- 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
- Links to Websites
- Distribution Maps
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PicturesTop of page
IdentityTop of page
Preferred Scientific Name
- Alexandrium minutum Halim, 1960
Other Scientific Names
- Alexandrium ibericum Balech, 1985
- Alexandrium lusitanicum Balech, 1985
Summary of InvasivenessTop of page
A. minutum is a photosynthetic dinoflagellate that, like many species in its genus, is responsible for outbreaks of Paralytic Shellfish Poisoning (PSP). This phytoplankton species can also form extremely dense blooms that have the capacity to kill finfish, in addition to their PSP toxin production. As this species forms a tough resting cyst, it is easily transport by ballast water and in translocated shellfish, and it has been reported from most continents and every ocean. Control appears to be impossible.
Taxonomic TreeTop of page
- Domain: Eukaryota
- Kingdom: Protista
- Phylum: Protozoa
- Class: Dinophyceae
- Order: Gonyaulacales
- Family: Gonyaulacaceae
- Genus: Alexandrium
- Species: Alexandrium minutum
Notes on Taxonomy and NomenclatureTop of page
Among the armoured dinoflagellates, the genus Alexandrium has an even more confusing taxonomic history than most. Species in this genus have been classified in the genera Gessnerium Halim, Goniodoma Stein, Gonyaulax Diesing, Protogonyaulax Taylor and Pyrodinium Plate. Of those species that are now classified as Alexandrium, the first to be described was Goniodoma ostenfeldii, by Paulsen (1904). He then transferred this species to Gonyaulax in 1949. It was not until the 1970s that a distinct group was recognized within Gonyaulax, then referred to as “Gonyaulax of the tamarensis or catenella group”.
It was in 1960 that Halim discovered a small dinoflagellate in Alexandria harbour, Egypt, for which he created the new genus Alexandrium. Balech (1989) recognized that species in the “tamarensis group” were very similar to the species in Halim’s genera, of which Alexandrium had chronological priority, and transferred them into this genus.
According to Balech (1995), species in this genus are characterized by their Kofoidian thecal plate formula Po, 4’, 6”, 5”’, 2””, 6c and 9-10s. Most species have thin and smooth thecal plates, though Alexandrium minutum can sometimes have a reticulated hypotheca (Montresoret al., 1990; Balech, 1995).
There are two distinct genotypes that comprise the morphospecies A. minutum, one of which occurs in Europe and Western Australia, the other in the north- and south-Pacific. There is some speculation that the Pacific genotype of A. minutum may be an altogether different species (Montresoret al., 2004). However, as they have not yet been separated, both genotypes are here treated as one species.
DescriptionTop of page
Plant TypeTop of page Aquatic
DistributionTop of page
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.
|Continent/Country/Region||Distribution||Last Reported||Origin||First Reported||Invasive||Reference||Notes|
|Atlantic, Antarctic||Absent, intercepted only||Lilly et al., 2005|
|Atlantic, Eastern Central||Absent, intercepted only||Lilly et al., 2005|
|Atlantic, Northeast||Widespread||Native||Lilly et al., 2005||Portugal to Norway|
|Atlantic, Northwest||Present, few occurrences||Lilly et al., 2005|
|Atlantic, Southeast||Localised||Lilly et al., 2005||South Africa|
|Atlantic, Southwest||Absent, intercepted only||Lilly et al., 2005|
|Atlantic, Western Central||Absent, intercepted only||Lilly et al., 2005|
|Indian Ocean, Antarctic||Absent, intercepted only||Lilly et al., 2005|
