. We present an overview of the plankton studies conducted during the last 25 years in the epipelagic offshore waters of the Mediterranean Sea. This quasi-enclosed sea is characterized by a rich and complex physical dynamics with distinctive traits, especially in regard to the thermohaline circulation. Recent investigations have basically confirmed the long-recognised oligotrophic nature of this sea, which increases along both the west-east and the north-south directions. Nutrient availability is low, especially for phosphorous (N:P up to 60), though this limitation may be buffered by inputs from highly populated coasts and from the atmosphere. Phytoplankton biomass, as chl a, generally displays low values (less than 0.2 μg chl a l−1) over large areas, with a modest late winter increase. A large bloom (up to 3 μg l−1) is observed throughout the late winter and spring exclusively in the NW area. Relatively high biomass values are recorded in fronts and cyclonic gyres. A deep chlorophyll maximum is a permanent feature for the whole basin, except during the late winter mixing. It is found at increasingly greater depths ranging from 30 m in the Alboran Sea to 120 m in the easternmost Levantine basin. Primary production reveals a west-east decreasing trend and ranges between 59 and 150 g C m−2 y−1 (in situ measurements). Overall, the basin is largely dominated by small autotrophs, microheterotrophs and egg-carrying copepod species. The microorganisms (phytoplankton, viruses, bacteria, flagellates and ciliates) and zooplankton components reveal a considerable diversity and variability over spatial and temporal scales, although the latter is poorly studied. Examples are the wide diversity of dinoflagellates and coccolithophores, the multifarious role of diatoms or picoeukaryotes, and the distinct seasonal or spatial patterns of the species-rich copepod genera or families which dominate the basin. Major dissimilarities between western and eastern basins have been highlighted in species composition of phytoplankton and mesozooplankton, but also in the heterotrophic microbial components and in their relationships. Superimposed to these longitudinal differences, a pronounced biological heterogeneity is also observed in areas hosting deep convection, fronts, cyclonic and anti-cyclonic gyres or eddies. In such areas, the intermittent nutrient enrichment promotes a switching between a small-sized microbial community and diatom-dominated populations. A classical food web readily substitutes the microbial food web in these cases. These switches, likely occurring within a continuum of trophic pathways, may greatly increase the flux towards higher trophic levels, in spite of the apparent heterotrophy. Basically, the microbial system seems to be both bottom-up and top-down controlled. A "multivorous web" is shown by the great variety of feeding modes and preferences and by the significant and simultaneous grazing impact on phytoplankton and ciliates by mesozooplankton.

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Plankton in the open Mediterranean Sea: a review

Atlas of modern organic-walled dinoflagellate cyst distribution

Plankton primary production of the World Ocean.

The brackish Baltic Sea hosts species of various origins and environmental tolerances. These immigrated to the sea 10,000 to 15,000 years ago or have been introduced to the area over the relatively recent history of the system. The Baltic Sea has only one known endemic species. While information on some abiotic parameters extends back as long as five centuries and first quantitative snapshot data on biota (on exploited fish populations) originate generally from the same time, international coordination of research began in the early twentieth century. Continuous, annual Baltic Sea-wide long-term datasets on several organism groups (plankton, benthos, fish) are generally available since the mid-1950s. Based on a variety of available data sources (published papers, reports, grey literature, unpublished data), the Baltic Sea, incl. Kattegat, hosts altogether at least 6,065 species, including at least 1,700 phytoplankton, 442 phytobenthos, at least 1,199 zooplankton, at least 569 meiozoobenthos, 1,476 macrozoobenthos, at least 380 vertebrate parasites, about 200 fish, 3 seal, and 83 bird species. In general, but not in all organism groups, high sub-regional total species richness is associated with elevated salinity. Although in comparison with fully marine areas the Baltic Sea supports fewer species, several facets of the system's diversity remain underexplored to this day, such as micro-organisms, foraminiferans, meiobenthos and parasites. In the future, climate change and its interactions with multiple anthropogenic forcings are likely to have major impacts on the Baltic biodiversity.

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Status of Biodiversity in the Baltic Sea

Several Dinophysis species produce diarrhoetic toxins (okadaic acid and dinophysistoxins) and pectenotoxins, and cause gastointestinal illness, Diarrhetic Shellfish Poisoning (DSP), even at low cell densities (<103 cells·L−1). They are the main threat, in terms of days of harvesting bans, to aquaculture in Northern Japan, Chile, and Europe. Toxicity and toxin profiles are very variable, more between strains than species. The distribution of DSP events mirrors that of shellfish production areas that have implemented toxin regulations, otherwise misinterpreted as bacterial or viral contamination. Field observations and laboratory experiments have shown that most of the toxins produced by Dinophysis are released into the medium, raising questions about the ecological role of extracelular toxins and their potential uptake by shellfish. Shellfish contamination results from a complex balance between food selection, adsorption, species-specific enzymatic transformations, and allometric processes. Highest risk areas are those combining Dinophysis strains with high cell content of okadaates, aquaculture with predominance of mytilids (good accumulators of toxins), and consumers who frequently include mussels in their diet. Regions including pectenotoxins in their regulated phycotoxins will suffer from much longer harvesting bans and from disloyal competition with production areas where these toxins have been deregulated.

