Is the invasion of Prorocentrum minimum (Dinophyceae) related to the nitrogen enrichment of the Baltic Sea?

The abundance of the invasive, bloom-forming dinoflagellate Prorocentrum minimum (Pavillard) Schiller and triangular, oval, and oval-round cell shapes were examined relative to salinity, temperature, and nutrient concentrations at the selected sites in the Baltic Sea. Based on the multiple regression and multivariate statistical analysis, all cell shapes of P. minimum had highly similar distribution relative to these environmental parameters as well as chlorophyll-a, nitrite + nitrate, ammonium, total nitrogen, phosphate, total phosphorus, and silicate. The species was related positively to total nitrogen, and negatively to salinity, temperature, nitrite + nitrate, and silicate:total nitrogen ratio. The results suggest that P. minimum could well adapt to low salinity and temperature and occurred particularly in coastal waters, rich in total nitrogen relative to silicate or other inorganic nutrients. These results indicate that the recent invasion of P. minimum into the Baltic Sea could have been enhanced by the DON enrichment. The results also support the suggestion that P. minimum is one morphospecies with no distinct subtaxa.     

Ecological niche partitioning of the invasive dinoflagellate Prorocentrum minimum and its native congeners in the Baltic Sea

This study analyses three decades of the peculiar bloom-formation history of the potentially toxic invasive planktonic dinoflagellates Prorocentrum minimum (Pavillard) Schiller in the SW Baltic Sea. We tested a research hypothesis that the unexpectedly long delay (nearly two decades) in population development of P. minimum prior to its first bloom was caused by competition with one or several closely related native dinoflagellate species due to ecological niche partitioning which hampered the spread and bloom-forming potential of the invader. We applied the ecological niche concept to a large, long-term phytoplankton database and analysed the invasion history and population dynamics of P. minimum in the SW Baltic Sea coastal waters using the data on phytoplankton composition, abundance and biomass. The ecological niche dimensions of P. minimum and its congener P. balticum were identified as the optimum environmental conditions for the species during the bloom events based on water temperature, salinity, pH, concentration of nutrients (PO₄³⁻; total phosphorus, TP; total nitrogen, TN; SiO₄⁴⁻), TN/TP-ratio and habitat type. The data on spatial distribution and ecological niche dimensions of P. minimum have contributed to the development of the “protistan species maximum concept”. High microplankton diversity at critical salinities in the Baltic Sea may be considered as a possible reason for the significant niche overlap and strong competitive interactions among congeners leading to prolonged delay in population growth of P. minimum preceding its first bloom in the highly variable brackishwater environment.     

Occurrences of Prorocentrum minimum (Pavillard) in México

Prorocentrum minimum is a planktonic dinoflagellate known to produce red tides that can be harmful. To recognize localities and understand occurrences of Prorocentrum minimum blooms in Mexico, published data of plankton from 1942 to present, as well as unpublished data from the authors, were reviewed. Studies and reports covered marine and coastal waters of México during different periods. Presence of P. minimum were reported in the Pacific coast, Gulf of California, Gulf of México, and the Caribbean, but blooms have been only reported since 1990. Thirteen bloom events were recorded. Six occurred in shrimp ponds and seven near aquaculture regions or coastal areas where intensive agriculture is practiced. Most of the blooms can be associated with damage to the surrounding marine biota either in aquaculture ponds or open waters. Direct toxicity has not been fully evaluated, but data suggest that low oxygen may not easily explain all of the damage. Interestingly, for yet unknown reasons, cells belonging to the triangular morphotype have seldom been reported in México.     

The dinoflagellate genus Prorocentrum along the coasts of the Mexican Pacific

We surveyed the dinoflagellate genus Prorocentrum Ehrenberg in Mexican Pacific waters, where it is rather common and sometimes causes red tides in coastal areas or shrimp farms. Material collected from Baja California and the Gulf of California was analyzed. Thirteen species were identified, all of them planktonic (although P. mexicanum is also epiphytic). All species are described by light microscopy, and most are also described by scanning electron microscopy; comments on morphology, taxonomy and distribution are made. Red tides were caused by P. dentatum, P. minimum and P. triestinum. Prorocentrum mexicanum and P. minimum were suspected of being toxic. Four species, previously reported in the Gulf of California, were not found. A total of 18 species, including the new records P. dactylum and P. lebourae have been to date reported from the Mexican Pacific.     

