Emergence of brown tides caused by Aureococcus anophagefferens Hargraves et Sieburth in China

Large-scale blooms suspected to be “brown tides” occurred in early summer for three consecutive years from 2009 to 2011 in the coastal waters of Qinhuangdao, China, and had significant negative impacts on the shellfish mariculture industry. To identify the causative species of the blooms, phytoplankton samples were collected from regions with and without bloom in the coastal waters of Qinhuangdao in 2011, and clone libraries were built using eukaryote-specific 18S ribosomal RNA gene (18S rDNA). Altogether 50 clones, including 17 clones from bloom area and 33 clones from nearby regions without bloom were amplified. Blasted in GenBank, 17 clones amplified from the bloom area were assigned to Pelagophyceae (8 clones), Mediophyceae (2 clones), Cryptophyta (2 clones), Dinophyceae (2 clones) and unidentified eukaryotic species (3 clones). Those from the non-bloom site were assigned to Cryptophyta, Eustigmatophyceae, Prasinophyceae, Coscinodiscophyceae, Mediophyceae, Raphidophyceae and Dinophyceae, but not Pelagophyceae. All 8 pelagophyte clones from the bloom area were 99.7–100% similar to a single species, Aureococcus anophagefferens Hargraves et Sieburth, the causative species of brown tides on the east coast of USA. For nearly the entire length of the 18S rDNA, there were 0–6 base pair differences between the 8 amplicons and those of A. anophagefferens from USA. Furthermore, all of the 8 clones were clustered into the same well-supported clade with A. anophagefferens (posterior probability = 0.99) in a phylogenetic tree established for pelagophytes and other related microalgae. In our previous studies, the causative species of the bloom was tentatively identified as a pelagophyte, haptophyte or silicoflagellate, based on the pigment profile of the size-fractioned phytoplankton samples. Based on this study, we conclude that blooms in the coastal waters of Qinhuangdao of the Bohai Sea were brown tides caused by A. anophagefferens. As far as we know, this is the first report of brown tide events caused by A. anophagefferens in China, which is the third country in the world reporting A. anophagefferens blooms in addition to USA and South Africa.     

Spatial-temporal variation of Aureococcus anophagefferens blooms in relation to environmental factors in the coastal waters of Qinhuangdao,China

The brown tides occurring in the coastal scallop cultivation area of Qinhuangdao, China, in recent years are caused by Aureococcus anophagefferens and significantly impact the scallop industry and the marine ecosystem in this region. Long-term investigations of phytoplankton and hydrological variables in the Qinhuangdao sea area were conducted in this study to understand the spatial-temporal variations of A. anophagefferens in relation to environmental factors. Samples were collected during twelve cruises from July 2011 to December 2013 and were analyzed for the temperature, salinity, dissolved oxygen (DO), nutrients and phytoplankton pigments. All diagnostic pigments of A. anophagefferens, such as chlorophyll c₃ (Chl c₃), Chl c₂, 19′-butanoyloxyfucoxanthin (But-fuco), fucoxanthin (Fuco), and diadinoxanthin (Diad), were detected in the surface water by using high-performance liquid chromatography (HPLC). The highest concentrations of But-fuco (5.64 μg L⁻¹), Fuco (37.94 μg L⁻¹) and chlorophyll a (Chl a, 17.25 μg L⁻¹) occurred in different seasons and sampling sites. The A. anophagefferens bloom (as indicated by But-fuco) usually expanded from the south to the north of the Qinhuangdao sea area, close to scallop-culturing regions. The bloom unusually starts in May, reaches its peak in June and almost disappears in August, with the temperature ranging from ca. 19 °C to 23 °C. The redundancy analysis (RDA) indicated that relatively high salinity (>29) and low inorganic nutrients were suitable for the development of the A. anophagefferens bloom. The ratios of diagnostic pigments to Chl a were not constant during different cruises and generally obeyed two different linear relationships, thus indicating the co-occurrence of the blooms of A. anophagefferens and other species, such as Minutocellus polymorphus. In summary, our work reports the long-term variation of A. anophagefferens blooms based on diagnostic pigments and environmental controls, which may provide more insights into the formation mechanisms of the brown tide in this region.     

