GENETIC, MORPHOLOGICAL, AND TOXICOLOGICAL VARIATION AMONG GLOBALLY DISTRIBUTED STRAINS OF NODULARIA (CYANOBACTERIA)

Morphological, toxicological, and genetic variation was examined among 19 strains of Nodularia. The strains examined could be morphologically discriminated into four groups corresponding to N. spumigena Mertens, N. sphaerocarpa Bornet et Flahault, and two strains that did not clearly correspond to currently accepted Nodularia species. Genetic variation was examined using nucleotide sequencing of the phycocyanin intergenic spacer region (cpcBA-IGS) and RAPD-PCR. The PCR-RFLP of the cpcBA-IGS differentiated four genotypes corresponding to the four morphological groups. However, nucleotide sequencing of 598 bp of the 690-bp fragment showed that one of the three strains corresponding to N. sphaerocarpa (PCC 7804) was genetically divergent from the other two, suggesting that it constitutes a distinct species. Nucleotide variation within the morphospecies groups was limited (<1%), and all 14 Australian strains of N. spumigena possessed identical cpcBA-IGS sequences. The RAPD-PCR differentiated the same groups as the cpcBA sequencing and discriminated each of the seven different Australian populations of N. spumigena. Strains from within a bloom appeared genetically identical; however, strains isolated from different blooms could be separated into either a western or a southeastern Australian cluster, with one strain from western Australia showing considerable genetic divergence. The pattern of variation suggests that individual blooms of N. spumigena are clonal but also that Australian N. spumigena populations are genetically distinct from each other. Examination of genetic distance within and between blooms and within and between morphological groups showed clear genetic dicontinuities that, in combination with the cpcBA-IGS data, suggest that Nodularia contains genetically distinct morphospecies rather than a continuous cline of genetic variation. Furthermore, these morphospecies are genetically variable, exhibiting hierarchical patterns of genetic variation on regional and global scales. Production of the hepatotoxin nodularin was not restricted to one genetic lineage but was distributed across three of the five genotypic groups. A strain of N. spumigena from a nontoxic Australian population was found to fall within the range of genetic variation for other toxic Australian strains and appears to be a unique nontoxic strain that might have arisen by loss of toxin production capacity.     

Extensive genetic diversity and rapid population differentiation during blooms of Alexandrium fundyense (Dinophyceae) in an isolated salt pond on Cape Cod,MA, USA

In Massachusetts, paralytic shellfish poisoning (PSP) is annually recurrent along the coastline, including within several small embayments on Cape Cod. One such system, the Nauset Marsh System (NMS), supports extensive marshes and a thriving shellfishing industry. Over the last decade, PSP in the NMS has grown significantly worse; however, the origins and dynamics of the toxic Alexandrium fundyense (Balech) populations that bloom within the NMS are not well known. This study examined a collection of 412 strains isolated from the NMS and the Gulf of Maine (GOM) in 2006–2007 to investigate the genetic characteristics of localized blooms and assess connectivity with coastal populations. Comparisons of genetic differentiation showed that A. fundyense blooms in the NMS exhibited extensive clonal diversity and were genetically distinct from populations in the GOM. In both project years, genetic differentiation was observed among temporal samples collected from the NMS, sometimes occurring on the order of approximately 7 days. The underlying reasons for temporal differentiation are unknown, but may be due, in part, to life‐cycle characteristics unique to the populations in shallow embayments, or possibly driven by selection from parasitism and zooplankton grazing; these results highlight the need to investigate the role of selective forces in the genetic dynamics of bloom populations. The small geographic scale and limited connectivity of NMS salt ponds provide a novel system for investigating regulators of blooms, as well as the influence of selective forces on population structure, all of which are otherwise difficult or impossible to study in the adjacent open‐coastal waters or within larger estuaries.     

