Justus van Beusekom

Diversity and dynamics of rare and of resident bacterial populations in coastal sands

Coastal sands filter and accumulate organic and inorganic materials from the terrestrial and marine environment, and thus provide a high diversity of microbial niches. Sands of temperate climate zones represent a temporally and spatially highly dynamic marine environment characterized by strong physical mixing and seasonal variation. Yet little is known about the temporal fluctuations of resident and rare members of bacterial communities in this environment. By combining community fingerprinting via pyrosequencing of ribosomal genes with the characterization of multiple environmental parameters, we disentangled the effects of seasonality, environmental heterogeneity, sediment depth and biogeochemical gradients on the fluctuations of bacterial communities of marine sands. Surprisingly, only 3–5% of all bacterial types of a given depth zone were present at all times, but 50–80% of them belonged to the most abundant types in the data set. About 60–70% of the bacterial types consisted of tag sequences occurring only once over a period of 1 year. Most members of the rare biosphere did not become abundant at any time or at any sediment depth, but varied significantly with environmental parameters associated with nutritional stress. Despite the large proportion and turnover of rare organisms, the overall community patterns were driven by deterministic relationships associated with seasonal fluctuations in key biogeochemical parameters related to primary productivity. The maintenance of major biogeochemical functions throughout the observation period suggests that the small proportion of resident bacterial types in sands perform the key biogeochemical processes, with minimal effects from the rare fraction of the communities.

Distinct flavobacterial communities in contrasting water masses of the North Atlantic Ocean

Members of the class Flavobacteria in the phylum Bacteroidetes are among the most abundant picoplankton in coastal and polar oceans. Their diversity is high in marine waters. However, quantitative information about distribution patterns of flavobacterial clades is scarce. We analyzed the diversity and clade-specific abundances of individual Flavobacteria in different oceanic provinces in the North Atlantic Ocean. Samples were taken along the 30°W meridian between the East Greenland current and the North Atlantic subtropical gyre. Comparative sequence analysis of 16S ribosomal RNA (rRNA) gene libraries revealed high diversity and significant spatial variability within the class Flavobacteria. Published and newly designed oligonucleotide probes were used to enumerate eleven flavobacterial clades by catalyzed reporter deposition fluorescence in situ hybridization (CARD-FISH). We found that different provinces harbor distinct flavobacterial communities. Clade DE2 accounted for a substantial fraction of total Flavobacteria only in the Polar Biome (BPLR), whereas the VISION clades VIS1 and VIS4 significantly increased in the Arctic (ARCT) province. Members of the genus Polaribacter were the most abundant clade in all the water masses analyzed, with highest absolute numbers in BPLR and ARCT. We improved the CARD-FISH protocol to quantify the rare clades VIS2, VIS3, VIS5 and VIS6, which were present in abundances below 0.5%. They all showed pronounced regional distribution patterns. Microscopic analysis proved a specific enrichment of Flavobacteria in the phycosphere of nanophytoplankton of BPLR and ARCT. Our results suggest that different marine flavobacterial clades have distinct niches and different life strategies.

Time- and sediment depth-related variations in bacterial diversity and community structure in subtidal sands.

Bacterial community structure and microbial activity were determined together with a large number of contextual environmental parameters over 2 years in subtidal sands of the German Wadden Sea in order to identify the main factors shaping microbial community structure and activity in this habitat. Seasonal changes in temperature were directly reflected in bacterial activities and total community respiration, but could not explain variations in the community structure. Strong sediment depth-related patterns were observed for bacterial abundances, carbon production rates and extracellular enzymatic activities. Bacterial community structure also showed a clear vertical variation with higher operational taxonomic unit (OTU) numbers at 10–15 cm depth than in the top 10 cm, probably because of the decreasing disturbance by hydrodynamic forces with sediment depth. The depth-related variations in bacterial community structure could be attributed to vertical changes in bacterial abundances, chlorophyll a and NO3−, indicating that spatial patterns of microbes are partially environmentally controlled. Time was the most important single factor affecting microbial community structure with an OTU replacement of up to 47% over 2 years and a contribution of 34% to the total variation. A large part of this variation was not related to any environmental parameters, suggesting that temporal variations in bacterial community structure are caused by yet unknown environmental drivers and/or by stochastic events in coastal sand habitats. Principal ecosystem functions such as benthic oxygen consumption and extracellular hydrolysis of organic matter were, however, at a high level at all times, indicating functional redundancy in the microbial communities.

