Pseudo-cryptic species Arenicola defodiens and Arenicola marina (Polychaeta: Arenicolidae) in Wadden Sea,North Sea and Skagerrak: Morphological and molecular variation

The polychaete Arenicola defodiens, or black lug, was recently described as a morphologically highly similar species alongside the blow lug Arenicola marina. So far it was only known from the British Isles. A double spawning peak was observed earlier in lugworms of the western Wadden Sea. Here, we test the hypothesis that the two spawning peaks represent the two species in sympatry. This hypothesis is refuted on the basis of both morphological and mitochondrial DNA data; both spawning peaks are attributed to A. marina. In spite of this, we confirm, on the basis of new collections as well as re-examination of museum collections, the presence of A. defodiens in the western Wadden Sea, North Sea and also the Skagerrak; its distribution is restricted to habitats which are either submerged or have short emersion times, have relatively coarse sediments and high and stable salinities. Sympatry is common. The species differ strongly in the mitochondrial DNA fragment examined; the observed 14% uncorrected minimum difference amounts to an estimated 33–63 million years since sequence divergence. The amount of intraspecific molecular variation is larger for A. defodiens than for A. marina. This is evidence to suggest that A. marina may have undergone more recent and/or more severe population size bottlenecks.     

Population dynamics of phytoplankton, heterotrophic bacteria, and viruses during the spring bloom in the western subarctic Pacific

We characterized the community composition of phytoplankton in the western subarctic Pacific from the pre-bloom to the decline phase of the spring bloom with special reference to decreases in the silicic acid concentration in surface waters as an index for diatom bloom development. Furthermore, responses of heterotrophic bacteria and viruses to the spring bloom were also concomitantly investigated. Under pre-bloom conditions when nutrients were abundant but the surface mixed layer depth was relatively deep, chlorophyll (Chl) a concentrations were consistently low and green algae (chlorophytes and prasinophytes), cryptophytes, and diatoms were predominant in the phytoplankton assemblages as estimated by algal pigment signatures. Together with the shallowing of the mixed layer depth and the decrease in silicic acid concentration, diatoms bloomed remarkably in the Oyashio region, though the magnitude of the bloom in the Kuroshio-Oyashio transition (hereafter Transition) region was relatively small. A total of 77 diatom species were identified, with the bloom-forming diatoms mainly consisting of Thalassiosira, Chaetoceros, and Fragilariopsis species. It has become evident that the carotenoid fucoxanthin can serve as a strong indicator of the diatom carbon biomass during the spring diatom bloom. Differences in the species richness of diatoms among stations generally enabled us to separate the Oyashio bloom stations from the Transition and the Oyashio pre-bloom stations. Relatively high values of the Shannon-Wiener index for the diatom species were also maintained during the Oyashio bloom, indicating that a wide variety of species then shared dominance. In the decline phase of the Oyashio bloom when surface nutrient concentrations decreased, senescent diatom cells increased, as inferred from the levels of chlorophyllide a. Although the cell density of heterotrophic bacteria changed little with the development of the diatom bloom, viral abundance increased toward the end of the bloom, suggesting an increased likelihood of mortality among organisms including diatoms resulting from viral infection. This is the first report on the microbial trophodynamics, including viruses, during the spring diatom bloom in the western subarctic Pacific.     

Long-term changes in the seasonality of selected diatoms related to grazers and environmental conditions

