Biogenic and taphonomic processes affecting the development of shell assemblages: an actuopaleontological case study from mussel banks on North Sea tidal flats

Summary From spring 1997 to winter 2000, the composition and structure of epibenthic bivalve clusters and patches on a backbarrier tidal flat of the East Frisian Wadden Sea were continoously analyzed. The studied bivalve aggregates developed from intense and approximately simultaneous spatfalls of Mytilus edulis and Cerastoderma edule in 1996, after a severe winter (1995/96) with 64 days of driftice coverage. Spatfall occurred on a bare sandy tidal flat, whereas adjacent shell layers situated at the southern margin of the spatfall site were not colonized. Usually, such shell layers are the most common secondary hard substrates in the backbarrier area favorable for initial settlement of Mytilus larvae. The shell layers predominantly originate from erosion of former Mya arenaria and C. edule populations. So far, the studied bivalve aggregates are composed of approximately equal proportions of living specimens of both species, M. edulis and C. edule. The incorporated cockles were prevented from burrowing into the rediment by the coherent byssal mesh. In contrast to the results of previous studies, the great bulk of epibenthic living C. edule did survive for several years within the clusters. Therefore, the common terms ‘Mytilus banks’, ‘Mytilus patches’ or ‘Mytilus clumps’ may be too general and may erroneously imply a dominance of (or mono-specific composition by) M. edulis. In the fossil record, i.e. in diagenetically consolidated shell layers, the enforced epibenthic life-mode of C. edule and the remarkable species composition as well as the temporal development of the aggregates would not be recognizable.     

Land reclamation as a cause of fine-grained sediment depletion in backbarrier tidal flats (Southern North Sea)

Grain size distribution patterns along the East Frisian coast (German Wadden Sea) were evafuated with the aim of identifying potential effects of man-made structures (e.g. dikes) on the textural composition and sediment budget of the area. It was found that the distribution of various grain size fractions and settling velocities revealed a well-defined cross-shore energy gradient. Adjacent to the mainland dike the sediment is dominated by the 3.0–3.5 phi sand fraction (>70% by weight). In comparison to the sedimentary sequences normally observed in unobstructed backbarrier depositional environments, the sediments along the land-ward margin are in this case conspicuously depleted in grain sizes <3.5 phi, the corresponding size fractions contributing <20% by weight. This suggests that the dike interrupts the normal energy gradient, thereby inhibiting the deposition of sediments with settling velocities <0.5 cm s⁻¹. This interpretation is confirmed when comparing individual cross-shore transects. The shorter the transect between the barrier island shore and the dike, the greater the fine sediment depletion. Since many faunal assemblages are grain-size specific, it is further suggested that the backbarrier ecosystem must have undergone significant modifications as a result of the substantial reduction in mud flats since the onset of dike construction some 1000 years BP. It is predicted that in the wake of the persisting and possibly accelerating sealevel rise this process of fine sediment depletion will continue and also affect progressively coarser sediments.     

Calcium carbonate precipitation in cyanobacterial mats from sandy tidal flats of the North Sea

Precipitated calcium carbonate was found in annual cyanobacterial mats developing on the beaches of the North Sea barrier island Schiermonnikoog (the Netherlands). A variety of different calcium carbonate morphs were found in the cyanobacterial mucous secretions and identified by light- and scanning electron microscopy and X-ray diffraction. Most of the calcium carbonate seemed to be associated with degrading extracellular polymer. It is conceived that supersaturation of calcium carbonate resulted from the periodic evaporation of the mats and from the release of calcium from the cyanobacterial mucous as a result of its degradation. The analysis of the carbon stable isotopic composition of the calcium carbonate showed only a slight depletion of ¹³C, indicating that it did not in major part originated from the decomposition of organic matter. The δ¹⁸O values of the calcium carbonate confirmed the temperature differences between spring and summer but excluded the possibility that excessive evaporative events controlled precipitation. The precipitation of calcium carbonate could represent a potential factor enhancing the stabilization of intertidal siliciclastic sediments through cementing the sand. The discovery of massive calcium carbonate precipitation in these cyanobacterial mats was unexpected and hitherto unknown.     

