Diatoms as a proxy in reconstructing the Holocene environmental changes in the south-western Baltic Sea: the lower Rega River Valley sedimentary record

This study focuses on diatom assemblages occurring in cores of Late-glacial and Holocene deposits retrieved from the mouth of the lower Rega River valley, of the southern coast of the Baltic Sea. Sediment samples from four cores were the subject of the present study. Diatom-inferred environmental characteristics, e.g., water level; water salinity (conductivity), trophic status and pH, within each core are presented. Diatom assemblage zones (DAZ) were distinguished, based on differences in the distribution of particular ecological groups. Each DAZ appears to be related to environmental changes during the deposition of a given sediment interval. The Late-glacial (Allerød) sediments originated in a shallow lake with increasing concentrations of solutes and nutrients. The Holocene record begins in the early Atlantic Chronozone and the diatoms point to weakly brackish-water sediments deposited in a shallow water environment. During the period of 8,500–5,800 cal year BP sedimentation took place in a shallow embayment and/or lagoon. From ca. 5,800 cal year BP onwards sedimentation took place in a peat bog environment alternating with Aeolian deposition. Changes in diatom community structure imply a close relationship with the climate-controlled eustatic rise of the ocean level and its consequence Littorina transgression. As with other southern Baltic Sea localities, brackish-water diatoms appear in the sediments, signaling the onset of marine transgression somewhat earlier than previously accepted. Differences and similarities in diatom assemblages and the palaeogeographic development of nearby regions within the Baltic Sea basin and lagoons (coastal areas) from different geographic regions, are also discussed.     

Population and size-specific distribution of Atlantic salmon Salmo salar in the Baltic Sea over five decades

Population-specific assessment and management of anadromous fish at sea requires detailed information about the distribution at sea over ontogeny for each population. However, despite a long history of mixed-stock sea fisheries on Atlantic salmon, Salmo salar, migration studies showing that some salmon populations feed in different regions of the Baltic Sea and variation in dynamics occurs among populations feeding in the Baltic Sea, such information is often lacking. Also, current assessment of Baltic salmon assumes equal distribution at sea and therefore equal responses to changes in off-shore sea fisheries. Here, we test for differences in distribution at sea among and within ten Atlantic salmon Salmo salar populations originating from ten river-specific hatcheries along the Swedish Baltic Sea coast, using individual data from >125,000 tagged salmon, recaptured over five decades. We show strong population and size-specific differences in distribution at sea, varying between year classes and between individuals within year classes. This suggests that Atlantic salmon in the Baltic Sea experience great variation in environmental conditions and exploitation rates over ontogeny depending on origin and that current assessment assumptions about equal exploitation rates in the offshore fisheries and a shared environment at sea are not valid. Thus, our results provide additional arguments and necessary information for implementing population-specific management of salmon, also when targeting life stages at sea.     

Holocene environmental changes and the seal (Phocidae) fauna of the Baltic Sea: coming,going and staying

• 1 The occurrence of different species of seal (Phocidae) during the Holocene in north‐eastern Europe was influenced by a changing geological situation over the last 12 000 years, characterized by alternating freshwater, brackish and marine stages and by spatially and temporally limited opportunities for migration of marine species into the Baltic basin. The patterns of immigration by ringed seal Phoca hispida, harp seal Phoca groenlandica, grey seal Halichoerus grypus and harbour seal Phoca vitulina are reviewed here in order to understand present distributions and population sizes. Furthermore, the future of their populations is analysed.
• 2 Phoca hispida immigrated into the Baltic basin during its brackish Yoldia Stage at the Late Glacial/Early Holocene boundary and has persisted in some disjunct populations until now. A second immigration followed at the beginning of the Littorina Transgression – which resulted in the present stage of the Baltic Sea, obviously caused by the strong cooling event that occurred 6200 calibrated ¹⁴C years BC. Its present status is not threatened, but global warming will restrict suitable habitat.
• 3 Phoca groenlandica had a mid‐Holocene intermezzo in the Baltic Sea. Its later disappearance was probably caused by hunting of subadult animals by humans.
• 4 Halichoerus grypus and Phoca vitulina also immigrated at the beginning of the present stage of the Baltic Sea. Whereas H. grypus has been common from this beginning point until modern times, the population of P. vitulina persisted at a low level for millennia. Only since the 18th century has its population increased and spread over the whole Baltic Sea.