The decoupling of trophic interactions could be one of the severe consequences of climate warming in aquatic systems. The timing of phytoplankton blooms, in particular, can affect competition within the plankton community as well as food-web interactions with zooplankton and fish. Using long-term data from Helgoland Roads in the southern North Sea, we examine diatom seasonality, using three representative diatom species combined with environmental and copepod time series over the last four decades. The long-term annual abundances of Guinardia delicatula, Thalassionema nitzschioides and Odontella aurita exhibited interannual variations and dissimilar cyclic patterns during the time period under study (1962-2008). Of the three diatoms, G. delicatula showed a significant trend towards earlier bloom timings for 1962-2008 and a later decline of its abundance over time was found. Grazing and water transparency explained most of the bloom timing fluctuations of the diatoms considered. The annual timing of occurrences of each diatom species was correlated with their preceding concentrations. Earlier bloom timings occurred when autumn/winter concentrations were higher than average and later bloom timings occurred when autumn/winter concentrations were lower than average. Different environmental and predation variables related to the diatom bloom timings were found suggesting that climate warming might not affect the onset of the blooms of the three diatom species in the same manner. The results of the multiple linear regression analyses showed that the timings of decline of the three diatoms were mainly correlated with decreasing nutrient concentrations. Sunshine duration could prolong the duration of the blooms of T. nitzschioides and O. aurita provided that enough nutrients were available. In the case of G. delicatula, however, sunshine duration was negatively correlated with its end of the growth period. G. delicatula and T. nitzschioides showed later decreases in abundances under warmer spring and summer temperatures. Such species specific differences in the sensitivity to the forcing variables could lead to shifts in community structure and could ultimately have wider implications to the overall ecosystem health of the North Sea.     

白令海特征区域的表层沉积硅藻分布及其古海洋学意义

对白令海表层沉积物样品进行硅藻分析,共鉴定硅藻30属56种(含变种),并确定17种硅藻新记录,其中包括Kisseieviella carina等3种化石种。白令海表层沉积物中的硅藻优势种为Neodenticula seminae,Fragilariopsis cylindrus和Fragilariopsis oceanica,在白令海北部陆坡深水区附近以Neodenticula seminae为主,而在白令海北部陆架以Fragilariopsis cylindrus和Fragilariopsis oceanica为主。对硅藻结果进行聚类分析,可以划分出3个硅藻组合,硅藻组合Ⅰ代表海冰种硅藻组合,组合Ⅱ代表受阿拉斯加流影响的大洋浮游硅藻组合,组合Ⅲ代表上述两个硅藻组合之间的过渡组合。Fossula arctica是17种硅藻新记录之一,首次记录于白令海表层沉积物中,其百分含量分布趋势与Fragilariopsis cylindrus和Fragilariopsis oceanica的相近,在白令海北部陆架为11.7%~17.1%,而在陆坡深水区附近明显减少,是继Fragilariopsis cylindrus和Fragilariopsis oceanica之后白令海又一海冰指示种,并有望成为一种有效的海冰变化替代物运用于晚第四纪以来白令海海冰进退历史研究。     

Dynamics of Phaeocystis pouchetii blooms in the Wadden Sea of Sylt (German Bight,North Sea)

The colony-forming alga Phaeocystis pouchetii (Haptophyceae) forms regular blooms in spring and early summer in the German Wadden Sea. This paper gives a summary of observations of Phaeocystis blooms in 1975, 1976, and 1981 in the Wadden Sea of Sylt (German Bight, North Sea).The onset of Phaeocystis pouchetii blooms started after the decline of diatoms at low inorganic nutrient concentrations, when particularly inorganic phosphate had dropped to minimal concentrations. Phaeocystis blooms did not start unless water temperatures were above 7°C in the investigation area, and the admixture of organic substances seemed to be an important factor in controlling the onset and the size of Phaeocystis blooms.As diatom populations increased again at the end of the Phaeocystis blooms, a causal relationship between the successive development of both algal groups, possibly in combination with other abiotic factors, cannot be excluded.     

