Spring phytoplankton blooms contribute a substantial part to annual production, support pelagic and benthic secondary production and influence biogeochemical cycles in many temperate aquatic systems. Understanding environmental effects on spring bloom dynamics is important for predicting future climate responses and for managing aquatic systems. We analyzed long-term phytoplankton data from one coastal and one offshore station in the Baltic Sea to uncover trends in timing, composition and size of the spring bloom and its correlations to environmental variables. There was a general trend of earlier phytoplankton blooms by 1-2 weeks over the last 20 years, associated with more sunshine and less windy conditions. High water temperatures were associated with earlier blooms of diatoms and dinoflagellates that dominate the spring bloom, and decreased diatom bloom magnitude. Overall bloom timing, however, was buffered by a temperature and ice related shift in composition from early blooming diatoms to later blooming dinoflagellates and the autotrophic ciliate Mesodinium rubrum. Such counteracting responses to climate change highlight the importance of both general and taxon-specific investigations. We hypothesize that the predicted earlier blooms of diatoms and dinoflagellates as a response to the expected temperature increase in the Baltic Sea might also be counteracted by more clouds and stronger winds. A shift from early blooming and fast sedimenting diatoms to later blooming groups of dinoflagellates and M. rubrum at higher temperatures during the spring period is expected to increase energy transfers to pelagic secondary production and decrease spring bloom inputs to the benthic system, resulting in lower benthic production and reduced oxygen consumption.

Assessing the relative importance of environmental conditions and community interactions is necessary for evaluating the sensitivity of biological communities to anthropogenic change. Phytoplankton communities have a central role in aquatic food webs and biogeochemical cycles, therefore, consequences of differing community sensitivities may have broad ecosystem effects. Using two long-term time series (28 and 20 years) from the Baltic Sea, we evaluated coastal and offshore major phytoplankton taxonomic group biovolume patterns over annual and monthly time-scales and assessed their response to environmental drivers and biotic interactions. Overall, coastal phytoplankton responded more strongly to environmental anomalies than offshore phytoplankton, although the specific environmental driver changed with time scale. A trend indicating a state shift in annual biovolume anomalies occurred at both sites and the shift's timing at the coastal site closely tracked other long-term Baltic Sea ecosystem shifts. Cyanobacteria and the autotrophic ciliate Mesodinium rubrum were more strongly related than other groups to this trend with opposing relationships that were consistent across sites. On a monthly scale, biotic interactions within communities were rare and did not overlap between the coastal and offshore sites. Annual scales may be better able to assess general patterns across habitat types in the Baltic Sea, but monthly community dynamics may differ at relatively small spatial scales and consequently respond differently to future change.

In late autumn/winter 2007, high abundances of the alternate-stage Prymnesium polylepis were observed in many Baltic Sea areas, attaining bloom concentrations in spring 2008. To understand long-term variability in Prymnesiales density, we analysed changes in the abundance of different size classes of Prymnesiales using a long-term time series (1985-2008) for an inshore-offshore gradient in the northern Baltic proper. Further, to understand environmental conditions contributing to the P. polylepis bloom, we examined environmental factors associated with the increase in Prymnesiales abundances by placing the bloom dynamics within the context of temporal and spatial environmental variability over the last decades in the Baltic Sea. Significantly increasing abundances were found for larger size classes of Prymnesiales (6-10 mu m and >10 mu m) but not for the smaller species (Prymnesiales < 6 mu m). Our analyses indicate that high spring sea surface temperatures favor development of larger Prymnesiales (6-10 mu m and > 10 mu m) that tolerate low annual NO2+NO3 concentrations. Thus, long-term trends in increase in abundance, as well as changes in the relative contributions of species within these potentially toxic haptophytes, are likely to be the result of interacting changes in temperature and nutrient conditions since the late 1980s.

