Occurrences of harmful algal blooms (HABs) associated with ocean environments in the South China Sea

Harmful algal blooms (HABs) occur frequently in the South China Sea (SCS), causing enormous economic losses in aquaculture. We analyzed historical HAB records during the period from 1980 to 2003 in SCS. We found that HABs-affected areas have expanded and the frequency of HABs varied during this period. The seasonal and annual variations, as well as causative algal species of HABs are different among the four regions. Areas with frequent HABs include the Pearl River Estuary (China), the Manila Bay (the Philippines), the Masinloc Bay (the Philippines), and the western coast of Sabah (Malaysia). HABs occurred frequently during March–May in the northern region of SCS, May–July in the eastern region, July in the western region, and year-round in the southern region. Among the species that cause HABs, Noctiluca scintillans dominated in the northern region, and Pyrodinium bahamense in the southern and eastern regions. Causative species also varied in different years for the entire SCS. Both P. bahamense and N. scintillans were the dominant species during 1980–2003. Some species not previously recorded formed blooms during 1991–2003, including Phaeocystis globosa, Scrippsiella trochoidea, Heterosigma akashiwo, and Mesodinium rubrum. Variations in HABs are related to various regional conditions, such as a reversed monsoon wind in the entire SCS, river discharges in the northern area, upwelling in Vietnam coastal waters during southwest winds and near Malaysia coastal waters during northeast winds, and eutrophication from coastal aquaculture in the Pearl River estuary, Manila Bay, and Masinloc Bay. Handling editor: D. Hamilton     

Harmful phytoplankton diversity and dynamics in an upwelling region (Sagres,SW Portugal) revealed by ribosomal RNA microarray combined with microscopy

The study region in Sagres, SW Portugal, is subject to natural eutrophication of coastal waters by wind-driven upwelling, which stimulates high primary productivity facilitating the recent economic expansion of bivalve aquaculture in the region. However, this economic activity is threatened by harmful algal blooms (HAB) caused by the diatoms Pseudo-nitzschia spp., Dinophysis spp. and other HAB dinoflagellates, all of which can produce toxins, that can induce Amnesic Shellfish Poisoning (ASP), Diarrhetic Shellfish Poisoning (DSP) and Paralytic Shellfish Poisoning (PSP). This study couples traditional microscopy with 18S/28S rRNA microarray to improve the detection of HAB species and investigates the relation between HAB and the specific oceanographic conditions in the region. Good agreement was obtained between microscopy and microarray data for diatoms of genus Pseudo-nitzschia and dinoflagellates Dinophysis spp., Gymnodinium catenatum and raphidophyte Heterosigma akashiwo, with less effective results for Prorocentrum. Microarray provided detection of flagellates Prymnesium spp., Pseudochattonella spp., Chloromorum toxicum and the important HAB dinoflagellates of the genera Alexandrium and Azadinium, with the latter being one of the first records from the study region. Seasonality and upwelling induced by northerly winds were found to be the driving forces of HAB development, with Pseudo-nitzschia spp. causing the risk of ASP during spring and summer upwelling season, and dinoflagellates causing the risk of DSP and PSP during upwelling relaxation, mainly in summer and autumn. The findings were in agreement with the results from toxicity monitoring of shellfish by the Portuguese Institute for Sea and Atmosphere and confirm the suitability of the RNA microarray method for HABs detection and aquaculture management applications.     

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.     

