Invasive Species Compendium

Detailed coverage of invasive species threatening livelihoods and the environment worldwide


Potamopyrgus antipodarum
(New Zealand mudsnail)



Potamopyrgus antipodarum (New Zealand mudsnail)


  • Last modified
  • 27 September 2018
  • Datasheet Type(s)
  • Invasive Species
  • Preferred Scientific Name
  • Potamopyrgus antipodarum
  • Preferred Common Name
  • New Zealand mudsnail
  • Taxonomic Tree
  • Domain: Eukaryota
  •   Kingdom: Metazoa
  •     Phylum: Mollusca
  •       Class: Gastropoda
  •         Subclass: Caenogastropoda
  • Summary of Invasiveness
  • P. antipodarum is an aquatic snail native to New Zealand. It has been introduced to Europe, North America, Australia, Iraq, Turkey and Japan. In several ecosystems it is considered invasive because it becomes h...

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Potamopyrgus antipodarum (New Zealand mudsnail); dorsal view.
TitleDorsal view
CaptionPotamopyrgus antipodarum (New Zealand mudsnail); dorsal view.
Copyright©U.S. Fish & Wildlife Service - Pacific Region/Dan Gustafson - CC BY 2.0
Potamopyrgus antipodarum (New Zealand mudsnail); dorsal view.
Dorsal viewPotamopyrgus antipodarum (New Zealand mudsnail); dorsal view.©U.S. Fish & Wildlife Service - Pacific Region/Dan Gustafson - CC BY 2.0
Potamopyrgus antipodarum (New Zealand mudsnail); lateral view.
TitleLateral view
CaptionPotamopyrgus antipodarum (New Zealand mudsnail); lateral view.
Copyright©Michal Maňas, via wikipedia - CC BY 4.0
Potamopyrgus antipodarum (New Zealand mudsnail); lateral view.
Lateral viewPotamopyrgus antipodarum (New Zealand mudsnail); lateral view.©Michal Maňas, via wikipedia - CC BY 4.0
Potamopyrgus antipodarum (New Zealand mudsnail); a colection of empty shells. Note scale in mm.
TitleEmpty shells
CaptionPotamopyrgus antipodarum (New Zealand mudsnail); a colection of empty shells. Note scale in mm.
Copyright©Mike Gangloff/ - CC BY-NC 3.0 US
Potamopyrgus antipodarum (New Zealand mudsnail); a colection of empty shells. Note scale in mm.
Empty shellsPotamopyrgus antipodarum (New Zealand mudsnail); a colection of empty shells. Note scale in mm.©Mike Gangloff/ - CC BY-NC 3.0 US


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Preferred Scientific Name

  • Potamopyrgus antipodarum (J. E. Gray, 1853)

Preferred Common Name

  • New Zealand mudsnail

Other Scientific Names

  • Hydrobia jenkinsi (Synonym) E. A. Smith 1884
  • Potamopyrgus jenkinsi (synonym) E. A. Smith 1889

International Common Names

  • English: jenkins' spire shell; New Zealand mudsnail
  • Spanish: caracol del cieno de Nueva Zelanda

EPPO code

  • POTAAN (Potamopyrgus antipodarum)

Summary of Invasiveness

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P. antipodarum is an aquatic snail native to New Zealand. It has been introduced to Europe, North America, Australia, Iraq, Turkey and Japan. In several ecosystems it is considered invasive because it becomes highly abundant, impacting the structure and function of the invaded ecosystems. Females are parthenogenetic, meaning they can reproduce without males, so a population can be founded by a single female. Most the non-native populations are female. There can be up to six generations per year, with an average number of 230 offspring per adult per year. P. antipodarum can also tolerate desiccation for several days, which allows for rapid spread (such as by birds and fishing tools) throughout different aquatic ecosystems. In several countries, including Spain, USA and Australia, it is considered as an invasive species.

Taxonomic Tree

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  • Domain: Eukaryota
  •     Kingdom: Metazoa
  •         Phylum: Mollusca
  •             Class: Gastropoda
  •                 Subclass: Caenogastropoda
  •                     Order: Littorinimorpha
  •                         Unknown: Rissooidea
  •                             Family: Hydrobiidae
  •                                 Genus: Potamopyrgus
  •                                     Species: Potamopyrgus antipodarum

Notes on Taxonomy and Nomenclature

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Potamopyrgus antipodarum is an aquatic snail of the family Hydrobiidae which is placed in the clade Littorinimorpha of the Caenogastropoda according to teh classification of Bouchet and Rocroi (2005). This is a cosmopolitan family comprising over 100 genera of small snails (Kabat and Hershler, 1993). Other names given to this species include Potamopyrgus jenkinsi (E. A. Smith, 1889) and Hydrobia jenkinsi (E. A. Smith, 1884). The common name is the New Zealand mudsnail. Several clones of this species have been identified in the invaded areas of Europe, America and Japan (Jensen and Forbes, 2001; Hershler et al., 2010; Hamada et al., 2013a).


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P. antipodarum is a prosobranch snail (Tateidae, Mollusca). In its non-native range it has a maximum shell size of 6-7 mm, but shell size can be up to 12 mm in its native New Zealand (Winterbourn, 1970). P. antipodarum has a solid operculum (i.e. a cover in the shell aperture) (Alonso and Castro-Díez, 2008) and its shell colour ranges from light to dark brown. Both males and females are morphologically very similar, but females have developing embryos in their reproductive systems (Jokela et al., 1997). The surface of the shell is characterized by right-handed coiling of 5-6 whorls. Some individuals have spines in the middle of each shell whorl.


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P. antipodarum is native to New Zealand and adjacent islands (Ponder, 1988). It has been introduced to Europe, Iraq, Turkey, Japan, the Americas and Australia.  