     

Late- and postglacial history of lakes of the Karelian Isthmus

The Late Pleistocene and Holocene history of five lakes in the central part of Karelian Isthmus, south of the present Vuoksi River, are described on the basis of sediment stratigraphical investigations. Two of the Lakes, Michurinskoe (94 m a.s.l.) and Uzornoe (55 m a.s.l.) are situated in an upland area that remained dry land after the deglaciation even during the early high water stages of the Baltic Sea (Baltic Ice Lake until c. 10 000 yr BP and Ancylus Lake 9500–8800 BP). The low-lying central parts of the Isthmus were flooded by the outflow of Lake Ladoga that took place across this area until the formation of Neva River, c. 3 100 yr BP, and further by the waters of River Vuoksi that started flowing into the area from the NW c. 5000 yr BP as a new outlet of Lake Saimaa. The basins of the lakes Krasnoe and Vishnevskoe (both 16 m a.s.l.) became isolated when River Neva was formed and Lake Ladoga sank to its present level. Lake Rakovoe (12 m a.s.l.) was on the level of River Vuoksi up until the mid-19th century, when water level in the central stretch of the river was artificially lowered. Each of the lakes has been variously affected by hydrological and climatic changes and consequences of human activities, e.g. eutrophication due to intensified land use. As a consequence to artificial lowering, done in order to gain field and meadowland, the large, shallow Lake Rakovoe has been largely overgrown by macrophytic vegetation.     

Major sedimentary cycles in the corallian beds (Oxfordian) of Southern England

The Corallian Beds of southern England consist of four asymmetric, upward shallowing cycles. Each cycle is separated from the next by a non-sequence, usually an erosion surface, which marks a marine transgression. Transgression was the result of a sudden rise in sea level, the period of rise being perhaps less than 7000 years. There is evidence of the same non-sequences occurring elsewhere in Britain and abroad, suggesting that the transgressions were the result of world-wide, eustatic changes in sea level.     

Forecasting shifts in habitat suitability of three marine predators suggests a rapid decline in inter‐specific overlap under future climate change

Understanding how environmental and climate change can alter habitat overlap of marine predators has great value for the management and conservation of marine ecosystems. Here, we estimated spatiotemporal changes in habitat suitability and inter‐specific overlap among three marine predators: Baltic gray seals (Halichoerus grypus), harbor seals (Phoca vitulina), and harbor porpoises (Phocoena phocoena) under contemporary and future conditions. Location data (>200 tagged individuals) were collected in the southwestern region of the Baltic Sea; one of the fastest‐warming semi‐enclosed seas in the world. We used the maximum entropy (MaxEnt) algorithm to estimate changes in total area size and overlap of species‐specific habitat suitability between 1997–2020 and 2091–2100. Predictor variables included environmental and climate‐sensitive oceanographic conditions in the area. Sea‐level rise, sea surface temperature, and salinity data were taken from representative concentration pathways [RCPs] scenarios 6.0 and 8.5 to forecast potential climate change effects. Model output suggested that habitat suitability of Baltic gray seals will decline over space and time, driven by changes in sea surface salinity and a loss of currently available haulout sites following sea‐level rise in the future. A similar, although weaker, effect was observed for harbor seals, while suitability of habitat for harbor porpoises was predicted to increase slightly over space and time. Inter‐specific overlap in highly suitable habitats was also predicted to increase slightly under RCP scenario 6.0 when compared to contemporary conditions, but to disappear under RCP scenario 8.5. Our study suggests that marine predators in the southwestern Baltic Sea may respond differently to future climatic conditions, leading to divergent shifts in habitat suitability that are likely to decrease inter‐specific overlap over time and space. We conclude that climate change can lead to a marked redistribution of area use by marine predators in the region, which may influence local food‐web dynamics and ecosystem functioning.     