|Indian Ocean, Eastern||Localised||Lilly et al., 2005|
|Indian Ocean, Western||Absent, intercepted only||Lilly et al., 2005|
|Mediterranean and Black Sea||Widespread||Native||Lilly et al., 2005||Widespread in the Mediterranean|
|Pacific, Antarctic||Localised||Native||Lilly et al., 2005||Great Australian Bight|
|Pacific, Eastern Central||Absent, intercepted only||Lilly et al., 2005|
|Pacific, Northeast||Absent, intercepted only||Lilly et al., 2005|
|Pacific, Northwest||Localised||Native||Lilly et al., 2005||Japan|
|Pacific, Southeast||Absent, intercepted only||Lilly et al., 2005|
|Pacific, Southwest||Localised||Native||Lilly et al., 2005||New Zealand, Australia|
|Pacific, Western Central||Localised||Native||Lilly et al., 2005||Vietnam, Thailand, Malaysia|
|China||Present, few occurrences||Liu and Liang, 2008|
|India||Present, few occurrences||Godhe et al., 2001|
|-Karnataka||Localised||Godhe et al., 2001||Offshore Mangalore|
|Japan||Widespread||Lilly et al., 2005|
|Kuwait||Widespread||2001||Glibert et al., 2002|
|Malaysia||Localised||Usup et al., 2002|
|-Peninsular Malaysia||Localised||Usup et al., 2002|
|Taiwan||Widespread||Lilly et al., 2005|
|Thailand||Localised||Piumsomboon et al., 2001||Prakan River Estauary|
|Vietnam||Localised||2000||Yoshida et al., 2000|
|South Africa||Localised||2003||McCauley et al., 2008||Cape Town Habour|
|Tunisia||Localised||Dally et al., 2001|
|-New York||Localised||Balech, 1995||Minneola|
Central America and Caribbean
|Jamaica||Localised||Hansen et al., 2003|
|Croatia||Localised||Marasovic et al., 1995||Kastela Bay|
|Denmark||Present||Hansen et al., 2003||Korsor Nor|
|Ireland||Localised||Hansen et al., 2003|
|Italy||Widespread||Montresor et al., 1990|
|Norway||Localised||Balech and Tangen, 1985||Oslofjord|
|Portugal||Localised||Franco et al., 1995|
|Spain||Widespread||Vila et al., 2001|
|-Balearic Islands||Localised||Forteza et al., 1998||Palma de Mallorca|
|Sweden||Localised||Hansen et al., 2003|
|UK||Localised||Nascimento et al., 2005|
|Australia||Widespread||de et al., 2001|
|-New South Wales||Localised||Emmerik MJvan, 1999|
|-South Australia||Localised||Bolch et al., 1991|
|-Western Australia||Localised||Swan River Trust, 1998|
|New Zealand||Widespread||Hansen et al., 2003|
History of Introduction and SpreadTop of page
In Australia, the first record for A. minutum is that of Hallegraeff (1988), closely followed by the work of Jean Cannon (1990), on the discovery of this species in the Port River (Adelaide, South Australia). Shortly after this its resting cyst was described (Bolchet al., 1991), and in a survey of the genus in south-eastern Australian waters, reported only from this locality (Hallegraeff et al., 1991).
Risk of IntroductionTop of page
HabitatTop of page
Habitat ListTop of page
|Mud flats||Present, no further details|
|Intertidal zone||Present, no further details|
|Lagoons||Secondary/tolerated habitat||Harmful (pest or invasive)|
|Inshore marine||Principal habitat||Harmful (pest or invasive)|
|Inshore marine||Principal habitat||Natural|
|Pelagic zone (offshore)||Principal habitat|
|Benthic zone||Secondary/tolerated habitat|
Biology and EcologyTop of page Genetics
Several ribosomal RNA gene sequences of A. minutum can be found in GenBank. They fall broadly into two genotypes, described by Lilly et al. (2005) as the “Global” and “Pacific” clades. They form a monophyletic group, indicating either that they are all one species, or if they should be separated into distinct species, they still would be genetically more closely related to each other than to any other species.