The houting Coregonus oxyrinchus(L.)(Salmoniformes: Coregonidae), a globally extinct species from the North Sea basin

Coregonus oxyrinchus is redescribed and a neotype is designated (BMNH 1862.11.20.1.). It was restricted to south England and the lower parts of the Rivers Rhine, Meuse and Schelde. It is now globally extinct; the last individual was caught in 1940. Coregonus oxyrinchus is distinguished from other coregonids by having 38–46 gill rakers and a long, pointed snout.     

Community structure of the macroinfauna inhabiting tidal flats characterized by the presence of different species of burrowing bivalves in Southern Chile

Several species of bivalves coexist at the lower intertidal of large tidal flats located in the enclosed or inland coast of the northern area of the Nord-Patagonic archipelagos on the Chilean coast (ca. 40–42°S): Tagelus dombeii (Lamarck), Mulinia edulis (King & Broderip), Venus antiqua King & Broderip, Semele solida (Gray), Gari solida (Gray) and Diplodonta insconspicua Philippi. To explore possible spatial variation in the community structure of the macroinfauna inhabiting sediments with different assemblages of these bivalves, seasonal sampling was carried out during 2003–2004 at two tidal flats of that area. Higher species richness and specimen densities of the macroinfauna occurred in sediments with the higher densities of bivalves, especially in sediments where the deep burrower T. dombeii reaches its greatest abundances. Our results suggest that, apart from presence of bivalves, the burrowing depth of these organisms is also important in promoting the abundance of macroinfauna. Our results are in contrast with earlier conceptualizations for community organization of the soft bottom macroinfauna inhabiting intertidal flats, related to biological interactions occurring among different phyletic groups, such as that arguing that suspension feeding bivalves (such as T. dombeii and V. antiqua) will negatively affect the recruitment of species with planktonic larvae, by filtering them before they become established in the substrate. Thus, it is concluded that beneficial effects of bivalve bioturbation overcome that negative effects on the macroinfauna, although detrimental effects may well occur at bivalve densities higher than those studied here. Electronic supplementary material Supplementary material is available for this article at and accessible for authorised users     

Historic mussel beds (Mytilus edulis) in the sedimentary record of a back-barrier tidal flat near Spiekeroog Island, southern North Sea

Mussel beds show irregular cycles of appearance, disappearance and reappearance at preferred distinct locations on intertidal flats. These cycles are documented in the depth profile by the presence of shell-rich sediments intercalated with sandy layers. Such mussel bed layers were regularly found over the past 12 years on the Swinnplate, a back-barrier tidal flat south of Spiekeroog Island, southern North Sea. They are characterised by the occurrence of sub-articulated pairs of Mytilus shells. Based on life span considerations, a period of at least 35–40 years over which mussel beds may have repeatedly established themselves is suggested. A survey in spring 2000 revealed a reduced occurrence of old, embedded beds on the Swinnplate. The present results, based on core profiles from 1992 to 2000 and amino acid age determination, show that alternations of shell layers indicative of former Mytilus beds and sediment layers provide insight into the historic development of tidal flat environments. The frequent occurrence of sub-articulated valve pairs in the shell layers documents that the embedding of the mussel beds took place soon after the death of the organisms without prior physical disturbance. Successions of historic Mytilus beds provide evidence that mussel beds are a regular, but not necessarily permanent, faunal feature of back-barrier tidal flats of the North Sea. Electronic Publication     

Reinterpreting two regime shifts in North Sea plankton communities through the lens of functional traits

Aim

The so-called regime shifts in North Sea plankton communities provide an important historical case study to understand marine regime shifts. Previous studies characterized regime shifts using a variety of community metrics (e.g., indicator species abundances, taxonomic composition and chlorophyll biomass) but left the functional traits of plankton unassessed. Here, we explicitly re-assess the historically recognized North Sea regime shifts through the lens of plankton functional traits to gain a better understanding of these events.