Interactions between planktonic microalgae and protozoan grazers

For an algal bloom to develop, the growth rate of the bloom-forming species must exceed the sum of all loss processes. Among these loss processes, grazing is generally believed to be one of the more important factors. Based on numerous field studies, it is now recognized that microzooplankton are dominant consumers of phytoplankton in both open ocean and coastal waters. Heterotrophic protists, a major component of microzooplankton communities, constitute a vast complex of diverse feeding strategies and behavior which allow them access to even the larger phytoplankton species. A number of laboratory studies have shown the capability of different protistan species to feed and grow on bloom-forming algal species. Because of short generation times, their ability for fast reaction to short-term variation in food conditions enables phagotrophic protists to fulfill the function of a heterotrophic buffer, which might balance the flow of matter in case of phytoplankton blooms. The importance of grazing as a control of microalgae becomes most apparent by its failure; if community grazing controls initial stages of bloom development, there simply is no bloom. However, if a certain algal species is difficult to graze, e.g. due to specific defense mechanisms, reduced grazing pressure will certainly favor bloom development. The present contribution will provide a general overview on the interactions between planktonic microalgae and protozoan grazers with special emphasis on species-specific interactions and algal defense strategies against protozoan grazers.     

3,000 km and 1,500‐year presence of Aureococcus anophagefferens reveals indigenous origin of brown tides in China

The nonmotile, spherical, picoplanktonic (2‐μm‐sized) pelagophyte Aureococcus anophagefferens has caused numerous harmful blooms (“brown tides”) across global marine ecosystems. Blooms have developed along the east coast of the USA since 1985, a limited number of times in South Africa around 1997, and frequently in China since 2009. As a consequence, the harmful blooms have caused massive losses in aquaculture and coastal ecosystems, particularly mortalities in cultured shellfish. Therefore, whether A. anophagefferens was recently introduced to China via natural/artificial transport of resting stage cells or has been an indigenous species has become a question of profound ecological significance and broad interest, which motivated our extensive investigation on the geographic and historical presence of this species in the seas of China. We applied a combined approach of extensive PCR‐based detection and sequencing, germination experiments and monoclonal antibody staining of germlings to samples of surface sediment and sediment core (dated via combined isotopic measurements) collected from all four seas of China, and searched the supplementary data set of a recent Science publication. We discovered that A. anophagefferens does have a resting stage in the sediment, but it also has a wide geographic distribution both in China (covering a range of ~30° in latitude, ~15.7° in longitude and 2.5–3,456 m in water depth; temperate to tropical and coastal to open oceans) and in almost all oceans of the world and a historical presence of >1,500 years in the Bohai Sea, China. The work revealed that A. anophagefferens is not a recently introduced, but an indigenous species in China and has in fact a globally cosmopolitan distribution.     

Assessment of brown tide blooms, caused by Aureococcus anophagefferens, and contributing factors in New Jersey coastal bays: 2000–2002

A 3 year study (2000–2002) in Barnegat Bay-Little Egg Harbor (BB/LEH), New Jersey (USA), was conducted by the New Jersey Department of Environmental Protection, Division of Science Research and Technology (DSRT) in cooperation with several partners to assess brown tide blooms in coastal waters in NJ. Water samples were collected by boat and helicopter at coastal stations from 2000 to 2002 along with field measurements. Aureococcus anophagefferens were enumerated and associated environmental factors were analyzed. A. anophagefferens abundances were classified using the Brown Tide Bloom Index and mapped, along with salinity and temperature parameters, to their geo-referenced location using the ArcView GIS. The highest A. anophagefferens abundances (>10⁶ cells ml⁻¹), including category 3 blooms (≥200,000 cells ml⁻¹) and category 2 blooms (≥35,000 to ≤200,000 cells ml⁻¹), recurred during each of the 3 years of sampling and covered significant geographic areas of the estuary, especially in Little Egg Harbor. While category 3 blooms were generally associated with warmer water temperatures (>16 °C) and higher salinity (>25–26 ppt), these factors were not sufficient alone to explain the timing or distribution of A. anophagefferens blooms. There was no significant relationship between brown tide abundances and dissolved organic nitrogen measured in 2002 but this was consistent with other studies. Extended drought conditions, with corresponding low freshwater inputs and elevated bay water salinities, occurring during this time were conducive to blooms. A. anophagefferens abundances were well above the reported levels that have been reported to cause negative impacts on shellfish. It was shown that over 50% of the submerged aquatic vegetation (SAV) habitat located in Barnegat Bay/Little Egg Harbor was categorized as having a high frequency of category 2 or 3 blooms for all 3 years.     