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Dinophysis Toxins: Causative Organisms, Distribution and Fate in Shellfish

Evaluation of a 45-year data set of primary production (PP), a 30-year data set of phytoplankton biomass, and a 51-year data set of species composition shows an increase of phytoplankton biomass and abundance in the period from the mid-1980s to the mid-1990s. Phytoplankton biomass showed bimodal seasonal cycles, with winter and spring maxima, which did not change over the past 30 years. Diatoms were the most abundant functional group and they prevailed during the colder part of the year while the dinoflagellate contribution to the phytoplankton community increased in the warmer period from May to August. Diatoms showed a significant negative correlation with sea surface temperature (SST), while dinoflagellates were positively correlated with SST. An increase of phytoplankton abundance, particularly dinoflagellate, in the period from the mid-1980s to the mid-1990s coincided with years characterized by a high North Atlantic Oscillation (NAO) index. Primary production and chlorophyll a concentration in the spring period were negatively correlated with the NAO winter (DJFM) index, probably caused by increased precipitation associated with a low or negative NAO index. PP in winter during the mixing period was positively related to the NAO winter index associated with higher temperatures and dry conditions which brought more clear days and increased input of solar radiation.

Inter-decadal Variability in Phytoplankton Community in the Middle Adriatic (Kaštela Bay) in Relation to the North Atlantic Oscillation

. Long-term time series of physical and chemical parameters collected between 1960 and 2010 along the Palagruža Sill transect, Middle Adriatic Sea, have been investigated in terms of average water properties and their variability. Nutrients, especially orthophosphates, reached rather high levels of concentration below the euphotic zone between 1991 and 1998, the highest levels in the investigated period. Simultaneously, the N:P ratio, which is normally higher than 25:1, decreased to values less than 16:1 in the euphotic zone, indicating a switch from typical phosphorus to nitrogen- limited preconditioning for the primary production. Higher-than-usual nutrient levels peaking in the mid-1990s, coupled with lower-than-usual temperature, salinity, dissolved oxygen and pH values, are presumably related to the flow of the nutrient richer Western Mediterranean waters to the Adriatic below the euphotic layer. These waters, which keep their footprint in the N:P ratio, enter the Adriatic during the anticyclonic phase of the Bimodal Adriatic-Ionian Oscillation (BiOS), which has been uniquely strengthened by the Eastern Mediterranean Transient occurring in the early 1990s. This hypothesis should be confirmed through targeted research and modelling exercises, as it is highly relevant for the biogeochemistry of the Adriatic Sea.

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Changes in the Adriatic oceanographic properties induced by the Eastern Mediterranean Transient

Eutrophication influence on phytoplankton community composition in three bays on the eastern Adriatic coast

Long-term changes of basic biological and chemical parameters at two stations in the middle Adriatic

Variability in the bottom-up (BU) and top-down (TD) regulation of bacteria was analysed on trophic and temporal (seasonal and inter-annual) scales in the middle Adriatic Sea dur- ing 1997-2006 using 3 empirical models. The analyses showed the tendency for bacteria to be TD controlled in oligotrophic open sea stations, and BU controlled in more eutrophic coastal stations. However, temporal variability in BU and TD controls was much stronger, with periods of both strong BU and strong TD controls being observed at all studied stations, independently of their trophic sta- tus. Decomposition of the time series was performed to identify seasonal and inter-annual changes in the relative importance of the BU and TD controls of bacteria. At all stations, BU control dominated during colder periods of the year, whereas TD control dominated during warmer periods. Non- seasonal fluctuations in the relative importance of BU and TD controls of bacteria pointed to a few periods when one or the other type of control was very strong. These periods coincided with some specific meteorological and hydrographic conditions—the strong influence of North Adriatic Dense Water in 1997, the strong Levantine Intermediate Water ingression in 2004, and the extremely warm winter and the Po River runoff in 2000-2001.

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Grozdan Kušpilić

Papers

Inter-decadal Variability in Phytoplankton Community in the Middle Adriatic (Kaštela Bay) in Relation to the North Atlantic Oscillation

Changes in the Adriatic oceanographic properties induced by the Eastern Mediterranean Transient

Eutrophication influence on phytoplankton community composition in three bays on the eastern Adriatic coast

Long-term changes of basic biological and chemical parameters at two stations in the middle Adriatic

Variability in the bottom-up and top-down controls of bacteria on trophic and temporal scales in the middle Adriatic Sea