Spatial, seasonal and species variations of harmful algal blooms in the South Yellow Sea and East China Sea

The occurrences of harmful algal blooms (HABs), in terms of frequency and area in the Chinese coastal waters, have been increasing since 1980s and caused considerable economic losses. In the present study, we have analyzed spatial and seasonal characteristics of HAB events in the southern Yellow Sea and East China Sea along Chinese coast from 1933 to 2004. With a total 435 HAB records, the most frequent HAB occurrence area (FHA) is off the Yangtze River mouth and another two FHA areas are located south of the Yangtze River estuary along about isobaths of 30–60 m coastal water in the East China Sea. The time of HAB occurrence shifted during our study period: from autumn (August–October) before 1980s to July–August in 1980s, during May–July in 1990s, and May–June for the period of 2000–2004. Causative species were found to be different: Noctiluca scintillans and Skeletonema costatum were dominant causative species prior to 2000; and Prorocentrum donghaiense Lu was dominant from 2000 to 2004 and also caused large blooms in May. Trichodesmium sp. caused many HABs in autumn (August–October) prior to 1980s with only one HAB between 1980 and 2004. The changes of the dominant HAB species may have affected the timings of HAB occurrence, as well as the increasing HAB-affected areas in recent years.     

Vulnerability of coastal ecosystems to changes in harmful algal bloom distribution in response to climate change: projections based on model analysis

Harmful algal blooms (HABs), those proliferations of algae that can cause fish kills, contaminate seafood with toxins, form unsightly scums, or detrimentally alter ecosystem function have been increasing in frequency, magnitude, and duration worldwide. Here, using a global modeling approach, we show, for three regions of the globe, the potential effects of nutrient loading and climate change for two HAB genera, pelagic Prorocentrum and Karenia, each with differing physiological characteristics for growth. The projections (end of century, 2090–2100) are based on climate change resulting from the A1B scenario of the Intergovernmental Panel on Climate Change Institut Pierre Simon Laplace Climate Model (IPCC, IPSL‐CM4), applied in a coupled oceanographic‐biogeochemical model, combined with a suite of assumed physiological ‘rules’ for genera‐specific bloom development. Based on these models, an expansion in area and/or number of months annually conducive to development of these HABs along the NW European Shelf‐Baltic Sea system and NE Asia was projected for both HAB genera, but no expansion (Prorocentrum spp.), or actual contraction in area and months conducive for blooms (Karenia spp.), was projected in the SE Asian domain. The implications of these projections, especially for Northern Europe, are shifts in vulnerability of coastal systems to HAB events, increased regional HAB impacts to aquaculture, increased risks to human health and ecosystems, and economic consequences of these events due to losses to fisheries and ecosystem services.     

Reconstruction of Indian monsoon variability over the past 230,000 years: Planktic foraminiferal evidence from the NW Arabian Sea open-ocean upwelling area

We determined the faunal composition and total number of tests (#/g) of planktic foraminifera (> 125 μm) in core KH00-05 GOA 6 near Oman in order to decipher monsoon-induced variability of oceanographic productivity in the open-ocean upwelling area in the northwest Arabian Sea. The core contains a continuous record of sedimentation over the last 230 kyr, with the age model based on oxygen isotope and accelerator mass spectrometry ¹⁴C dates. We focused on species (Globigerina bulloides and Globigerinita glutinata) typical for SW monsoonal upwelling and species typical for NE monsoon conditions (Neogloboquadrina incompta, Neogloboquadrina dutertrei, Globigerinoides ruber, and Globigerinoides sacculifer). The changes in relative abundance of these monsoonal indicators suggest that the open-ocean upwelling area was dominated by the SW monsoon during interglacial periods, but by the NE monsoon during glacial periods.Increases in total test abundance during glacial periods confirmed that the NE monsoon rather than SW monsoon contributes largely to planktic foraminiferal productivity in this area. We argue that three types of circumstances resulted in high productivity, with nine high productivity events occurring at a 23-kyr frequency. The first type caused high productivity events at 102 and 199 ka (interglacial periods), characterized by the dominance of upwelling species, indicating high productivity during strong SW monsoons, correlated with high July insolation at 45° N. An exceptional high productivity event occurred at 37 ka during interglacial marine isotope stage (MIS) 3, with contributions from both SW and NE monsoons. The second type of high productivity event occurred at 61, 147, and 175 ka, during glacial periods, characterized by dominance of NE monsoon species, and correlated with low January insolation at 45° N. In addition, a high productivity event at 85 ka (interglacial period) also was induced by enhanced NE monsoons. The last two high productivity events occurred during transitional periods from glacial to interglacial (MIS 6/5.5 and 2/1), were characterized by the replacement of NE monsoon species with upwelling species, and corresponded to abrupt climate warming, suggesting that they are related to both accelerated SW monsoon systems and reduced NE monsoon systems.     