Thiamin dynamics during the adult life cycle of Atlantic salmon (Salmo salar)

Thiamin is an essential water-soluble B vitamin known for its wide range of metabolic functions and antioxidant properties. Over the past decades, reproductive failures induced by thiamin deficiency have been observed in several salmonid species worldwide, but it is unclear why this micronutrient deficiency arises. Few studies have compared thiamin concentrations in systems of salmonid populations with or without documented thiamin deficiency. Moreover, it is not well known whether and how thiamin concentration changes during the marine feeding phase and the spawning migration. Therefore, samples of Atlantic salmon (Salmo salar) were collected when actively feeding in the open Baltic Sea, after the sea migration to natal rivers, after river migration, and during the spawning period. To compare populations of Baltic salmon with systems without documented thiamin deficiency, a population of landlocked salmon located in Lake Vänern (Sweden) was sampled as well as salmon from Norwegian rivers draining into the North Atlantic Ocean. Results showed the highest mean thiamin concentrations in Lake Vänern salmon, followed by North Atlantic, and the lowest in Baltic populations. Therefore, salmon in the Baltic Sea seem to be consistently more constrained by thiamin than those in other systems. Condition factor and body length had little to no effect on thiamin concentrations in all systems, suggesting that there is no relation between the body condition of salmon and thiamin deficiency. In our large spatiotemporal comparison of salmon populations, thiamin concentrations declined toward spawning in all studied systems, suggesting that the reduction in thiamin concentration arises as a natural consequence of starvation rather than to be related to thiamin deficiency in the system. These results suggest that factors affecting accumulation during the marine feeding phase are key for understanding the thiamin deficiency in salmonids.     

Impact of 21st century climate change on the Baltic Sea fish community and fisheries

The Baltic Sea is a large brackish semienclosed sea whose species-poor fish community supports important commercial and recreational fisheries. Both the fish species and the fisheries are strongly affected by climate variations. These climatic effects and the underlying mechanisms are briefly reviewed. We then use recent regional – scale climate – ocean modelling results to consider how climate change during this century will affect the fish community of the Baltic and fisheries management. Expected climate changes in northern Europe will likely affect both the temperature and salinity of the Baltic, causing it to become warmer and fresher. As an estuarine ecosystem with large horizontal and vertical salinity gradients, biodiversity will be particularly sensitive to changes in salinity which can be expected as a consequence of altered precipitation patterns. Marine-tolerant species will be disadvantaged and their distributions will partially contract from the Baltic Sea; habitats of freshwater species will likely expand. Although some new species can be expected to immigrate because of an expected increase in sea temperature, only a few of these species will be able to successfully colonize the Baltic because of its low salinity. Fishing fleets which presently target marine species (e.g. cod, herring, sprat, plaice, sole) in the Baltic will likely have to relocate to more marine areas or switch to other species which tolerate decreasing salinities. Fishery management thresholds that trigger reductions in fishing quotas or fishery closures to conserve local populations (e.g. cod, salmon) will have to be reassessed as the ecological basis on which existing thresholds have been established changes, and new thresholds will have to be developed for immigrant species. The Baltic situation illustrates some of the uncertainties and complexities associated with forecasting how fish populations, communities and industries dependent on an estuarine ecosystem might respond to future climate change.     

Quantifying an overlooked aspect of partial migration using otolith microchemistry

For the first time, an overlooked aspect of partial migration was quantified using otolith microchemistry and brown trout, Salmo trutta, as a model species. Relative contributions of freshwater resident and anadromous female brown trout to mixed-stock sea trout populations in the Baltic Sea were estimated. Out of 236 confirmed wild sea trout sampled around the coast of Estonia 88% were of anadromous maternal origin and 12% were of resident maternal origin. This novel finding underscores the importance of the resident contingent in maintaining the persistence and resilience of the migratory contingent.     

Climatic change and the Wadden Sea,The Netherlands

The predicted increase in atmospheric carbon dioxide and the effects of global warming will influence the Wadden Sea, The Netherlands, an area of exceptional ecological value. The direct effect of elevated atmospheric CO₂ on terrestrial coastal ecosystems is either marginal or unknown. The slight acidification of the sea which is predicted might have an impact on primary producers and juvenile animals. The effect of CO₂ fertilization on marine primary production remains to be elucidated. Profound changes will occur if sea level rises at the predicted rate of 60 cm per century, as sedimentation rates will be insufficient to maintain the salt marshes on the barrier islands. The marshes of the mainland coast will be impoverished, as high and low marshes are not expected to continue to coexist at the same locations. As sediment supply to the Wadden Sea is sufficient to compensate for sea level rise, the estuarine character of the Wadden Sea, with sand- and mudflats, is expected to remain largely unchanged.     