Transport and accumulation of sediment and contaminants in the Lagoon of Kuršiųmarios (Lithuania) and Baltic Sea

In this paper an overview is given of earlier work on transport, deposition AND transformation of suspended matter, and contaminants (metals, hydrocarbons) in Lithuanian coastal waters as well as in other parts of the Baltic Sea. The Lithuanian river Nemunas dischargesc. 600.10⁶ kg suspended matter into the Kuršių Marios Lagoon annually, of which two thirds are accumulated in the lagoon. The remainder is exported to the Baltic Sea through the Klaipéda Strait influencing biological activity in the coastal zone. Sedimentation rate in Kuršių Marios Lagoon is estimated at 3.2 mm y⁻¹. In the last 50 years the surface area of the lagoon decreased by 0.7 km² y⁻¹. Concentration profiles of metals and hydrocarbons were measured in the Kuršių Marios Lagoon, the Baltic proper, and in the Gulf of Finland. Among hydrocarbons both anthropogenic and compounds from natural sources are present. It is argued that correct interpretation of concentration horizons in sediment may be hampered by various processes,e. g. early sedimentogenesis and diagenesis, and cyclic salinity changes in the Baltic Sea.     

Distribution Patterns of Isomorphic Cold-Water Dinoflagellates (Scrippsiella/Woloszynskia Complex) Causing ‘red tides’ in the Baltic Sea

During the latest years medium-sized (15–30 μm), single-celled dinoflagellates have been reported to form blooms in the northern Baltic Proper and the Gulf of Finland in winter and spring. Recent studies (Kremp et al., 2003. Proceedings of the 7th International conference of Modern and Fossil Dinoflagellates, September 21–25, Nagasaki, Japan, 66 pp.) indicate that those blooms are caused by two isomorphic species – Scrippsiella hangoei (Schiller) Larsen, and a new species, tentatively belonging to the genus Woloszynskia. Until now there has been no report on how widely distributed these phytoplankton species are in the Baltic Sea. In this study, the occurrence of Scrippsiella/Woloszynskia complex in the entire Baltic Sea was investigated, by using monitoring data from 1997 to 2003. The species occurred in a salinity range from 2 to 8 PSU. Highest concentrations were observed at salinity 4.5–6.5 PSU. Maximum cell densities of Scrippsiella/Woloszynskia complex in the water column were mainly obtained in April or in the beginning of May by the water temperature <3 °C prior to stratification was formed. In the central Gulf of Finland, the second maximum was found in 1999 and 2002 by the temperature >6 °C. Bloom formations in the Baltic Proper and in the Gulf of Finland may not only be explained by optimum temperature and salinity, but also with other factors e.g. high nutrient concentrations and good seeding conditions from the sediments.     

Decadal-scale changes of dinoflagellates and diatoms in the anomalous baltic sea spring bloom

The algal spring bloom in the Baltic Sea represents an anomaly from the winter-spring bloom patterns worldwide in terms of frequent and recurring dominance of dinoflagellates over diatoms. Analysis of approximately 3500 spring bloom samples from the Baltic Sea monitoring programs revealed (i) that within the major basins the proportion of dinoflagellates varied from 0.1 (Kattegat) to >0.8 (central Baltic Proper), and (ii) substantial shifts (e.g. from 0.2 to 0.6 in the Gulf of Finland) in the dinoflagellate proportion over four decades. During a recent decade (1995-2004) the proportion of dinoflagellates increased relative to diatoms mostly in the northernmost basins (Gulf of Bothnia, from 0.1 to 0.4) and in the Gulf of Finland, (0.4 to 0.6) which are typically ice-covered areas. We hypothesize that in coastal areas a specific sequence of seasonal events, involving wintertime mixing and resuspension of benthic cysts, followed by proliferation in stratified thin layers under melting ice, favors successful seeding and accumulation of dense dinoflagellate populations over diatoms. This head-start of dinoflagellates by the onset of the spring bloom is decisive for successful competition with the faster growing diatoms. Massive cyst formation and spreading of cyst beds fuel the expanding and ever larger dinoflagellate blooms in the relatively shallow coastal waters. Shifts in the dominant spring bloom algal groups can have significant effects on major elemental fluxes and functioning of the Baltic Sea ecosystem, but also in the vast shelves and estuaries at high latitudes, where ice-associated cold-water dinoflagellates successfully compete with diatoms.     

