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ICES Cooperative Research Report No. 313 5. PHYTOPLANKTON AND MICROBIAL PLANKTON OF THE BALTIC SEA Norbert Wasmund, Sirpa Lehtinen, Anetta Ameryk, Günther Nausch, and Marie Johansen The Baltic Sea is a classic example of a shallow (Helsinki Commission, HELCOM) in 1974. One intracontinental shelf sea, with a mean depth of of its aims was to investigate long-term trends 52 m. It is connected to the North Sea only by the in trophic conditions by the Baltic Monitoring shallow and narrow Danish Straits, which limit Programme (BMP, later the COMBINE programme), water exchange between these regions. The Kattegat, which has been conducted since 1979 according Sound (Øresund), and Belt Sea (Great Belt, Little to a coordinated sampling schedule and binding Belt, Kiel Bight, and Mecklenburg Bight) represent methodology. The data collected by the riparian the transitional area between the North Sea and the countries in the frame of the HELCOM monitoring Baltic Proper. The Baltic Proper stretches from Darss are stored in the ICES databank and form the basis Sill to the entrances to the gulfs of Riga, Finland, for this analysis. and Bothnia, and comprises different basins that are stratified by a permanent halocline (at depth Already in previous periods, the data have been of 60–80 m in the deepest basins) and a summer analysed for trends by the HELCOM community, thermocline (at 10–30 m depth). Owing to a limited for example in periodic assessments (e.g. HELCOM, exchange with the North Sea, the Baltic Sea is 2002) or in thematic assessments (e.g. on climate a brackish sea with not only a vertical, but also a change or eutrophication; see HELCOM, 2007, horizontal, salinity gradient. The salinity of surface 2009). Specific papers on trends in phytoplankton water decreases to the east and north. A detailed had to take into account differences between the survey of the state and evolution of the Baltic Sea is subareas of the Baltic Sea. In fact, trends in the given by Feistel et al. (2008). different regions may even be opposite. For example, trends in chlorophyll a concentrations were found Large rivers import nutrients from the densely to decrease in Mecklenburg Bight, but increase populated and intensively cultivated catchment in the Baltic Proper in spring from 1979 to 2005 area (Bergström and Carlsson, 1994), resulting in (Wasmund and Siegel, 2008). Wasmund et al. (1998) an increase in phytoplankton biomass, primary discovered a strong decrease in diatom biomass in production, and turbidity in the euphotic zone and spring blooms in the southern Baltic Proper, but oxygen deficit in deep-water layers (Elmgren and not in Mecklenburg Bight. This sudden diatom Larsson, 2001; Rönnberg and Bonsdorff, 2004). decrease at the end of the 1980s was later identified The riparian countries recognized the increasing as a regime shift by Alheit et al. (2005). The decrease environmental problems and agreed to establish the in spring diatoms is compensated by an increase Baltic Marine Environment Protection Commission in dinoflagellates (Wasmund and Uhlig, 2003). ICES Phytoplankton and Microbial Plankton Status Report 2009/2010 Figure 5.1 Locations of the Baltic Sea plankton monitoring areas (Sites 10–29) plotted on a map of average chlorophyll concentration. Blue stars indicate locations of sites described in the adjacent North Sea and English Channel region (see Section 6). The diatoms and dinoflagellates demonstrated The analysis presented here generally agrees with alternating oscillations: strong spring blooms of these previous analyses. Its data basis is, however, diatoms occurred in the 1980s and since 2000, but more complete than in the older papers, because those of dinoflagellates in the 1990s (Wasmund et it involves all data available from the specific al., 2011). sea areas and is not restricted to selected shorter periods (as in Klais et al., 2011) or selected seasons Suikkanen et al. (2007) analysed especially the (as in Suikkanen et al., 2007). In contrast to the summer phytoplankton communities in the earlier study of Suikkanen et al. (2007), for example, northern Baltic Proper and the Gulf of Finland. cyanophytes and chlorophytes did not increase in They found increasing chrysophyte and chlorophyte the Gulf of Finland or in the northern Baltic Proper biomass in both areas, increasing dinoflagellates in the analysis in this report. in the northern Baltic Proper, and increasing cyanobacteria in the Gulf of Finland, together Since the start of the monitoring programme in with an increase in chlorophyll concentrations, 1979, the biomass in many phytoplankton groups temperature, and winter DIN (dissolved inorganic has increased. Changes in many smaller groups are nitrogen) concentrations. statistically significant, but the ecological importance of these groups is rather low owing to their small A complex trend analyses with monitoring data biomass. From the ecologically important groups, of the whole Baltic Sea was performed by Klais the dinoflagellates are increasing, whereas