Experimental Evidence on Nutrient and Substrate Limitation of Baltic Sea sea-ice Algae and Bacteria

The effect of nutrient limitation on Baltic Sea ice algae, and substrate and nutrient limitation on ice bacteria, was studied in a series of in situ -experiments conducted during the winter of 2002 in northern Baltic Sea. Community level changes in algal biomass (chlorophyll a) and productivity, and bacterial thymidine and leucine incorporation were followed for one week after the addition of nutrient and/or organic carbon rich filtered seawater to the experimental units. The results showed the major contribution of snow cover to the algal responses during the beginning of the ice-covered season. Algal communities were able to grow even in January if no snow was present. Nutrient addition did occasionally have an effect on algal biomass and productivity in the ice. Surprisingly, seeding effect from the ice to the underlying water was negatively affected by the nutrient availability in March. Bacterial limitation varied between nutrient (phosphorus) and substrate limitations. The results showed, that limitation in both algal and bacterial communities changed periodically in the northern Baltic Sea ice.     

Geographical variation in Fucus vesiculosus morphology in the Baltic and North Seas

Effects of stress and disturbance on morphology, reproductive effort, size and sex ratio were studied for Fucus vesiculosus populations from the Baltic Sea at Askö and the North Sea on the west coast of Sweden at Tjäm[otilde]. High morphological variation was found between Fucus populations, with significant differences in length and weight of individuals, thallus breadth, number of branches and receptacles and receptacle weight, not only between Baltic and North Sea populations but also between populations within the same area, differing in wave exposure. With increasing disturbance, individuals in both studied populations were smaller and less branched. Differences were observed in plant size, with longer, broader and more branched plants being found in Askö compared with Tjärnö. Fucus populations at Tjämö allocated more biomass to reproduction and had longer, heavier receptacles than at Askö. Although the observed morphological changes may be partly explained by differences in wave exposure and salinity between the two sites, it is not possible to rule out genetic differences between the Baltic and North Sea populations. However, it is unlikely that the variations observed within the populations and between populations from the same area are genetically determined.     

Biostratigraphy and <Superscript>14</Superscript>C dating of a lake sediment sequence on the north-west Estonian carbonaceous plateau,interpreted in terms of human impact in the surroundings

The sediment stratigraphy of a medium-sized mixotrophic lake (Ruila) situated below the highest shoreline of the Baltic Ice Lake in the West-Estonian Lowland is described. The lake is without natural inlets our outlets. The reconstruction of vegetation and land-use history based on pollen data, combined with available archaeological data and detailed ¹⁴C dating allows us to give a provisional reconstruction of the temporal and spatial pattern of natural and human induced environmental changes in north-west Estonia during the Holocene. Both radiocarbon dates derived from terrestrial macrofossil dating by accelerator mass spectrometry (AMS) and conventional dating of bulk lake sediment are discussed. The isolation of the lake basin from the Yoldia Sea took place ca. 9700 cal B. C. The Ancylus Lake transgression at ca. 8400 cal B. C. did not reach the basin, but caused a ground water rise, seen in the sediment stratigraphy of the lake. The first signs of human impact on the pollen record appear ca. 5400 cal B. C. (Late Mesolithic). The history of arable farming has been divided into three periods: 1) introduction of crop cultivation and animal husbandry (1500 cal B. C. – A. D. 500); 2) establishment of animal husbandry A. D. 500–1000) and 3) establishment of crop cultivation and intensive cattle breeding (A. D. 1000–today). Due to unfavourable eda-phic conditions the introduction of arable farming was delayed for more than 1000 years compared with elsewhere on the north coast of Esotnia, and intensity of land-use never reached the same proportion as in these areas. Received August 15, 2001 / Accepted August 5, 2002 Correspondence to: Leili Saarse     

Letters to the editor

A decrease in salinity and temperature over the past 3000 years has presented the marine algae of the Baltic Sea with very considerable problems in adaptation. The effects of salinity upon a number of Baltic algae have been measured. The results showed cell mortality to be severe in 0, 68 and 102‰, and minimal in 6 and 11‰: there was most variation in tolerance to 34 and 51‰. The salt tolerances of Baltic marine algae have proved more hyposaline than those of British intertidal algae. Water uptake and loss in tissues of Chorda filum and Fucus vesiculosus from Baltic and British populations have been measured in response to salinity changes. The results revealed significant population differences in both live and killed tissues. Receptacle development and oogonial maturation have been observed in Baltic and British F. vesiculosus, and found to differ in seasonality. Some observations were associated with local sea temperatures but differences in the timing of receptacle initiation and in oogonial size were not. Th depauperate thallus, commonly ascribed to the effects of low salinity, was found to be a complicated phenomenon, comprising numerous attributes which are combined differently in different taxa. The morphological differences between Baltic and British marine algae were usually striking.