Location

The North Sea (NW European shelf).

Time period

1958–2018, focusing on the 1980s and 1996–2003 regime shifts.

Major taxa studied

Marine phyto- and zooplankton.

Methods

We compute trait spaces for both phyto- and zooplankton of the North Sea using traits from the literature and a Gower's distance-based method. Using abundance data from the Continuous Plankton Recorder Survey, we then compute monthly time series of the centroids of the communities, an indicator of functional composition. We then use principal component analysis on the centroids to assess the main temporal changes in plankton functional composition associated with the 1980s and 1996–2003 regime shifts.

Results

Little change in plankton functional composition was associated with the 1980s regime shift. In contrast, the functional composition of plankton communities changed markedly after the 1996–2003 regime shift, with an increase in the summer relative abundance of non-motile autotrophs (i.e., diatoms) and the spring relative abundance of meroplankton.

Main conclusions

The North Sea regime shifts were not associated systematically with changes in functional composition, calling into question the definition of regime shifts and illustrating the importance of taking different metrics into account to interpret ecological events accurately. Taking into account functional composition, we interpret the 1980s so-called regime shift as a latitudinal shift in communities that was insufficient to impact functional composition and the 1996–2003 so-called regime shift as a period of change in bentho-pelagic coupling.     

Macroparasite community in molluscs of a tidal basin in the Wadden Sea

We provide a quantitative inventory of macroparasites in intertidal molluscs from a tidal basin in the Wadden Sea (eastern North Sea). Gastropods and bivalves contained a species rich macroparasite community consisting of trematodes (26 species), turbellarians (1), nematodes (1), copepods (2) and polychaetes (1) in 3,800 host individuals from 10 host species. Highest parasite burdens were observed in the gastropods Hydrobia ulvae and Littorina littorea and in the bivalves Cerastoderma edule and Mytilus edulis. In contrast, only one parasite species and no trematodes were found in Crepidula fornicata. The parasite community in the molluscs was similar to other Western European localities but some parasite species showed obvious differences, related to the large-scale distribution of intermediate and final hosts. Parasitism seems to be a common phenomenon in molluscs of the Wadden Sea and hence the detrimental effects observed in experiments can be expected to frequently happen in the field.     

Spatial variability of epifaunal communities in the North Sea in relation to sampling effort

Benthic epifauna was sampled in six areas from the German Bight towards the Norwegian Sea using a 2-m standard beam trawl. Nine replicates were taken in each area and year from 1999 to 2006. This data set (60–67 replicates per area) was used to describe the spatial variability in local species composition and to assess the effect of increasing sampling effort on species richness and community structure. Our results confirmed the importance of the 50-m depth contour for the separation of benthic fauna in the North Sea. Low species richness, sparse sessile fauna and high abundances of scavenging species such as Asterias rubens, Liocarcinus holsatus, Astropecten irregularis and Ophiura albida were characteristic of the low-water area south of the 50-m contour. Differences in community structure were less conspicuous in deeper waters north of the 50-m contour, but distribution patterns and abundances of single species such as Echinus elegans, Hyalinoecia tubicola, Ophiothrix fragilis, Scaphander lignarius as well as several hermit crabs resulted in well-defined epifaunal communities. One replicate caught 17–28% of the species found in 60–67 samples and was sufficient to separate the community in the German Bight from those in the central and northern North Sea by using multidimensional scaling. Nine replicates sampled a proportion of 53–60% and provided additional information on the spatial variability of community structure in the central and northern North Sea. Our study indicates that appropriate replication enhances the quality of the data and can partly overcome the constraints of sampling with a 2-m beam trawl. This might be helpful for future monitoring programmes.     

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.     

