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In the Baltic Sea, the dinoflagellates Apocalathium malmogiense , Biecheleria baltica , and Gymnodinium corollarium are important contributors to the spring bloom. However, their relative contribution to the bloom community cannot be unambiguously determined by conventional light microscopy due to a lack of resolution of distinctive morphological features of the three species. Here, we describe a molecular approach based on a quantitative real-time polymerase chain reaction (qPCR) primer and probe system, targeting the ITS1 and ITS2 regions of the rRNA gene for all three species and enabling their quantification. The specificity of the method was demonstrated using monocultures of A. malmogiense , B. baltica , G. corollarium as well as three other dinoflagellate species co-occurring in the Baltic Sea during spring and validated using field-collected phytoplankton samples., Competing Interests: The authors declare that the research was conducted in the absence of any commercial or financial relationships that could be construed as a potential conflict of interest., (Copyright © 2024 Brink, Kremp and Gorokhova.)

Phytoplankton spring blooms are typical features in coastal seas and provide heterotrophic bacteria with a rich blend of dissolved substrates. However, they are difficult to study in coastal seas in-situ. Here, we induced a phytoplankton spring bloom and followed its fate for 37 days in four 600 L-mesocosms. To specifically investigate the significance of phytoplankton-born dissolved organic carbon (DOC) we used artificial seawater with low DOC background and inoculated it with a 100 µm-prefiltered plankton community from the North Sea. A biphasic bloom developed, dominated by diatoms and Phaeocystis globosa respectively. In between, bacterial numbers peaked, followed by a peak in virus-like particles, implying that virus infection caused the collapse. Concentrations of dissolved free amino acids exhibited rapid changes, in particular during the diatom bloom and until the peak in bacterial abundance. Dissolved combined amino acids and neutral monosaccharides accumulated continuously, accounting for 22% of DOC as a mean and reaching levels as high as 44%. Bacterial communities were largely dominated by Bacteroidetes, especially the NS3a marine group (family Flavobacteriaceae), but Rhodobacteraceae and Gammaproteobacteria were also prominent members. Our study shows rapid organic matter and community composition dynamics that are hard to trace in natural coastal ecosystems., (© The Author(s) 2023. Published by Oxford University Press on behalf of FEMS.)

Net growth of microbial populations, that is, changes in abundances over time, can be studied using 16S rRNA fluorescence in situ hybridization (FISH). However, this approach does not differentiate between mortality and cell division rates. We used FISH-based image cytometry in combination with dilution culture experiments to study net growth, cell division, and mortality rates of four bacterial taxa over two distinct phytoplankton blooms: the oligotrophs SAR11 and SAR86, and the copiotrophic phylum Bacteroidetes , and its genus Aurantivirga . Cell volumes, ribosome content, and frequency of dividing cells (FDC) co-varied over time. Among the three, FDC was the most suitable predictor to calculate cell division rates for the selected taxa. The FDC-derived cell division rates for SAR86 of up to 0.8/day and Aurantivirga of up to 1.9/day differed, as expected for oligotrophs and copiotrophs. Surprisingly, SAR11 also reached high cell division rates of up to 1.9/day, even before the onset of phytoplankton blooms. For all four taxonomic groups, the abundance-derived net growth (-0.6 to 0.5/day) was about an order of magnitude lower than the cell division rates. Consequently, mortality rates were comparably high to cell division rates, indicating that about 90% of bacterial production is recycled without apparent time lag within 1 day. Our study shows that determining taxon-specific cell division rates complements omics-based tools and provides unprecedented clues on individual bacterial growth strategies including bottom-up and top-down controls. IMPORTANCE The growth of a microbial population is often calculated from their numerical abundance over time. However, this does not take cell division and mortality rates into account, which are important for deriving ecological processes like bottom-up and top-down control. In this study, we determined growth by numerical abundance and calibrated microscopy-based methods to determine the frequency of dividing cells and subsequently calculate taxon-specific cell division rates in situ . The cell division and mortality rates of two oligotrophic (SAR11 and SAR86) and two copiotrophic ( Bacteroidetes and Aurantivirga ) taxa during two spring phytoplankton blooms showed a tight coupling for all four taxa throughout the blooms without any temporal offset. Unexpectedly, SAR11 showed high cell division rates days before the bloom while cell abundances remained constant, which is indicative of strong top-down control. Microscopy remains the method of choice to understand ecological processes like top-down and bottom-up control on a cellular level., Competing Interests: The authors declare no conflict of interest.