Regulation of blue-green algal buoyancy and bloom formation by light,inorganic nitrogen,C02, and trophic level interactions

In highly eutrophic ponds, buoyancy of the gas-vacuolate blue-green alga Anabaenopsis Elenkinii (Miller) was regulated by complex interactions between chemical and physical parameters, as well as by biological interactions between various trophic levels. Algal buoyancy and surface bloom formation were enhanced markedly by decreased light intensity, and to a lesser extent by decreased CO₂ availability and increased availability of inorganic nitrogen. In the absence of dense populations of large-bodied Cladocera, early season blooms of diatoms and green algae reduced light availability in the ponds thus creating conditions favorable for increased buoyancy and bloom formation by A. Elenkinii. The appearance of blue-green algal blooms could be prevented by a reduced density of planktivorous fish, which allowed development of dense cladoceran populations. The cladocerans limited the growth of precursory blooms of diatoms and green algae, and given the resulting clear-water conditions, buoyancy of A. Elenkinii was reduced, and blue-green algal blooms never appeared.     

Morphological and molecular characterization of free-floating and attached green macroalgae Ulva spp. in the Yellow Sea of China

During the summer of 2008 and 2009, massive algal blooms repeatedly broke out in the Yellow Sea of China. These were undoubtedly caused by the accumulations of one or more species in the macroalgal genus Ulva. In previous reports, morphological observation indicated that the species involved in this phenomenon is Ulva prolifera but molecular analyses indicated that the species belongs to an Ulva linza–procera–prolifera (LPP) clade. Correct identification of the bloom species is required to understand and manage the blooms, but the taxonomic status of the bloom species remains unclear. In the current study, the taxonomic status of 22 selected specimens from the Yellow Sea was assessed by using both morphological and molecular (ITS and rbcL sequences) data. In addition, 5S rDNA analyses were performed for those samples clustering in the LPP clade, and phylogenetic tree and ribotype analyses were constructed for determining the possible origin of the bloom. Three free-floating and two attached Ulva species were distinguished and described: Ulva compressa Linnaeus and Ulva pertusa Kjellman were found in free-floating samples; U. linza Linnaeus was found on rocks; and U. prolifera O.F. Müller was found in both habitats. Diversity in free-floating Ulva of the Yellow Sea appears to be greater than previously thought. The dominant free-floating Ulva species, U. prolifera, was not closely related to local populations attached to rocks but was closely related to populations from Japan.     

Abiotic factors influencing cyanobacterial bloom development in the Gulf of Finland (Baltic Sea)

Blooms of cyanobacteria are a recurrent phenomenon in the Baltic Sea, including the Gulf of Finland. The spatial extension, duration, intensity and species composition of these blooms varies widely between years. Alg@line data collected regularly from ferries as well as weather service and marine monitoring data from 1997 to 2005 are analysed to determine the main abiotic factors influencing the intensity and species composition of cyanobacterial blooms in the Gulf of Finland. It is demonstrated that the development of the Nodularia spumigena Mertens bloom is highly dependent on weather conditions such as photosynthetically active radiation and water temperature. Nutrient conditions, especially the surplus of phosphorus (according to Redfield ratio) related to the pre-bloom upwelling events in the Gulf, affect the intensity of Aphanizomenon sp. (L.) Ralfs blooms. Differences in bloom timing and duration indicate that, if the preconditions (like nutrient ratio/concentration and weather conditions) for bloom formation are favourable, then the Aphanizomenon bloom starts earlier, the overall bloom period is longer and the Nodularia peak might appear in a wider time window. Handling editor: K. Martens     

Examination of Ulva bloom species richness and relative abundance reveals two cryptically co-occurring bloom species in Narragansett Bay,Rhode Island

Blooms caused by the green macroalga Ulva pose a serious threat to coastal ecosystems around the world. Despite numerous studies of the causes and consequences of these blooms, we still have a limited understanding of Ulva bloom species richness and abundance due to difficulties in identifying Ulva species using morphological features. Along the northeastern U.S. coastline, all blooms of distromatic Ulva blades were previously identified as Ulva lactuca. Recent molecular sequencing, however, discovered the presence of additional distromatic Ulva species. Therefore, in order to determine the relative abundance of Ulva species within blooms, we conducted monthly surveys at four Narragansett Bay, RI, sites representing a gradient of bloom severity. We found that the biomass of Ulva within blooms was a mix of Ulva compressa and Ulva rigida, not U. lactuca as previously reported. In contrast, sites not impacted by blooms that were located near the mouth of Narragansett Bay were dominated by U. lactuca. We also observed spatial and temporal differences in Ulva and total macroalgal diversity between bloom-impacted sites, indicating that Ulva bloom composition can be radically different between similar sites within close proximity. We discuss our results in the context of Ulva blooms worldwide, highlighting the need to definitively determine bloom species composition in order to fully understand bloom dynamics.     