Filamentous, nitrogen-fixing cyanobacteria form extensive summer blooms in the Baltic Sea. Their ability to fix dissolved N-2 allows cyanobacteria to circumvent the general summer nitrogen limitation, while also generating a supply of novel bioavailable nitrogen for the food web. However, the fate of the nitrogen fixed by cyanobacteria remains unresolved, as does its importance for secondary production in the Baltic Sea. Here, we synthesize recent experimental and field studies providing strong empirical evidence that cyanobacterial nitrogen is efficiently assimilated and transferred in Baltic food webs via two major pathways: directly by grazing on fresh or decaying cyanobacteria and indirectly through the uptake by other phytoplankton and microbes of bioavailable nitrogen exuded from cyanobacterial cells. This information is an essential step toward guiding nutrient management to minimize noxious blooms without overly reducing secondary production, and ultimately most probably fish production in the Baltic Sea.

It is commonly accepted that summer cyanobacterial blooms cannot be efficiently utilized by grazers due to low nutritional quality and production of toxins; however the evidence for such effects in situ is often contradictory. Using field and experimental observations on Baltic copepods and bloom-forming diazotrophic filamentous cyanobacteria, we show that cyanobacteria may in fact support zooplankton production during summer. To highlight this side of zooplankton-cyanobacteria interactions, we conducted: (1) a field survey investigating linkages between cyanobacteria, reproduction and growth indices in the copepod Acartia tonsa; (2) an experiment testing relationships between ingestion of the cyanobacterium Nodularia spumigena (measured by molecular diet analysis) and organismal responses (oxidative balance, reproduction and development) in the copepod A. bifilosa; and (3) an analysis of long term (1999-2009) data testing relationships between cyanobacteria and growth indices in nauplii of the copepods, Acartia spp. and Eurytemora affinis, in a coastal area of the northern Baltic proper. In the field survey, N. spumigena had positive effects on copepod egg production and egg viability, effectively increasing their viable egg production. By contrast, Aphanizomenon sp. showed a negative relationship with egg viability yet no significant effect on the viable egg production. In the experiment, ingestion of N. spumigena mixed with green algae Brachiomonas submarina had significant positive effects on copepod oxidative balance, egg viability and development of early nauplial stages, whereas egg production was negatively affected. Finally, the long term data analysis identified cyanobacteria as a significant positive predictor for the nauplial growth in Acartia spp. and E. affinis. Taken together, these results suggest that bloom forming diazotrophic cyanobacteria contribute to feeding and reproduction of zooplankton during summer and create a favorable growth environment for the copepod nauplii.

The effects of an extensive bloom of the potentially toxic Prymnesium polylepis (Haptophyta) on breeding eiders (Somateria mollissima) in the Baltic Sea were analysed. Increasing abundances of the alternate stage P. polylepis was detected by a marine monitoring programme in the autumn 2007. The bloom peaked between March and May 2008 in the southern, central and northwestern Baltic Proper and abundances of up to 5 x 10(6) cells l(-1) were recorded. At several sites P. polylepis constituted between 30 and 90% of the total phytoplankton biovolume. The flagellate was only recorded in low numbers in the northeastern Baltic Proper and Gulf of Finland. The abundances were low in 2007, 2009 and 2010. In 28 eider colonies situated in the southern and central Baltic Proper, sharp and synchronous declines in the number of nesting eiders were observed from 2007 to 2008. In colonies on Gotland in the central Baltic Proper, a 76% decrease, from 6650 nests to 1620 nests, was followed by increases in 2009 and 2010, although not up to numbers observed in 2007. At Utklippan and Ertholmene in the southern Baltic Proper, the observed decreases of 55%, from 144 to 65 nests, and 36%, from 1660 to 1060 nests, respectively, between 2007 and 2008, were followed by increases in 2009 and 2010 up to the level observed in 2007. By contrast, no general decline of the number of nesting eiders was observed from 2007 to 2008 in 75 colonies in the northeastern Baltic Proper and Gulf of Finland. Hence, the spatial distribution of the P. polylepis bloom in 2008 closely matched the observed distribution of extensive non-breeding of female eiders. We suggest that the intensive spring bloom of P. polylepis, either through a toxic or non-toxic pathway, affected the main benthic food of eiders, i.e. blue mussels (Mytilus trossulus x Mytilus edulis), at pre-breeding foraging sites close to the breeding sites, and, subsequently, the body condition of adult female eiders and their breeding propensity.