Phylogenetically, the species A. minutum as a whole forms part of what is known as the A. minutum group within the Alexandrium clade, which also includes A. insuetum, A. tamutum, A. ostenfeldii and A. andersoni (Lilly et al., 2005).Reproductive Biology
A. minutum has a typical, sexual dinoflagellate life cycle with a resting cyst capable of persisting in sediments for many years (Bolch et al., 1991). This species also has a complex mating system (Blackburn and Parker, 2005) in which certain strains only mate with other, compatible strains (complex heterothallic). This probably reflects recognition of more than one surface protein.Environmental Requirements
Given the degree of genetic differentiation between the two clades of A. minutum, it is not surprising that this species shows a vast range of environmental tolerances, from cold-temperate to fully tropical, and spanning a wide range of salinities and nutrient conditions. As stated above, this species is present in a range of water temperatures, though it seems to prefer lower salinity conditions present in estuarine systems.
Water TolerancesTop of page
|Parameter||Minimum Value||Maximum Value||Typical Value||Status||Life Stage||Notes|
|Depth (m b.s.l.)||Optimum||Photic zone|
|Salinity (part per thousand)||15||35||Optimum||Estuarine and coastal areas|
|Water temperature (ºC temperature)||15||25||Optimum||7-35 tolerated, found from tropical to cool-temperate waters|
Notes on Natural EnemiesTop of page
Means of Movement and DispersalTop of page Natural Dispersal (Non-Biotic)
This species is primarily an inhabitant of environments with a high terrestrial influence, such as lagoons and estuaries where nutrient levels are high, the water column is stratified, and mechanical disturbance low. In spite of this, A. minutum is able to grow in coastal areas where land-derived influences are low. As such it is capable of dispersal along natural currents downstream from seed populations in estuaries, lagoons and bays.Vector Transmission (Biotic)
No evidence for vector-mediated introduction exists for this species.Accidental Introduction
Translocated shellfish stock originating from infested areas can act as a transmission vector (Scholin et al., 1995), and PCR detection methods have been developed to detect this species in contaminated mussels (Galluzzi et al., 2005).
Ballast water contamination is the single largest risk factor (Hallegraeff and Bolch, 1992), which could introduce A. minutum to new areas.Intentional Introduction
No evidence for intentional human-mediated introduction exists for this species.
Pathway CausesTop of page
Pathway VectorsTop of page
Impact SummaryTop of page
Economic ImpactTop of page
Environmental ImpactTop of page
Social ImpactTop of page
The reduction of shellfish consumption following outbreaks of Paralytic Shellfish Poisoning is significant and has a negative impact on the industry in addition to the damage caused by the cessation of shellfish harvesting.
Risk and Impact FactorsTop of page Invasiveness
- Invasive in its native range
- Has a broad native range
- Abundant in its native range
- Highly adaptable to different environments
- Highly mobile locally
- Long lived
- 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
- Competition - smothering
- Interaction with other invasive species
- Rapid growth
- Highly likely to be transported internationally accidentally
- Highly likely to be transported internationally deliberately
- Highly likely to be transported internationally illegally
- Difficult to identify/detect as a commodity contaminant
- Difficult to identify/detect in the field
- Difficult/costly to control
UsesTop of page Economic Value
There is a low potential for developing toxin standards from strains that produce large amounts of a single toxin, and for the production of saxitoxin as a chemical weapon.
Uses ListTop of page
- Laboratory use
- Research model
- Miscellaneous materials
Detection and InspectionTop of page
- Microscopical examination of sediment and plankton samples
- PCR detection
- rRNA-targeted fluorescent in-situ hybridisation and sandwich hybridisation.
Similarities to Other Species/ConditionsTop of page
Prevention and ControlTop of page
Ballast water containment. Shellfish translocation prevention or quarantine. May already be established (sometimes below detection levels) in most places.