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.     

Monitoring Karenia brevis blooms in the Gulf of Mexico using satellite ocean color imagery and other data

Harmful algal blooms (HABs) of Karenia brevis are a recurrent problem in the Gulf of Mexico, with nearly annual occurrences on the Florida southwest coast, and fewer occurrences on the northwest Florida and Texas coasts. Beginning in 1999, the National Oceanic and Atmospheric Administration has issued the Gulf of Mexico HAB Bulletins to support state monitoring and management efforts. These bulletins involve analysis of satellite imagery with field and meteorological station data. The effort involves several components or models: (a) monitoring the movement of an algal bloom that has previously been identified as a HAB (type 1 forecast); (b) detecting new blooms as HAB or non-HAB (type 2); (c) predicting the movement of an identified HAB (type 3); (d) predicting conditions favorable for a HAB to occur where blooms have not yet been observed (type 4). The types 1 and 2 involve methods of bloom detection requiring routine remote sensing, especially satellite ocean color imagery and in situ data. Prediction (types 3 and 4) builds on the monitoring capability by using interpretative and numerical modeling. Successful forecasts cover more than 1000 km of coast and require routine input of remotely sensed and in situ data.The data sources used in this effort include ocean color imagery from the Sea-Viewing Wide Field-of-View Sensor/OrbView-2 satellite and processed using coastal-specific algorithms, wind data from coastal and offshore buoys, field observations of bloom location and intensity provided by state agencies, and forecasts from the National Weather Service. The HAB Bulletins began in coordination with the state of Florida in autumn of 1999 and included K. brevis bloom monitoring (type 1), with limited advisories on transport (type 3) and the detection of blooms in new areas (type 2). In autumn 2000, we improved both the transport forecasts and detection capabilities and began prediction of conditions favorable for bloom development (type 4). The HAB Bulletins have had several successes. The state of Florida was advised of the potential for a bloom to occur at the end of September 2000 (type 4), and the state was alerted to the position of blooms in January 2000 and October 2001 in areas that had not been previously sampled (type 3). These successful communications of HAB activity allowed Florida agencies responsible for shellfish management and public health to respond to a rapidly developing event in a timely, efficient manner.     

POPULATION DYNAMICS OF GREEN NOCTILUCA SCINTILLANS (DINOPHYCEAE) ASSOCIATED WITH THE MONSOON CYCLE IN THE UPPER GULF OF THAILAND1

Population dynamics of Noctiluca scintillans (Macartney) Kof. et Swezy containing the photosynthetic endosymbiont Pedinomonas noctilucae (Subrahman.) Sweeney was investigated in relation to environmental conditions in the upper Gulf of Thailand. A clear association was observed between the abundance of N. scintillans and the monsoon cycle, with its blooms occurring during the southwest (SW) monsoon from May to September, and low abundance during the northeast (NE) monsoon from November to February. Nutrient concentrations were higher during the SW monsoon than during the NE monsoon due to the combined effect of increased river discharge into the northern upper gulf and the transport of the riverine inputs by the prevailing clockwise circulation of the water. These nutrient conditions favored the growth of both phytoplankton and the endosymbiont. Correlation analysis revealed that the higher abundance of N. scintillans in the SW monsoon was manifested primarily by higher growth through both sexual and asexual reproduction supported by phagotrophy. However, the dependence of N. scintillans on the nutrient concentration was not significant, probably because the nutrient supply for the endosymbiont was sufficient due to intracellular accumulation of nutrients within the host cells. Sexual reproduction occurred only during the SW monsoon, and its potential importance in population growth was suggested. These findings showed the bottom‐up control of the population dynamics of N. scintillans through growth of phytoplankton as prey. The seasonal shift in the circulation pattern associated with the monsoon cycle played a crucial role in blooming of N. scintillans by producing favorable food conditions.     