the et al. (2011). They found high variability in the diatoms are decreasing, especially if spring data are spring bloom biomass, with peaks in the Gulf considered, and the diatoms:diatoms+dinoflagellate of Finland, Gulf of Riga, Pomeranian Bight, and ratio during the spring bloom period is also the Danish coast of the Kattegat. Dinoflagellates decreasing. The sudden decrease at the end of the dominate in the northern Baltic Proper, but 1980s, known from the literature, is clearly visible, diatoms in coastal areas and in the western primarily in the Bornholm Sea and the Eastern Baltic. In the relatively short period considered Gotland Basin. The diatoms:diatoms+dinoflagellates (1995–2004), dinoflagellates decreased primarily ratio may be an important indicator of changes in in the Bornholm Sea and the Gulf of Riga, but the ecosystem, which may have high relevance for increased in the Gulf of Finland and Bothnian Bay. the food chain. 38/39 ICES Cooperative Research Report No. 313 The autotrophic ciliate Mesodinium rubrum A general increase in temperature was detected in establishes itself as a third ecologically important all sea areas, but was not significant in the Eastern group. It is strongly increasing in all seasons, Gotland Basin. The concentrations of dissolved especially in the 1990s, but part of this shift may inorganic nitrogen (DIN = NO2+NO3+NH4) are be an artefact because some contributors may have decreasing in the long-term dataseries in all open included this ciliate in the phytoplankton only after sea areas. This is interesting because nitrogen is the early 1990s. However, the sudden increase of the considered to be the limiting nutrient in the Baltic prymnesiophyte Chrysochromulina polylepis since Proper, but its decrease has no clear effect on the the end of 2007 is well documented. The only major phytoplankton biomass. The lack of correlations group that is decreasing is cyanobacteria. Only the between DIN and chlorophyll concentrations is summer period, when they form blooms, is relevant surprising and needs to be investigated. to this consideration. The decrease in cyanobacteria is especially prominent in July and/or August in the Arkona Sea and Bornholm Sea. ICES Phytoplankton and Microbial Plankton Status Report 2009/2010 5.1 Northern Baltic Sea (Sites 10–21) Sirpa Lehtinen Figure 5.1.1 Location of Finnish Environ- mental Institute (SYKE) phyto- plankton monitoring stations in the northern Baltic Sea (Sites 10–21), and their corresponding environmental summary plots (see Section 2.2.1). The yellow boxes and labels indicate the four subareas described in the report text. The Northern Baltic Sea consists of four subareas: manual and the HELCOM PEG (Phytoplankton Bothnian Bay, Bothnian Sea, the Gulf of Finland, Expert Group) taxa and biovolume list (Olenina and the northern Baltic Proper (Figure 5.1.1). et al., 2006; HELCOM, 2010). For the purposes of In the frame of the international COMBINE- this report, the twelve open-water stations have monitoring programme of the Helsinki Commission been combined into four geographical subareas: (HELCOM), the Marine Research Centre of the Bothnian Bay, Bothnian Sea, Gulf of Finland, and Finnish Environment Institute (SYKE) samples northern Baltic Proper. Additional time-series phytoplankton yearly in August at twelve open- results for the individual sites, as well as additional water stations (Sites 10–21) and in numerous SYKE coastal monitoring sites, are available online coastal stations (not considered in this report). at http://wgpme.net/time-series. This sampling has been carried out since 1979, with the data available in the main ICES database. Sampling, processing, and analysis of the samples have been carried out according to the compulsory 40/41 ICES Cooperative Research Report No. 313 5.1.1 Bothnian Bay (Sites 10–11) baltica and dinoflagellate Peridiniella catenata in spring, and photosynthetic ciliate Mesidinium The Bothnian Bay is the most northern part of rubrum after the spring bloom. the Baltic Sea, characterized by low salinity (ca. 2.9–3.5), the strong influence of river waters, and Since the start of the monitoring programme the long duration of ice cover (over 150 d). In in 1979, temperature and chlorophyll a have addition, Bothnian Bay is the only predominantly increased, whereas salinity and silicate have phosphorus-limited Baltic Sea subregion, most decreased. Diatoms, dinoflagellates, cryptophytes, notibly during summer (Tamminen and Andersen, and heterotrophs have decreased significantly, 2007). The maximum bottom depth is 148 m, with whereas chrysophytes and the endosymbiotic ciliate a mean depth of 40 m. The open-water sampling Mesodinium rubrum (not shown) have increased. stations in the Bothnian Bay are F2 (Site 10, 65°23'N 23°27'E) and BO3 (Site 11, 64°18'N 22°21'E). 5.1.3 Gulf of Finland (Sites 15–17) Seasonal and interannual trends (Figure 5.1.2) The surface salinity in the Gulf of Finland is ca. 5.4–6.1. The maximum depth of the Gulf of Finland Diatoms, especially Chaetoceros wighamii, dominate is 121 m.