Ballast water contamination is the single largest risk factor (Hallegraeff and Bolch, 1992). Filtration to remove this species (Cangelosi et al., 2007) would need to be under 25 µm, making it not feasible. Treatment with biocides is ineffective with cyst-forming species (Bolch and Hallegraeff, 1993; Gregg and Hallegraeff, 2007) and also poses corrosion issues. Heat treatment is a possibility though not yet implemented successfully in commercial situations (Hallegraeff, 1998).
Early warning systems
Possibility of automated quantitative detection with qPCR probes. Several have been developed (Galluzziet al., 2004; Galluzzi et al., 2005; Galluzziet al., 2006).Rapid response
Not applicable. Closure of shellfish harvesting at higher than legislated PSP levels in shellfish meat.
Prevention of ballast water, shellfish and sediment exchange between contaminated and non-contaminated areas may prevent non-natural dispersal. However this species is able to grow in coastal areas with low land influence, allowing fairly long-distance natural dispersal.
No management is feasible once a seed population is established.
Biological control is not currently developed for phytoplankton species. There are some natural parasites of this species (Erard-Le Denn et al., 2000), which may eventually be able to be developed for biological control. Recent work on marine viruses affecting a range of other species (Brussaard et al., 1996; Larsen et al., 2001; Brussaard et al., 2004a; Brussaard et al., 2004b; Baudoux and Brussaard, 2005; Nagasaki et al., 2005) open a potential avenue for future research in this direction.
Not feasible to target single cells.
Not applicable. Damage control by closing shellfish harvesting if bloom causes PSP levels to exceed legal limits.
Gaps in Knowledge/Research NeedsTop of page
ReferencesTop of page
Balech E; Tangen K, 1985. Morphology and taxonomy of toxic species in the tamarensis group (Dinophyceae) Alexandrium excavatum (Braarud) comb. nov. and Alexandrium ostenfeldii (Paulsen) comb. nov. Sarsia, 70:333-343.
Baudoux AC; Brussaard CPD, 2005. Characterization of different viruses infecting the marine harmful algal bloom species Phaeocystis globosa. Virology, 341(1):80-90. http://www.sciencedirect.com/science/journal/00426822
Belin C, 1993. Distribution of Dinophysis spp. and Alexandrium minutum along French coasts since 1984 and their DSP and PSP toxicity levels. , Elsevier, pp. In: Toxic Phytoplankton Blooms in the Sea [ed. by Smayda TJ, Shimizu Y] Amsterdam, Netherlands: Elsevier, 469-474.
Brussaard CPD; Short SM; Frederickson CM; Suttle CA, 2004. Isolation and phylogenetic analysis of novel viruses infecting the phytoplankton Phaeocystis globosa (Prymnesiophyceae). Applied and Environmental Microbiology, 70(6):3700-3705.
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
Cannon JA, 1990. Development and dispersal of red tides in the Port River, South Australia. New York, Elsevier. In: Toxic Marine Phytoplankton [ed. by Graneli E, Sundström B, Edler L, Anderson DM] New York, USA: Elsevier, 110-115.
Cembella AD; Sullivan JJ; Boyer GL; Taylor FJR; Anderson RJ, 1987. Variation in paralytic shellfish toxin composition within the Protogonyaulax tamarensis / catenella species complex: red tide dinoflagellates. Biochemical Systematics & Ecology, 15(2):171-186.
de Salas MF; van Emmerik MJ; Hallegraeff GM; Negri AP; Vaillancourt RE; Bolch CJS, 2001. Toxic Australian Alexandrium dinoflagellates: introduced or indigenous? In: Harmful Algal Blooms 2000 [ed. by Hallegraeff G, Bolch C, Blackburn S, Lewis , R] Paris, France: UNESCO, 214-217.
Delgado M; Estrada M; Camp J; Fernandez JJ; Santmarti M; Lleti C, 1990. Development of a toxic Alexandrium minutum Halim (Dinophyceae) bloom in the herbour of Sant Carles de la Ràpita (Ebro Delta, Northwestern Mediterranean). Scientia Marina, 54(1):1-7.
