Prymnesium parvum (golden algae)
- Summary of Invasiveness
- Taxonomic Tree
- Notes on Taxonomy and Nomenclature
- 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
- Impact Summary
- Economic Impact
- Environmental Impact
- Risk and Impact Factors
- 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
- Prymnesium parvum N.Carter
Preferred Common Name
- golden algae
Other Scientific Names
- Prymnesium patelliferum J.C.Green, D.J.Hibberd & R.N.Pienaar
Summary of InvasivenessTop of page
P. parvum is an algal species that forms harmful blooms in inland and coastal aquatic environments and is responsible for devastating fish kills causing ecological and economic damage. While blooms of P. parvum were documented in the eastern hemisphere since the early 1900s, the species has now spread widely, with blooms occurring in all southern regions of the USA and some northern regions. P. parvum is not on an alert list or listed as a regulated pest.
Taxonomic TreeTop of page
- Domain: Eukaryota
- Kingdom: Plantae
- Phylum: Haptophyta
- Class: Prymnesiophyceae
- Family: Prymnesiaceae
- Genus: Prymnesium
- Species: Prymnesium parvum
Notes on Taxonomy and NomenclatureTop of page
In the older literature there are reports of both Prymnesium parvum and Prymnesiumpatelliferum. They were distinguishable by minor details in scale morphology seen only using transmission electron microscopy (Green et al., 1982). More recently, evidence was provided that indicated P. parvum and P. patelliferum were the same species representing different forms (Larsen and Medlin, 1997; Larsen and Edvardsen, 1998; Larsen 1999). Consequently, the names were emended to P. parvum f. parvum and P. parvum f. patelliferum.
DescriptionTop of page
P. parvum is a microscopic (about 10 µm), flagellated alga believed to have four morphologically distinct forms. Two forms are bi-flagellated haploid cell types. One form is a bi-flagellated diploid cell type. Another form is non-motile with no flagella, which might be a resting stage. The flagellated forms have a haptonema, a specialized external needle-like structure that enables attachment to surfaces. The haptonema does not appear to aid in particle capture and phagocytosis in P. parvum f. patelliferum (Tillman 1998), unlike what has been observed for another haptophyte taxa (Kawachi et al. 1991). P. parvum f. parvum can be either haploid or diploid. P. parvum f. patelliferum is haploid (Larsen and Edvardsen, 1998). P. parvum cells have two saddle shaped chloroplasts that are usually yellow-green to olive in colour.Green et al. (1982) found that the flagella range from 12-15 micrometers long and the flexible, non-coiling haptonema ranges from 3-5 micrometers long. Each cell has body scales of two types found in two layers with scales of the outer layer having narrow inflexed rims and those of the inner layer having wide, strongly inflexed rims (Green et al., 1982).
DistributionTop of page
P. parvum is found worldwide (Edvardsen and Paasche, 1998; Graneli et al., 2012), including the Baltic Sea, Europe, China, Australia, the USA and Morocco. It is most often associated with estuarine or marine waters, but it can also occur in inland waters that have a relatively high mineral content. The type locality is a brackish water pond in the Isle of Wight, England, UK.
