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Special issue Marine Benthic Harmful Algal Blooms.-- 11 pages, 5 figures, supplementary materials https://doi.org/10.1016/j.hal.2022.102352.-- Data availability: Data will be made available on request, A new chytrid genus and species was isolated and cultured from samples obtained in the Baltic Sea during a dinoflagellate bloom event. This species is characterized by having a spherical sporangium without papillae and zoospores of 2–3 µm in diameter that are released through 3 discharge pores. Molecular phylogeny based on ribosomal operon showed its sister position to the Dinomyces cluster in Rhizophydiales. Zoospores lack fenestrated cisternae but contain a paracrystalline inclusion, found in a Rhizophydiales representative for the first time. Additionally, the kinetid features are uncommon for Rhizophydiales and only observed in Dinomyces representatives so far. These morphological features and its phylogenetic relationships justify the description of the new genus and speciesParadinomyces triforaminorum gen. nov. sp. nov. belonging to the family Dinomycetaceae. The chytrid was detected during a high-biomass bloom of the dinoflagellate Kryptoperidinium foliaceum. Laboratory experiments suggest this species is highly specific and demonstrate the impact it can have on HAB development. The chytrid co-occurred with three other parasites belonging to Chytridiomycota (Fungi) and Perkinsea (Alveolata), highlighting that parasitic interactions are common during HABs in brackish and marine systems, and these multiple parasites compete for similar hosts, AR and EG were funded by the Spanish MICINN Project SMART PID2020-112978GB-I00. This work was partly supported by the Russian Science Foundation grant no. 21–74–20089 (writing, microscopy analysis), the infrastructure of SAK's laboratory was supported by ZIN RAS program 1021051402849–1, and the C7 strain was incorporated in CCPP ZIN RAS collection supported by a grant 075–15–2021–1069 of the Ministry of Science and Higher Education of the Russian Federation. AK was supported by the Academy of Finland, grant 310,449. MK was supported by JSPS KAKENHI 19H05667. EA was supported by The Royal Society (UK) under the Newton International Fellowship (grant number NF170346, call 2017), With the institutional support of the ‘Severo Ochoa Centre of Excellence’ accreditation (CEX2019-000928-S)

In recent years, blooms of toxic Alexandrium ostenfeldii strains have been reported from around the world. In 2013, the species formed a red tide in a shallow lagoon in western Japan, which was the first report of the species in the area. To investigate the genetic relatedness of Japanese A. ostenfeldii and global isolates, the full-length SSU, ITS and LSU sequences were determined, and phylogenetic analyses were conducted for isolates from western and northern Japan and from the Baltic Sea. Genotyping and microsatellite sequence comparison were performed to estimate the divergence and connectivity between the populations from western Japan and the Baltic Sea. In all phylogenetic analyses, the isolates from western Japan grouped together with global isolates from shallow and low saline areas, such as the Baltic Sea, estuaries on the east coast of U.S.A. and from the Bohai Sea, China. In contrast, the isolates from northern Japan formed a well-supported separate group in the ITS and LSU phylogenies, indicating differentiation between the Japanese populations. This was further supported by the notable differentiation between the sequences of western and northern Japanese isolates, whereas the lowest differentiation was found between the western Japanese and Chinese isolates. Microsatellite genotyping revealed low genetic diversity in the western Japanese population, possibly explained by a recent introduction to the lagoon from where it was detected. The red tide recorded in the shallow lagoon followed notable changes in the salinity of the waterbody and phytoplankton composition, potentially facilitating the bloom of A. ostenfeldii.

The paralytic shellfish toxin (PST) producing dinoflagellate Alexandrium ostenfeldii forms dense, recurrent blooms during summer in shallow coastal areas of the Baltic Sea. We studied the intra-population variability of its allelochemical potency and the responses of co-occurring and potentially competing dinoflagellates to the allelochemicals. The lytic activity of 10 northern Baltic A. ostenfeldii strains was evaluated by their EC 50 values (i.e. the cell concentration yielding a 50% decline in cryptophyte density), which were found to vary between 236 and 1726 cells ml −1 . When co-occurring dinoflagellates ( Kryptoperidinium foliaceum , Levanderina fissa and Heterocapsa triquetra ) were exposed to filtrate of A. ostenfeldii , short-term ( K. foliaceum cells formed cysts, L. fissa cells lost their cell shape and lysed, whereas H. triquetra cells shed their thecae. After 24 h, K. foliaceum had returned into vegetative cells and the number of immotile L. fissa and H. triquetra cells had significantly decreased. The results indicate that A. ostenfeldii can disturb the growth of competing dinoflagellates by excreting allelochemicals at bloom concentrations and that co-occurring species may develop efficient means to escape and recover from the allelochemicals, allowing them to coexist with A. ostenfeldii .