The marine algae of the Baltic Sea have therefore diverged in a number of ways from their N. Atlantic counterparts. The naturally high variability of these taxa has enabled them to survive the period of increasingly strong selection pressure which followed the Littorina Sea episode. Divergence seems not to have advanced to the point where speciation may be said to have occurred. The Baltic may therefore be contrasted with the much older Mediterranean Sea, which contains a large number of endemic species. Nevertheless, the Baltic is a site of very considerable evolutionary importance.     

Zoobenthic communities of the Dagestan Region of the Caspian Sea

The results of bottom community studies of the Dagestan Region of the Caspian Sea were presented; key factors affecting their formation were distinguished. Special attention was paid to restoration of the amphipod Pontogammarus maeoticus, which was characteristic in the coastal zone community until it was destroyed after a rise in the sea-level. It was established that biomass of the benthos was much higher in the period of the rise in the sea level than in the years of a low sea level.Original Russian Text Copyright © 2005 by Biologiya Morya, Guseinov.     

Genetic diversity of mitochondrial D-LOOP sequences in the spotted scat (Scatophagus argus) from different geographical populations along the northern coast of the South China Sea

The genetic diversity of 289 spotted scat (Scatophagus argus) from seven populations along the northern coast of the South China Sea was studied by analyzing the full-length sequences of the mitochondrial control region (D-LOOP). The S. argus D-LOOP sequence was 1,004–1,010 bp long and contained 156 variant sites. The seven studied S. argus populations had a high degree of genetic diversity (haplotype diversity [Hd] = 0.99135; nucleotide diversity (π) = 0.01313). There was no obvious genetic differentiation among the seven geographical populations and gene exchange was frequent (Fst = −0.01867–0.01117, p > .05). Four distinct mitochondrial lineages were identified in the phylogenetic tree and the haplotype network. The between-lineage Fst was 0.71690–0.84940 (p < .001), but these lineages showed no obvious phylogeographic pattern. Based on D-LOOP mutation rates, we estimated that the four lineages diverged approximately 513,800–93,600 years ago, during the Eocene ice age, at which time falling sea levels may have led to population segregation. We estimated that S. argus population expansion occurred approximately 2.29–0.68 million years ago, during the late Pleistocene. During this period, sea levels rose again, allowing previously separated lineages to come into sympatry, which eventually gave rise to a highly genetically diverse population without pyhlogeographic structure. Here, we characterized the genetic structure and differentiation of seven S. argus populations from the northern coast of the South China Sea. Our results suggested that the seven S. argus populations from the northern coast of the South China Sea have a relatively low level of genetic variation and can be considered a single unit for the purposes of fishery development, utilization, and management.     

An environmental gradient dominates ecological and genetic differentiation of marine invertebrates between the North and Baltic Sea

Environmental gradients have emerged as important barriers to structuring populations and species distributions. We set out to test whether the strong salinity gradient from the marine North Sea to the brackish Baltic Sea in northern Europe represents an ecological and genetic break, and to identify life history traits that correlate with the strength of this break. We accumulated mitochondrial cytochrome oxidase subunit 1 sequence data, and data on the distribution, salinity tolerance, and life history for 28 species belonging to the Cnidaria, Crustacea, Echinodermata, Mollusca, Polychaeta, and Gastrotricha. We included seven non‐native species covering a broad range of times since introduction, in order to gain insight into the pace of adaptation and differentiation. We calculated measures of genetic diversity and differentiation across the environmental gradient, coalescent times, and migration rates between North and Baltic Sea populations, and analyzed correlations between genetic and life history data. The majority of investigated species is either genetically differentiated and/or adapted to the lower salinity conditions of the Baltic Sea. Species exhibiting population structure have a range of patterns of genetic diversity in comparison with the North Sea, from lower in the Baltic Sea to higher in the Baltic Sea, or equally diverse in North and Baltic Sea. Two of the non‐native species showed signs of genetic differentiation, their times since introduction to the Baltic Sea being about 80 and >700 years, respectively. Our results indicate that the transition from North Sea to Baltic Sea represents a genetic and ecological break: The diversity of genetic patterns points toward independent trajectories in the Baltic compared with the North Sea, and ecological differences with regard to salinity tolerance are common. The North Sea–Baltic Sea region provides a unique setting to study evolutionary adaptation during colonization processes at different stages by jointly considering native and non‐native species.     