Genetic diversity of Cladophora oligocladoidea forming a bloom in the coastal area of Korea

Green algal blooms by Cladophora species are primarily reported in freshwater and coastal regions and cause severe ecological problems. A taxonomic report of the geographic distribution of Cladophora blooms is essential to explore the cause and impact of macroalgal blooms and for eco-physiological studies. The identification of blooming species is necessary for monitoring and controlling algal blooms. Genetic information of DNA sequences is useful for identifying species in the genus Cladophora. In the coastal area of Sangrok, Korea, a large-scale Cladophora bloom was reported for the first time in September 2015. In the present study, we identified the taxonomic entity of Cladophora oligocladoidea (Ulvophyceae, Cladophorales) in Korea. We report for the first time a green algal bloom by this species globally. This is the second report about genetic diversity of C. oligocladoidea since the species was established in Japan. Four ribotypes in the internal transcribed spacer (ITS) region of the nuclear ribosomal DNA represent the genetic diversity of Korean C. oligocladoidea. The results from this taxonomic report and our analysis of the green algal bloom of Korean C. oligocladoidea can provide valuable data to understand the geographic distribution and the genetic diversity of the species.     

How does eutrophication affect the role of grazers in harmful algal bloom dynamics?

Population dynamics of harmful algal bloom species are regulated both from the “bottom-up” by factors that affect their growth rate and from the “top-down” by factors that affect their loss rates. While it might seem apparent that eutrophication would have the greatest impact on factors affecting growth rates of phytoplankton (nutrient supply, light availability) the roles of top-down controls, including grazers and pathogens, cannot be ignored in studies of harmful bloom dynamics. Lags between the growth of phytoplankton and zooplankton populations, or disruption of zooplankton populations by adverse environmental conditions may be important factors in the initiation of plankton blooms under eutrophic conditions. Grazers that avoid feeding on harmful species and actively graze on competing species may also play important roles in bloom initiation. Grazers that are not affected by phytoplankton toxins and have growth rates comparable to phytoplankton (e.g. protozoan grazers) may have the potential to control the initiation of blooms. If the inhibition of grazers varies with cell density for blooms of toxic phytoplankton, eutrophication may increase the chances of blooms reaching threshold densities for grazer inhibition. In addition, secondary effects of eutrophication, including hypoxia and change in pH may adversely affect grazer populations, and further release HAB species from top-down control. The Texas brown tide (Aureoumbra lagunensis) blooms provide evidence for the role of grazer disruption in bloom initiation and the importance of high densities of brown tide cells in continued suppression of grazers.     

The Diversity of Harmful Algal Bloom-Triggering Mechanisms and the Complexity of Bloom Initiation

Mechanisms influencing initiation of harmful algal blooms (HABs) are diverse, and are not likely to be mutually exclusive. Rather, initiation of HABs is a result of interactions between processes, which result in biological, physical, and chemical conditions optimal for a bloom. Due to the complexity of some bloom initiation processes, bloom-preventative management may be possible. Results from a modeling exercise and a laboratory experiment indicated that a phytoplankton bloom could be circumvented through manipulation of the nutrient-loading mode, i.e., pulsed vs. continuous loading. These findings, should they prove consistent in more robust field experiments, may provide insights for the development of new management approaches for some HABs. Optimal bloom conditions, however, vary between HAB species. Consequently, it is unlikely that a single management solution will exist. Preventative management efforts will require early warning of HAB initiation, perhaps even before the appearance of an HAB species. An indicator based on the dynamic nature of phytoplankton succession events and phytoplankton species diversity may prove useful for this purpose. Applying this index to an existing plankton data set showed that Microcystis blooms might have been predicted months before the start of the bloom.     