On the control of HAB species using low biosurfactant concentrations

Biosurfactants have been suggested as a method to control harmful algal blooms (HABs), but warrant further and more in-depth investigation. Here we have investigated the algicidal effect of a biosurfactant produced by the bacterium Pseudomonas aeruginosa on five diverse marine and freshwater HAB species that have not been tested previously. These include Alexandrium minutum (Dinophycaee), Karenia brevis (Dinophyceae), Pseudonitzschia sp. (Bacillariophyceae), in marine ecosystems, and Gonyostomum semen (Raphidophyceae) and Microcystis aeruginosa (Cyanophyecae) in freshwater. We examined not only lethal but also sub-lethal effects of the biosurfactant. In addition, the effect of the biosurfactant on Daphnia was tested. Our conclusions were that very low biosurfactant concentrations (5 μg mL⁻¹) decreased both the photosynthesis efficiency and the cell viability and that higher concentrations (50 μg mL⁻¹) had lethal effects in four of the five HAB species tested. The low concentrations employed in this study and the diversity of HAB genera tested suggest that biosurfactants may be used to either control initial algal blooms without causing negative side effect to the ecosystem, or to provoke lethal effects when necessary.     

Meta‐analysis reveals enhanced growth of marine harmful algae from temperate regions with warming and elevated CO2 levels

Elevated pCO₂ and warming may promote algal growth and toxin production, and thereby possibly support the proliferation and toxicity of harmful algal blooms (HABs). Here, we tested whether empirical data support this hypothesis using a meta‐analytic approach and investigated the responses of growth rate and toxin content or toxicity of numerous marine and estuarine HAB species to elevated pCO₂ and warming. Most of the available data on HAB responses towards the two tested climate change variables concern dinoflagellates, as many members of this phytoplankton group are known to cause HAB outbreaks. Toxin content and toxicity did not reveal a consistent response towards both tested climate change variables, while growth rate increased consistently with elevated pCO₂. Warming also led to higher growth rates, but only for species isolated at higher latitudes. The observed gradient in temperature growth responses shows the potential for enhanced development of HABs at higher latitudes. Increases in growth rates with more CO₂ may present an additional competitive advantage for HAB species, particularly as CO₂ was not shown to enhance growth rate of other non‐HAB phytoplankton species. However, this may also be related to the difference in representation of dinoflagellate and diatom species in the respective HAB and non‐HAB phytoplankton groups. Since the proliferation of HAB species may strongly depend on their growth rates, our results warn for a greater potential of dinoflagellate HAB development in future coastal waters, particularly in temperate regions.     

High-resolution spatio-temporal distribution of a coastal phytoplankton bloom using laser in situ scattering and transmissometry (LISST)

Development of optical observation technologies provides new insights into harmful algal bloom (HAB) detection and assessment of HAB species dynamics. Based on preliminary laboratory tests, a laser in situ scattering and transmissometry instrument (LISST-100X) was used to monitor a high-biomass phytoplankton proliferation in the field. Short-term spatial and temporal changes in particle size distribution were measured during a recurrent Alexandrium taylori outbreak. Since the bloom was not monospecific, a size-fraction method to discriminate particular species from LISST-100X measurements was proposed. The results were validated to simultaneous microscopic counts of phytoplankton, and the significantly positive correlation obtained between the two methodologies confirmed the instrument's ability to discriminate phytoplankton at the group and species level. The LISST-100X obtains high-resolution in situ data, and is therefore a better alternative than the traditional microscope for assessing temporal and spatial evolution of HABs. Field observations showed high variability over a short time scale associated with diel vertical migration of A. taylori and the whole phytoplankton population (nanoplankton and microplankton). A numerical circulation model was used to investigate the influence of beach hydrodynamics in the observed horizontal variability. Simulations of the model suggested an important role of daily coastal circulation in determining the distribution of A. taylori in coastal environments.     

