- Summary of Invasiveness
- Taxonomic Tree
- Notes on Taxonomy and Nomenclature
- Plant Type
- Distribution Table
- History of Introduction and Spread
- Risk of Introduction
- Habitat List
- Biology and Ecology
- Water Tolerances
- Notes on Natural Enemies
- Means of Movement and Dispersal
- Pathway Causes
- Pathway Vectors
- Impact Summary
- Economic Impact
- Environmental Impact
- Social Impact
- Risk and Impact Factors
- Uses List
- Detection and Inspection
- Similarities to Other Species/Conditions
- Prevention and Control
- Gaps in Knowledge/Research Needs
- Distribution Maps
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PicturesTop of page
IdentityTop of page
Preferred Scientific Name
- Gymnodinium catenatum
Summary of InvasivenessTop of page
G. catenatum is a photosynthetic dinoflagellate that is most notable as the only naked dinoflagellate known to be responsible for paralytic shellfish poisoning (PSP), a neurotoxic poisoning syndrome which affects human consumers of contaminated shellfish. Shellfish contamination by PSP affects thousands of people per year on a worldwide basis and the economic costs of harmful algal blooms in the United States alone are in excess of 50 million dollars annually (Hoagland et al., 2002). In Australia, the direct costs (toxin and plankton monitoring) of PSP contamination of shellfish exceed 1 million Australian dollars per year. Indirect costs to the shellfish industry are difficult to estimate, but include loss of income and markets due to harvest closures and the “halo effect” of reduced confidence in shellfish and other seafood products.
Australian populations form dense and extensive blooms confined largely to estuarine and coastal waters typically in calm conditions and stratified water columns after heavy rainfall (Hallegraeff et al., 1995). Coastal currents can disperse the organism along coastlines to suitable estuarine/coastal habitats. In some areas of the world populations appear to be also associated with continental shelf-edge upwelling areas (e.g. northwest Spain, New Zealand) that can facilitate rapid and widespread dispersal along coastlines (Mackenzie and Beauchamp, 2002; Sordo et al., 2001). Resistant resting stages (cysts) are formed during periods of nutrient stress which fall to the bottom sediments and may later germinate to establish a planktonic population when conditions are suitable. These cysts are highly resistant to physical and chemical attack, are long-lived (>10 years) under suitably anoxic conditions, and are therefore capable of long-distance (inter-continental) dispersal by a number of natural or anthropogenic vectors. The most notable vector is ballast water (Hallegraeff and Bolch, 1992); other vectors include equipment contaminated with marine water or sediment (e.g. dredges and fishing gear), movements of commercial shellfish or other marine animals, attachment to flotsam/jetsam and seaweed.
Taxonomic TreeTop of page
- Domain: Eukaryota
- Kingdom: Protista
- Phylum: Protozoa
- Class: Dinophyceae
- Order: Gymnodiniales
- Family: Gymnodiniaceae
- Genus: Gymnodinium
- Species: Gymnodinium catenatum
Notes on Taxonomy and NomenclatureTop of page
The family Gymnodiniales is a diverse and highly variable group of dinoflagellates, sometimes known as “naked” dinoflagellates, which do not possess the array of cellulose-like plates as part of their outer cell covering (amphiesma). The largest genus of the group, Gymnodinium, has in the past included a huge diversity of naked photosynthetic and non-photosynthetic forms. Molecular and ultra-structural studies over the past 10-15 years indicate that the genus contains many divergent or unrelated lineages and many species have been transferred to newly erected genera leaving the genus Gymnodinum more tightly defined as naked species that possess a horseshoe-shaped apical groove encircling the apex of the cell (Daugbjerg et al., 2000). Current classical and molecular taxonomic data indicate that the toxic dinoflagellate Gymnodinium catenatum is relatively closely related to the type species of the genus Gymnodinium fuscum. As of early 2008, there are no proposals to move G. catenatum out of the genus Gymnodinium. The toxic, chain forming G. catenatum is part of a group of 4 species known as the microreticulate-cyst species that include the non-chain forming, non-toxic species Gymnodinium nolleri (Ellegaard and Oshima 1998), Gymnodinium microreticulatum (Bolch et al., 1999) and Gymnodinium trapeziforme (Attaran-Fariman et al., 2007).