A large bloom of Prymnesium polylepis occurred in the Baltic Sea during the winter 2007 - spring 2008. Based on numerous reports of strong allelopathic effects on phytoplankton exerted by P. polylepis and its toxicity to grazers, we hypothesized that during this period negative correlations will be observed between P. polylepis and (1) main phytoplankton groups contributing to the spring bloom (i.e., diatoms and dinoflagellates), and (2) zooplankton growth and abundance. To test these hypotheses, we analyzed inter-annual variability in phytoplankton and zooplankton dynamics as well as growth indices (RNA: DNA ratio) in dominant zooplankton in relation to the Prymnesium abundance and biomass. Contrary to the hypothesized relationships, no measurable negative responses to P. polylepis were observed for either the total phytoplankton stocks or the zooplankton community. The only negative response, possibly associated with P. polylepis occurrence, was significantly lower abundance of dinoflagellates both during and after the bloom in 2008. Moreover, contrary to the expected negative effects, there were significantly higher total phytoplankton abundance as well as significantly higher winter abundance and winter-spring RNA: DNA ratio in dominant zooplankton species in 2008, indicating that P. polylepis bloom coincided with favourable feeding conditions for zooplankton. Thus, primary consumers, and consequently also zooplanktivores (e.g., larval fish and mysids), may benefit from haptophyte blooms, particularly in winter, when phytoplankton is scarce.

Scale is a key to determining which processes drive community structure. We analyse size distributions of phytoplankton to determine time scales at which we can observe either fixed environmental characteristics underlying communities structure or competition-driven size distributions. Using multiple statistical tests, we characterise size distributions of phytoplankton from 20-year time series in two sites of the Baltic Sea. At large temporal scales (5-20 years), size distributions are unimodal, indicating that fundamental barriers to existence are here subtler than in other systems. Frequency distributions of the average size of the species weighted by biovolume are multimodal over large time scales, although this is the product of often unimodal short-term (<1 year) patterns. Our study represents a much-needed structured, high-resolution analysis of phytoplankton size distributions, revealing that short-term analyses are necessary to determine if, and how, competition shapes them. Our results provide a stepping-stone on which to further investigate the intricacies of competition and coexistence.

Spring bloom composition in the Baltic Sea, a partially ice-covered brackish coastal waterbody, is shaped by winter-spring weather conditions affecting the relative dominance of diatoms and a heterogeneous assemblage of cold-water dinoflagellates, dominated by the chain-forming Peridiniella catenata and a complex of at least three medium-sized, single-celled species: Biecheleria baltica, Gymnodinium corollarium, and Scrippsiella hangoei. During the last decades, the bloom community has dramatically changed in several basins. We analyze here a 30 yr time series of quantitative phytoplankton data, as predicted by hindcast modeled ice thickness and storminess for three distinct Baltic Sea localities, to verify climate-driven mechanisms affecting the spring bloom composition. Thick (> 30 cm) and long-lasting ice cover favored diatom-dominated spring blooms, and mild winters, with storms and thin ice cover (10 to 20 cm), supported blooms of the B. baltica complex. Dispersal limitation plays an important role in the spatial extent of blooms of the B. baltica complex, caused by intricate interplay of local hydrodynamics and the dinoflagellate life cycle. Proportion peaks of key phytoplankton groups have shifted about 10 d earlier in the northwestern Baltic Sea (P. catenata and diatoms) and in the Gulf of Riga (P. catenata). The significant weather effects imply future shifts in spring bloom composition and consequent biogeochemical cycles, driven by the predicted changes in winter storminess and decrease in ice cover extent and duration in climate change models.