Satellite discrimination of Karenia mikimotoi and Phaeocystis harmful algal blooms in European coastal waters: Merged classification of ocean colour data

The detection of dense harmful algal blooms (HABs) by satellite remote sensing is usually based on analysis of chlorophyll-a as a proxy. However, this approach does not provide information about the potential harm of bloom, nor can it identify the dominant species. The developed HAB risk classification method employs a fully automatic data-driven approach to identify key characteristics of water leaving radiances and derived quantities, and to classify pixels into “harmful”, “non-harmful” and “no bloom” categories using Linear Discriminant Analysis (LDA). Discrimination accuracy is increased through the use of spectral ratios of water leaving radiances, absorption and backscattering. To reduce the false alarm rate the data that cannot be reliably classified are automatically labelled as “unknown”. This method can be trained on different HAB species or extended to new sensors and then applied to generate independent HAB risk maps; these can be fused with other sensors to fill gaps or improve spatial or temporal resolution. The HAB discrimination technique has obtained accurate results on MODIS and MERIS data, correctly identifying 89% of Phaeocystis globosa HABs in the southern North Sea and 88% of Karenia mikimotoi blooms in the Western English Channel. A linear transformation of the ocean colour discriminants is used to estimate harmful cell counts, demonstrating greater accuracy than if based on chlorophyll-a; this will facilitate its integration into a HAB early warning system operating in the southern North Sea.     

Predictive Ecological Modeling of Harmful Algal Blooms

We have thus far constructed a series of coupled ecological/physical models to predict the origin and fate of harmful algal blooms of the toxic dinoflagellate, Gymnodinium breve, on the West Florida shelf. We find that (1) the maximal population growth rate of G. breve must be ～0.80 day^?1 during initiation of a red tide, but as little as 0.08 day^?1 during its decay, (2) diatoms dominate when estuarine and shelf-break supplies of nitrate are made available to a model community of small and large diatoms, coccoid cyanophytes and Trichodesmium, non-toxic and red-tide dinoflagellates, microflagellates, and coccolithophores, (3) a numerical recipe for large red tides of G. breve requires DON supplies, mediated by iron-starved, nitrogen-fixers, while small blooms may persist on sediment sources of DON, (4) selective grazing must be exerted on the non-toxic dinoflagellates, and (5) vertical migration of G. breve in relation to seasonal changes of summer downwelling and fall/winter upwelling flow fields determines the duration and intensity of red tide landfalls along the barrier islands and beaches of the west coast of Florida, once other losses are specified. Given poorly known initial and boundary conditions and the expense of shipboard monitoring programs, however, bio-optical moorings or remote sensors are the most likely sources of model validation and improved HAB forecasts. Accordingly, the bio-optical implications of our ecological models must be included in future simulation analyses of HABs on both the West Florida shelf and within other coastal regions. Of particular importance for initiation of the coupled biophysical models is an improved understanding of the relationship of remotely sensed surface signals to shade-adapted dinoflagellates, aggregating below the first optical depth of the water column.     

A HAB warning system for shellfish harvesting in Portugal

The development of sustainable shellfish aquaculture is highly dependent on the provision of reliable monitoring and predictive information on the occurrence of harmful algal blooms (HABs). The Portuguese HAB early warning system and shellfish closures presented here is a prototype, developed in the ASIMUTH project. It relies on weekly monitoring data composed of observations of HAB species and toxin concentrations within shellfish, and ocean circulation forecasts generated by an operational oceanographic model. The shellfish harvesting areas comprise coastal areas, estuaries + rías and coastal lagoons. The weekly bulletin characterizes the current shellfish closure situation and next week's forecasts for potentially impacted areas. The period analyzed ranged from 27 July 2013 to 17 March 2014, and describes the first skill assessment of the warning system. The forecast accuracy was evaluated, considering the number of forecasts that were verified to be correct the following week (85%) as well as the number of events not forecasted (false negatives, 12%) and those expected but did not occur (false positives, 3%). Variations were most visible in the first weeks of bulletin implementation and during autumn–winter months. The complementary use of field data, remote sensing and operational models led to more accurate predictions of blooms and range of the event.     