DescriptionTop of page
G. catenatum is a large (30-40 µm cell width) photosynthetic, gymnodinoid dinoflagellate that forms distinctive long chains of cells. Cells of G. catenatum have a girdle groove (cingulum) that encircles the cell only once. Single cells are not uncommon and are difficult to distinguish from a range of other photosynthetic Gymnodinium species without detailed examination of intracellular features such as the shape and distribution of the chloroplasts and the nucleus (Hallegraeff et al., 1988; Rees and Hallegraeff, 1991). The resting cyst is a distinctive red-brown colour and 35-62 µm in diameter with complex microreticulate surface markings that reflect structures present on the planktonic cell (girdle and ventral groove and a loop-shaped apical groove).
Apart from its spectacular planktonic form, it is most notable as the only naked dinoflagellate known to produce paralytic shellfish toxins (PST, saxitoxins). It is a causative organism of paralytic shellfish poisoning (PSP), a neurotoxic poisoning syndrome which affects human consumers of contaminated shellfish. Shellfish contamination by PSP affects thousands of people per year on a worldwide basis and the economic costs of harmful algal blooms in the United States alone are in excess of 50 million dollars annually (Hoagland et al., 2002).
Plant TypeTop of page Perennial
DistributionTop of page
From combination of confirmed planktonic and benthic resting cyst reports, G. catenatum has now been documented within discrete areas along the coastlines of all continents (see Bolch and Reynolds, 2002; Bolch and Salas, 2007). These two works also clarify a number of earlier reports of G. catenatum that are now known to refer to related species or other morphologically similar species. G. catenatum is best described as widespread but rare or cryptic, ranging from tropical to cool-temperate waters. Distribution within most areas is typically localised within a few hundred kilometres of coastline; however, reports are typically scattered or confined to specific bloom areas/events. Localised distributions appear to be increasing over time and it is likely that local distributions are more widespread than currently documented.
In addition to the countries listed in the distribution table, G. catenatum is also present in South Africa and Palau (C Bolch, University of Tasmania, personal communication, 2008).
Distribution TableTop of page
The distribution in this summary table is based on all the information available. When several references are cited, they may give conflicting information on the status. Further details may be available for individual references in the Distribution Table Details section which can be selected by going to Generate Report.Last updated: 10 Jan 2020
|Continent/Country/Region||Distribution||Last Reported||Origin||First Reported||Invasive||Reference||Notes|
|Angola||Present, Localized||Rangel and Silva (2006)|
|Morocco||Present, Widespread||Tagmouti et al. (1995)|
|China||Present, Widespread||Qi et al. (1996)|
|India||Present, Localized||Godhe et al. (1996)|
|Indonesia||Present, Few occurrences||Holmes et al. (2002)|
|Japan||Present, Widespread||Matsuoka and Fukuyo (1994)|
|-Honshu||Present, Widespread||Matsuoka and Fukuyo (1994)|
|-Kyushu||Present, Widespread||Matsuoka and Fukuyo (1994)|
|-Shikoku||Present, Widespread||Matsuoka and Fukuyo (1994)|
|Kuwait||Present||Glibert et al. (2002)|
|Malaysia||Present||Anton and Mohamad-Noor (1996)|
|Philippines||Present||Fukuyo et al. (1993)|
|Singapore||Present, Few occurrences||Holmes et al. (2002)|
|South Korea||Present, Widespread||Kim et al. (1996)|
|Gibraltar||Present, Widespread||Gómez (2003)|
|Greece||Present||Giannakourou et al. (2005)|
|Portugal||Present, Widespread||Franca and Almeida (1989)|
|Russia||Present||CABI (Undated)||Present based on regional distribution.|
|-Russian Far East||Present, Localized||Orlova et al. (2004)|