Toxic cyanobacteria (blue-green algae) in Finnish fresh and coastal waters

A survey of the occurrence of toxic blooms of cyanobacteria in Finnish fresh and coastal waters was made during 1985 and 1986. Toxicity of the freeze-dried water bloom samples was tested by mouse-bioassay (i.p.). Forty-four per cent (83/188) of the bloom samples were found to be lethally toxic. Hepatotoxic blooms (54) were almost twice as common as neurotoxic ones (29). Anabaena was the most frequently found genus in toxic and non-toxic blooms and it was present in all neurotoxic samples. Statistical associations were found between hepatotoxicity and incidence of Microcystis aeruginosa, M. viridis, M. wesenbergii, Anabaena flos-aquae and Anabaena spiroides. Neurotoxicity was statistically associated with Anabaena lemmermannii, Anabaena flos-aquae and Gomphosphaeria naegeliana. Isolation of strains of cyanobacteria confirmed the occurrence of hepatotoxic and neurotoxic strains of Anabaena, as well as hepatotoxic strains of Microcystis and Oscillatoria species.Toxic blooms caused cattle poisonings at three different lakes during the study period. Toxic blooms also occurred in drinking water sources. Our study shows that toxic cyanobacteria are more common in Finnish lakes than would be expected on the basis of animal poisonings. The results of this study show the existence of toxic cyanobacteria in Finnish water supplies and the need for their continued study as agents of water based disease.     

Chytrid epidemics may increase genetic diversity of a diatom spring-bloom

Contrary to expectation, populations of clonal organisms are often genetically highly diverse. In phytoplankton, this diversity is maintained throughout periods of high population growth (that is, blooms), even though competitive exclusion among genotypes should hypothetically lead to the dominance of a few superior genotypes. Genotype-specific parasitism may be one mechanism that helps maintain such high-genotypic diversity of clonal organisms. Here, we present a comparison of population genetic similarity by estimating the beta-dispersion among genotypes of early and peak bloom populations of the diatom Asterionella formosa for three spring-blooms under high or low parasite pressure. The Asterionella population showed greater beta-dispersion at peak bloom than early bloom in the 2 years with high parasite pressure, whereas the within group dispersion did not change under low parasite pressure. Our findings support that high prevalence parasitism can promote genetic diversification of natural populations of clonal hosts.     