大亚湾拟菱形藻水华及其在生物群落中的生态地位

对2000年4~5月藻类水华高发期大亚湾澳头海域的拟菱形藻进行逐日调查,并对其与浮游植物、浮游动物的相关关系进行分析.调查初期,整个生物群落的种类组成数量都相对贫乏;拟菱形藻数量急剧增长,最高的测站密度可达2.93×106个/dm3.拟菱形藻水华的组成种类有拟极细拟菱形藻和尖刺拟菱形藻两种,以尖刺拟菱形藻稍占优势.水华形成中其他种类硅藻和甲藻在种类组成和数量上也有一定程度的增加.分析表明,拟菱形藻在浮游植物群落的种间竞争中具有优势,并成为春季硅藻水华的最重要的组成者.桡足类等浮游动物对拟菱形藻的下行控制并不能明显抑制拟菱形藻的增长及水华的产生,这与其偏爱以其他种类藻类为食有关.以上因素外加拟菱形藻对环境的广泛适应性促使其在该水域成为最常见的赤潮生物之一.     

Environmental controls on phytoplankton community composition in the Thames plume,U.K.

The aim of this study was to investigate controls on the phytoplankton community composition and biogeochemistry of the estuarine plume zone of the River Thames, U.K. using an instrumented moored buoy for in situ measurements and preserved sample collection, and laboratory-based measurements from samples collected at the same site. Instrumentation on the moored buoy enabled high frequency measurements of a suite of environmental variables including in situ chlorophyll, water-column integrated irradiance, macronutrients throughout an annual cycle for 2001 e.g. nitrate and silicate, and phytoplankton biomass and species composition. The Thames plume region acts as a conduit for fluvial nutrients into the wider southern North Sea with typical winter concentrations of 45 μM nitrate, 17 μM silicate and 2 μM phosphate measured. The spring bloom resulted from water-column integrated irradiance increasing above 60 W h m^− 2 d^− 1 and was initially dominated by a diatom bloom mainly composed of Nitzschia sp. and Odontella sinesis. The spring bloom then switched after ∼ 30 days to become dominated by the flagellate Phaeocystis reaching a maximum chlorophyll concentration of 37.8 μg L^− 1. During the spring bloom there were high numbers of the heterotrophic dinoflagellates Gyrodinium spirale and Katodinium glaucum that potentially grazed the phytoplankton bloom. This diatom–flagellate switch was predicted to be due to a combination of further increasing water-column integrated irradiance > 100 W h m^− 2 d^− 1 and/or silicate reaching potentially limiting concentrations (< 1 μM). Post spring bloom, diatom dominance of the lower continuous summer phytoplankton biomass occurred despite the low silicate concentrations (Av. 0.7 μM from June–August). Summer diatom dominance, generally due to Guinardia delicatula, was expected to be as a result of microzooplankton grazing, dominated by the heterotrophic dinoflagellate Noctiluca scintillans, controlling 0.7–5.0 μm ‘flagellate’ fraction of the phytoplankton community with grazing rates up to 178% of ‘flagellate’ growth rate. The Thames plume region was therefore shown to be an active region of nutrient and phytoplankton processing and transport to the southern North Sea. The use of a combination of moorings and ship-based sampling was essential in understanding the factors influencing nutrient transport, phytoplankton biomass and species composition in this shelf sea plume region.     

Distinctly variable mudscapes: Distribution gradients of intertidal macrofauna across the Dutch Wadden Sea

The Wadden Sea is a shallow coastal region, with a large area of sedimentary tidal flats that extends from The Netherlands to Denmark and has been declared a site of international importance in the Dutch and German parts (Ramsar status and UNESCO World Heritage Site). Benthic macrofauna are central to the ecosystem functioning of this area, as they recycle nutrients, decompose organic matter and are an important food source for many secondary consumers, like fish and waterbirds. Due to the environmental gradients characteristic of estuarine systems, it is expected that changes in assemblage composition will be observed across the physical and environmental gradients of the Wadden Sea. First, we explored the spatial variation in assemblage composition of benthic macrofauna across the intertidal part of the Dutch Wadden Sea using 3 years of biomass data. Then, we identified the relative importance of six environmental variables for explaining and predicting changes in assemblage composition across the intertidal areas of the Wadden Sea using generalised dissimilarity modelling (GDM). In accordance with the environmental gradients across this system, the biomass contributed by a few common species differed from west to east and were distinct in the Dollard. In the west, bivalves Mya arenaria, Cerastoderma edule and Ensis directus contributed a relatively large and equal share of the total biomass, whereas C. edule contributed the sole largest share of the total biomass towards the east. The polychaete Alitta succinea became a large share of the total biomass in the upper Ems and in the Dollard estuary, but contributed little elsewhere. Similar to the observed differences in species composition, the spatial patterns in assemblage composition, as predicted by the GDM models, identified the Dollard as distinct and that the prevalence of assemblage types in the west differed to the east. Median grain size, followed by microphytobenthic biomass, and exposure time were the most important variables describing differences in assemblage composition. That the Wadden Sea forms a heterogeneous landscape where assemblage composition varies across multiple gradients has repercussions for management and monitoring.     