Roles of mixotrophy in blooms of different dinoflagellates: Implications from the growth experiment

Studies over the last two decades suggested that mixotrophy could be an important adaptive strategy for some bloom-forming dinoflagellates. In the coastal waters adjacent to the Changjiang River estuary in the East China Sea, recurrent blooms of dinoflagellates Prorocentrum donghaiense, Karenia mikimotoi and Alexandrium catenella started to appear from the beginning of the 21 century, but roles of mixotrophy in the formation of dinoflagellate blooms were not well understood. In the current study, mixotrophy-based growth of four selected bloom-causative dinoflagellate species, i.e. K. mikimotoi, A. catenella, P. donghaiense and Prorocentrum micans, were studied. Dinoflagellates were co-cultured with different prey organisms, including bacterium Marinobacter sp., microalgae Isochrysis galbana and Hemiselmis virescens, under a variant of nutrient conditions. It was found that growth of dinoflagellate K. mikimotoi was significantly promoted with the presence of prey organisms. Growth of P. donghaiense and P. micans was only slightly improved. For A. catenella, the addition of prey organisms has no effects on the growth, while both of the two prey microalgae I. galbana and H. virescens were killed, probably by allelochemicals released from A. catenella. There was no apparent relationship between nutrient conditions and the mixotrophy-based growth of the tested dinoflagellates. Based on the results of the growth experiment, it is implicated that mixotrophy may play different roles in the growth and bloom of the four dinoflagellate species. It can be an important competitive strategy for K. mikimotoi. For the two Prorocentrum species and A. catenella, however, the role of mixotrophy is much limited. They may depend more on other competitive strategies, such as phototrophy-based growth and allelopathic effect, to prevail in the phytoplankton community and form blooms.     

Host-Specificity and Dynamics in Bacterial Communities Associated with Bloom-Forming Freshwater Phytoplankton

Many freshwater phytoplankton species have the potential to form transient nuisance blooms that affect water quality and other aquatic biota. Heterotrophic bacteria can influence such blooms via nutrient regeneration but also via antagonism and other biotic interactions. We studied the composition of bacterial communities associated with three bloom-forming freshwater phytoplankton species, the diatom Aulacoseira granulata and the cyanobacteria Microcystis aeruginosa and Cylindrospermopsis raciborskii. Experimental cultures incubated with and without lake bacteria were sampled in three different growth phases and bacterial community composition was assessed by 454-Pyrosequencing of 16S rRNA gene amplicons. Betaproteobacteria were dominant in all cultures inoculated with lake bacteria, but decreased during the experiment. In contrast, Alphaproteobacteria, which made up the second most abundant class of bacteria, increased overall during the course of the experiment. Other bacterial classes responded in contrasting ways to the experimental incubations causing significantly different bacterial communities to develop in response to host phytoplankton species, growth phase and between attached and free-living fractions. Differences in bacterial community composition between cyanobacteria and diatom cultures were greater than between the two cyanobacteria. Despite the significance, major differences between phytoplankton cultures were in the proportion of the OTUs rather than in the absence or presence of specific taxa. Different phytoplankton species favoring different bacterial communities may have important consequences for the fate of organic matter in systems where these bloom forming species occur. The dynamics and development of transient blooms may also be affected as bacterial communities seem to influence phytoplankton species growth in contrasting ways.     

Applicability of Hydrogen Peroxide in Brown Tide Control – Culture and Microcosm Studies

Brown tide algal blooms, caused by the excessive growth of Aureococcus anophagefferens, recur in several northeastern US coastal bays. Direct bloom control could alleviate the ecological and economic damage associated with bloom outbreak. This paper explored the effectiveness and safety of natural chemical biocide hydrogen peroxide (H₂O₂) for brown tide bloom control. Culture studies showed that H₂O₂ at 1.6 mg L⁻¹ effectively eradicated high density A. anophagefferens within 24-hr, but caused no significant growth inhibition in the diatoms, prymnesiophytes, green algae and dinoflagellates of >2–3 μm cell sizes among 12 phytoplankton species tested over 1-week observation. When applied to brown tide bloom prone natural seawater in a microcosm study, this treatment effectively removed the developing brown tide bloom, while the rest of phytoplankton assemblage (quantified via HPLC based marker pigment analyses), particularly the diatoms and green algae, experienced only transient suppression then recovered with total chlorophyll a exceeding that in the controls within 72-hr; cyanobacteria was not eradicated but was still reduced about 50% at 72-hr, as compared to the controls. The action of H₂O₂ against phytoplankton as a function of cell size and cell wall structure, and a realistic scenario of H₂O₂ application were discussed.     