Reflections on the ballast water dispersal—harmful algal bloom paradigm

The ballast water dispersal—HAB paradigm, increasingly invoked circumstantially to explain puzzling and unaccountable HAB species outbreaks when lacking the multiple tests of confirmation recommended by Bolch and de Salas (2007), is evaluated. The types and examples of natural dispersions and taxon cycles are compared to exotic species bloom behavior linked to ballast water vectoring. The regional spreading, bloom behavior and disjunct distributions of the brown tide pelagophyte Aureococcus anophagefferens and the toxic dinoflagellate Gymnodinium catenatum, attributed to ballast water vectoring, are used as representative examples to evaluate the general application of the ballast water—HAB paradigm and associated interpretative problems. Human-aided emigration has a seeding and colonization ecology that differs from bloom ecology. For self-sustaining blooms to occur, these two ecologies must be accommodated by habitat growth conditions. The three stages that a non-native species must pass through (pioneering, persistence, community entry) to achieve colonization, community maintenance, and to bloom, and the niche-related factors and role of habitat disturbance are discussed. The relevance of cryptic occurrences, cyst deposits, dormancy periods and bloom rhythms of HAB species to their blooms attributed to ballast water-assisted introductions is also sketched. The different forms of HAB species rarity, their impact on the ballast water dispersal—HAB paradigm, and the dispersion and blooms of specialist and generalist HAB species are discussed. The remarkable novel and, often, monospecific blooms of dinoflagellate HAB species are being paralleled by similar eruptive bloom behavior cutting across phylogenetic lines, and being found also in raphidophytes, haptophytes, diatoms, silicoflagellates, etc. These blooms cannot be explained only as seeding events. An ecological release of ‘old barriers’ appears to be occurring generally at coastal bloom sites, i.e. something significant is happening ecologically and embedded within the ballast water—HAB paradigm. There may be a relationship between Life Form type [Smayda, T.J., Reynolds, C.S., 2001. Community assembly in marine phytoplankton: application of recent models to harmful dinoflagellate blooms. J. Plankton Res. 23, 447–461] and mode of expatriation; HAB dinoflagellate species commonly reported to produce ballast water-assisted toxic blooms invariably are members of cyst-producing Life Forms IV, V, VI. Ballast water vectoring of Life Forms I, II, III is rarely reported, even though many produce cysts, and where their novel introductions do occur they are more likely to be ichthyotoxic and vectored in shellfish stock consignments. The relevance of, and need to distinguish between morphospecies and their geographic/ribotype clades are discussed based on the Alexandrium tamarense/catenella/fundyense complex. Morphospecies-level ballast water dispersions are probably minor compared to the dispersal of the different ribotypes (toxic/non-toxic clades) making up HAB morphospecies; the redistribution and admixture of genotypes should be the focus. Ballast water-assisted expatriations impact the global occurrence of HABs through the direct transfer of previously absent species or introduction of genetic strains from the donor habitat that are ecologically favored over resident strains. The hybridization of species may be of potentially greater impact, resulting from the (1) mating of individuals from the donor and recipient habitats, or (2) through the interbreeding of strains introduced from two different donor sites into the recipient site, and whose progeny have greater ecological fitness than indigenous strains. Exceptional ecological changes of some sort appear to be occurring globally which, in combination with the genetically altered ecophysiological behavior of HAB species linked to ballast water dispersion and admixture, underpins the global HAB phenomenon. The impact of ballast water and shellfish transplantation on HABs and phytoplankton community ecology, generally, is considerably greater than the current focus on HAB species distributions, vectoring, and blooms. The methodological, investigative and conceptual potential of the ballast water—HAB paradigm should be exploited by developing a GEOHAB type intiative to advance quantification of global HAB ecology.     

Diversity and dynamics of microcystin—Producing cyanobacteria in China's third largest lake, Lake Taihu

Microcystins (MC), the most prevalent group of harmful cyanobacterial hepatotoxins, are primarily produced by strains of cyanobacteria in Microcystis, Anabaena and Planktothrix. Lake Taihu, which is the third largest freshwater lake in China, is a hypertrophic shallow lake in eastern China that has experienced lake-wide cyanobacterial blooms annually during the last few decades. In this study, PCR-DGGE was used to evaluate the diversity of potential MC-producing cyanobacteria and real-time PCR was used to analyze the dynamics of this population based on the presence of the mcy gene in samples collected during a year long study. The results revealed that all MC-producing genotypes detected belonged to the genus Microcystis. In addition, the MC-producing genotype communities were more diverse during the bloom season than the non-bloom season, and the diversity in the late bloom period was lower than the diversity in the early bloom period. Furthermore, the abundance of MC-producing genotypes increased dramatically during the bloom development period, reaching its peak in late summer (September). The results also suggested that the highest mcy gene concentration lagged behind the highest MC concentration, and the potential MC-producing cyanobacterial community shift lagged behind the development of blooms.     

A puzzle with many pieces: the genetic structure and diversity of Phaeocystis antarctica Karsten (Prymnesiophyta)

Strong ocean current systems characterize the Southern Ocean. The genetic structure of marine phytoplankton species is believed to depend mainly on currents. Genetic estimates of the relatedness of populations of phytoplankton species therefore should provide a proxy showing to what extent different geographic regions are interconnected by the ocean current systems. In this study, spatial and temporal patterns of genetic diversity were studied in the circumpolar prymnesiophyte Phaeocystis antarctica Karsten using seven nuclear microsatellite loci. Analyses were conducted for 86 P. antarctica isolates sampled around the Antarctic continent between 1982 and 2007. The results revealed high genetic diversity without single genotypes recurring even among isolates within a bloom or originating from the same bucket of water. Populations of P. antarctica were significantly differentiated among the oceanic regions. However, some geographically distant populations were more closely related to each other than they were to other geographically close populations. Temporal haplotype turnover within regions was also suggested by the multilocus fingerprints. Our data suggest that even within blooms of P. antarctica genetic diversity and population sizes are large but exchange between different regions can be limited. Positive and significant inbreeding coefficients hint at further regional substructure of populations, suggesting that patches, once isolated from one another, may not reconnect. These data emphasize that even for planktonic species in a marine ecosystem that is influenced by strong currents, significant breaks in gene flow may occur.     