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|
|China||Present||Present based on regional distribution.|
|-Ningxia||Present||Introduced||Invasive||Guo et al., 1996||Inland waters|
|-Shandong||Present||Introduced||Invasive||Guo et al., 1996||Inland waters|
|-Shanxi||Present||Introduced||Invasive||Guo et al., 1996||Inland waters|
|-Tianjin||Present||Introduced||Invasive||Guo et al., 1996||Inland waters|
|-Zhejiang||Present||Introduced||Invasive||Guo et al., 1996||Inland waters|
|Israel||Present||Introduced||Invasive||Gordon and Colorni, 2008||Aquaculture ponds|
|Mongolia||Present||Introduced||Invasive||Guo et al., 1996||Inland waters|
|Morocco||Present||Introduced||Invasive||Sabour et al., 2000||Oued Mellah Lake|
|USA||Present||Present based on regional distribution.|
|-Alabama||Present||Introduced||Invasive||Tomas et al., 2004; Sager et al., 2008|
|-Arizona||Present||Introduced||Invasive||Sager et al., 2008|
|-Arkansas||Present||Introduced||Invasive||Tomas et al., 2004; Sager et al., 2008|
|-California||Present||Introduced||Invasive||Tiffany et al., 2007; Sager et al., 2008||Salton Sea|
|-Colorado||Present||Introduced||Invasive||Roelke et al., 2016|
|-Florida||Present||Introduced||Invasive||Sager et al., 2008|
|-Georgia||Present||Introduced||Invasive||Tomas et al., 2004|
|-Hawaii||Present||Introduced||Invasive||Roelke et al., 2016|
|-Louisiana||Present||Introduced||Invasive||Sager et al., 2008|
|-Maine||Present||Introduced||Invasive||Sager et al., 2008|
|-Mississippi||Present||Introduced||Invasive||Sager et al., 2008|
|-Nebraska||Present||Introduced||Invasive||Roelke et al., 2016|
|-New Mexico||Present||Introduced||Invasive||Sager et al., 2008|
|-North Carolina||Present||Introduced||Invasive||Tomas et al., 2004; Barreto et al., 2011||Artesian Aquaculture facility|
|-Oklahoma||Present||Introduced||2004||Invasive||Hambright et al., 2010||Lake Texoma|
|-Pennsylvania||Present||Introduced||2009||Invasive||Renner, 2009; Brooks et al., 2011||Dunkard Creek|
|-South Carolina||Present||Introduced||Invasive||Lewitus et al., 2003; Tomas et al., 2004||Golf course ponds|
|-Texas||Present||Introduced||Invasive||Tomas et al., 2004; Roelke et al., 2010; Dawson et al., 2015||Several lakes throughout central and western regions of state|
|-Washington||Present||Introduced||Invasive||Sager et al., 2008|
|-West Virginia||Present||Introduced||2009||Invasive||Renner, 2009; Brooks et al., 2011||Appalachian Stream|
|-Wyoming||Present||Introduced||Invasive||Sager et al., 2008|
|Germany||Present||Introduced||Invasive||Dietrich and Hesse, 1990||Coastal waters|
|Greece||Present||Introduced||Invasive||Moustaka-Gouni et al., 2004; Michaloudi et al., 2009||Lake Koronia|
|Hungary||Present||Introduced||Invasive||Vasas et al., 2012||Shallow lakes|
|Italy||Present||Introduced||Invasive||Mattioli and Simoni, 1999||Shallow lakes|
|Netherlands||Present||Liebert and Deerns, 1920; Otterstrøm and Steemann-Nielsen, 1940|
|Norway||Present||Johnsen and Lein, 1989; Eikrem and Throndsen, 1993||Coastal waters|
|Spain||Present||Introduced||Invasive||Comin and Ferrer, 1978||Ebro Delta|
|Sweden||Present||Introduced||Invasive||Johnsen et al., 2010||Ryfylkefjordane fjord|
|UK||Present||Introduced||Invasive||Holdway et al., 1978; Watson, 2001||Coastal waters|
|Australia||Present||Present based on regional distribution.|
|-Western Australia||Present||Introduced||Invasive||Hallegraeff, 1992||Coastal waters|
History of Introduction and SpreadTop of page
While blooms of P. parvum were documented in the eastern hemisphere since the early 1900s (Liebert and Deerns, 1920), documented blooms in the western hemisphere only started in the mid-1980’s, and then only localized in the south-central USA. The first confirmed blooms of P. parvum in North America were identified in Texas in 1985 on the Pecos River (James and Cruz, 1989), although based on physical characteristics of these blooms, Rhodes and Hubbs (1992) speculated that P. parvum blooms could have been responsible for fish kills in this river since the 1960s. A rapid proliferation of blooms in the western hemisphere occurred during the early 2000s, with Sager et al. (2008) reporting that the alga had invaded reservoirs and river systems in 15 other U.S. states, including Alabama, Arizona, Arkansas, California, Washington, Hawaii, New Mexico, Wyoming, North Carolina, South Carolina, Florida, and Georgia. Blooms now occur in all southern regions of the USA and some northern regions (Roelke et al., 2016).
Genetic analysis of P. parvum in different locations in the USA and from other locales around the world led Lutz-Carillo et al. (2010) to suggest that P. parvum in the USA originated from Europe, and that there may have been multiple invasion events.
Risk of IntroductionTop of page
Without knowledge of the origins of P. parvum in the eastern hemisphere, it is not possible to infer mechanisms that might have facilitated proliferation of blooms there. In the western hemisphere, multiple factors likely contributed to the spread of P. parvum blooms: these include changes in salinity, hydrology and nutrient concentration ratios.