Does climatic warming explain why an introduced barnacle finally takes over after a lag of more than 50 years

Invading alien species may have to await appropriate conditions before developing from a rare addition to the recipient community to a dominance over native species. Such a retarded invasion seems to have happened with the antipodean cirripede crustacean Austrominiusmodestus Darwin, formerly known as Elminius modestus, at its northern range in Europe due to climatic change. This barnacle was introduced to southern Britain almost seven decades ago, and from there spread north and south. At the island of Sylt in the North Sea, the first A. modestus were observed already in 1955 but this alien remained rare until recently, when in summer of 2007 it had overtaken the native barnacles Semibalanus balanoides and Balanus crenatus in abundance. At the sedimentary shores of Sylt, mollusc shells provide the main substrate for barnacles and highest abundances were attained on mixed oyster and mussel beds just above low tide level. A. modestus ranged from the upper intertidal down to the subtidal fringe. Its realized spatial niche was wider than that of the two natives. We suggest that at its current northern range in Europe a long series of mild winters and several warm summers in a row has led to an exponential population growth in A. modestus.

Interactive effects of global and regional change on a coastal ecosystem

Shallow waters and lowland meet at the same level in the Wadden Sea, but are separated by walls of coastal defense. What are the prospects of this coastal ecosystem in a warmer world? We focus on tidal waters and inshore sedimentary bottoms, expect nutrient supply from land to decline and species introductions, temperature and sea level to rise. The effects are interrelated and will have an increasing likelihood of abrupt and irreversible developments. The biotic interactions are hardly predictable but we anticipate the following changes to be more likely than others: blooms of phytoplankton will be weak mainly because of increasing pelagic and benthic grazing pressure, both facilitated by warming. Possibly birds feeding on mollusks will encounter decreasing resource availability while fish-eaters benefit. Extensive reefs of Pacific oysters could facilitate aquatic macrophytes. Sea level rise and concomitant hydrodynamics above tidal flats favor well-anchored suspension feeders as well as burrowing fauna adapted to dynamic permeable sand. With high shares of immigrants from overseas and the south, species richness will increase; yet the ecosystem stability may become lower. We suggest that for the next decades invasions of introduced species followed by warming and declining nutrient supply will be the most pressing factor on the changes in the Wadden Sea ecosystem, and the effects of sea level rise to be the key issue on the scale of the whole century and beyond.

Thalassiosira species (Bacillariophyceae, Thalassiosirales) in the North Sea at Helgoland (German Bight) and Sylt (North Frisian Wadden Sea) – a first approach to assessing diversity

Thalassiosira species are an important component of phytoplankton populations in pelagic waters around the islands of Helgoland and Sylt in the German North Sea. This taxonomic study revealed the presence of at least 27 species, 21 at Helgoland and 21 at Sylt. Fifteen species had not been recorded previously for Helgoland (T. aestivalis, T. angulata, T. concaviuscula, T. curviseriata, T. delicatula, T. diporocyclus, T. hendeyi, T. kuschirensis, T. minima, T. minuscula, T. oceanica, T. pacifica, T. proschkinae, T. tealata, T. tenera) and 16 for Sylt (T. aestivalis, T. angulata, T. concaviuscula, T. constricta, T. curviseriata, T. delicatula, T. guillardii, T. hendeyi, T. lundiana, T. mediterranea, T. minima, T. pacifica, T. partheneia, T. punctigera, T. tealata, T. tenera). We provide brief descriptions with illustrations and document the seasonal occurrence of these species at the two stations. We also discuss investigation methods used for different levels of species recognition, namely Utermöhl counts, light microscopical observations of living net-samples as well as scanning electron microscopy of net-samples and isolated cultures. Clonal cultures of some species were established to facilitate the taxonomic investigations and 18S ribosomal DNA sequencing was carried out to complement the morphological studies. A perspective for a routine species-level assessment strategy is given.

Consensus forecasting of intertidal seagrass habitat in the Wadden Sea

1. After the dramatic eutrophication-induced decline of intertidal seagrasses in the 1970s, theWadden Sea has shown diverging developments. In the northern Wadden Sea, seagrass bedshave expanded and become denser, while in the southern Wadden Sea, only small beds withlow shoot densities are found. A lack of documentation of historical distributions hampersconservation management. Yet, the recovery in the northern Wadden Sea provides opportunityto construct robust habitat suitability models to support management.2. We tuned habitat distribution models based on 17 years of seagrass surveys in the northernWadden Sea and high-resolution hydrodynamics and geomorphology for the entire WaddenSea using five machine learning approaches. To obtain geographically transferablemodels, hyperparameters were tuned on the basis of prediction accuracy assessed by non-random,spatial cross-validation. The spatial cross-validation methodology was combined with aconsensus modelling approach.3. The predicted suitability scores correlated amongst each other and with the hold-out observationsin the training area indicating that the models converged and were transferable acrossspace. Prediction accuracy was improved by averaging the predictions of the best models.4. We graphically examined the relationship between the consensus suitability score andindependent presence-only data from outside the training area using the area-adjusted seagrassfrequency per suitability class (continuous Boyce index). The Boyce index was positivelycorrelated with the suitability score indicating the adequacy of the prediction methodology.5. We used the plot of the continuous Boyce index against habitat suitability score to demarcatethree habitat classes – unsuitable, marginal and suitable – for the entire international WaddenSea. This information is valuable for habitat conservation and restoration management.6. Divergence between predicted suitability and actual distributions from the recent past indicatesthat unaccounted factors limit seagrass development in the southern Wadden Sea.7. Synthesis and applications. Our methodology and data enabled us to produce a robust andvalidated consensus habitat suitability model. We identified highly suitable areas where intertidalseagrass meadows may establish and persist. Our work provides scientific underpinningfor effective conservation planning in a dynamic landscape and sets monitoring priorities.