Ecology of phytoplankton communities dominated by Aureococcus anophagefferens: the role of viruses, nutrients, and microzooplankton grazing

We investigated the impact of viruses, nutrient loading, and microzooplankon grazing on phytoplankton communities in two New York estuaries that hosted blooms of the brown tide alga Aureococcus anophagefferens during 2000 and 2002. The absence of a bloom at one location during 2002 allowed for the fortuitous comparison of a bloom and non-bloom year at the same location as well as a comparison of two sites experiencing bloom and non-bloom conditions during the same year. During the study, blooms were found at locations with high levels of dissolved organic nitrogen and lower nitrate concentrations compared to a non-bloom location. Experimental additions of inorganic nitrogen and phosphorus yielded growth rates within the total phytoplankton community which significantly exceeded control treatments in 83% of experiments, while A. anophagefferens experienced significantly increased growth during only 20% of experimental inorganic nutrient additions. Consistent with prior research, these results suggest brown tides are not caused by eutrophication, but instead are more likely to occur when sources of labile DOM are readily available. Microzooplankton grazing rates on the total phytoplankton community during a bloom were lower than grazing rates at a non-bloom site, and grazing rates on A. anophagefferens were lower than grazing rates on the total community on some dates, suggesting that reduced grazing mortality may also promote brown tides. Mean densities of viruses during blooms (3 × 10⁸ ml⁻¹) were elevated compared to most estuarine environments and were twice the levels found at a non-bloom site. Experimental enrichment of the natural viral densities yielded a significant increase in A. anophagefferens growth rates relative to control treatments when background levels of viruses were low (<1.7 × 10⁸ ml⁻¹), suggesting that viruses may promote bloom occurrence by regenerating DOM or altering the composition of microbial communities.     

Expansion risk of the toxic dinoflagellate Gymnodinium catenatum blooms in Chinese waters under climate change

The paralytic shellfish poison toxin (PST)-producing dinoflagellate, Gymnodinium catenatum, frequently blooms in China, posing a threat to food safety and human health. To understand the drivers of G. catenatum blooms and predict potential habitats for G. catenatum under climate change, samples from occurrence localities and environmental datasets from multiple agencies were aggregated and used to model the habitat suitability of G. catenatum in the China Sea using a maximum entropy model (Maxent). The accumulated variable contributions for the Maxent model were defined to measure the importance of key predictors in the model. The most important environmental variables were distance to the coastline, depth of seawater, and long-term average of the minimum annual temperature. This highlights the main reasons why G. catenatum blooms always occur in coastal waters. Occurrence probabilities higher than 0.66 were defined as habitats with high suitability for shellfish management and aquaculture. Projected habitats with high suitability in Haizhou Bay, coastal waters along the western Taiwan Strait, and Bohai Bay remained stable with increasing temperature by 2100, regardless of the IPCC Representative Concentration Pathways (RCPs). However, those in the China Sea would be reduced overall, leading to a northward movement of the center of integrated habitats. Habitats with a spatial area of >6000 km² in the Bohai Sea, Yellow Sea, and South China Sea and >23,000 km² in the East China Sea would be exposed to high risk under low greenhouse gas emission scenarios (RCP2.6).     

Temporal variations in phytoplankton, particulates, and colored dissolved organic material based on optical properties during a Long Island brown tide compared to an adjacent embayment

Since 1985, the coastal embayments of Long Island, New York, have been plagued with recurrent blooms, aptly called brown tides, of the pelagophyte Aureococcus anophagefferens. The distinct ocean color observed during these blooms suggests that optical methods can be used as a tool to study, detect, and track brown tides. Thus, the goal of our project was to compare the optical properties and pigment composition during bloom and non-bloom conditions and assess temporal variations in the phytoplankton and other constituents in the seawater associated with bloom development. From 17 May to 8 June 2000, we measured a time series of particle size distributions and concentrations as well as size-fractioned algal pigments and optical properties in two Long Island embayments where brown tides are known to occur. During our study, A. anophagefferens represented an insignificant contribution to the algal community in West Neck Bay (WNB), whereas a bloom developed in Quantuck Bay (QB). Initially, temperature and salinity were similar at the two locations; however, bulk optical properties, chlorophyll, and particle concentrations were nearly a factor of 2 greater at QB. Bulk optical properties remained constant at WNB, yet increased exponentially at QB as the bloom developed. The composition of particulates, including phytoplankton, varied little at QB, and the optical properties suggested the dominance of A. anophagefferens (confirmed by microscopy). The largest temporal variations were observed in the colored dissolved organic material (CDOM); the colloidal (0.2–0.7 μm) fraction, exhibiting a strong protein-like signal, increased dramatically at the height of the bloom. At WNB particle sizes and algal composition varied despite the invariant bulk optical properties; CDOM variations were minimal. Overall, the optical properties in the two bays demonstrated that at QB temporal variations were dominated by biomass and colloidal protein changes, whereas shifts in the algal community occurred at WNB. This study demonstrates the utility of in situ optical observations to resolve temporal changes in the ecological conditions associated with algal bloom development.     