A key phenological event in the annual cycle of many pelagic ecosystems is the onset of the spring algal bloom (OAB). Descriptions of the factors controlling the OAB in temperate to polar lakes have been limited to isolated studies of single systems and conceptual models. Here we present a validated modelling approach that, for the first time, enables a quantitative prediction of the OAB and a systematic assessment of the processes controlling its timing on a continental scale. We used a weather-driven, one-dimensional lake model to simulate the seasonal dynamics of the underwater light climate in 16 lake types characterized by the factorial combination of four lake depths with four levels of water transparency. We did so at 1962 locations across Western Europe and over 31 years (1979-2009). Assuming that phytoplankton production is light-limited in winter, we identified four patterns of OAB control across lake types and climate zones. OAB timing is controlled by (i) the timing of ice-off in ice-covered clear or shallow lakes, (ii) the onset of thermal stratification in sufficiently deep and turbid lakes and (iii) the seasonal increase in incident radiation in all other lakes, except for (iv) ice-free, shallow and clear lakes in the south, where phytoplankton is not light-limited. The model predicts that OAB timing should respond to two pervasive environmental changes, global warming and browning, in opposite ways. OAB timing should be highly sensitive to warming in lakes where it is controlled by either ice-off or the onset of stratification, but resilient to warming in lakes where it is controlled by incident radiation. Conversely, OAB timing should be most sensitive to browning where it is controlled by incident radiation, but resilient to browning where it is controlled by ice-off or the onset of stratification. Available lake data are consistent with our findings., (© 2021 The Authors. Global Change Biology published by John Wiley & Sons Ltd.)

The spring blooms of phytoplankton play a key role in the functioning of marine ecosystems in the polar regions. A spring bloom in the Subarctic White Sea was observed in order to determine the effect of ice cover on the distribution, composition, and temporal changes of phytoplankton communities. The obtained results clearly show that in the White Sea, as in other freezing Arctic seas, ice melting and ice removal both play an essential role in the onset and development of spring phytoplankton blooms. This facilitates the release of ice algae and ice-pelagic algae into the water, as well as the rapid development of true planktonic taxa within the euphotic zone. One major peak of algal biomass is associated with ice removal while the other is recorded in the early summer. Comparison of our results with earlier data from 1960s to 1990s indicated strong year-to-year variation in terms of ice removal, the onset of the spring bloom, and the abundance and composition of the dominant phytoplankton taxa., (Copyright © 2020 Elsevier Ltd. All rights reserved.)

Diarrhetic shellfish poisoning (DSP) toxins and pectenotoxins (PTX) produced by endemic species of the genus Dinophysis, mainly D. acuta and D. acuminata, pose a big threat to public health, artisanal fisheries and the aquaculture industry in Southern Chile. This work reports the first detection of lipophilic toxins, including pectenotoxin-2 (PTX-2) and gymnodimine-A (GYM-A), in hard razor clam (Tagelus dombeii) associated with an unprecedented spring bloom -38.4 × 10 3 cells L -1 in integrated hose sampler (0-10 m) - of Dinophysis acuminata in coastal waters of central Chile. The socio-economic challenges to small-scale fisheries are discussed. The study points to the pressing need for sound policies to face unexpected HAB event, probably due to biogeographical expansions, with a focus on fisheries management, participation of stakeholders, and development of adaptive capacities., (Copyright © 2020 Elsevier Ltd. All rights reserved.)