Observations of a mass occurrence of Macoma balthica larvae in midsummer

In 1995 the seasonal development of concentrations of both phytoplankton and larvae of the bivalve Macoma balthica was studied in the coastal zone behind the back-barrier island of Spiekeroog (German Wadden Sea). In July/August larvae reached maximum concentrations of about 1000 to 4200 ind. m⁻³ (depending on the sampling site and tidal period), probably in relation to a phytoplankton bloom in July. This observation of an unusually late maximum of Macoma larvae is discussed in detail in connection with the recent literature available about spawning of M. balthica in the Wadden Sea.     

Overlapping distributions of cryptic Scoloplos cf. armiger species in the western Wadden Sea

Unknowingly lumping cryptic species in biological monitoring studies hinders progress in understanding their functioning in a wide range of research fields including population dynamics, ecophysiology and community ecology. The common polychaete worm Scoloplos cf. armiger is a cryptic species complex comprising entirely different developmental modes: holobenthic and pelago-benthic development. In the northeast Atlantic, three putative species have been described on the basis of molecular data and a breeding study. We report on the development of a fast genotyping assay and on the occurrence and distribution of different molecular types in the western Wadden Sea. The genotyping assay consists of PCR-RFLP analysis by two enzymes of a mitochondrial (cox3-trnQ-nad6) DNA segment. A new, fourth type was observed and this was the only type whose geographical distribution was not uniform but instead skewed toward the eastern part of the study area. All three hitherto known types were also observed and these three displayed a significant difference in depth distribution within the study area. This is the first ecological difference reported for the ‘Subtidal Clade’ (SC) versus the ‘Type Locality Clade’ (TLC). The new type (‘Intertidal Clade 2’) had a similar distribution to what is known as the Intertidal Clade (IC). However, the most striking observation is the large degree of overlap between the depth distributions in the western Wadden Sea, where the clade names ‘Intertidal’ and ‘Subtidal’ do not necessarily reflect local species composition. The necessity to genotype Scoloplos cf. armiger in monitoring programs is emphasized.     

Annual dynamics of pelagic primary production and respiration in a shallow coastal basin

The Wadden Sea (North Sea, Europe) is a shallow coastal sea with high benthic and pelagic primary production rates. To date, no studies have been carried out in the Wadden Sea that were specifically designed to study the relation between pelagic respiration and production by comparable methods. Because previous studies have suggested that the import of primary-produced pelagic organic matter is important for benthic Wadden Sea carbon budgets, we hypothesised that on an annual average the northern Wadden Sea water column is autotrophic. To test this hypothesis, we studied annual dynamics of primary production and respiration at a pelagic station in a shallow tidal basin (List Tidal Basin, northern Wadden Sea). Since water depth strongly influences production estimates, we calculated primary production rates per unit area in two ways: on the basis of the mean water depth (2.7 m) and on the basis of 1 m depth intervals and their respective spatial extent in the List Tidal Basin. The latter more precise estimate yielded an annual primary production of 146 g C m^− 2 y^− 1. Estimates based on the mean water depth resulted in a 40% higher annual rate of 204 g C m^− 2 y^− 1. The total annual pelagic respiration was 50 g C m^− 2 y^− 1. The P/R ratio varied between seasons: from February to October the water column was autotrophic, with the highest P/R ratio of 4–5 during the diatom spring bloom in April/May. In autumn and winter the water column was heterotrophic. On an annual average, the water column of the List Tidal Basin was autotrophic (P/R 3). We suggest that a large fraction of the pelagic produced organic matter was respired locally in the sediment.     