|Spain||Present, Widespread||Estrada et al. (1984)|
|-Canary Islands||Present, Widespread||Targarona et al. (1999)|
|Costa Rica||Present||Viquez and Hargraves (1995)|
|Cuba||Present||Leal et al. (2003)|
|Mexico||Present, Localized||Band-Schmidt et al. (2006)|
|Australia||Present||CABI (Undated)||Present based on regional distribution.|
|-New South Wales||Present, Localized||Bolch and Reynolds (2002)|
|-South Australia||Present, Localized||Bolch and Reynolds (2002)|
|-Tasmania||Present, Widespread||Hallegraeff et al. (1988)|
|-Victoria||Present, Widespread||Sonneman and Hill (1997)|
|Mediterranean and Black Sea||Present, Widespread||Gómez (2003)|
|Argentina||Present, Localized||Balech (1964)|
|Brazil||Present, Localized||Oliveira-Proença et al. (2001)|
|Uruguay||Present, Widespread||Mendez and Brazeiro (1993)|
|Venezuela||Present, Localized||Barbera-Sánchez and Gamboa-Maruez (2001)|
History of Introduction and SpreadTop of page
Prior to the early 1970s, reports of G. catenatum cells were rare. Since its description from the Gulf of Mexico in 1939 (Graham, 1943), it had only been reported from Argentina in 1962 (Balech, 1964) and Japan in 1967 (Hada, 1967; as Gymnodinium sp. A3). Recent 210Pb-dated cyst studies have since shown that G. catenatum has been present in southern and western Japan since at least 1700 (Matsuoka et al., 2006). Scientific attention on the species was heightened after it was discovered as the cause of human PSP poisonings in Spain in 1976 (Estrada et al., 1984) and Mexico in 1979 (Mee et al., 1986). The geographic extent of G. catenatum reports increased rapidly during the 1980s and 1990s. The discovery of a distinctive microreticulate resting cyst (Anderson et al., 1988) prompted others to search for resting cysts and many reports of similarly patterned cysts emerged, often in areas where the species had never been seen in the plankton, such as northern Europe (Dale et al., 1993; Ellegaard et al., 1993; Nehring, 1995). A number of these reports have since been attributed to the related non-toxic species G. nolleri (Ellegaard and Oshima, 1998) and G. microreticulatum (Bolch et al., 1999; Bolch and Reynolds, 2002); however, it is clear that G. catenatum has established populations on the coast of all continents except Antarctica (Bolch and Reynolds, 2002; Bolch and Salas, 2007).
While a number of populations around the globe are suspected of being introduced, strong supporting evidence only exists for the introduction of Australasian G. catenatum from southern Japan, most likely via resting cysts contained in ballast water discharged from bulk cargo vessels entering Tasmanian waters from the early 1970s (McMinn et al., 1997; Bolch and Salas, 2007). Blooms in Australian waters were first identified in the Derwent Estuary (Hobart, Tasmania) in September 1985 (Hallegraeff et al., 1988). Massive blooms in Huon Estuary in southern Tasmania during September 1985 to March 1986 led to widespread closure of the local shellfish industry for several months (Hallegraeff and Sumner, 1986). Examination of historical plankton samples established its presence in southern Tasmania since at least 1980 (Hallegraeff et al., 1989). Sediment 210Pb-dating studies of resting cysts indicate that G. catenatum had been present in southern Tasmania since the early 1970s (McMinn et al., 1997). During the 1990s G. catenatum was discovered at a number of mainland Australian sites. In 1993, abundant resting cysts were found at several sites along the southern coast of Victoria (Sonneman and Hill, 1997) and a single culture was established from Port Philip Bay (since lost). Populations are also known from Port Lincoln in South Australia (Bolch and Reynolds, 2002), a suspected secondary translocation of Tasmanian populations (McMinn et al., 2001), and the Hawkesbury River Estuary in New South Wales (Bolch and Reynolds, 2002).