COASTAL DIATOM–ENVIRONMENT RELATIONSHIPS FROM THE GULF OF FINLAND,BALTIC SEA1

Eutrophication of the Baltic Sea has become a serious concern in recent decades. To provide a potential means for quality assessments of coastal waters in this area, we collected a data set of 49 embayments in the Gulf of Finland, and explored the relationship between surface sediment diatom assemblages and 15 environmental variables, with special emphasis on nutrients. Total dissolved nitrogen, total phosphorus, depth, and salinity all accounted for significant and independent fractions of variation in the diatom data and explained 34% of the total variation. There were clear changes in diatom assemblage structures along the nutrient gradients. Although these changes were gradual, we could identify a number of taxa that were more abundant in a particular nutrient environment. These taxa could be used as potential indicators of the quality of coastal waters in the Baltic Sea. Diatom assemblages that were least affected by nutrient enrichment included a variety of benthic species and a relatively high species richness. Small planktonic taxa such as Cyclotella atomus Hustedt, Cyclotella meneghiniana Kützing and Thalassiosira pseudonana Hasle and Heimdal were good indicators of highly elevated nutrient concentrations (>600 lg·L^?1 total dissolved nitrogen and 60 lg·L^?1 total phosphorus) together with low species richness. The first appearance of these small planktonic taxa in regular monitoring could be used as an early warning sign for deteriorating water quality. Diatoms could be applied to water quality classification and monitoring purposes in the coastal waters of the Baltic Sea area using techniques such as weighted‐averaging regression and calibration.     

Chinese mitten crab <Emphasis Type="Italic">Eriocheir sinensis</Emphasis> in the Baltic Sea—a supply-side invader?

Although the Chinese mitten crab Eriocheir sinensis (H. Milne-Edwards, 1853) (Crustacea, Decapoda, Varunidae) invaded the Baltic Sea about 80 years ago, published information on its present distribution and abundance in this region is lacking. We provide here information on its Baltic-wide distribution and long-term population dynamics. The species has been found all over the coastal Baltic Sea and also in some adjacent rivers and lakes. The Chinese mitten crab appears to have increased in abundance in recent years in the northeastern part of the Baltic Sea (Gulf of Finland, Gulf of Riga, northern Baltic Proper). Higher catch rates were observed in spring (April–June) and autumn (September–November). The size variation of crabs in different samples was low (mean carapace width 6.1–6.3 cm). Despite findings of gravid females, the reproduction of the mitten crab in the central, northern and eastern Baltic region is considered unlikely due to low salinity and the individuals caught are assumed to actively migrate into the region from the species’ main European distribution area (southeastern North Sea), certainly over 1500 km migration distance. Thus, the dynamics of the North Sea population is probably regulating, at least in part, the occurrence of the Chinese mitten crab in the Baltic Sea area.     

HETEROCAPSA ARCTICA SUBSP. FRIGIDA SUBSP. NOV. (PERIDINIALES,DINOPHYCEAE)—DESCRIPTION OF A NEW DINOFLAGELLATE AND ITS OCCURRENCE IN THE BALTIC SEA1

Characteristics important in identification of Heterocapsa species (i.e., thecal plate pattern, body scale structure, and shape and position of the nucleus and pyrenoid) are practically identical in the dinoflagellate investigated here and in Heterocapsa arctica T. Horig. described from the Canadian Arctic. Analysis of internal transcribed spacer (ITS) sequences confirms that the two dinoflagellates are very closely related; however, there is a clear difference in their size and shape. Our experiments show that the low‐salinity Baltic Sea brackish water does not reduce the size of the marine H. arctica to match that of the Baltic Sea morphotype. On the basis of these dissimilarities in general morphology and its geographic isolation in the Baltic Sea, we consider our material sufficiently differentiated from the typical H. arctica to warrant the status of a new subspecies, H. arctica subsp. frigida subsp. nov. Being of a distinct cell shape, the occurrence of subsp. frigida has been recorded in Algaline phytoplankton monitoring data collected since 1993. Although it has never been responsible for high biomass blooms, it commonly occurs in spring in the Northern Baltic Proper and in the western Gulf of Finland, when the water temperatures are <5°C.     