Free-living marine nematodes from the Southern Bight of the North Sea II. Notes on species of the Trefusiidae Gerlach, 1966

Three species of the Trefusiidae Gerlach, 1966 are described from the Southern Bight of the North Sea. Trefusia litoralis (Allgén, 1932) is redescribed and information is given on the intraspecific variation. Trefusia curvispiculosa sp.nov. is characterized by its typically curved spicular apparatus, up to now not found in other Trefusia species. Rhabdocoma riemanni Jayasree & Warwick, 1977 is synonymized with Rhabdocoma americana Cobb, 1920 and that species is redescribed.     

Climate change‐related regime shifts have altered spatial synchrony of plankton dynamics in the North Sea

During the 1980s, the North Sea plankton community underwent a well‐documented ecosystem regime shift, including both spatial changes (northward species range shifts) and temporal changes (increases in the total abundances of warmer water species). This regime shift has been attributed to climate change. Plankton provide a link between climate and higher trophic‐level organisms, which can forage on large spatial and temporal scales. It is therefore important to understand not only whether climate change affects purely spatial or temporal aspects of plankton dynamics, but also whether it affects spatiotemporal aspects such as metapopulation synchrony. If plankton synchrony is altered, higher trophic‐level feeding patterns may be modified. A second motivation for investigating changes in synchrony is that the possibility of such alterations has been examined for few organisms, in spite of the fact that synchrony is ubiquitous and of major importance in ecology. This study uses correlation coefficients and spectral analysis to investigate whether synchrony changed between the periods 1959–1980 and 1989–2010. Twenty‐three plankton taxa, sea surface temperature (SST), and wind speed were examined. Results revealed that synchrony in SST and plankton was altered. Changes were idiosyncratic, and were not explained by changes in abundance. Changes in the synchrony of Calanus helgolandicus and Para‐pseudocalanus spp appeared to be driven by changes in SST synchrony. This study is one of few to document alterations of synchrony and climate‐change impacts on synchrony. We discuss why climate‐change impacts on synchrony may well be more common and consequential than previously recognized.     

Marine molluscs and fish as biomarkers of pollution stress in littoral regions of the Red Sea, Mediterranean Sea and North Sea

The intensive development of industry and urban structures along the seashores of the world, as well as the immense increase in marine transportation and other activities, has resulted in the deposition of thousands of new chemicals and organic compounds, endangering the existence of organisms and ecosystems. The conventional single biomarker methods used in ecological assessment studies cannot provide an adequate base for environmental health assessment, management and sustainability planning. The present study uses a set of novel biochemical, physiological, cytogenetic and morphological methods to characterize the state of health of selected molluscs and fish along the shores of the German North Sea, as well as the Israeli Mediterranean and Red Sea. The methods include measurement of activity of multixenobiotic resistance-mediated transporter (MXRtr) and the system of active transport of organic anions (SATOA) as indicators of antixenobiotic defence; glutathione S-transferase (GST) activity as an indicator of biotransformation of xenobiotics; DNA unwinding as a marker of genotoxicity; micronucleus test for clastogenicity; levels of phagocytosis for immunotoxicity; cholinesterase (ChE) activity and level of catecholamines as indicators of neurotoxicity; permeability of external epithelia to anionic hydrophilic probe, intralysosomal accumulation of cationic amphiphilic probe and activity of non-specific esterases as indicators of cell/tissue viability. Complete histopathological examination was used for diagnostics of environmental pathology. The obtained data show that the activity of the defensive pumps, MXRtr and SATOA in the studied organisms was significantly higher in the surface epithelia of molluscs from a polluted site than that of the same species from control, unpolluted stations, providing clear evidence of response to stress. Enhanced frequency of DNA lesions (alkaline and acidic DNA unwinding) and micronucleus-containing cells was significantly higher in samples from polluted sites in comparison to those from the clean sites that exhibited genotoxic and clastogenic activity of the pollutants. In all the studied molluscs a negative correlation was found between the MXRtr levels of activity and the frequency of micronucleus-containing hemocytes. The expression of this was in accordance with the level of pollution. The complete histopathological examination demonstrates significantly higher frequencies of pathological alterations in organs of animals from polluted sites. A strong negative correlation was found between the frequency of these alterations and MXRtr activity in the same specimens. In addition to these parameters, a decrease in the viability was noted in molluscs from the polluted sites, but ChE activities remained similar at most sites. The methods applied in our study unmasked numerous early cryptic responses and negative alterations of health in populations of marine biota sampled from the polluted sites. This demonstrates that genotoxic, clastogenic and pathogenic xenobiotics are present and act in the studied sites and this knowledge can provide a reliable base for consideration for sustainable development. Received: 2 March 1999 / Received in revised form: 2 August 1999 / Accepted: 3 August 1999     