IntroductionsTop of page
Risk of IntroductionTop of page
Once established on a coastline, there are multiple means by which G. catenatum may spread to other suitable habitats along the coast. Gradual dispersal of blooms or resting cysts via coastal currents is inevitable. Longer range (hundreds of kilometres) secondary dispersal by shipping, recreational or other vessels is possible, and is the suspected vector for dispersal from southern Tasmania to Port Lincoln, South Australia, between port environments. Any activity that transfers untreated seawater or marine sediment at ambient temperatures (ranging from 4.0 to 45°C) to unaffected marine environments has the potential to spread G. catenatum to new environments (e.g. marine equipment such as dredges and fishing gear, commercial shellfish or other marine animals and fauna).
HabitatTop of page
Australian populations are confined largely to coastal estuaries and sheltered embayments (Hallegraeff et al., 1989) and cells die when dispersed offshore, away from protected shorelines and estuaries. Coastal currents can disperse the organism along coastlines to suitable estuarine/coastal habitats. In a number of other areas of the world (e.g. Japan, Korea, northwest Spain, New Zealand), planktonic populations appear to be also associated with continental shelf-edge upwelling areas (Mackenzie and Beauchamp, 2002; Sordo et al., 2001).
Habitat ListTop of page
|Mud flats||Present, no further details||Natural|
|Intertidal zone||Present, no further details||Natural|
|Inshore marine||Principal habitat||Natural|
|Pelagic zone (offshore)||Principal habitat||Natural|
|Benthic zone||Principal habitat||Natural|
Biology and EcologyTop of page
Like most planktonic dinoflagellates, the main lifecycle phase is the motile haploid cell phase. Benthic resting cysts are the diploid phase of the sexual lifecycle.
The primary form of population growth is through mitotic binary division of planktonic cells. Cell division occurs typically along a dorso-ventral diagonal plane, with the right side daughter cell migrating to a position below the other daughter cell and remaining attached to form chains (Blackburn et al., 1989). Other forms of division (sexual and non-sexual) have been noted in laboratory cultures (Figueroa et al., 2006) but the significance and relative frequency of these in natural populations is unclear. Chains may form up to 64 cells in length; however, longer chains (>120 cells) have been seen in some natural bloom populations (C Bolch, University of Tasmania, personal communication). Division of cells in chains is semi-synchronous, beginning at the anterior end of a chain and progressing along the chain such that cells in longer chains have a staggered arrangement (Blackburn et al., 1989).
Laboratory and bloom studies indicate that different G. catenatum populations have diverse environmental tolerances (see Hallegraeff and Fraga, 1998); however, few populations have been studied in extensive detail. Australian populations typically bloom in warmer months from August through to March-April (Hallegraeff et al., 1989, 1995), but since the late 1990s those in Tasmania have bloomed predominantly in winter (M. de Salas, University of Tasmania, Australia, personal communication, 2011). However, persistent and widespread blooms have also been noted in from May to August. Blooms occur at water temperatures in excess of 12°C (Hallegraeff et al., 1989), and appear correlated with stratified water columns promoted by heavy early-summer rain followed by low winds (Hallegraeff et al., 1995). Laboratory studies of Singapore strains indicate optimum growth at 26-29°C and poor growth below 25°C, suggesting that some tropical and sub-tropical populations are adapted to higher water temperatures. Mexican strains from the Gulf of California also appear to have tolerance of higher water temperatures (Band-Schmidt et al., 2006).