Forteza V; Quetglas G; Delgado M; Reyero MI; Fraga S; Franco JM; Cacho E, 1998. Toxic Alexandrium minutum bloom in Palma de Mallorca Harbour, (Balearic Islands, Western Mediterranean). In: Harmful Algae [ed. by Reguera B, Blanco J, Fernandez ML, Wyatt T] Santiago de Compostela, Spain: Xunta de Galicia and IOC of UNESCO, 58-59.
Franco JM; Fraga S; Zapata M; Bravo I; Fernandez P; Ramilo I, 1995. Comparison between strains of genus Alexandrium of the minutum group. In: Harmful Marine Algal Blooms [ed. by Lassus P, Arzul G, Erard E, Gentien P, Marcaillou C] Paris, France: Lavoisier/Intercept, 3-13.
Fukuyo Y, 1985. Morphology of Protogonyaulax tamarensis (Lebour) Taylor and Protogonyaulax catenella (Whedon and Kofoid) Taylor from Japanese coastal waters. Bulletin of Marine Science, 37(2):529-537.
Fukuyo Y; Pholpunthin Y, 1990. Alexandrium cohorticula (Balech) Balech. In: Red Tide Organisms in Japan - an Illustrated Guide [ed. by Fukuyo Y, Takano H, Chihara M, Matsuoka K] Tokyo, Japan: Uchida Rokakuho Co. Ltd, 88-89.
Fukuyo Y; Pholpunthin Y, 1990. Alexandrium leei (Balech) Balech. In: Red Tide Organisms in Japan - an Illustrated Guide [ed. by Fukuyo Y, Takano H, Chihara M, Matsuoka K] tokyo, Japan: Uchida Rokakuho Co. Ltd, 92-93.
Galluzzi L; Bertozzini E; Campo Adel; Penna A; Bruce IJ; Magnani M, 2006. Capture probe conjugated to paramagnetic nanoparticles for purification of Alexandrium species (Dinophyceae) DNA from environmental samples. Journal of Applied Microbiology, 101(1):36-43. http://www.blackwell-synergy.com/doi/pdf/10.1111/j.1365-2672.2006.02952.x
Galluzzi L; Penna A; Bertozzini E; Giacobbe MG; Vila M; Garcés E; Prioli S; Magnani M, 2005. Development of a qualitative PCR method for the Alexandrium spp. (Dinophyceae) detection in contaminated mussels (Mytilus galloprovincialis). Harmful Algae, 4(6):973-983. http://www.sciencedirect.com/science/journal/15689883
Galluzzi L; Penna A; Bertozzini E; Vila M; Garcés E; Magnani M, 2004. Development of a real-time PCR assay for rapid detection and quantification of Alexandrium minutum (a dinoflagellate). Applied and Environmental Microbiology, 70(2):1199-1206.
Garces E; Bravo I; Vila M; Figueroa RI; Maso M; Sampedro N, 2004. Relationship between vegetative cells and cyst production during Alexandrium minutum bloom in Arenys de Mar harbour (NW Mediterranean). Journal of Plankton Research, 26(6):637-645.
Giacobbe MG; Oliva FD; Maimone G, 1996. Environmental factors and seasonal occurrence of the dinoflagellate Alexandrium minutum, a PSP potential producer, in a Mediterranean lagoon. Estuar. Coast. Shelf Sci, 42:539-549.
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.
Godhe A; Otta SK; Rehnstam-Holm AS; Karunasagar I; Karunasagar I, 2001. Polymerase chain reaction in detection of Gymnodinium mikimotoi and Alexandrium minutum in field samples from southwest India. Marine Biotechnology, 3(2):152-162.
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, 1998. Transport of toxic dinoflagellates via ships ballast water - bioeconomic risk assessment and efficacy of possible ballast water management strategies. Marine Ecology-Progress Series, 168:297-309.