Eutrophication and harmful algal blooms: A scientific consensus

In January 2003, the US Environmental Protection Agency sponsored a “roundtable discussion” to develop a consensus on the relationship between eutrophication and harmful algal blooms (HABs), specifically targeting those relationships for which management actions may be appropriate. Academic, federal, and state agency representatives were in attendance. The following seven statements were unanimously adopted by attendees based on review and analysis of current as well as pertinent previous data:

• (1)Degraded water quality from increased nutrient pollution promotes the development and persistence of many HABs and is one of the reasons for their expansion in the U.S. and other nations;
• (2)The composition–not just the total quantity–of the nutrient pool impacts HABs;
• (3)High-biomass blooms must have exogenous nutrients to be sustained;
• (4)Both chronic and episodic nutrient delivery promote HAB development;
• (5)Recently developed tools and techniques are already improving the detection of some HABs, and emerging technologies are rapidly advancing toward operational status for the prediction of HABs and their toxins;
• (6)Experimental studies are critical to further the understanding about the role of nutrients in HABs expression, and will strengthen prediction and mitigation of HABs; and
• (7)Management of nutrient inputs to the watershed can lead to significant reduction in HABs.

Supporting evidence and pertinent examples for each consensus statement are provided herein.     

Genomewide investigation of adaptation to harmful algal blooms in common bottlenose dolphins (Tursiops truncatus)

Harmful algal blooms (HABs), which can be lethal in marine species and cause illness in humans, are increasing worldwide. In the Gulf of Mexico, HABs of Karenia brevis produce neurotoxic brevetoxins that cause large‐scale marine mortality events. The long history of such blooms, combined with the potentially severe effects of exposure, may have produced a strong selective pressure for evolved resistance. Advances in next‐generation sequencing, in particular genotyping‐by‐sequencing, greatly enable the genomic study of such adaptation in natural populations. We used restriction site‐associated DNA (RAD) sequencing to investigate brevetoxicosis resistance in common bottlenose dolphins (Tursiops truncatus). To improve our understanding of the epidemiology and aetiology of brevetoxicosis and the potential for evolved resistance in an upper trophic level predator, we sequenced pools of genomic DNA from dolphins sampled from both coastal and estuarine populations in Florida and during multiple HAB‐associated mortality events. We sequenced 129 594 RAD loci and analysed 7431 single nucleotide polymorphisms (SNPs). The allele frequencies of many of these polymorphic loci differed significantly between live and dead dolphins. Some loci associated with survival showed patterns suggesting a common genetic‐based mechanism of resistance to brevetoxins in bottlenose dolphins along the Gulf coast of Florida, but others suggested regionally specific mechanisms of resistance or reflected differences among HABs. We identified candidate genes that may be the evolutionary target for brevetoxin resistance by searching the dolphin genome for genes adjacent to survival‐associated SNPs.     

Zooming on dynamics of marine microbial communities in the phycosphere of Akashiwo sanguinea (Dinophyta) blooms