Water TolerancesTop of page
|Parameter||Minimum Value||Maximum Value||Typical Value||Status||Life Stage||Notes|
|Salinity (part per thousand)||28||32||Optimum||15-35 tolerated|
|Water temperature (ºC temperature)||12||30||Optimum||4-40 tolerated|
Notes on Natural EnemiesTop of page
A wide range of grazers and other heterotrophic protists appear to be capable of grazing on G. catenatum cells with little apparent impact. Parasites of dinoflagellates (e.g. Amoebophyra) are known, although little is known about specific pathogens and parasites of G. catenatum. Algicidal bacteria have been described that attack G. catenatum and other gymnodinoid cells (e.g. Lovejoy et al., 1998); however, at this stage, the mechanisms and dynamics of these interactions are not sufficiently understood to be exploited as a form of biological control.
Means of Movement and DispersalTop of page
Natural Dispersal (Non-Biotic)
Blooms of G. catenatum populations appear to be confined to coastal areas and/or continental shelves, therefore long-range dispersal across oceanic basins and boundaries appears unlikely over human timescales. However, once established on a coastline, gradual dispersal of blooms or resting cysts via coastal currents appears inevitable.
Vector Transmission (Biotic)
Planktonic cells are fragile and unlikely to survive longer than several days without light, and are killed by anoxia or sustained temperatures of 35°C or more. Resting cysts are more robust and may survive temperatures of up to 45-50°C for short periods (Bolch and Hallegraeff, 1993), are unaffected by anoxia, and survive potentially several years without light. Therefore, transport of resting cysts presents the biggest risk vector transmission. Longer range (100s of kilometres) secondary dispersal by shipping, recreational or other vessels is possible, and is the suspected vector for dispersal from southern Tasmania to Port Lincoln, South Australia, between port environments (see Bolch and Salas, 2007). Any activity that transfers untreated seawater at ambient temperature (10-20°C) or marine sediment at temperatures between 4.0 to 45°C, to unaffected marine environments has the potential to spread G. catenatum. For example, cells or cysts may be dispersed by contaminated marine equipment such as dredges and fishing gear, commercial shellfish or other marine animals and fauna. Given the relative hardiness of resting cysts, dispersal by birds or flying insects (Proctor, 1966) is also possible, but as yet not investigated for dinoflagellate cysts.
The few cases of suspected introduction (i.e. for which evidence is available) indicate introduction has been accidental and likely to be ballast-water-mediated (Bolch and Salas, 2007).
Pathway CausesTop of page
Pathway VectorsTop of page
Impact SummaryTop of page
Economic ImpactTop of page
The primary impact of G. catenatum is due to its role in paralytic shellfish poisoning (PSP), a neurotoxic poisoning syndrome which affects human consumers of contaminated shellfish. Shellfish contamination by PSP affects thousands of people per year on a worldwide basis and the economic costs of harmful algal blooms in the United States alone are in excess of 50 million dollars annually (Hoagland et al., 2002). In Australia, the direct costs (toxin and plankton monitoring) of PSP contamination of shellfish exceed 1 million Australian dollars per year. Human heath associated costs are relatively low due to extensive seafood toxin and phytoplankton monitoring programs in many affected countries. Human poisonings are therefore rare. Indirect costs to the shellfish industries are difficult to estimate, but include loss of income and markets due to harvest closures and the “halo effect” of reduced confidence in shellfish and other seafood products.
Environmental ImpactTop of page
Environmental impacts of G. catenatum are largely unknown.
Impact on Biodiversity
Social ImpactTop of page
Social impacts in developed countries are primarily reduction or loss of shellfish industries and associated employment during extended periods of harvest closure due to PST contamination. For example, some heavily affected areas of southern Tasmania (e.g. Huon Estuary) appear to no longer be commercially viable areas for oyster and/or mussel production due to the risk of extended periods of closure due to PST contamination. Human health impacts (intoxication and/or death) from G. catenatum-induced shellfish poisoning events can be much more severe in less developed areas that lack extensive toxin and phytoplankton monitoring.