In the south-central USA, where P. parvum blooms appear to have first occurred in the western hemisphere, salinity increases likely played a strong role enabling blooms. During the period of late 1990’s through early 2000s drought persisted in this region and salinity rose to a range of ~2 to ~4 psu in many lake and river systems. Of the many environmental variables measured during monitoring studies, elevated salinity related most strongly to P. parvum blooms (Roelke et al., 2012; Patiño et al., 2014; VanLandeghem et al., 2014; Hambright et al., 2015).
Lower river inflows to lakes during this period of drought might also have allowed P. parvum blooms to get established in the south-central USA. Experimentally, through deployment of in-lake mesocosm experiments, inflow events were shown to prevent bloom development (Hayden et al., 2012). In addition, field-monitoring studies showed that inflow events caused abrupt termination of blooms (Roelke et al., 2010, 2011; Jones et al., 2013). This notion was reinforced during follow-on, in-lake experiments (Hayden et al., 2012).
Long-term changes in the ratio between nutrients might have also played a role in enabling P. parvum blooms in the south-central USA. Laboratory experiments have shown that stoichiometric imbalance between nitrogen (N) and phosphorus (P) enhances toxicity to various organisms, which is probably the result of increased toxin production. When N:P was low or high, toxicity increased, with toxicity increasing more so in cultures studied when P was most limiting (Dafni et al., 1972; Johansson and Granéli, 1999; Granéli and Johansson, 2003b; Barreiro et al., 2005; Uronen et al., 2005; Valenti et al., 2010; Hambright et al., 2014).
HabitatTop of page
P. parvum occurs in brackish inland and coastal aquatic environments encompassing rivers, lakes, ponds, estuaries, lagoons and coastal oceans.
Habitat ListTop of page
|Lakes||Principal habitat||Harmful (pest or invasive)|
|Reservoirs||Principal habitat||Harmful (pest or invasive)|
|Rivers / streams||Principal habitat||Harmful (pest or invasive)|
|Rivers / streams||Principal habitat||Natural|
|Ponds||Principal habitat||Harmful (pest or invasive)|
|Estuaries||Principal habitat||Harmful (pest or invasive)|
|Lagoons||Principal habitat||Harmful (pest or invasive)|
|Inshore marine||Principal habitat||Harmful (pest or invasive)|
|Inshore marine||Principal habitat||Natural|
Biology and EcologyTop of page
P. parvum is a planktonic, single-celled, ellipsoid-shaped eukaryote. Its life cycle involves bi-flagellated forms and an unflagellated form. The flagellated forms are haploid and diploid. The unflagellated form might be a resting stage.
P. parvum supports population growth through autotrophy, where energy is obtained from sunlight, but also through heterotrophy, where energy is obtained from dissolved and particulate organic matter, i.e., saprophytic and phagocytic modes of nutrition (Fistarol et al., 2003; Granéli and Johansson, 2003a; Skovgaard and Hansen, 2003; Tillmann, 2003). Some chemicals produced by P. parvum are allelopathic and toxic to many other plankton, often resulting in death of other algae and zooplankton. The remains of killed organisms are then utilized by P. parvum. The killing effect of these chemicals also leads to P. parvum blooms that are near-monospecific with waters taking on a golden colour. For this reason, colloquially P. parvum is referred to as ‘golden algae’. It has been suggested that without the ecological benefits of its allelopathic and toxic chemicals, P. parvum appears to be a poor competitor and blooms do not occur (Roelke et al., 2007; Errera et al., 2008), although the role of allelopathy is disputed by some authors (Jonsson et al., 2009).
P. parvum is found at a wide range of salinities, being present in both coastal and inland waters. Studying one Scottish and two Israeli strains, McLaughlin (1958) found optimal NaCl concentrations of 0.3-6%, with growth possible at 0.2-10%. Larsen and Bryant (1998) reported that the Norwegian, Danish and English P. parvum strains they tested grew over a wide range of salinities each with different optimum growth concentrations, and that all three strains survived salinities from 3 to 30 psu (or 0.3%-3%). Dickson and Kirst (1987) speculated that the success of P. parvum in variable saline environments may be due to its ability to synthesize compatible solutes. Hambright et al. (2015) have found in Lake Texoma on the Oklahoma-Texas border that P. parvum presence and blooms are related positively to salinity and negatively to TN:TP ratio.