Characterizing ecological relationships between viruses, bacteria and phytoplankton in the ocean is critical to understanding the ecosystem; however, these relationships are infrequently investigated together. To understand the dynamics of microbial communities and environmental factors in harmful algal blooms (HABs), we examined the environmental factors and microbial communities during Akashiwo sanguinea HABs in the Jangmok coastal waters of South Korea by metagenomics. Specific bacterial species showed complex synergistic and antagonistic relationships with the A. sanguinea bloom. The endoparasitic dinoflagellate Amoebophrya sp. 1 controlled the bloom dynamics and correlated with HAB decline. Among nucleocytoplasmic large DNA viruses (NCLDVs), two Pandoraviruses and six Phycodnaviruses were strongly and positively correlated with the HABs. Operational taxonomic units of microbial communities and environmental factors associated with A. sanguinea were visualized by network analysis: A. sanguinea–Amoebophrya sp. 1 (r = .59, time lag: 2 days) and A. sanguinea–Ectocarpus siliculosus virus 1 in Phycodnaviridae (0.50, 4 days) relationships showed close associations. The relationship between A. sanguinea and dissolved inorganic phosphorus relationship also showed a very close correlation (0.74, 0 day). Microbial communities and the environment changed dynamically during the A. sanguinea bloom, and the rapid turnover of microorganisms responded to ecological interactions. A. sanguinea bloom dramatically changes the environments by exuding dissolved carbohydrates via autotrophic processes, followed by changes in microbial communities involving host‐specific viruses, bacteria and parasitoids. Thus, the microbial communities in HAB are composed of various organisms that interact in a complex manner.     

In vitro interactions between several species of harmful algae and haemocytes of bivalve molluscs

Harmful algal blooms (HABs) can have both lethal and sublethal impacts on shellfish. To understand the possible roles of haemocytes in bivalve immune responses to HABs and how the algae are affected by these cells (haemocytes), in vitro tests between cultured harmful algal species and haemocytes of the northern quahog (= hard clam) Mercenaria mercenaria, the soft-shell clam Mya arenaria, the eastern and Pacific oysters Crassostrea virginica and Crassostrea gigas and the Manila clam Ruditapes philippinarum were carried out. Within their respective ranges of distribution, these shellfish species can experience blooms of several HAB species, including Prorocentrum minimum, Heterosigma akashiwo, Alexandrium fundyense, Alexandrium minutum and Karenia spp.; thus, these algal species were chosen for testing. Possible differences in haemocyte variables attributable to harmful algae and also effects of haemolymph and haemocytes on the algae themselves were measured. Using microscopic and flow cytometric observations, changes were measured in haemocytes, including cell morphology, mortality, phagocytosis, adhesion and reactive oxygen species (ROS) production, as well as changes in the physiology and the characteristics of the algal cells, including mortality, size, internal complexity and chlorophyll fluorescence. These experiments suggest different effects of the several species of harmful algae upon bivalve haemocytes. Some harmful algae act as immunostimulants, whereas others are immunosuppressive. P. minimum appears to activate haemocytes, but the other harmful algal species tested seem to cause a suppression of immune functions, generally consisting of decreases in phagocytosis, production of ROS and cell adhesion and besides cause an increase in the percentage of dead haemocytes, which could be attributable to the action of chemical toxins. Microalgal cells exposed to shellfish haemolymph generally showed evidence of algal degradation, e.g. loss of chlorophyll fluorescence and modification of cell shape. Thus, in vitro tests allow a better understanding of the role of the haemocytes and the haemolymph in the defence mechanisms protecting molluscan shellfish from harmful algal cells and could also be further developed to estimate the effects of HABs on bivalve molluscs in vivo.     

Harmful algal blooms: combining excitability and competition

Harmful algal blooms (HABs) characterized by a large concentration of toxic species appear rather rarely, but have a severe impact on the whole ecosystem. To study on possible trigger mechanisms for the emergence of HABs, we consider a nutrient-phytoplankton-zooplankton model to find the conditions under which a toxic phytoplankton species is able to form a bloom by winning the competition against its nontoxic competitor. The basic mechanism is related to the excitability of the system, i.e., the ability to develop a large response on certain perturbations. In a large class of models, a HAB results from a combined effect of nutrient enrichment and selective predation on different phytoplankton populations by zooplankton. We show that the severity of HAB is controlled by nutrient enrichment and zooplankton abundance, while the frequency of its occurrence depends on the strength of selectivity of predation. Thereby the intricate interplay between excitability, competition, and selective grazing pressure builds the backbone of the mechanism of the emergence of HABs.