A comparative study on recurrent blooms of Alexandrium minutum in two Mediterranean coastal areas

Alexandrium minutum is a toxic dinoflagellate widespread along the Mediterranean coasts. This species is frequently detected year-round at low concentrations within the Mediterranean basin. However, it only proliferates recurrently in some localities. Two affected areas are the Catalan and Sicilian coasts. In order to identify the factors determining the A. minutum blooms in the Mediterranean Sea, we compare the bloom conditions in two harbours: Arenys de Mar (Catalan coast, Spain) and Syracuse (Sicily, Italy), during 2002–2003. Arenys de Mar harbour is a fishing and leisure harbour and receives an input of freshwater rich in nutrients. Likewise, the Syracuse harbour – located on the Ionian coast of Sicily – is subject to freshwater inputs. Some points of this site are used for productive activities such as shellfish farming. A. minutum from the two areas studied were morphologically and genetically identical. In both sites, recurrent blooms take place from winter to spring. Surface water temperatures and salinities during A. minutum bloom events were 12–14.5 °C and 32–38, and 16–24 °C and 32–37.7 for Arenys and Syracuse, respectively. During the blooms, the spatial distribution of A. minutum in the two harbours, the physicochemical characteristics and the phytoplankton community were studied. Similarities in composition of the phytoplankton community were evidenced, with a clear dominance of dinoflagellates over the other taxa. In Arenys, the second dominant species was Prorocentrum micans followed by Scrippsiella spp. and Dinophysis sacculus. The same species were found in Syracuse although P. triestinum, and alternatively Lingulodinium polyedrum, reached cell densities much higher than the other dinoflagellates giving marked water discolourations.     

First occurrence of Cochlodinium blooms in Sabah, Malaysia

Harmful algal blooms (HABs) resulting in red discoloration of coastal waters in Sepanggar Bay, off Kota Kinabalu, Sabah, East Malaysia, were first observed in January 2005. The species responsible for the bloom, which was identified as Cochlodinium polykrikoides, coincided with fish mortalities in cage-cultures. Determinations of cell density between January 2005 and June 2006 showed two peaks that occurred in March–June 2005 and June 2006. Cell abundance reached a maximum value of 6 × 10⁶ cells L⁻¹ at the fish cage sampling station where the water quality was characterized by high NO₃–N and PO₄–P concentrations. These blooms persisted into August 2005, were not detected during the north–east monsoon season and occurred again in May 2006. Favorable temperature, salinity and nutrient concentrations, which were similar to those associated with other C. polykrikoides blooms in the Asia Pacific region, likely promoted the growth of this species. Identification of C. polykrikoides as the causative organism was based on light and scanning microscopy, and confirmed by partial 18S ribosomal DNA sequences of two strains isolated during the bloom event (GenBank accession numbers DQ915169 and DQ915170).     

CONFIRMATION OF DOMOIC ACID PRODUCTION BY PSEUDONITZSCHIA AUSTRALIS (BACILLARIOPHYCEAE) CULTURES1

Single done isolates of Pseudonitzschia australis Frenguelli (= Nitzschia pseudoseriata Hasle) isolated from a toxic bloom in Monterey Bay, California produced domoic acid in culture. Although long-term historical records do not indicate previous blooms of this species on the Pacific coast, this is probably because it has been often misidentified as Nitzschia seriata Hasle; previous evidence for toxicity is lacking. Hydrographic data suggest that areas such as Monterey Bay might be “hot spots” for domoic acid-producing blooms.     

