Wide ecological niches ensure frequent harmful dinoflagellate blooms.

Harmful algal blooms (HABs) and their consequences cause multiple devastating effects in various freshwater, brackish and marine ecosystems. However, HAB species at moderate population densities have positive ecological roles as primary producers of organic matter and food for zooplankton and fish. They also enhance benthic-pelagic coupling and participate in the biogeochemical cycles. The consequences of HABs are transported across the conventional environmental boundaries by numerous cascade effects in the food webs and beyond. Meanwhile, forecasts of bloom events are still limited, largely because of scarcity of reliable information on ecological niches of the bloom-forming algae. To fill up this knowledge gap, this study focused on dinoflagellates, a diverse group of mostly photosynthesizing protists (unicellular eukaryotes) capable of mixotrophy, since they play a key role in primary production and formation of blooms in marine and brackish waters worldwide. In this study, ecological niches of 17 abundant bloom-forming dinoflagellate species from coastal regions of the southern Baltic Sea were identified for the first time. It was hypothesized that wider ecological niches ensure more frequent dinoflagellate blooms compared to the species with narrower niches. This hypothesis was verified using the long-term (44 years) database on phytoplankton abundance and physical-chemical characteristics of the environment. It were analyzed 4534 datasets collected from 1972 to 2016. Fourteen abiotic parameters (water temperature, salinity, Secchi depth, pH, Chl a, and concentration of basic nutrients) were considered as ecological niche dimensions. The Principal Component Analysis presented the dissolved inorganic nitrogen, total nitrogen, Chl a, and temperature as principal niche dimensions of dinoflagellates. The algal bloom criteria were refined. It was for the first time proved statistically that HAB frequency of dinoflagellate species robustly correlated with the width of their ecological niches.

Chaos theory discloses triggers and drivers of plankton dynamics in stable environment.

Despite the enticing discoveries of chaos in nature, triggers and drivers of this phenomenon remain a classical enigma which needs irrefutable empirical evidence. Here we analyze results of the yearlong replicated mesocosm experiment with multi-species plankton community that allowed revealing signs of chaos at different trophic levels in strictly controlled abiotic environment. In mesocosms without external stressors, we observed the "paradox of chaos" when biotic interactions (internal drivers) were acting as generators of internal abiotic triggers of complex plankton dynamics. Chaos was registered as episodes that vanished unpredictably or were substituted by complex behaviour of other candidates when longer time series were considered. Remarkably, episodes of chaos were detected even in the most abiotically stable conditions. We developed the Integral Chaos Indicator to validate the results of the Lyapunov exponent analysis. These findings are essential for modelling and forecasting behaviour of a variety of natural and other global systems.

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.

Macroalgal diversity along the Baltic Sea salinity gradient challenges Remane's species-minimum concept.

Remane's species-minimum concept, which states that the lowest number of taxa occurs at the horohalinicum (5-8psu), was tested by investigating macroalgal diversity on hard substrates along the natural salinity gradient in the Baltic Sea. Field data on species occurrence and abundance were collected by SCUBA diving along 10 transects of the Finnish, Swedish and German coasts, covering a salinity range from 3.9 to 27psu. Macroalgal species numbers declined steadily with salinity, decreasing until 7.2psu was reached, but in the horohalinicum, a marked reduction of species number and a change in diversity were indicated by the Shannon index and evenness values. The non-linear decrease in macroalgal diversity at 5-8psu and the lack of increase in species numbers at salinities below 5psu imply a restricted applicability of Remane's species-minimum concept to macroalgae.

Status of biodiversity in the Baltic Sea.

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.

Principal processes within the estuarine salinity gradient: a review.

The salinity gradient is one of the main features characteristic of any estuarine ecosystem. Within this gradient in a critical salinity range of 5-8 PSU the major biotic and abiotic processes demonstrate non-linear dynamics of change in rates and directions. In estuaries, this salinity range acts as both external ecological factor and physiological characteristics of internal environment of aquatic organisms; it divides living conditions appropriate for freshwater and marine faunas, separates invertebrate communities with different osmotic regulation types, and defines the distribution range of high taxa. In this paper, the non-linearity of biotic processes within the estuarine salinity gradient is illustrated by the data on zooplankton from the Baltic estuaries. The non-tidal Baltic Sea provides a good demonstration of the above phenomena due to gradual changes of environmental factors and relatively stable isohalines. The non-linearity concept coupled with the ecosystem approach served the basis for a new definition of an estuary proposed by the authors.

Plankton of the Baltic estuarine ecosystems with emphasis on Neva Estuary: a review of present knowledge and research perspectives.

Different levels of eutrophication and pollution in the Baltic coastal waters in general, and in estuaries particularly define variations in the buffering capacity of ecosystems of these important filter zones between rivers and the adjacent Baltic Proper. Phyto- and zooplankton in the estuaries form important components of the pelagic food webs that participate in producing and structuring the matter, energy, and information fluxes in the ecosystems. Due to salinity gradients, estuaries provide a large variety of aquatic habitats for native populations of marine, brackish and fresh water plankton species. Non-indigenous planktonic species that expand their bio-geographical range and invade the Baltic coastal ecosystems also find favourable environments, establish permanent populations there, and impact aboriginal ecosystems. The dynamics of abundance, species diversity, population structure, productivity of planktonic communities, and peculiarities of trophic interactions in different Baltic estuaries are influenced by a number of common environmental factors. Being measured on the unified basis and monitored, changes (or stability) of structural and functional parameters of plankton communities can serve as indicators of alterations to the ecosystems. Comparative analyses and quantitative estimates of long-term changes in phyto- and zooplankton can contribute to evaluation of functional response of Baltic coastal ecosystems to anthropogenic stress. Both theoretical (modeling) and empirical studies of interactions within plankton communities aimed at the assessment of the role of planktonic organisms in water quality regulation, and applicability of these data for the evaluation of the status of ecosystems and their management can be considered as major goals for the future investigations in Baltic estuaries.