Copepod feeding behaviour and egg production during a dinoflagellate bloom in the North Sea

Feeding strategies of copepods were studied during a dinoflagellate-dominated bloom in the North Sea in August 2001. The aim of this study was to evaluate the importance of mesozooplankton grazing as a biological loss factor of harmful algal blooms under natural conditions. Therefore, ingestion, egestion and egg production experiments were performed with the most abundant copepod species Calanus helgolandicus, Temora longicornis and Acartia sp. feeding on the natural phytoplankton community. Dinophysis norvegica and Ceratium furca were the most abundant dinoflagellate species at the time of the experiments. Grazing experiments as well as examination of fecal pellet content revealed C. helgolandicus fed efficiently on D. norvegica. Ingestion rates up to 47 cells female⁻¹ h⁻¹ were measured and a large proportion of the C. helgolandicus fecal pellets contained intact D. norvegica cells. Dinophysis cells were rarely seen in fecal pellets produced by T. longicornis, and never observed in pellets produced by Acartia sp. The ingestion rate of C. furca, which was the dominating Ceratium species, mimicked that of D. norvegica. C. helgolandicus grazed significantly on C. furca (16 cells female⁻¹ h⁻¹), while the ingestion rate of T. longicornis was low and Acartia sp. was not able to graze on C. furca. Egg production experiments revealed that 92% of the C. helgolandicus females produced eggs. The specific egg production rate and the proportion of females producing eggs among T. longicornis were low. This field experiment clearly shows that some copepod species feed efficiently on D. norvegica and C. furca under natural conditions, which may affect the bloom development of these dinoflagellates.     

Occurrence of pelagic stingray Pteroplatytrygon violacea (Bonaparte, 1832) in the North Sea

A mature male pelagic stingray Pteroplatytrygon violacea (99 cm total length, 33·5 cm disc length and 42 cm disc width; 2·5 kg) was caught by commercial Nephrops trawl in the North Sea (at c . 54º49' N; 0º57' W) on 14 November 2005. Records of P. violacea in the north-east Atlantic are reviewed, with only two vagrant specimens known from the North Sea, representing the most northerly records of this species in the Atlantic Ocean.     

Seasonal variability of inorganic and organic nitrogen in the North Sea

This study considers the cycling of nitrogen in the waters of the North Sea, particularly focussing on organic nitrogen. Dissolved inorganic nitrogen (DIN), dissolved organic nitrogen (DON) and particulate organic nitrogen (PON) were measured in the North Sea over a one-year period (autumn 2004–summer 2005). The surface water concentrations of nitrate, ammonium, DON and PON during the present study ranged from <0.1–7.2 μM, <0.1–2.0 μM, 1.9–11.2 μM and 0.3–5.6 μM, respectively, with DON the dominant fraction of total nitrogen at all times. These nutrients concentrations were significantly lower compared to previous studies in the southern North Sea. The seasonal variations showed high mean surface concentrations of nitrate (4.7 ± 0.6 μM) and DON (8.9 ± 0.9 μM), low ammonium (<0.1 μM) and PON (0.8 ± 0.1 μM) in winter, shifting to low nitrate (0.3 ± 0.3 μM) and DON (4.2 ± 1.2 μM) in summer, with high ammonium (0.8 ± 0.4 μM) in autumn and PON (2.5 ± 1.2 μM) in spring. Highest mean surface DON concentration was measured in winter and may be due to resuspension of the organic matter from the bottom sediments. For autumn and spring, phytoplankton DON release was likely to be the most significant source of DON as shown by high concentrations of low molecular weight (LMW) DON and its positive correlation to chlorophyll a. Low total and LMW DON concentrations during summer were likely to be due to the uptake of the LMW DON fraction by phytoplankton and bacteria and the stratification of the water column. DON is therefore shown to be a potentially important source of nitrogen in shelf seas especially after the spring bloom has depleted nitrate to limiting concentrations. Handling editor: L. Naselli-Flores     