The impacts of grazing by meso- and microzooplankton on phytoplankton primary production (PP) was investigated in the surface layer of the western North Atlantic during spring. Shipboard experiments were performed on a latitudinal transect at three stations that differed in mixed layer depth, temperature, and mesozooplankton taxonomic composition. The mesozooplankton community was numerically dominated by Calanus finmarchicus at the northern and central station, with Calanus hyperboreus also present at the northern station. The southern station was >10 °C warmer than the other stations and had the most diverse mesozooplankton assemblage, dominated by small copepods including Paracalanus spp. Microzooplankton grazing was detected only at the northern station, where it removed 97% of PP. Estimated clearance rates by C. hyperboreus and C. finmarchicus suggested that at in-situ abundance these mesozooplankton were not likely to have a major impact on phytoplankton abundance, unless locally aggregated. Although mesozooplankton grazing impact on total phytoplankton was minimal, these grazers completely removed the numerically scarce > 10 µm particles, altering the particle-size spectrum. At the southern station, grazing by the whole mesozooplankton assemblage resulted in a removal of 14% of PP, and its effect on net phytoplankton growth rate was similar irrespective of ambient light. In contrast, reduction in light availability had an approximately 3-fold greater impact on net phytoplankton growth rate than mesozooplankton grazing pressure. The low mesozooplankton grazing impact across stations suggests limited mesozooplankton-mediated vertical export of phytoplankton production. The constraints provided here on trophic transfer, as well as quantitative estimates of the relative contribution of light and grazer controls of PP and of grazer-induced shifts in particle size spectra, illuminate food web dynamics and aid in parameterizing modeling-frameworks assessing global elemental fluxes and carbon export., Competing Interests: Susanne Menden Deuer is an Academic Editor for PeerJ. The other authors declare they have no competing interests., (©2020 Morison et al.)

Azaspiracids (AZA) are the most recently discovered group of lipophilic marine biotoxins of microalgal origin, and associated with human incidents of shellfish poisoning. They are produced by a few species of Amphidomataceae, but diversity and occurrence of the small-sized dinophytes remain poorly explored for many regions of the world. In order to analyze the presence and importance of Amphidomataceae in a highly productive area of Argentinean coastal waters (El Rincón area, SW Atlantic), a scientific cruise was performed in 2015 to sample the early spring bloom. In a multi-method approach, light microscopy was combined with real-time PCR molecular detection of Amphidomataceae, with chemical analysis of AZA, and with the establishment and characterization of amphidomatacean strains. Both light microscopy and PCR revealed that Amphidomataceae were widely present in spring plankton communities along the El Rincón area. They were particularly abundant offshore at the shelf front, reaching peak densities of 2.8 × 10 5 cells L -1 , but no AZA were detected in field samples. In total, 31 new strains were determined as Az. dalianense and Az. spinosum, respectively. All Az. dalianense were non-toxigenic and shared the same rRNA sequences. The large majority of the new Az. spinosum strains revealed for the first time the presence of a non-toxigenic ribotype of this species, which is otherwise the most important AZA producer in European waters. One of the new Az. spinosum strains, with a particular slender shape and some other morphological peculiarities, clustered with toxigenic strains of Az. spinosum from Norway and, exceptionally for the species, produced only AZA-2 but not AZA-1. Results indicate a wide diversity within Az. spinosum, both in terms of sequence data and toxin profiles, which also will affect the qualitative and quantitative performance of the specific qPCR assay for this species. Overall, the new data provide a more differentiated perspective of diversity, toxin productivity and occurrence of Amphidomataceae in a poorly explored region of the global ocean., (Copyright © 2019 Elsevier B.V. All rights reserved.)

Small photosynthetic eukaryotes play important roles in oceanic food webs in coastal regions. We investigated seasonal changes in the communities of photosynthetic picoeukaryotes (PPEs) of the class Mamiellophyceae, including the genera Bathycoccus, Micromonas and Ostreococcus, in Ofunato Bay, which is located in northeastern Japan and faces the Pacific Ocean. The abundances of PPEs were assessed over a period of one year in 2015 at three sampling stations, KSt. 1 (innermost bay area), KSt. 2 (middle bay area) and KSt. 3 (bay entrance area) at depths of 1 m (KSt. 1, KSt. 2 and KSt. 3), 8 m (KSt. 1) or 10 m (KSt. 2 and KSt. 3) by employing MiSeq shotgun metagenomic sequencing. The total abundances of Bathycoccus, Ostreococcus and Micromonas were in the ranges of 42-49%, 35-49% and 13-17%, respectively. Considering all assayed sampling stations and depths, seasonal changes revealed high abundances of PPEs during the winter and summer and low abundances during late winter to early spring and late summer to early autumn. Bathycoccus was most abundant in the winter, and Ostreococcus showed a high abundance during the summer. Another genus, Micromonas, was relatively low in abundance throughout the study period. Taken together with previously suggested blooming periods of phytoplankton, as revealed by chlorophyll a concentrations in Ofunato Bay during spring and autumn, these results for PPEs suggest that greater phytoplankton blooming has a negative influence on the seasonal occurrences of PPEs in the bay., (Copyright © 2018 Elsevier B.V. All rights reserved.)