The ability of a field population of diatoms to discriminate between phosphate and arsenate

Dissolved inorganic arsenic and phosphate levels present in an enclosed experimental ecosystem have been studied during the course of a natural spring diatom bloom. Whilst phosphate levels dropped rapidly to almost zero as the bloom developed, arsenic levels remained constant even when arsenate levels exceeded the dissolved phosphate. Certain diatom species (Thalassiosira spp., Chaetoceras spp. and Skeletonema costatum) therefore appear to be capable of discriminating, either by uptake or excretion, between phosphate and arsenate. The implications of such findings are discussed.     

Differential reproductive strategies of two bivalves in the Dutch Wadden Sea

Cerastoderma edule and Mya arenaria are two common bivalve species in European waters. Longevity and maximum size are much greater in the latter species. Because comparison of species life-history strategies states that a long life span (i.e. high annual survival) generally goes with lower fecundity, we hypothesise that reproductive output would be lower in M. arenaria than in C. edule. In the present paper, we studied the reproductive strategies of these two species in an intertidal and a subtidal area of the western Dutch Wadden Sea, by following seasonal changes in absolute and relative weights of somatic and gonadal tissues in these bivalves. Starting of spawning was similar in the two species, around May, except for intertidal M. arenaria, which initiated spawning in August. Individual energy investment in reproduction was similar for the two species but, unlike M. arenaria, C. edule spawned completely, releasing all energy of gonadal mass in the form of gametes. Mya arenaria used the gonad not only for reproduction but also for storage. In the intertidal area, we found a trade-off between longevity and reproduction, i.e. maximum reproductive output (expressed as a proportion of body mass) was higher in C. edule than in M. arenaria. However, since body size is larger and life span longer in M. arenaria than in C. edule, mean lifetime reproductive output per individual must be higher in the first than in the latter. Based on the differences in reproductive strategies of these two species, we hypothesise that the negative effects of warming climate on bivalve population dynamics in the Wadden Sea will be stronger in C. edule than in M. arenaria.     

Facilitative effects of introduced Pacific oysters on native macroalgae are limited by a secondary invader,the seaweed Sargassum muticum

Introduced habitat-providing organisms such as epibenthic bivalves may facilitate the invasion and expansion of further non-native species which may modify the effects of the primary invader on the native system. In the sedimentary intertidal Wadden Sea (south-eastern North Sea) introduced Pacific oysters (Crassostrea gigas) have overgrown native blue mussel beds (Mytilus edulis). These oyster beds are now providing the major attachment substratum for macroalgae. Recently, oysters have expanded their distribution into the shallow subtidal zone of the Wadden Sea, and there support a rich associated species community including the Japanese seaweed Sargassum muticum, which has been presumably introduced together with the oysters. With a block designed field experiment, we explored the effects of S. muticum on the associated community of soft-bottom C. gigas beds in the shallow subtidal. Replicated oyster plots of 1 m² were arranged with a density of 0, 7, 15 or 45 S. muticum m^? 2, respectively. We found no effects of different S. muticum densities on associated epi- and endobenthic community compositions associated to the oyster plots. However, the overall coverage of sessile organisms settling on the oyster shells was significantly reduced at high S. muticum densities. The occurrence of abundant native macro-algal species such as Polysiphonia nigrescens, Antithamnion plumula and Elachista fucicola decreased with increasing S. muticum densities. Sessile invertebrates, by contrast, were only marginally affected and we found no effects of S. muticum canopy on diversity and abundance of endofauna organisms. We conclude that increasing densities of S. muticum on C. gigas beds in the shallow subtidal zone of the Wadden Sea limit the occurrence of native macroalgae which otherwise would benefit from the additional hard substratum provided by the oysters. Thus, a secondary invader may abolish the effects of the primary invader for native species by occupying the new formed niche.     