Risk and Impact FactorsTop of page Invasiveness
- Proved invasive outside its native range
- Highly adaptable to different environments
- Highly mobile locally
- Fast growing
- Has high reproductive potential
- Has propagules that can remain viable for more than one year
- Reproduces asexually
- Has high genetic variability
- Monoculture formation
- Negatively impacts cultural/traditional practices
- Negatively impacts human health
- Negatively impacts livelihoods
- Negatively impacts aquaculture/fisheries
- Negatively impacts tourism
- Reduced amenity values
- Competition - monopolizing resources
- Rapid growth
- Highly likely to be transported internationally accidentally
- Difficult to identify/detect as a commodity contaminant
- Difficult to identify/detect in the field
- Difficult/costly to control
UsesTop of page Economic Value
Uses ListTop of page
- Laboratory use
- Research model
Detection and InspectionTop of page
Detection in the environment or in ballast water currently relies on water/sediment collection and concentration, followed by microscopic analysis for the planktonic cell and resting stages. Molecular detection approaches for the G. catenatum have been attempted (Bolch, 2001; Patil et al., 2005), including quantitative PCR detection and enumeration (J Patil, CSIRO Marine Research, Australia, personal communication), but none have reached the stage of commercial deployment as of April 2011 (M. de Salas, University of Tasmania, Australia, personal communication, 2011).
Similarities to Other Species/ConditionsTop of page
Cells of G. catenatum can be distinguished from the related microreticulate species by cell size and chain formation. Cells of G. microreticulatum (Bolch et al., 1999) and G. trapeziforme (Attaran-Fariman et al., 2007) do not form chains and the cells are considerably smaller; G. nolleri is also much smaller and forms chains of only 2 cells (Ellegaard et al., 1993). Attaran-Fariman et al. (2007) give a detailed description of features separating the species in the group. G. catenatum may also be confused with other chain-forming species, especially in fixed samples due to poor preservation of fine cell structure (e.g. Lugol’s iodine and formalin).
Prevention and ControlTop of page
Due to the variable regulations around (de)registration of pesticides, your national list of registered pesticides or relevant authority should be consulted to determine which products are legally allowed for use in your country when considering chemical control. Pesticides should always be used in a lawful manner, consistent with the product's label.
The primary form of long range dispersal is thought to be through uptake and transit of resting cysts in cargo vessel ballast water (Hallegraeff and Bolch, 1992). To combat this vector a number of voluntary measures and international guidelines have been implemented by the international shipping industry (International Maritime Organisation (IMO)) to limit the uptake, transport across oceanic basins, and release of large quantities of viable organisms in ballast water (see Gollasch et al., 2007 for details). A variety of treatment options for ballast water are currently being investigated. Filtration to remove organisms >25 µm is potentially achievable (Cangelosi et al., 2007) and may reduce the concentration of G. catenatum resting cysts in discharged ballast; however, considerably more research is required before this approach is taken up on a larger scale. Treatment of ballast water with biocides or other chemicals is either ineffective or not economically feasible at effective concentrations (Bolch and Hallegraeff, 1993; Gregg and Hallegraeff, 2007). There are also serious safety and/or corrosion issues associated with the practical application of biocides on vessels (Gollasch et al., 2007).
Eradication of established populations is not currently feasible or achievable.
Local containment does not appear to be feasible. Institution of measures controlling shellfish transfers and managing ballast water movement/discharge (Gollasch et al. 2007) may reduce the risk of primary intercontinental or secondary transfer within regions.
See comments under Containment/zoning.
Biological control methods for phytoplankton species are currently not feasible, and too little is known of pathogen/parasite interactions to devise specific control strategies.
Chemical control is not feasible as it is impossible to confine the effects of any chemical application specifically to G. catenatum.