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.     

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.     

Can Humic Water Discharge Counteract Eutrophication in Coastal Waters?

A common and established view is that increased inputs of nutrients to the sea, for example via river flooding, will cause eutrophication and phytoplankton blooms in coastal areas. We here show that this concept may be questioned in certain scenarios. Climate change has been predicted to cause increased inflow of freshwater to coastal areas in northern Europe. River waters in these areas are often brown from the presence of high concentrations of allochthonous dissolved organic carbon (humic carbon), in addition to nitrogen and phosphorus. In this study we investigated whether increased inputs of humic carbon can change the structure and production of the pelagic food web in the recipient seawater. In a mesocosm experiment unfiltered seawater from the northern Baltic Sea was fertilized with inorganic nutrients and humic carbon (CNP), and only with inorganic nutrients (NP). The system responded differently to the humic carbon addition. In NP treatments bacterial, phytoplankton and zooplankton production increased and the systems turned net autotrophic, whereas the CNP-treatment only bacterial and zooplankton production increased driving the system to net heterotrophy. The size-structure of the food web showed large variations in the different treatments. In the enriched NP treatments the phytoplankton community was dominated by filamentous >20 µm algae, while in the CNP treatments the phytoplankton was dominated by picocyanobacteria <5 µm. Our results suggest that climate change scenarios, resulting in increased humic-rich river inflow, may counteract eutrophication in coastal waters, leading to a promotion of the microbial food web and other heterotrophic organisms, driving the recipient coastal waters to net-heterotrophy.     

Are changes in N:P ratios in coastal waters the key to increased red tide blooms?

There is mounting evidence of a global increase in nutrient levels of coastal waters through riverine and sewage inputs, and in both the numbers and frequency(as well as the species composition) of red tides. However, it is still not possible to conclude the extent to which the increase in red tides in coastal waters can be attributed to the increase in nutrient levels, since so many other factors are involved. Undoubtedly, a relationship exists between red tides and the N and P load of coastal waters, and many nutrient enrichment experiments have shown that marine phytoplankton blooms are often nutrient limited. What is now becoming clear, however, is that although in classical Liebigian terms minimum amounts can be limiting, nutrient ratios (such as N:P and Si:P) are far more important regulators. This paper reviews evidence collected by the authors from Tolo Harbour, Hong Kong together with data collected in Japanese and North European coastal waters by various authors, which indicates that both long term and relatively short term changes in the N:P ratio are accompanied by increased blooms of non-siliceous phytoplankton groups and, furthermore,that the growth of most red tide causative organisms in Hong Kong coastal water is optimized at a low N:P(atomic) ratio of between 6 and 15. This revised version was published online in August 2006 with corrections to the Cover Date.     

HPLC pigment records provide evidence of past blooms of Aureococcus anophagefferens in the Coastal Bays of Maryland and Virginia, USA

Concentrations of the accessory phytoplankton pigment 19′-butanoyloxyfucoxanthin (but-fuco), derived from archived high performance liquid chromatography (HPLC) data from the Atlantic coastal bays of Maryland and Virginia (1993–1995 and 1999–2002), were used to determine the presence of Aureococcus anophagefferens at 18 stations. Paired data of direct cell counts of A. anophagefferens and but-fuco concentration data from 2000 to 2002 were linearly regressed (R² = 0.78). This regression was used to estimate historical cell densities from 1994 to 1995 and to improve the spatial resolution from 1999 to 2002. Although the HPLC method used did not permit quantification of but-fuco before 1994, the records indicate that qualitatively A. anophagefferens was present in 1993. Quantitative measurements grouped into bloom index categories showed that annually, peak densities occurred in May to July. Severe Category 3 blooms (>200,000 cells ml⁻¹) were found in 1995, 2001, and 2002. Spatially, concentrations of but-fuco were higher in the northern extent of the study region than in the lower Chincoteague Bay, and along the western shore of Chincoteague Bay than on the eastern side. On an interannual basis, it appeared that A. anophagefferens became more geographically widespread and blooms more intense throughout the study period.     