Abrupt emergence of a large pockmark field in the German Bight, southeastern North Sea

A series of multibeam bathymetry surveys revealed the emergence of a large pockmark field in the southeastern North Sea. Covering an area of around 915 km2, up to 1,200 pockmarks per square kilometer have been identified. The time of emergence can be confined to 3 months in autumn 2015, suggesting a very dynamic genesis. The gas source and the trigger for the simultaneous outbreak remain speculative. Subseafloor structures and high methane concentrations of up to 30 μmol/l in sediment pore water samples suggest a source of shallow biogenic methane from the decomposition of postglacial deposits in a paleo river valley. Storm waves are suggested as the final trigger for the eruption of the gas. Due to the shallow water depths and energetic conditions at the presumed time of eruption, a large fraction of the released gas must have been emitted to the atmosphere. Conservative estimates amount to 5 kt of methane, equivalent to 67% of the annual release from the entire North Sea. These observations most probably describe a reoccurring phenomenon in shallow shelf seas, which may have been overlooked before because of the transient nature of shallow water bedforms and technology limitations of high resolution bathymetric mapping.

Long-Term Ecological Change in the Northern Wadden Sea

The Wadden Sea is a shallow coastal region in the south eastern North Sea. Karl Mobius started ecological research in the northern Wadden Sea about 150 years ago studying the extensive oyster beds. With the foundation of a field station of the Biologische Anstalt Helgoland in List/Sylt in 1924 biological research in the Wadden Sea was continued to date. Several time series were initiated between the 1970s and the 1990s including a bi-weekly phytoplankton and zooplankton program and an observation program on macrobenthos. Three factors dominating the changes observed during the past decades are a rise in temperature, decreasing nutrients, and increasing invasions of non-native species. Phytoplankton blooms gradually decrease due to the combined effect of decreasing nutrient loads and increasing winter temperatures. Mean annual zooplankton abundance is stimulated by higher winter temperatures. Recently, invading species are increasingly dominating native mussel beds. For several invaders, a positive effect of temperature was shown. We expect that major pressures of change during the next years will be further species introductions, temperature increase, and reduced nutrient loads. On the long run (21st century), we expect sea level rise to be the key factor of coastal change through a loss of habitats with fine-grained sediments and intertidal sediments in general. A major challenge for coastal research will be to disentangle the interactive effects of these pressures on the long-term development of the Wadden Sea.

Challenges of achieving Good Environmental Status in the Northeast Atlantic

The sustainable exploitation of marine ecosystem services is dependent on achieving and maintaining an adequate ecosystem state to prevent undue deterioration. Within the European Union, the Marine Strategy Framework Directive (MSFD) requires member states to achieve Good Environmental Status (GEnS), specified in terms of 11 descriptors. We analyzed the complexity of social-ecological factors to identify common critical issues that are likely to influence the achievement of GEnS in the Northeast Atlantic (NEA) more broadly, using three case studies. A conceptual model developed using a soft systems approach highlights the complexity of social and ecological phenomena that influence, and are likely to continue to influence, the state of ecosystems in the NEA. The development of the conceptual model raised four issues that complicate the implementation of the MSFD, the majority of which arose in the Pressures and State sections of the model: variability in the system, cumulative effects, ecosystem resilience, and conflicting policy targets. The achievement of GEnS targets for the marine environment requires the recognition and negotiation of trade-offs across a broad policy landscape involving a wide variety of stakeholders in the public and private sectors. Furthermore, potential cumulative effects may introduce uncertainty, particularly in selecting appropriate management measures. There also are endogenous pressures that society cannot control. This uncertainty is even more obvious when variability within the system, e.g., climate change, is accounted for. Also, questions related to the resilience of the affected ecosystem to specific pressures must be raised, despite a lack of current knowledge. Achieving good management and reaching GEnS require multidisciplinary assessments. The soft systems approach provides one mechanism for bringing multidisciplinary information together to look at the problems in a different light.