Plankton in the open Mediterranean Sea: a review.

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 that includes unique thermohaline features, particular multilayer circulation, topographic gyres, and meso- and sub-mesoscale activity. Recent investigations have basically confirmed the long-recognised oligotrophic character of this sea, which enhances along both the west-east, and the north-south directions. Nutrient availability is low, especially for phosphorous (N:P up to 60), although limitation may be relaxed by inputs from highly populated coasts and from the atmosphere. Phytoplankton biomass as chl-a, generally displays low values (less than 0.2 µgchl-al^-1) over large areas, with a modest late winter increase. A large bloom (up to 3 µgl^-1) throughout the late winter and early spring is only observed in the NW area. Relatively high biomass peaks are also 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 progressively deepens from the Alboran Sea (30 m) to the easternmost Levantine basin (120 m). Primary production reveals a similar west-east decreasing trend and ranges from 59 to 150 gCm^-2y^-1 (in situ measurements). Overall the basin is largely dominated by small-sized autotrophs, microheterotrophs and egg-carrying copepod species. The phytoplankton, the microbial (both autotrophic and heterotrophic) and the zooplankton components reveal a considerable diversity and variability over spatial and temporal scales, the latter less explored though. 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-reach copepod genera or families which dominate in the basin. Major dissimilarities between western and eastern basins have been highlighted in species composition of phytoplankton and mesozooplankton, but also in the 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. There, the intermittent nutrient enrichment promotes switches from a small-sized microbial community to diatom-dominated populations. A classical food web is ready to substitute the microbial food web in these cases. These switches, likely occurring within a continuum of trophic pathways, may greatly enhance the flux towards high 5 trophic levels, in spite of an apparent heterotrophy. Basically, the system seems to be top-down controlled and characterised by a multivorous web, as shown by the great variety of feeding modes and preferences and by the significant and simultaneous grazing impact on phytoplankton and ciliates by mesozooplankton. [ABSTRACT FROM AUTHOR]