen Periode und während eines minimalen Süßwasserzuflusses aus der Elbe. Vergiftungserscheinungen an Seetieren wurden bei Helgoland nicht festgestellt.

     

Systematics and Ecology of Protodrilus helgolandicus sp.n., an Interstitial Polychaete (Protodrilidae) from Subtidal Sands off Helgoland, German Bight

Protodrilus helgolandicus sp.n. is described and compared to similar species. Indirect sperm transmission by spermatophores and egg deposition are reported. Some data on life cycle, population dynamics and ecology are given.     

Five new species of the nanoflagellatePirsonia in the German Bight,North Sea,feeding on planktic diatoms

The structure, development, and host range of five newly detected species of the phagotrophic nanoflagellatePirsonia are described. They feed on planktic diatoms common in the North Sea:P. verrucosa sp. nov. (onRhizosolenia delicatula), P. formosa sp. nov. (onRhizosolenia setigera), P. diadema sp. nov. (onCoscinodiscus granii andC. wailesii), P. eucampiae (onEucampia zodiacus) andP. mucosa sp. nov. (onRhizosolenia shrubsolei). The occurrence of resting cysts inPirsonia guinardiae is reported. The impact ofPirsonia on phytoplankton communities is discussed.     

Seasonal variability and inter-specific differences in hatching of calanoid copepod resting eggs from sediments of the German Bight (North Sea)

Hatching experiments were conducted on sediment samples collectedon seven cruises between March 2002 and February 2003, at fourstations near Helgoland (54°11'N, 7°53'E). Samples wereincubated for up to 12 months and examined weekly for calanoidcopepod nauplii. 12807 nauplii hatched in total. Of these 36.43%were stage N1 (pooled), 44.08% were Temora longicornis (stageN2 and older), and 17.96% were Centropages hamatus (stage N2and older). Some Acartia spp. and an unidentified species werefound, too. Hatching varied significantly between sampling dates.At three stations, counts of all nauplii peaked in samples takenin April, while at one station the maximum was recorded in thesample collected in October. The seasonal pattern of hatchingwas similar in T. longicornis and C. hamatus. In March, April,October and February numbers of all nauplii were initially low,but increased over the following 2–3 weeks. In June, Augustand December, however, average numbers were comparatively highat first, but declined thereafter. In three of the four samplesthat were incubated for 12 months nauplii hatched from the sedimentfor the entire experimental period. Temora longicornis and C.hamatus displayed clearly distinct patterns of hatching in theselong-term incubations.     

Parasites of flounder Platichthys flesus (L.) from the German Bight, North Sea, and their potential use in ecosystem monitoring

The analysis of fish-parasite community structure and the use of ecological richness and diversity measurements are commonly used for the evaluation of environmental stress in aquatic ecosystems. As part of an integrated biological-effect monitoring, the parasite community of flounder Platichthys flesus (L.) was investigated for various locations in the German Bight during spring and autumn of 1995–2000, using established ecological methods. Although the parasite-community composition was very similar at the component-community level, the number of component species as well as the species accumulation curves showed clear differences among the sites. On the infra-community level, all of the ecological measurements showed significantly lower values in flounder from the Elbe estuary, the most polluted site, than in flounder from Helgoland. This was seen during a single season or during both seasons. When the data were pooled over the years, gradual differences between the sites, which were seldom detected at individual sampling periods, became evident for different measurements of species richness and species diversity and corresponded to a contamination gradient established between Elbe > Inner Eider, Outer Eider > Helgoland. Despite seasonal variations, which were observed in almost all measurements, these gradual differences were found in both seasons.