To assess protistan grazing impact and temperature sensitivity on plankton population dynamics, we measured bulk and species-specific phytoplankton growth and herbivorous protist grazing rates in Disko Bay, West Greenland in April-May 2011. Rate estimates were made at three different temperatures in situ (0 °C), +3 °C and +6 °C over ambient. In situ Chlorophyll a (Chl a ) doubled during the observation period to ∼12  µg Chl a L -1 , with 60-97% of Chl a in the >20 µm size-fraction dominated by the diatom genus Chaetoceros. Herbivorous dinoflagellates comprised 60-80% of microplankton grazer biomass. At in situ temperatures, phytoplankton growth or grazing by herbivorous predators <200 µm was not measurable until 11 days after observations commenced. Thereafter, phytoplankton growth was on average 0.25 d -1 . Phytoplankton mortality due to herbivorous grazing was only measured on three occasions but the magnitude was substantial, up to 0.58 d -1 . Grazing of this magnitude removed ∼100% of primary production. In short-term temperature-shift incubation experiments, phytoplankton growth rate increased significantly (20%) at elevated temperatures. In contrast, herbivorous protist grazing and species-specific growth rates decreased significantly (50%) at +6 °C. This differential response in phytoplankton and herbivores to temperature increases resulted in a decrease of primary production removed with increasing temperature. Phaeocystis spp. abundance was negatively correlated with bulk grazing rate. Growth and grazing rates were variable but showed no evidence of an inherent, low temperature limitation. Herbivorous protist growth rates in this study and in a literature review were comparable to rates from temperate waters. Thus, an inherent physiological inhibition of protistan growth or grazing rates in polar waters is not supported by the data. The large variability between lack of grazing and high rates of primary production removal observed here and confirmed in the literature for polar waters implies larger amplitude fluctuations in phytoplankton biomass than slower, steady grazing losses of primary production., Competing Interests: Susanne Menden-Deuer is an Academic Editor for PeerJ.

An indoor mesocosm experiment was carried out to investigate the combined effects of ocean acidification and warming on the species composition and biogeochemical element cycling during a winter/spring bloom with a natural phytoplankton assemblage from the Kiel fjord, Germany. The experimental setup consisted of a "Control" (ambient temperature of ~4.8 °C and ~535 ± 25 μatm p CO 2 ), a "High-CO 2 " treatment (ambient temperature and initially 1020 ± 45 μatm p CO 2 ) and a "Greenhouse" treatment (~8.5 °C and initially 990 ± 60 μatm p CO 2 ). Nutrient replete conditions prevailed at the beginning of the experiment and light was provided at in situ levels upon reaching p CO 2 target levels. A diatom-dominated bloom developed in all treatments with Skeletonema costatum as the dominant species but with an increased abundance and biomass contribution of larger diatom species in the Greenhouse treatment. Conditions in the Greenhouse treatment accelerated bloom development with faster utilization of inorganic nutrients and an earlier peak in phytoplankton biomass compared to the Control and High CO 2 but no difference in maximum concentration of particulate organic matter (POM) between treatments. Loss of POM in the Greenhouse treatment, however, was twice as high as in the Control and High CO 2 treatment at the end of the experiment, most likely due to an increased proportion of larger diatom species in that treatment. We hypothesize that the combination of warming and acidification can induce shifts in diatom species composition with potential feedbacks on biogeochemical element cycling.