Distribution,population structure and salinity tolerance of the invasive amphipod Gmelinoides fasciatus (Stebbing) in the Neva Estuary (Gulf of Finland,Baltic Sea)

In the early 1970s, the Baikalian amphipod Gmelinoides fasciatus (Stebbing) was intentionally introduced into several lakes in the Gulf of Finland basin in order to enhance fish production. By 1996, G. fasciatus successfully colonized the littoral zone of Lake Ladoga and, via the Neva River, invaded the Neva Bay, the freshwater part of the Neva Estuary. In 1999, G. fasciatus was first registered in the inner Neva Estuary, the very first record of the Baikalian amphipod in brackish waters of the Baltic Sea. Distribution, abundance, reproduction and population structure of G. fasciatus in the Neva Estuary were studied during 1998–2000. In some locations of the Neva Estuary, maximum densities of G. fasciatus reached 3500 ind. m⁻². In general, density and biomass of G. fasciatus in the freshwater part of the Neva Estuary were higher (around 1.5 fold) than in the brackish-water part. Fecundity of this amphipod averaged 10–20 eggs per female, depending on body size of females and season. In order to assess the possibility of further spread of G. fasciatus in the Baltic Sea, the salinity tolerance of this species was determined in a series of laboratory experiments. Our results showed that the invasive amphipod G. fasciatus is potentially able to colonize shallow coastal habitats of, for example, the Gulf of Bothnia, Gulf of Riga and other parts of the Baltic Sea with water salinities ranging from 1 to 5 psu.

     

Direct evidence for production of microcystins by Anabaena strains from the Baltic Sea

Anabaena is a filamentous, N(2)-fixing, and morphologically diverse genus of cyanobacteria found in freshwater and brackish water environments worldwide. It contributes to the formation of toxic blooms in freshwater bodies through the production of a range of hepatotoxins or neurotoxins. In the Baltic Sea, Anabaena spp. form late summer blooms, together with Nodularia spumigena and Aphanizomenon flos-aquae. It has been long suspected that Baltic Sea Anabaena may produce microcystins. The presence of microcystins has been reported for the coastal regions of the Baltic proper, and a recent report also indicated the presence of the toxin in the open Gulf of Finland. However, at present there is no direct evidence linking Baltic Sea Anabaena spp. to microcystin production. Here we report on the isolation of microcystin-producing strains of the genus Anabaena in the open Gulf of Finland. The dominant microcystin variants produced by these strains included the highly toxic MCYST-LR as well as [d-Asp(3)]MCYST-LR, [d-Asp(3)]MCYST-HtyR, MCYST-HtyR, [d-Asp(3),Dha(7)]MCYST-HtyR, and [Dha(7)]MCYST-HtyR variants. Toxic strains were isolated from the coastal Gulf of Finland as well as from the easternmost open-sea sampling station, where there were lower salinities than at other stations. This result suggests that lower salinity may favor microcystin-producing Anabaena strains. Furthermore, we sequenced 16S rRNA genes and found evidence for pronounced genetic heterogeneity of the microcystin-producing Anabaena strains. Future studies should take into account the potential presence of microcystin-producing Anabaena sp. in the Gulf of Finland.     

The influence of seasonal and year-to-year variability of water discharge from the Lake Ladoga—Neva River system on the salinity regime of the Baltic Sea

The study is based on multi-year monthly and yearly averaged time series of the Neva River discharge, monthly salinity observations from near-shore hydrographic and hydrometeorological stations in the Gulf of Finland, and several years of data from international seasonal salinity observations in the open region of NE Baltic Sea. There is a good coherence between seasonal variations of the Neva River flow and the salinity regime in the upper layer of the Baltic Sea. Near the shores periods of low, normal and high salinity correspond to the periods of high, normal and low river runoff, and the same trends are observed with some time lag in the open parts of the Sea as well. The impacts of the Neva River discharge oscillations on salinity of the Gulf of Finland surface waters are analyzed in terms of time regression probability models. The obtained river flow-marine salinity transfer functions summarize well the observation data on both multiannual and seasonal scales of variability. Some seasonal periods with different anomalies of the influence are distinguished.     

Prorocentrum minimum (Pavillard) Schiller: A review of a harmful algal bloom species of growing worldwide importance