Temporal and spatial changes in benthic invertebrate trophic networks along a taxonomic richness gradient

Species interactions underlie most ecosystem functions and are important for understanding ecosystem changes. Representing one type of species interaction, trophic networks were constructed from biodiversity monitoring data and known trophic links to assess how ecosystems have changed over time. The Baltic Sea is subject to many anthropogenic pressures, and low species diversity makes it an ideal candidate for determining how pressures change food webs. In this study, we used benthic monitoring data for 20 years (1980–1989 and 2010–2019) from the Swedish coast of the Baltic Sea and Skagerrak to investigate changes in benthic invertebrate trophic interactions. We constructed food webs and calculated fundamental food web metrics evaluating network horizontal and vertical diversity, as well as stability that were compared over space and time. Our results show that the west coast of Sweden (Skagerrak) suffered a reduction in benthic invertebrate biodiversity by 32% between the 1980s and 2010s, and that the number of links, generality of predators, and vulnerability of prey have been significantly reduced. The other basins (Bothnian Sea, Baltic Proper, and Bornholm Basin) do not show any significant changes in species richness or consistent significant trends in any food web metrics investigated, demonstrating resilience at a lower species diversity. The decreased complexity of the Skagerrak food webs indicates vulnerability to further perturbations and pressures should be limited as much as possible to ensure continued ecosystem functions.     

Aquatic biodiversity under anthropogenic stress: an insight from the Archipelago Sea (SW Finland)

The Archipelago Sea in the northern Baltic has been subjected to large-scale cultural, economic and ecological changes, especially during the last three decades. Environmental threats originate from both basin-wide sources, affecting the whole Baltic Sea, and from local sources, such as nutrient loading from nearby river outflows, intense agriculture, fish farming, ships' traffic, boating, and man's physical impacts on the landscape and seascape. Both the Åland archipelago and the Archipelago Sea have been listed as hot-spots by HELCOM, Baltic Marine Environment Protection Commission, eutrophication being the main threat to the aquatic environment. In this study we review how biological communities have reacted to an increase in man-induced multisource stresses. Changes in plankton, benthic animals, macroalgal assemblages and fish communities have been documented in most parts of the Baltic Sea since the 1970s. What remains to be understood is the importance of these structural changes for the functioning of the Archipelago Sea ecosystem under various levels of human impact.     

Prorocentrum minimum (Dinophyceae) in the Baltic Sea: morphology, occurrence—a review

The potentially toxic dinoflagellate Prorocentrum minimum (Pavillard) Schiller has successfully established in the Baltic Sea in the last two decades. A review of the invasion history is presented as well as new data on the spatial and inter-annual variability of this species and its relation to salinity, temperature, and nutrient concentrations. A short literature review of the morphological characters of the Baltic P. minimum is also included.From 1993 to 2002, P. minimum was a regular component of the summer and autumn plankton flora of the Baltic Sea proper and the Gulf of Finland. Its abundance varied considerably inter-annually and did not show any clear trends during the period. Abundance of P. minimum was significantly higher in the nutrient-enriched Bay of Mecklenburg (German coast) and the southern Baltic proper than in the central and northern Baltic proper and the Gulf of Finland, where its abundance was mostly sparse. In coastal waters P. minimum occasionally reached densities of several million cells per litre and dominated phytoplankton biomass (>90%).Abundance of the Baltic P. minimum was generally not related to salinity or temperature. It could be a dominant species at both high and low salinity (over 15 and 4.8 PSU), and its temperature range was broad (from 2.7 to 26.4 °C). However, dense populations usually occurred from July to October at temperatures above 10 °C.Further, there appears to be a positive correlation between the success of P. minimum in the Baltic Sea and high concentrations of total phosphorus and nitrogen.This tolerant and morphologically variable dinoflagellate seems to be a morphospecies without subtaxa, which can expand its range in the Baltic Sea, especially in nutrient-rich coastal waters.