Biological control of short-term variations in the concentration of DMSP and DMS during a Phaeocystis spring bloom

In the spring of 1995, short-term variations in the concentration of particulate and dissolved dimethylsulfoniopropionate (DMSP) and dimethylsulfide (DMS) were monitored in the western Wadden Sea, a shallow coastal region in open connection with the North Sea. Significant correlations were found between abundance of Phaeocystis globosa and particulate DMSP; concentrations increased rapidly from 100 to 1650 nM in the middle of April. Highest DMS concentrations were found during the initial phase of the exponential growth of the bloom. DMS production and loss rates of DMSP and DMS were estimated experimentally during various phases of the bloom. DMS production and consumption were roughly in balance, with production only slightly exceeding consumption at the start of the bloom. Rates of production and consumption were highest during the exponential growth phase of Phaeocystis and declined in the course of the bloom (from 300–375 to less than 5 nmol dm⁻³ d⁻¹). Demethylation of DMSP increased during the bloom (from 11 to 1300 nmol dm⁻³ d⁻¹); it accounted for up to 100% of the DMSP loss at the end of the bloom. The shift from DMSP cleavage to demethylation in the course of a Phaeocystis bloom implies that DMS concentrations are not necessarily highest at the peak or towards the end of blooms.     

Tendencies in Coccolithophorid Blooms in Some Marine Environments of the Northern Hemisphere according to the Data of Satellite Observations in 1998–2013

Based on the method developed for the delineation of E. huxleyi blooms, a new technique is achieved for (1) the automated detection of E. huxleyi blooms among coexisting massive blooms of microalgae species of other phytoplankton groups and (2) quantifying the boom surface of this type of coccolithophores. As a result, according to the data of the Climate Change Initiative Ocean Colour (OC CCI) for 1998–2013, we have obtained multiyear time series of variability in both the incidence of E. huxleyi bloom and its area in the North, Norwegian, Greenland, Barents, and Bering seas. It is found that E. huxleyi blooms propagate within the intra-annual cycle from the studied middle-latitude marine areas towards the northern areas of the Northern Atlantic Ocean (NAO) and the Arctic Ocean (AO) following the pathways of the main Gulfstream and its branches. It is also found that E. huxleyi blooms are formed annually, initially in the vicinity of the British Islands; then they successively emerge in the northward direction following the western coast of the Great Britain, turn over its northern extremity to reach, firstly, the North Sea (in May), the Norwegian Sea, and finally the Greenland Sea (in June). Then they burst out in the Barents Sea, where the typical period of blooming lasts until late August and, in some years, even to mid-September. We determine the patterns of maximal rates and duration of blooms for each of the seas studied in the Atlantic and Arctic Oceans. As for the Bering Sea, the temporal and spatial variability in the growth of E. huxleyi has an irregular pattern: after a period of remarkably high expression of this phenomenon in 1998–2001, there was an abrupt decrease in both the number and, especially, extent of bloom areas.     

The oxygen isotopic composition of the fraction less than 2 microns (clay fraction) in Recent sediments from western Europe

The oxygen isotopic composition of the clay fraction in Recent argillaceous sediments in western Europe shows regional differences, the δ¹⁸_SMOW values varying between + 15.8 and + 21.7‰. This phenomenon is used for sediment-transport studies. In the Rhine-Meuse estuary a landward transport of marine sediments is observed. The clay fraction of the sediments in the Wadden Sea is only for a minor part derived from the rivers Rhine and Meuse.     