Prorocentrum minimum (Pavillard) Schiller, a common, neritic, bloom-forming dinoflagellate, is the cause of harmful blooms in many estuarine and coastal environments. Among harmful algal bloom species, P. minimum is important for the following reasons: it is widely distributed geographically in temperate and subtropical waters; it is potentially harmful to humans via shellfish poisoning; it has detrimental effects at both the organismal and environmental levels; blooms appear to be undergoing a geographical expansion over the past several decades; and, a relationship appears to exist between blooms of this species and increasing coastal eutrophication. Although shellfish toxicity with associated human impacts has been attributed to P. minimum blooms from a variety of coastal environments (Japan; France; Norway; Netherlands; New York, USA), only clones isolated from the Mediterranean coast of France, and shellfish exposed to P. minimum blooms in this area, have been shown to contain a water soluble neurotoxic component which killed mice. Detrimental ecosystem effects associated with blooms range from fish and zoobenthic mortalities to shellfish aquaculture mortalities, attributable to both indirect biomass effects (e.g., low dissolved oxygen) and toxic effects. P. minimum blooms generally occur under conditions of high temperatures and incident irradiances and low to moderate salinities in coastal and estuarine environments often characterized as eutrophic, although they have been found under widely varying salinities and temperatures if other factors are conducive for growth. The physiological flexibility of P. minimum in response to changing environmental parameters (e.g., light, temperature, salinity) as well as its ability to utilize both inorganic and organic nitrogen, phosphorus, and carbon nutrient sources, suggest that increasing blooms of this species are a response to increasing coastal eutrophication.     

GYMNODINIUM COROLLARIUM SP. NOV. (DINOPHYCEAE)—A NEW COLD‐WATER DINOFLAGELLATE RESPONSIBLE FOR CYST SEDIMENTATION EVENTS IN THE BALTIC SEA1

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.     

Diversity of luciferase sequences and bioluminescence production in Baltic Sea Alexandrium ostenfeldii

The toxic dinoflagellate Alexandrium ostenfeldii is the only bioluminescent bloom-forming phytoplankton in coastal waters of the Baltic Sea. We analysed partial luciferase gene (lcf) sequences and bioluminescence production in Baltic A. ostenfeldii bloom populations to assess the distribution and consistency of the trait in the Baltic Sea, and to evaluate applications for early detection of toxic blooms. Lcf was consistently present in 61 Baltic Sea A. ostenfeldii strains isolated from six separate bloom sites. All Baltic Sea strains except one produced bioluminescence. In contrast, the presence of lcf and the ability to produce bioluminescence did vary among strains from other parts of Europe. In phylogenetic analyses, lcf sequences of Baltic Sea strains clustered separately from North Sea strains, but variation between Baltic Sea strains was not sufficient to distinguish between bloom populations. Clustering of the lcf marker was similar to internal transcribed spacer (ITS) sequences with differences being minor and limited to the lowest hierarchical clusters, indicating a similar rate of evolution of the two genes. In relation to monitoring, the consistent presence of lcf and close coupling of lcf with bioluminescence suggests that bioluminescence can be used to reliably monitor toxic bloom-forming A. ostenfeldii in the Baltic Sea.     

Growth of enterobacteria on algal mats in the eastern part of the Gulf of Finland

Model experiments on a possibility that pathogenic enterobacteria Salmonella enteritidis (Gartneri) can grow on decaying algal mats with prevalence of the filamentous algae Cladophora glomerata (L.) Kütz were carried out. Samples of algal mats have been collected in the eastern part of the Gulf of Finland in the Baltic Sea. A bacterial culture of Salmonella enteritidis was placed into tubes containing samples of mats. The intensive growth of salmonella was noted in alga samples collected in the freshwater zone (salinity 0.2–1.5‰); growth was practically absent in the samples of algae collected in a zone with salinity 2–3‰, while salmonella remained viable in the control tubes with water without algae. The growth of coliform enterobacteria initially inhabited in the algal mats was discovered in all experiments. Studies carried out in 2009 show that the thickness of the algal mats in the costal zones of the Gulf of Finland reached 20 cm and their biomass reached a few tons per 1 km². These experiments showed that dead algal mats stimulate the growth of enterobacteria in the littoral zone of the Baltic Sea, especially in the freshwater part, and can promote the development of these pathogenic microorganisms.     

Impacts and potential effects due to Prorocentrum minimum blooms in Chesapeake Bay

The dinoflagellate Prorocentrum minimum (P. minimum) can be found in all seasons and over a broad range of habitat conditions in the Chesapeake Bay and its tributaries. Blooms (>3000 cells ml⁻¹), locally referred to as ‘mahagony tides’, were restricted to salinities of 4.5–12.8 psu, water temperatures of 12–28 °C, and occurred most frequently in April and May. P. minimum blooms have been detected at routine water quality monitoring stations located in the main channel of the Bay and tidal tributaries. Nearshore investigations of bloom events, however, have accounted for the majority of events recorded in excess of 10⁵ cells ml⁻¹. Mahogany tides were associated with widespread harmful impacts including anoxic/hypoxic events, finfish kills, aquaculture shellfish kills and submerged aquatic vegetation losses. We summarize the state of knowledge regarding physical and chemical factors related to P. minimum blooms, their abundance, distribution and frequency, and ecological effects in Chesapeake Bay.     