Long-term changes of intertidal and subtidal sediment compositions in a tidal basin in the northern Wadden Sea (SE North Sea)

Wadden Sea tidal flats are highly dynamic regarding the spatial distribution and the grain size composition of their sediments. From 2003 to 2006 surface sediments have been surveyed in an intertidal and a subtidal area within the tidal inlet Königshafen (south-eastern North Sea, northern Wadden Sea, island of Sylt) with the goal to gain information on short-term development trends in the grain size composition. The average grain size (Mean) becomes finer in the sheltered part of the intertidal survey area whereas a coarsening tendency can be observed in the more exposed part of the intertidal and especially in the subtidal survey area. The trend of the most frequent grain size (first Mode) shows the same spatial distribution pattern but is far less distinct. Thus, the changing Mean must be related to an increase in the deposition of fines in the sheltered part of the intertidal Königshafen as well as a general removal of fine-grained material in the exposed intertidal and subtidal Königshafen. In order to see long-term trends the surveys of 2003–2006 were compared to earlier studies conducted in 1932/1933, 1981 and 1989. A significant depletion of mud can be observed in the entire survey area. It is concluded that primarily changed hydrodynamics that may accompany ongoing climate change are responsible for this. However, the loss of fine-grained sediments is additionally amplified by a reduced vegetation cover and coastal protection measures.     

Community structure and functional roles of meiofauna in the North Sea

Knowiedge on community structure of North Sea meiofauna has greatly increased recently. A quasisynoptic picture of meiofauna densities and copepod community structure from 171 stations of the southern North Sea, sampled in April–May 1986, has been obtained during the North Sea Benthos Survey. Latitudinal patterns in meiofauna abundance and copepod weight, abundance and diversity exist in an area between 51°30′N and 58°30′N. Using TWINSPAN-classification five major groups of copepod species can be recognized which are related to sediment type, latitude and depth. The part of the meiofauna in total benthic energy flow, their role in the benthic food web and in biogeochemical cycles is discussed based on existing literature. There are still considerable gaps in knowledge and the field is not progressing rapidly. Publication no. 599 Netherlands Institute of Ecology, Centre for Estuarine and Coastal Ecology, Yerseke, The Netherlands.     

Candidatus Abditibacter,a novel genus within the Cryomorphaceae,thriving in the North Sea

Coastal phytoplankton blooms are frequently followed by successive blooms of heterotrophic bacterial clades. The class Flavobacteriia within the Bacteroidetes has been shown to play an important role in the degradation of high molecular weight substrates that become available in the later stages of such blooms. One of the flavobacterial clades repeatedly observed over the course of several years during phytoplankton blooms off the coast of Helgoland, North Sea, is Vis6. This genus-level clade belongs to the family Cryomorphaceae and has been resistant to cultivation to date. Based on metagenome assembled genomes, comparative 16S rRNA gene sequence analyses and fluorescence in situ hybridization, we here propose a novel candidate genus Abditibacter, comprising three novel species Candidatus Abditibacter vernus, Candidatus Abditibacter forsetii and Candidatus Abditibacter autumni. While the small genomes of the three novel photoheterotrophic species encode highly similar gene repertoires, including genes for degradation of proteins and algal storage polysaccharides such as laminarin, two of them – Ca. A. vernus and Ca. A. forsetii – seem to have a preference for spring blooms, while Ca. A. autumni almost exclusively occurs in late summer and autumn.