Hallegraeff GM; Bolch CJ, 1992. Transport of diatom and dinoflagellate resting spores in ships' ballast water: implications for plankton biogeography and aquaculture. Journal of Plankton Research, 14(8):1067-1084.
Hansen G; Daugbjerg N; Franco JM, 2003. Morphology, toxin composition and LSU rDNA phylogeny of Alexandrium minutum (Dinophyceae) from Denmark, with some morphological observations on other European strains. Harmful Algae, 2(4):317-335.
Honsell G, 1993. First report of Alexandrium minutum in northern Adriatic waters (Mediterranean Sea). In: Toxic Phytoplankton Blooms in the Sea [ed. by Smayda TJ, Shimizu Y] Amsterdam, Netherlands: Elsevier, 127-132.
Hwang DengFwu; Tsai YungHsiang; Liao HsuanJung; Matsuoka K; Noguchi T; Jeng SenShyong, 1999. Toxins of the dinoflagellate Alexandrium minutum Halim from the coastal waters and aquaculture ponds in southern Taiwan. Fisheries Science, 65(1):171-172.
Larsen A; Castberg T; Sandaa RA; Brussaard CPD; Egge J; Heldal M; Paulino A; Thyrhaug R; van Hannen EJ; Bratbak G, 2001. Population dynamics and diversity of phytoplankton, bacteria and viruses in a seawater enclosure. Marine Ecology-Progress Series, 22:147-157.
Lilly EL; Halanych KM; Anderson DM, 2005. Phylogeny, biogeography, and species boundaries within the Alexandrium minutum group. Harmful Algae, 4(6):1004-1020. http://www.sciencedirect.com/science/journal/15689883
Marasovic I; Nincevic Z; Odzak N, 1995. The effect of temperature on blooms for Lingulodinium polyedrum and Alexandrium minutum in Kastela Bay. In: Harmful Marine Algal Blooms [ed. by Lasus PA, Erard-Le-Denn G, Gentien P, Maraillou-Le-Baut C] Paris, France: Lavoisier, 187-192.
McCauley LAR; Erdner DL; Nagai S; Richlen ML; Anderson DM, 2008. Biogeographic analysis of the globally distributed harmful algal bloom species Alexandrium minutum (Dinophyceae) based on rRNA gene sequences and microsatellite markers. Journal of Phycology.
Nagasaki K; Shirai Y; Takao Y; Mizumoto H; Nishida K; Tomaru Y, 2005. Comparison of genome sequences of single-stranded RNA viruses infecting the bivalve-killing dinoflagellate Heterocapsa circularisquama. Applied and Environmental Microbiology, 71(12):8888-8894.
Nascimento SM; Purdie DA; Lilly EL; Larsen J; Morris S, 2005. Toxin profile, pigment composition, and large subunit rDNA phylogenetic analysis of an Alexandrium minutum (Dinophyceae) strain isolated from the Fleet Lagoon, United Kingdom. Journal of Phycology, 41:343-353.
Oshima Y; Hirota M; Yasumoto T; Hallegraeff G; Blackburn S; Steffensen D, 1989. Production of paralytic shellfish toxins by the dinoflagellate Alexandrium minutum Halim from Australia. Nipp. Suis. Gakk, 55:9-25.
Piumsomboon A; Songroop C; Kungsuwan A; Polpunthin P, 2001. Species of the dinoflagellate genus Alexandrium (Gonyaulacales) in the Gulf of Thailand. In: Harmful Algal Blooms 2000 [ed. by Hallegraeff GM, Blackburn SI, Bolch CJ, Lewis RJ] Hobart, Australia: Intergovernmetal Oceanographic Commission of UNESCO, 5-12.
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ContributorsTop of page
30/05/08 Original text by:
Miguel de Salas, University of Tasmania, School of Plant Science, Life Sciences Building, Australia
Distribution MapsTop of page
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