Dinoflagellates of the Alexandrium ostenfeldii complex (A. ostenfeldii, A. peruvianum) are capable of producing different types of neurotoxins: paralytic shellfish toxins (PSTs), spirolides and gymnodimines, depending on the strain and its geographic origin. While Atlantic and Mediterranean strains have been reported to produce spirolides, strains originating from the brackish Baltic Sea produce PSTs. Some North Sea, USA and New Zealand strains contain both toxins. Causes for such intraspecific variability in toxin production are unknown. We investigated whether salinity affects toxin production and growth rate of 5 A. ostenfeldii/peruvianum strains with brackish water (Baltic Sea) or oceanic (NE Atlantic) origin. The strains were grown until stationary phase at 7 salinities (6–35), and their growth and toxin production was monitored. Presence of saxitoxin (STX) genes (sxtA1 and sxtA4 motifs) in each strain was also analyzed. Salinity significantly affected both growth rate and toxicity of the individual strains but did not change their major toxin profile. The two Baltic Sea strains exhibited growth at salinities 6–25 and consistently produced gonyautoxin (GTX) 2, GTX3 and STX. The two North Sea strains grew at salinities 20–35 and produced mainly 20-methyl spirolide G (20mG), whereas the strain originating from the northern coast of Ireland was able to grow at salinities 15–35, only producing 13-desmethyl spirolide C (13dmC). The effects of salinity on total cellular toxin concentration and distribution of toxin analogs were strain-specific. Both saxitoxin gene motifs were present in the Baltic Sea strains, whereas the 2 North Sea strains lacked sxtA4, and the Irish strain lacked both motifs. Thus sxtA4 only seems to be specific for PST producing strains. The results show that toxin profiles of A. ostenfeldii/peruvianum strains are predetermined and the production of either spirolides or PSTs cannot be induced by salinity changes. However, changes in salinity may lead to changed growth rates, total cellular toxin concentrations as well as relative distribution of the different PST and spirolide analogs, thus affecting the actual toxicity of A. ostenfeldii/peruvianum populations.

Many marine dinoflagellates, including several species of the genus Alexandrium, Gymnodinium catenatum, and Pyrodinium bahamense are known for their capability to produce paralytic shellfish toxins (PST), which can cause severe, most often food-related poisoning. The recent discovery of the first PST biosynthesis genes has laid the foundation for the development of molecular detection methods for monitoring and study of PST-producing dinoflagellates. In this study, a probe-based qPCR method for the detection and quantification of the sxtA4 gene present in Alexandrium spp. and Gymnodinium catenatum was designed. The focus was on Alexandrium ostenfeldii, a species which recurrently forms dense toxic blooms in areas within the Baltic Sea. A consistent, positive correlation between the presence of sxtA4 and PST biosynthesis was observed, and the species was found to maintain PST production with an average of 6 genomic copies of sxtA4. In August 2014, A. ostenfeldii populations were studied for cell densities, PST production, as well as sxtA4 and species-specific LSU copy numbers in Foglo, Aland, Finland, where an exceptionally dense bloom, consisting of 6.3×106cellsL-1, was observed. Cell concentrations, and copy numbers of both of the target genes were positively correlated with total STX, GTX2, and GTX3 concentrations in the environment, the cell density predicting toxin concentrations with the best accuracy (Spearman's ρ=0.93, p

A B S T R A C T In the past years, some coastal areas in the central and northern Baltic Sea have experienced recurrent blooms of the potentially toxic dinoflagellate Alexandrium ostenfeldii. The population dynamics and spatial distribution of the species were studied in the Fogloarchipelago, a bloom area in Aland, northern Baltic Sea. During a two-year survey, cell and toxin concentrations of A. ostenfeldii were recorded in summer phytoplankton communities and related to environmental parameters. The studies revealed that A. ostenfeldii blooms were restricted to a small area in a narrow sound, where cells of this species were detected from May to September in 2009 and 2010. A. ostenfeldii proliferated to bloom concentrations (1.7-2.1 � 10 5 cells L � 1 ) only at water temperatures around 20 8C during warm periods in July and August. Abundance of A. ostenfeldii did not significantly correlate with dissolved inorganic nutrient concentrations (p = 0.4-0.6) but was related to high concentrations of resting cysts in the sediment (p < 0.001). The blooms were not monospecific but A. ostenfeldii was a part of a diverse productive phytoplankton community forming 30-60% of the total biomass during the abundance peaks. This study provides the first evidence of paralytic shellfish poisoning (PSP) toxins produced by dinoflagellate blooms in the Baltic Sea: PSP toxin dynamics correlated with the progression of the A. ostenfeldii bloom with toxin peaks mirroring the species abundance peaks. No other potentially toxic species were identified from the phytoplankton community. PSP toxin concentrations of up to 2.3 mg L � 1 measured in the A. ostenfeldii cell fraction suggest that blooms may have toxic effects on co-occurring biota.

In the past years, late summer blooms of the bioluminescent dinoflagellate Alexandrium ostenfeldii have become a recurrent phenomenon in coastal waters of the central and Northern Baltic Sea. This paper reports exceptionally high cell concentrations (105 to 106 cells L−1) of the species found during bioluminescent blooms in 2003 and 2004 in a shallow embayment of the Aland archipelago at the SW coast of Finland. Clonal cultures were established for morphological, molecular, toxicological and ecophysiological investigations to characterize the Finnish populations and compare them to other global A. ostenfeldii isolates. The Finnish isolates exhibited typical morphological features of A. ostenfeldii such as large size, a prominent ventral pore and an orthogonally bent first apical plate. However, unambiguous differentiation from closely related Alexandrium peruvianum was difficult due to considerable variation of sulcal anterior plate shapes. The Finnish strains were genetically distinct from other isolates of the species, but phylogenetic analyses revealed a close relationship to isolates from southern England and an A. peruvianum morphotype from the Spanish Mediterranean. Together these isolates formed a distinct clade which was separated from a clade containing other Northern European, North American and New Zealand populations. Toxin analyses confirmed the presence of the PSP toxins GTX2, GTX3 and STX in both Finnish isolates with GTX3 being the dominant toxin. Total relative PSP toxin contents were moderate, ranging from approximately 6 to 15 fmol cell−1 at local salinities of 5 and 10 psu, respectively. Spirolides were not detected. Salinity tolerance experiments showed that the Finnish isolates were well adapted to grow at the low salinities of the Baltic Sea. With a salinity range of approximately 6 to 20–25 psu, Baltic populations are physiologically distinct from their marine relatives. Vigorous production of different cyst types in the cultures suggest that cysts may play a crucial role in the survival and retainment of A. ostenfeldii populations in the Baltic Sea.