Low reproductive success for copepods during a bloom of the non‐aldehyde‐producing diatom Cerataulina pelagica in the North Adriatic Sea

Egg production rates and/or hatching success in the copepods Acartia clausi, Calanus helgolandicus and Temora longicornis were negatively affected by a late spring (May–June 2003) phytoplankton bloom in the North Adriatic Sea, dominated mainly by the large diatom Cerataulina pelagica. Highest total concentrations of 3.3·10⁴ cells·ml^?1 were located in the vicinity of the Po River, which also corresponded to the area where the highest numbers of phaeophorbides were measured (0.779, 0.528 and 0.419 μg·l^?1, respectively, compared to an average of the remaining stations of 0.183 ± 0.049 SD), suggesting some grazing on the bloom. Phytoplankton biomass in terms of carbon was dominated by diatoms, representing on average 42% of total phytoplankton carbon and more than 80% at several stations. Cerataulina pelagica, Cyclotella spp., Chaetoceros spp. and small unidentified centric diatoms dominated the diatom community numerically but C. pelagica was by far the dominant diatom in terms of carbon due to its large cell size. This species represented more than 60% of the diatom biomass at nine of the 14 stations sampled, and was absent only at one station, which was the most offshore station sampled during the cruise. Although polyunsaturated aldehydes (PUAs) were not detected, other oxylipins which are hydroxy and keto derivatives of eicosapentaenoic and docosahexaenoic acids that affect copepod reproduction were found in these samples. Hence, we can attribute the negative impact of diatoms not only to PUAs, as previously believed, but also to these compounds. This is the first direct evidence of the presence of oxylipins other than PUAs in marine blooms dominated by diatoms.     

Diatoms in surface sediments of the Sea of Okhotsk and the northwest Pacific Ocean

The present investigation was targeted at diatom composition studies in the surface sediments (0–1 cm) sampled in the Sea of Okhotsk and the northwest Pacific in the depth range from 130 to 6110 m. The taxonomic analysis, as well as the quantitative (the diatom cell abundance per sediment dry weight unit) content and ecological group definition, was applied. Ten diatom taxa are the main body (80–100%) of the diatom assemblages: Bacterosira bathyomphala, Chaetoceros spp. (spores), Actinocyclus curvatulus, Thalassiosira latimarginata (group), T. antarctica (spores), Neodenticula seminae, Rhizosolenia hebetata f. hiemalis, Thalassiothrix longissima, Coscinodiscus marginatus, Coscinodiscus oculus iridis. The relative content of these species reflects the sedimentation conditions for different parts of the sea: the shelf, the continental slope, the open sea, and the ocean. The highest diatom content (45.6.3–60.0 10⁶ per g of dry weight) was found for the surface sediments in the central part of the Sea of Okhotsk and the continental slope of western Kamchatka.     

Seasonality of microzooplankton grazing in the northern Wadden Sea

A sequence of nine dilution experiments was conducted according to Landry and Hassett [Landry, M.R., Hassett, R.P., 1982. Estimating the grazing impact of marine microzooplankton. Mar. Biol. 67, 283–288] in the northern Wadden Sea from March until October 2004 to investigate the seasonality of microzooplankton grazing. From March until April, no grazing was observed. Microzooplankton grazing started in May (0.66 d^− 1) and increased until August (1.22 d^− 1). In October microzooplankton grazing was low again (0.17 d^− 1). Phytoplankton growth rates varied between 0 and 1.1 d^− 1. Since the reliability of dilution experiments is still frequently discussed in literature, we tested if our data obtained by dilution experiments reflected short-term in situ phytoplankton dynamics of the study site. We scaled experimental growth rates to water column irradiance, calculated short-term chlorophyll-a dynamics and compared the results to in situ measured chlorophyll-a concentrations. Calculated chlorophyll-a concentrations correlated significantly with in situ measured chlorophyll-a concentrations but slightly overestimated the in situ measured chlorophyll-a. This overestimation was in the range of phytoplankton assimilation reported for the Wadden Sea benthos. We will show that microzooplankton grazing had a large impact during the Phaeocystis bloom and during summer suggesting that a large proportion of phytoplankton biomass remained the pelagic food web. Microzooplankton grazing did not impact the diatom spring bloom and its demise.