Benthic cyanobacterial bloom impacts the reefs of South Florida (Broward County, USA)

Benthic cyanobacteria of the genus Lyngbya can form prominent mats and blooms in tropical and subtropical coral reef and seagrass habitats worldwide. A Lyngbya bloom on the reef tract offshore of Broward County, Florida, was first noted in 2002, and although it is seasonally variable in its distribution and abundance, it has persisted and spread over the past 3 years. In this study, the most abundant species of Lyngbya found in the blooms have been identified and compared to other species of Lyngbya by morphological and molecular methods. The most common species of Lyngbya is consistent with the properties of Lyngbya confervoides C. Agardh. The 16S ribosomal DNA sequence shares 88–92% identity with other known Lyngbya sequences, suggesting that this bloom consists primarily of a new, previously unsequenced species of Lyngbya. The second most common Lyngbya in the bloom is consistent with Lyngbya polychroa. This persistent bloom is a concern because it smothers octocorals and other invertebrates and negatively impacts these southeastern Florida reefs.     

INTERSTRAIN VARIABILITY IN PHYSIOLOGY AND GENETICS OF HETEROSIGMA AKASHIWO (RAPHIDOPHYCEAE) FROM THE WEST COAST OF NORTH AMERICA1

High levels of intraspecific variability are often associated with HAB species, and this variability is likely an important factor in their competitive success. Heterosigma akashiwo (Hada) Hada ex Y. Hara et M. Chihara is an ichthyotoxic raphidophyte capable of forming dense surface‐water blooms in temperate coastal regions throughout the world. We isolated four strains of H. akashiwo from fish‐killing northern Puget Sound blooms in 2006 and 2007. By assessing numerous aspects of biochemistry, physiology, and toxicity, we were able to describe distinct ecotypes that may be related to isolation location, source population, or bloom timing. Contrasting elements among strains were cell size, maximum growth and photosynthesis rates, tolerance of low salinities, amino acid use, and toxicity to the ciliate grazer Strombidinopsis acuminatum (Fauré‐Fremiet). In addition, the rDNA sequences and chloroplast genome of each isolate were examined, and while all rDNA sequences were identical, the chloroplast genome identified differences among the strains that tracked differences in ecotype. H. akashiwo strain 07A, which was isolated from an unusual spring bloom, had a significantly higher maximum potential photosynthesis rate (28.7 pg C · cell^?1 · h^?1) and consistently exhibited the highest growth rates. Strains 06A and 06B were not genetically distinct from one another and were able to grow on the amino acids glutamine and alanine, while the other two strains could not. Strain 07B, which is genetically distinct from the other three strains, exhibited the only nontoxic effect. Thus, molecular tools may support identification, tracking, and prediction of strains and/or ecotypes using distinctive chloroplast gene signatures.     

Seasonality of North Atlantic phytoplankton from space: impact of environmental forcing on a changing phenology (1998–2012)

Seasonal pulses of phytoplankton drive seasonal cycles of carbon fixation and particle sedimentation, and might condition recruitment success in many exploited species. Taking advantage of long‐term series of remotely sensed chlorophyll a (1998–2012), we analyzed changes in phytoplankton seasonality in the North Atlantic Ocean. Phytoplankton phenology was analyzed based on a probabilistic characterization of bloom incidence. This approach allowed us to detect changes in the prevalence of different seasonal cycles and, at the same time, to estimate bloom timing and magnitude taking into account uncertainty in bloom detection. Deviations between different sensors stressed the importance of a prolonged overlap between successive missions to ensure a correct assessment of phenological changes, as well as the advantage of semi‐analytical chlorophyll algorithms over empirical ones to reduce biases. Earlier and more intense blooms were detected in the subpolar Atlantic, while advanced blooms of less magnitude were common in the Subtropical gyre. In the temperate North Atlantic, spring blooms advanced their timing and decreased in magnitude, whereas fall blooms delayed and increased their intensity. At the same time, the prevalence of locations with a single autumn/winter bloom or with a bimodal seasonal cycle increased, in consonance with a poleward expansion of subtropical conditions. Changes in bloom timing and magnitude presented a clear signature of environmental factors, especially wind forcing, although changes on incident photosynthetically active radiation and sea surface temperature were also important depending on latitude. Trends in bloom magnitude matched changes in mean chlorophyll a during the study period, suggesting that seasonal peaks drive long‐term trends in chlorophyll a concentration. Our results link changes in North Atlantic climate with recent trends in the phenology of phytoplankton, suggesting an intensification of these impacts in the near future.