The gulf of Bothnia: The northernmost part of the Baltic Sea

Summary The Baltic Sea, one of the largest brackish water areas in the world, can be characterized as a young, cold sea containing an impoverished ecosystem due to salinity stress. The present Baltic Sea was formed as late as 2000 to 2500 years ago when the Danish sounds became more narrow and shallow. The inflow of freshwater from the surrounding land areas caused the Baltic to gradually attain its brackish character. Today the Baltic covers an area of some 366,000 km² as a series of basins separated by shallower areas and filled with about 22,000 km³ of brackish water. These basins are, from north to south, the Gulf of Bothnia, the Gulf of Finland, the Gotland Sea and the Bornholm Sea. The climate gradient ranges from almost arctic conditions in the extreme north to a more maritime climate in the southern parts. The North Sea salt water is connected to the Baltic through the shallow Kattegat and the sills in the Danish sounds. The inflow of salt water occurs in two different ways,viz. as a continuous flow along the bottom due to the salinity gradient and as pulses of salt water generated by the distribution of air pressure and the direction of the wind. The freshwater input (500 km³) from mainly the large rivers equals roughly the net outflow and stresses the south-bound current along the Swedish coast that also compensates for the salt water inflow. Tidal movements can be seen in the southern Baltic, but are of minor importance for the system. The residence time of the total water mass is 25 years and the hydrographical conditions within the different basins are stable and dominated by a permanent halocline, and a thermocline developing every spring. The salinity ranges from about 1–2 per mille in the innermost part of the Gulf of Bothnia to 10–15 per mille in the Bornholm Sea. Total vertical mixing takes place during winter in at least the northern parts of the sea. Due to the climate-gradient, the ice condition differs from about four months of total ice-cover in the inner parts of the Gulf of Bothnia to one month or less of coastal ice in the southern part of the Baltic. Thus, the seasonal effect is more pronounced in the northern parts.The living systems of the Baltic are reduced and adapted to these varying conditions. When comparing the deeper soft bottoms of the Gulf of Bothnia to the rest of the Baltic, the following pattern can be seen. The pelagic primary productivity increases by a factor 6 from north to south. The southern parts of the sea show a pronounced spring peak, while in the north the spring development is delayed or replaced by a summer maximum. The total increase of the macrofauna biomass is striking, from about 1 g.m^–2 (w.wt) in the north to 100 g.m^–2 (w.wt) or more in the south. The meiofauna and the zooplankton biomasses show less variability. The meiofauna increases by a factor of 2–4, giving a biomass of about twice that of the macrofauna in the northernmost part. The extremely low salinity of this area causes the exclusion of bivalves (filter-feeders) from the fauna. Available data, pooled with the high metabolic rate of the meiofauna, roughly follow the changes in primary productivity within the Baltic Sea. The changing ratio of macro- to meiofauna, as well as results from intensive studies of the macrobenthic amphipodPontoporeia affinis (Lindström), suggest that the macrofauna is regulated mainly by food limitation and that the benthic and pelagic systems are closely coupled.     

LOCAL ADAPTATION OF CERAMIUM TENUICORNE (CERAMIALES,RHODOPHYTA) WITHIN THE BALTIC SEA1

Ceramium tenuicorne (Kützing) Wærn is a red alga that is widely distributed in the brackish Baltic Sea. We studied the growth response of Ceramium to low salinity and nutrient enrichment, using isolades from two regions of the Baltic Sea where the alga approaches its inner distribution limit. Ecotypic differentiation was observed in that differences in growth response among isolates corresponded to salinity conditions in their regions of origin. Isolates from the Gulf of Bothnia (4 psu) had inherently lower growth rates that were not increased when transferred to higher salinity, but were better adapted to very low salinity levels than isolates from the Baltic Proper (7 psu). Further, the results indicate that Ceramium from different regions of the Baltic Sea vary in their responses to nutrient enrichment. The observed differences may be best described as a quantitative difference in the proportion of isolates with hyposaline adaptation. The results indicate that the wide distribution of Ceramium in the Baltic Sea is better explained by the occurrence of locally adapted genotypes than by a generalist life strategy, and provide example of adaptive differentiation in a marine edge environment.