Previous studies conducted on Daya Bay implied that the bay had been undergoing potential phosphorus limitation. In this context, alkaline phosphatase activity (APA) and the associated microbes were investigated in three different seasons in Daya Bay, South China Sea. Both bulk-community (fractioned into dissolved and particulate) and single-cell assays of APA were used to estimate the P status of phytoplankton at the community and species level. Unexpected high potential APA (V max ) was observed in Daya Bay. Bulk APA showed that the maximum value in the spring (mean 583.26 nM h -1 ) corresponded well to low phosphate concentration. Furthermore, particulate APA (P-APA) showed an inverse hyperbolic relationship with phosphate, implying the coexistence of both constitutive and inducible AP; meanwhile, a threshold phosphate concentration for the transition from high to low APA was found around 0.2 μM in our study. P-APA and dissolved APA (D-APA) exhibited a tight link with phytoplankton and bacteria, which indicated that both of them were two main carriers of the enzyme. During the spring cruise, we encountered a small-scaled bloom of Gymnodinium that was probably at a declining phase. Extreme high levels of bulk and D-APA were characterized at this spring bloom event, and we suspected that bacteria especially active bacteria played an important role in APA production and partitioning at the post-bloom phase. In Daya Bay, diatoms were the dominant phytoplankton groups and percentages of ELF (Enzyme Labelled Fluorescence) labelled diatoms followed the same seasonal fluctuation as bulk APA, which suggested that diatoms were responsible for major variations of the bulk AP activity except for the spring bloom. Taken together, we considered that phytoplankton may be experiencing more P stress in spring and that the mineralization of organic P via alkaline phosphatase may help phytoplankton overcome P deficiency., (Copyright © 2018 Elsevier Ltd. All rights reserved.)

Carbohydrates represent an important fraction of labile and semi-labile marine organic matter that is mainly comprised of exopolymeric substances derived from phytoplankton exudation and decay. This study investigates the composition of total combined carbohydrates (tCCHO; >1 kDa) and the community development of free-living (0.2-3 μm) and particle-associated (PA) (3-10 μm) bacterioplankton during a spring phytoplankton bloom in the southern North Sea. Furthermore, rates were determined for the extracellular enzymatic hydrolysis that catalyzes the initial step in bacterial organic matter remineralization. Concentrations of tCCHO greatly increased during bloom development, while the composition showed only minor changes over time. The combined concentration of glucose, galactose, fucose, rhamnose, galactosamine, glucosamine, and glucuronic acid in tCCHO was a significant factor shaping the community composition of the PA bacteria. The richness of PA bacteria greatly increased in the post-bloom phase. At the same time, the increase in extracellular β-glucosidase activity was sufficient to explain the observed decrease in tCCHO, indicating the efficient utilization of carbohydrates by the bacterioplankton community during the post-bloom phase. Our results suggest that carbohydrate concentration and composition are important factors in the multifactorial environmental control of bacterioplankton succession and the enzymatic hydrolysis of organic matter during phytoplankton blooms.

Early spring phytoplankton blooms can occur at very low water temperatures but they are often decoupled from bacterial growth, which is assumed to be often temperature controlled. In a previous mesocosm study with Baltic Sea plankton communities, an early diatom bloom was associated with a high relative abundance of Glaciecola sequences ( Gammaproteobacteria ), at both low (2°C) and elevated (8°C) temperatures, suggesting an important role for this genus in phytoplankton-bacteria coupling. In this study, the temperature-dependent dynamics of free-living Glaciecola spp. during the bloom were analyzed by catalyzed reporter deposition fluorescence in situ hybridization using a newly developed probe. The analysis revealed the appearance of Glaciecola spp. in this and in previous spring mesocosm experiments as the dominating bacterial clade during diatom blooms, with a close coupling between the population dynamics of Glaciecola and phytoplankton development. Although elevated temperature resulted in a higher abundance and a higher net growth rate of Glaciecola spp. (Q 10 ∼ 2.2), their growth was, in contrast to that of the bulk bacterial assemblages, not suppressed at 2°C and showed a similar pattern at 8°C. Independent of temperature, the highest abundance of Glaciecola spp. (24.0 ± 10.0% of total cell number) occurred during the peak of the phytoplankton bloom. Together with the slightly larger cell size of Glaciecola , this resulted in a ∼30% contribution of Glaciecola to total bacterial biomass. Overall, the results of this and previous studies suggest that Glaciecola has an ecological niche during early diatom blooms at low temperatures, when it becomes a dominant consumer of phytoplankton-derived dissolved organic matter.