Changes in community structure and photosynthetic activities of total phytoplankton species during the growth,maintenance, and dissipation phases of a Prorocentrum donghaiense bloom

The potential interactions between the bloom-forming dinoflagellates and other phytoplankton during the algal bloom cycle are interesting, while the causes for the phytoplankton community changes were not fully understood. We hypothesized that phytoplankton community structure and photosynthetic activities of total phytoplankton have their special characteristics in different phases of the algal blooms. To test this hypothesis, a survey covering the process of a Prorocentrum donghaiense bloom in coastal waters between Dongtou and Nanji Islands was carried out from 9 to 20 May 2016, and the changes in the phytoplankton community and photosynthetic activities of total phytoplankton were determined. Surface seawater was sampled for microscopic analysis of phytoplankton composition and pulse amplitude modulated (PAM) chlorophyll fluorescence analysis of photosynthetic activities of the total phytoplankton species. A total of 25, 31, and 19 phytoplankton species were identified in its growth (9–12 May), maintenance (13–18 May) and dissipation phases (19–20 May), respectively. Diatoms were dominant in terms of species number while dinoflagellates were predominant at cell abundance. Dinoflagellates were the major dominant species during three phases of the bloom based on the dominance (Y) value, whereas the dominant species extended to dinoflagellates and diatoms including P. donghaiense, Coscinodiscus argus, Gonyaulax spinifera, Cyclotella sp. and Scrippsiella trochoidea in the dissipation phase. In the maintenance phase, the average cell abundances of the total phytoplankton and P. donghaiense were consistent with the chlorophyll a (Chla) concentration in the seawater; for the diversity indices of total phytoplankton species, Simpson index (C) was the highest while Shannon index (H′) and Pielou evenness index (J′) were the lowest. Furthermore, photosynthetic activities of the total phytoplankton species represented by the effective quantum yield (F_q'/F_m') and the maximum relative electron transport rate (rETR_max) in the maintenance phase were higher than those in the growth and dissipation phases. The results indicated that the characteristics of phytoplankton community structure and photosynthetic activities could be regarded as criteria in predicting the phases of algal blooms.     

Comparative composition and dynamics of harmful dinoflagellates in Mediterranean salt marshes and nearby external marine waters

The taxonomic structure of phytoplankton populations in two Mediterranean coastal lagoons were compared with those of nearby marine waters (external waters). Mediterranean confined lagoons remain isolated for most the year and concentrate phytoplankton to a very high biomass. Coastal lagoons on the Mediterranean may, therefore, act as accumulators of neritic phytoplankton (including species related to harmful algal blooms). We examined whether coastal lagoons act as concentrators of marine toxic dinoflagellates during confinement periods, and the common environmental factors that favour growth of specific harmful species in the two ecosystems considered: coastal lagoons and external waters. An alternation between the dominance of diatoms and dinoflagellates was observed, coinciding with that described in Margalef's mandala, occurring in external waters as well as in coastal lagoons. Moreover, the temporal patter was different in the two ecosystems. Dinoflagellate species composition and their bloom period were highly variable in time and space, thus, species had to be analysed individually. Most of the dinoflagellate species found in this study were potentially harmful and high biomass producers. Harmful dinoflagellate species performed well in both, external waters and lagoons, but the specific species-dependent affinity to each of these environments determined which organisms bloom there. Thus, expansion of harmful algal blooms (HAB) to inland waters is not likely and some environmental factors such as the oxidised state of available nitrogen, became determinant to the success and bloom of a species in the coastal lagoon ecosystem.     

Flora of planktonic microalgae of Amursky Bay, Sea of Japan

Long-term studies of the species composition of phytoplankton in Amursky Bay (Sea of Japan) carried out during the period from 1991 to 2006 revealed a total of 357 species of planktonic microalgae from eight divisions: Cyanophyta (8 species), Chrysophyta (8), Bacillariophyta (157), Cryptophyta (5), Dinophyta (143), Raphidophyta (3), Euglenophyta (11), and Chlorophyta (22 species). An annotated checklist of species is presented. In comparison with the late 1960s and early 1970s, the species richness of phytoplankton increased markedly; a greater number of bloom-forming species was recorded.