Gaps in Knowledge/Research NeedsTop of page
The bulk of available published research on G. catenatum is confined to populations in northwest Spain, and the introduced Australian populations, and to a lesser extent, the populations of southern Japan. Considerably more information is required from a wider range of regional populations to begin to understand the biogeography and dispersal of this species. Sediment dating studies of resting cysts from a range of affected areas would allow a better appraisal of the introduced or native status of particular populations. Considerably more basic biological and ecological and genetic information on this species may allow improved detection and understanding of factors controlling bloom initiation and decline, and the development/control of cellular toxicity.
ReferencesTop of page
Attaran-Fariman G; Salas MF de; Negri AP; Bolch CJS, 2007. Morphology and phylogenetic affinities of Gymnodinium trapeziforme sp. nov. (Dinophyceae): A new dinoflagellate from the southeast coast of Iran that forms microreticulate resting cysts. Phycologia, 46:644-656.
Band-Schmidt C; Bustillos-Guzman J; Morquecho L; Garate-Lizarraga I; Alonso-Rodriguez R; Reyes-Salinas A; Erler K; Luckas B, 2006. Variations of PSP toxin profiles during different growth phases in Gymnodinium catenatum (Dinophyceae) strains isolated from three locations in the Gulf of California, Mexico. J. Phycol, 42:757-768.
Barbera-Sánchez A la; Gamboa-Maruez JF, 2001. Distribution of Gymnodinium catenatum Graham and shellfish toxicity on the coast of Sucre State, Venezuela, from 1989 to 1998. Journal of Shellfish Research, 20:1257-1261.
Barbera-Sanchez A; Hall S; Ferraz-Reyes E, 1993. Alexandrium sp., Gymnodinium catenatum and PSP in Venezuela. In: Toxic Phytoplankton Blooms in the Sea [ed. by Smayda TJ, Shimizu Y] New York, USA: Elsevier Science Publishers BV, 281-285.
Blackburn SI; Bolch CJS; Haskard KA; Hallegraeff GM, 2001. Reproductive compatibility among four global populations of the toxic dinoflagellate Gymnodinium catenatum (Dinophyceae). Phycologia, 40:78-87.
Blackburn SI; Hallegraeff GM; Bolch CJ, 1989. Vegetative reproduction and sexual life cycle of the toxic dinoflagellate Gymnodinium catenatum Graham from Tasmania, Australia. J. Phycol, 25(3):577-590.
Bolch CJ; Negri A; Hallegraeff GM, 1999. Gymnodinium microreticulatum sp. nov. (Dinophyceae): a naked, microreticulate cyst producing dinoflagellate, distinct from Gymnodinium catenatum Graham and Gymnodinium nolleri Ellegaard et Moestrup. Phycologia, 38:301-313.
Bolch CJS; Salas MF de, 2007. A review of the molecular evidence for ballast water introduction of the toxic dinoflagellates Gymnodinium catenatum and the Alexandrium "tamarensis complex" to Australasia. Harmful Algae, 6(4):465-485. http://www.sciencedirect.com/science/journal/15689883
Cangelosi AA; Mays NL; Balcer MD; Reavie ED; Reid DM; Sturtevant R; Gao XQ, 2007. The response of zooplankton and phytoplankton from the North American Great Lakes to filtration. Harmful Algae, 6(4):547-566. http://www.sciencedirect.com/science/journal/15689883
Dale B; Madsen A; Nordberg K; Thorsen TA, 1993. Evidence for prehistoric "blooms" of the toxic dinoflagellate Gymnodinium catenatum in the Kattegat-Skagerrak region of Scandinavia. In: Toxic Phytoplankton Blooms in the Sea [ed. by Smayda TJ, Shimizu Y] New York, USA: Elsevier Science Publishers B.V, 47-52.
Daugbjerg N; Hansen G; Larsen J; Moestrup Ø, 2000. Phylogeny of some of the major genera of dinoflagellates based on ultrastructure and partial LSU rDNA sequence data, including the erection of three new genera of unarmoured dinoflagellates. Phycologia, 39:302-317.