Shilo and Aschner (1953) observed that temperatures greater than 30°C were inhibitory to the growth of P. parvum, and 35°C resulted in lysis. McLaughlin (1958) reported erratic growth of Scottish and Israeli strains of P. parvum above 32°C.
Water TolerancesTop of page
|Parameter||Minimum Value||Maximum Value||Typical Value||Status||Life Stage||Notes|
|Depth (m b.s.l.)||Optimum||Surface. Blooms occur in photo zone|
|Salinity (part per thousand)||0.5||35||22||Optimum||Experimentally determined|
|Water pH (pH)||7||Optimum||pH below 7 ameliorates toxicity and organism loses its competitive edge|
|Water temperature (ºC temperature)||10||35||27||Optimum||Experimentally determined|
Notes on Natural EnemiesTop of page
Experimental evidence suggests that P. parvum may be vulnerable to viruses, algicidal bacteria and some zooplankton. In virus-free waters, P. parvum growth was better during in-lake mesocosm studies, suggesting a negative association between viral presence and P. parvum (Schwierzke et al., 2010). In bacteria-free waters, P. parvum growth was better during in-estuary mesocosm studies, but only during some seasons. This suggests an intermittent negative association between the bacteria and P. parvum (Lundgren et al., 2015). Regarding zooplankton, some (few) ciliates, rotifers and copepods were able to co-exist and even feed on toxic P. parvum (Koski et al., 1999; Rosetta and McManus, 2003; Michaloudi et al., 2009; Hambright et al., 2010; Schwierzke et al., 2010; Roelke et al., 2012; Davis et al., 2015; Lundgren et al., 2015).
Means of Movement and DispersalTop of page
The mechanism of P. parvum dispersal from the eastern hemisphere to the western is unknown. Within the western hemisphere, the mechanisms of dispersal are also unknown, but could involve facilitated invasions through interbasin water transfers, interbasin transportation on/in boats from recreational boaters, interbasin fish stocking activities, attachment to feathers of birds migrating across watersheds, or long distance transport during severe storms after becoming airborne (Roelke et al., 2016).
Impact SummaryTop of page
Economic ImpactTop of page
Some of the chemicals produced by P. parvum disrupt the function of gill tissue (Ulitzur and Shilo, 1966; Shilo 1967) leading to wreckage of fish and shellfish populations. Blooms often result in fish kills numbering in the 100,000s of individuals and costing local economies in the tens of millions of dollars (USD) (Southard et al. 2010). Losses to local economies in Texas during 2001 winter fish kills were estimated to exceed $18 million (Texas Parks and Wildlife Department, 2002).
Watson (2001) lists mass fish mortalities from many countries, both in natural waters and in aquaculture ponds. Guo et al. (1996) report recurrent kills in carp ponds in China due to P. parvum. The alga is also a problem in Tilapia culture in Israel, and barramundi culture in Australia (Seger et al., 2015). Hallegraeff (1992) suggest that caged fish are more vulnerable than wild fish stocks to P. parvum toxins as they cannot swim away from toxic areas.
Environmental ImpactTop of page
Ecological damages to fish communities from P. parvum blooms over long periods are variable, as populations in some watersheds appear to have recovered from recurrent blooms (Zamor et al., 2014), while in other watersheds they have not (VanLandeghem et al., 2013). Differences in fish recoveries between river systems are probably due to differing watershed attributes, e.g., rock and soil types, vegetation, precipitation and land-use practices, as well as varying lake morphologies (VanLandeghem et al., 2013).
In the 1986 event in the Pecos River, about 200 blue suckers (Cycleptus elongatus) and 3600 Rio Grande darters (Etheostoma grahami) were killed: both of these fish are on the Texas list of threatened animals (Texas Parks and Wildlife Department, 2002). Several state or federally endangered or threatened fish occur in areas of previous golden algal fish kills, and future blooms may affect these protected animals (Texas Parks and Wildlife Department, 2002).