In temperate systems, phytoplankton spring blooms deplete inorganic nutrients and are major sources of organic matter for the microbial loop. In response to phytoplankton exudates and environmental factors, heterotrophic microbial communities are highly dynamic and change their abundance and composition both on spatial and temporal scales. Yet, most of our understanding about these processes comes from laboratory model organism studies, mesocosm experiments or single temporal transects. Spatial-temporal studies examining interactions of phytoplankton blooms and bacterioplankton community composition and function, though being highly informative, are scarce. In this study, pelagic microbial community dynamics (bacteria and phytoplankton) and environmental variables were monitored during a spring bloom across the Baltic Proper (two cruises between North Germany to Gulf of Finland). To test to what extent bacterioplankton community composition relates to the spring bloom, we used next generation amplicon sequencing of the 16S rRNA gene, phytoplankton diversity analysis based on microscopy counts and population genotyping of the dominating diatom Skeletonema marinoi. Several phytoplankton bloom related and environmental variables were identified to influence bacterial community composition. Members of Bacteroidetes and Alphaproteobacteria dominated the bacterial community composition but the bacterial groups showed no apparent correlation with direct bloom related variables. The less abundant bacterial phyla Actinobacteria, Planctomycetes, and Verrucomicrobia, on the other hand, were strongly associated with phytoplankton biomass, diatom:dinoflagellate ratio, and colored dissolved organic matter (cDOM). Many bacterial operational taxonomic units (OTUs) showed high niche specificities. For example, particular Bacteroidetes OTUs were associated with two distinct genetic clusters of S. marinoi. Our study revealed the complexity of interactions of bacterial taxa with inter- and intraspecific genetic variation in phytoplankton. Overall, our findings imply that biotic and abiotic factors during spring bloom influence bacterial community dynamics in a hierarchical manner.

Most of phytoplankton influence is barely understood at the sub meso scale and daily scale because of the lack of means to simultaneously assess phytoplankton functionality, dynamics and community structure. For a few years now, it has been possible to address this objective with an automated in situ high frequency sampling strategy. In order to study the influence of environmental short-term events (nutrients, wind speed, precipitation, solar radiation, temperature, and salinity) on the onset of the phytoplankton bloom in the oligotrophic Bay of Villefranche-sur-Mer (NW Mediterranean Sea), a fully remotely controlled automated flow cytometer (CytoSense) was deployed on a solar-powered platform (EOL buoy, CNRS-Mobilis). The CytoSense carried out single-cell analyses on particles (1-800 μm in width, up to several mm in length), recording optical pulse shapes when analyzing several cm(3). Samples were taken every 2 h in the surface waters during 2 months. Up to 6 phytoplankton clusters were resolved based on their optical properties (PicoFLO, Picoeukaryotes, Nanophytoplankton, Microphytoplankton, HighSWS, HighFLO). Three main abundance pulses involving the 6 phytoplankton groups monitored indicated that the spring bloom not only depends on light and water column stability, but also on short-term events such as wind events and precipitation followed by nutrient pulses. Wind and precipitation were also determinant in the collapse of the clusters' abundances. These events occurred within a couple of days, and phytoplankton abundance reacted within days. The third abundance pulse could be considered as the spring bloom commonly observed in the area. The high frequency data-set made it possible to study the phytoplankton cell cycle based on daily cycles of forward scatter and abundance. The combination of daily cell cycle, abundance trends and environmental pulses will open the way to the study of phytoplankton short-term reactivity to environmental conditions.

A naked dinoflagellate with a unique arrangement of chloroplasts in the center of the cell was isolated from the northern Baltic proper during a spring dinoflagellate bloom (March 2005). Morphological, ultrastructural, and molecular analyses revealed this dinoflagellate to be undescribed and belonging to the genus Gymnodinium F. Stein. Gymnodinium corollarium A. M. Sundström, Kremp et Daugbjerg sp. nov. possesses features typical of Gymnodinium sensu stricto, such as nuclear chambers and an apical groove running in a counterclockwise direction around the apex. Phylogenetic analyses based on partial nuclear-encoded LSU rDNA sequences place the species in close proximity to G. aureolum, but significant genetic distance, together with distinct morphological features, such as the position of chloroplasts, clearly justifies separation from this species. Temperature and salinity experiments revealed a preference of G. corollarium for low salinities and temperatures, confirming it to be a cold-water species well adapted to the brackish water conditions in the Baltic Sea. At nitrogen-deplete conditions, G. corollarium cultures produced small, slightly oval cysts resembling a previously unidentified cyst type commonly found in sediment trap samples collected from the northern and central open Baltic Sea. Based on LSU rDNA comparison, these cysts were assigned to G. corollarium. The cysts have been observed in many parts of the Baltic Sea, indicating the ecologic versatility of the species and its importance for the Baltic ecosystem., (© 2009 Phycological Society of America.)