Ellegaard M; Christensen NF; Moestrup Ø, 1993. Temperature and salinity effects on growth of a non-chain-forming strain of Gymnodinium catenatum (Dinophyceae) established from a cyst from Recent sediments in The Sound (Oresund), Denmark. J. Phycol, 29:418-426.
Ellegaard M; Oshima Y, 1998. Gymnodinium nolleri Ellegaard et Moestrup sp. ined. (Dinophyceae) from Danish waters, a new species producing Gymnodinium catenatum-like cysts: molecular and toxicological comparisons with Australian and Spanish strains of Gymnodinium catenatum. Phycologia, 37:369-378.
Franca S; Almeida JF, 1989. Paralytic shellfish poisons in bivalve molluscs on the Portugese coast caused by a bloom of the dinoflagellate Gymnodinium catenatum. In: Red Tides: Biology, Envionmental Science and Toxicology [ed. by Okaichi T, Anderson DM, Nemoto T] New York, USA: Elsevier Science Publishers BV, 93-96.
Fukuyo Y; Kodama M; Ogata T; Ishimaru T; Matsuoka K; Okaichi T; Maala AM; Ordones JA, 1993. Occurrence of Gymnodinium catenatum in Manila Bay, the Philippines. In: Toxic Phytoplankton Blooms in the Sea [ed. by Smayda TJ, Shimizu Y] New York, USA: Elsevier Science Publishers BV, 875-880.
Giannakourou A; Orlova TY; Assimakopoulou G; Pagou K, 2005. Dinoflagellate cysts in recent marine sediments from Thermaikos Gulf, Greece: Effects of resuspension events on vertical cyst distribution. Cont. Shelf Res, 25:2585-2596.
Glibert PM; Landsberg JH; Evans JJ; Al-Sarawi MA; Muna Faraj; Al-Jarallah MA; Haywood A; Shahnaz Ibrahem; Klesius P; Powell C; Shoemaker C, 2002. A fish kill of massive proportion in Kuwait Bay, Arabian Gulf, 2001: the roles of bacterial disease, harmful algae, and eutrophication. Harmful Algae, 1(2):215-231.
Gollasch S; David M; Voigt M; Dragsund E; Hewitt C; Fukuyo Y, 2007. Critical review of the IMO international convention on the management of ships' ballast water and sediments. Harmful Algae, 6(4):585-600. http://www.sciencedirect.com/science/journal/15689883
Gregg MD; Hallegraeff GM, 2007. Efficacy of three commercially available ballast water biocides against vegetative microalgae, dinoflagellate cysts and bacteria. Harmful Algae, 6(4):567-584. http://www.sciencedirect.com/science/journal/15689883
Hallegraeff GM; Fraga S, 1998. Bloom dynamics of the toxic dinoflagellate Gymnodinium catenatum, with emphasis on Tasmanian and Spanish coastal waters. In: Physiological Ecology of Harmful Algal Blooms [ed. by Anderson DM, Cembella AD, Hallegraeff GM] Heidelberg, Germany: Springer-Verlag, 59-80.
Hallegraeff GM; Stanley SO; Bolch CJ; Blackburn SI, 1989. Gymnodinium catenatum blooms and shellfish toxicity in Southern Tasmania, Australia. In: Red Tides: Biology, Envionmental Science and Toxicology [ed. by Okaichi T, Anderson DM, Nemoto T] New York, USA: Elsevier Science Publishers BV, 77-80.
Holmes MJ; Bolch CJS; Green DH; Cembella AD; Ming Teo SL, 2002. Singapore isolates of the dinoflagellate Gymnodinium catenatum (Dinophyceae) produce a unique profile of paralytic shellfish poisoning toxins. J. Phycol, 38:96-106.
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19/12/07 Original text by:
Christopher Bolch, University of Tasmania, Australia
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