Long-term ecological damages to shellfish populations from P. parvum blooms are less studied. Die-offs of several bivalve species were documented in the south-central USA where P. parvum blooms originated in the western hemisphere (James and De La Cruz 1989). In the 1986 kill in the Pecos River, all species in the area were affected, and after the bloom no live Corbicula fluminea (Asiatic clam, previously common in the river) were observed. In addition, well-established mussel beds have been destroyed in the area of the West Virginia-Pennsylvania border, part of P. parvum’s northward expansion (Brooks et al., 2011).
Risk and Impact FactorsTop of page Invasiveness
- Proved invasive outside its native range
- Highly adaptable to different environments
- Is a habitat generalist
- Tolerant of shade
- Highly mobile locally
- Reproduces asexually
- Altered trophic level
- Damaged ecosystem services
- Ecosystem change/ habitat alteration
- Modification of natural benthic communities
- Modification of nutrient regime
- Modification of successional patterns
- Monoculture formation
- Negatively impacts animal health
- Negatively impacts livelihoods
- Negatively impacts aquaculture/fisheries
- Negatively impacts tourism
- Reduced amenity values
- Reduced native biodiversity
- Threat to/ loss of endangered species
- Threat to/ loss of native species
- Antagonistic (micro-organisms)
- Highly likely to be transported internationally accidentally
- Difficult to identify/detect as a commodity contaminant
- Difficult/costly to control
UsesTop of page
Detection and InspectionTop of page
Traditional inverted microscopic techniques (bright field, phase contrast) are effective for P. parvum enumeration. To differentiate forms of P. parvum, transmission electron microscopy is required. Molecular techniques have also been developed for detection of P. parvum and activation of genes responsible for production of some toxins (La Claire, 2006; Medlin et al., 2010). Quantitative real-time PCR (qPCR) has been found to be an effective tool for detecting and monitoring P. parvum, particularly at pre-bloom densities (Zamor et al, 2012).
Similarities to Other Species/ConditionsTop of page
Within the Prymnesium genus there are seven species that are distinguishable by body scale morphology and/or DNA-sequences. All are of similar size, bi-flagellated and with haptonema. They are Prymnesium annuliferum, P. calathiferum, P. faveolatum, P. lepailleurii, P. nemametecum, P. parvum and P. zebrinum. The body scales on P. parvum cells are a diagnostic feature that can be used to distinguish it from related algal species (Green et al., 1982). The flagella-to-cell length ratio and the haptonema-to-cell length ratio are also important diagnostic features that aid in identifying this species (Chang and Ryan, 1985).
There are five additional species within the Prymnesium genus that have not been well characterized (Edvardsen and Imai, 2006) and are scarcely observed. These are P. czosnowskii, P. gladiociliatum,P. minutum, P. papillatum and the type species P. saltan.
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.
Landscape-scale reductions in nutrient loading and hydrologic manipulations would likely be required for management of P. parvum blooms at the ecosystem level (Hambright et al., 2010; Roelke et al., 2010, 2011). This has not occurred. Nor have attempts to control P. parvum blooms occurred at smaller scales, with the exception of aquaculture ponds (Barkoh and Fries, 2005; Barkoh et al., 2010; Kurten et al., 2007, 2010, 2011). From in-lake and laboratory experiments, there are promising approaches that include nutrient enrichment (Roelke et al., 2007; Errera et al., 2008), ammonium additions (Grover et al., 2013), pH lowering (Prosser et al., 2012), pulsed hydraulic flushing (Hayden et al., 2012; Lundgren et al., 2013), and herbicide application (Umphres et al., 2012, 2013). Chinese carp breeders have used suspended solids (mud), the addition of fertilizer (manure), and reduced salinity with varying degrees of success (Guo et al., 1996). Barkoh et al. (2004) point out that concentrations of ammonium sulfate required to control P. parvum may produce ammonia levels toxic to some fish.
Gaps in Knowledge/Research NeedsTop of page
While there have been great strides in developing approaches to manage P. parvum blooms, demonstration projects are still needed before significant funds will be committed to preventing blooms. In addition, information on trophic interactions between some of the organisms that appear unaffected by P. parvum’s toxins is needed. Finally, research directed at identifying toxic chemicals produced by P. parvum and under what environmental conditions is needed, as there is still much uncertainty in this area (Henrikson et al., 2010; Bertin et al., 2012a, b, 2014; Manning and La Claire, 2013; Blossom et al., 2014a, b).
ReferencesTop of page
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11/05/2016 Original text by:
Professor Dan Roelke, Texas A&M University, USA
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