Distribution Table

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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/RegionDistributionLast ReportedOriginFirst ReportedInvasiveReferenceNotes

Sea Areas

Atlantic, NortheastWidespreadIntroduced1887Grant and Briggs, 1998; Leppäkoski and Olenin, 2000Baltic Sea


IraqLocalisedIntroducedNaser and Son, 2009Garmat Ali River
JapanWidespreadIntroduced1990Shimada and Urabe, 2003; Masuda, 2010
-HonshuLocalisedIntroducedShimada and Urabe, 2003; Katayama and Ryoji, 2004Moriyama Channel
TurkeyLocalisedIntroduced2003Kalyoncu et al., 2008Akçapinar and Akyaka Kadin Azmagi streams

North America

CanadaPresentPresent based on regional distribution.
-British ColumbiaLocalisedIntroduced2007Davidson et al., 2008Port Alberni, Vancouver Island
-OntarioWidespreadIntroduced1991Zaranko et al., 1997Lake Ontario
USARestricted distributionEPPO, 2014
-ArizonaWidespreadIntroduced1995 Not invasive Cross et al., 2010; EPPO, 2014Colorado River just below Lake Powell in Glen Canyon
-CaliforniaWidespreadIntroducedHerbst et al., 2008; EPPO, 2014Upper Owens River
-IdahoWidespreadIntroducedRichards et al., 2001Higher densities than native snails. Snake River
-MontanaWidespreadIntroducedKerans et al., 2005; EPPO, 2014Lower colonisation of substrates by native species. Madison river basin (Montana and Wyoming)
-New YorkWidespreadIntroduced1991Zaranko et al., 1997; EPPO, 2014Lake Ontario
-OregonLocalisedIntroduced Not invasive Brenneis et al., 2010Columbia river estuary
-UtahPresentIntroduced Invasive Vinson and Baker, 2008It causes a reduction of fish health. Green River
-WashingtonLocalisedIntroduced2002Davidson et al., 2008Long Beach
-WyomingWidespreadIntroduced Invasive Hall et al., 2006; Riley et al., 2008; Arango et al., 2009; EPPO, 2014Increase in secondary productivity of invertebrate community. Polecat Creek


AustriaLocalisedIntroducedZieritz and Waringer, 2008Weidlingbach River near Vienna
DenmarkWidespreadIntroduced Invasive Thomsen et al., 2009West, north-east and south-east open and estuarine waters. A likely impact of this species but not confirmed
FinlandWidespreadIntroducedCarlsson, 2000Lakes of the Aland Islands
FranceWidespreadIntroducedMouthon and Dubois, 2001; Gerard et al., 2003; Zettler and Richard, 2004Domination of gastropod communities. Basin of Mont St-Michel Bay.
GermanyLocalisedIntroducedWagner, 2000; Arle and Wagner, 2013Main River
GreeceLocalisedIntroducedRadea et al., 2008Dominant among the gastropod community where it dwells. central ad western Greece: littoral zone of Lake Trichonis, permanent, slowly flowing stream discharging into Trichonis near Dougri
ItalyWidespreadIntroducedCianfanelli et al., 2007Large distribution in Italy
NetherlandsLocalisedIntroducedDorgelo, 1987; Berg et al., 1997Lake Veluwemeer and Lake Wolderwijd
PolandWidespreadIntroduced Invasive Brzezinski and Kolodziejczyk, 2001; Lewin and Smolinski, 2006Domination of gastropod communities. The Katowicka Upland, Upper Silesia
PortugalLocalisedIntroducedSimoes, 1988; Sousa et al., 2005Freshwater tidal area of River Minho estuary
Russian FederationLocalisedIntroducedSon et al., 2008Don River and in a spring in the Botanic Garden of Southern Federal University (Rostov-on-Don) at the Russian part of the Azov Sea - Black Sea basin
SpainWidespreadIntroduced Not invasive Hinz et al., 1994; Alonso, 2006; Soler, 2006; Múrria et al., 2008Vallvidrera stram (Barcelona province)
SwitzerlandWidespreadIntroducedWalter, 1980; Schmidlin et al., 2012It causes changes to the native invertebrates that dwell in the hard bottom. Lake Neuchatel
UKPresentIntroducedBoycott, 1936; Heywood and Edwards, 1962; Grant and Briggs, 1998; EPPO, 2014Norfolk (sea)


AustraliaPresentIntroducedSchreiber et al., 2002Tarra River (south-eastern Australia)
-TasmaniaPresentIntroducedPonder, 1988
-VictoriaPresentIntroducedQuinn et al., 1998; Schreiber et al., 1998; Schreiber et al., 2003Lake Purrumbete
New ZealandWidespreadNative Not invasive Winterbourn, 1969; EPPO, 2014Widespread in lakes, ponds, rivers and streams

History of Introduction and Spread

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P. antipodarum is native to New Zealand and adjacent islands (Ponder, 1988). Its first occurrence in Europe was in England in 1859 (Ponder, 1988). In Australia it was first reported in Tasmania in 1872 and in continental Australia in 1895 (Ponder, 1988). In America it was first recorded in 1987 in middle reaches of the Snake River, Idaho, probably escaped from a fish farm (Bowler, 1991). The species was recorded in Lake Ontario in 1991 and in the Columbia River, Oregon, in 1997, where it probably arrived via ballast water from commercial ships (Zaranko et al., 1997). The species has also been recorded in Japan (Shimada and Urabe, 2003), Iraq (Nasser and Son, 2009) and Turkey (Kalyoncu, et. al, 2008). The causes of introductions to new ecosystems are not very clear, but ballast waters and fish farms are among the most commonly cited (Zaranko et al., 1997; Bowler, 1991).


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Introduced toIntroduced fromYearReasonIntroduced byEstablished in wild throughReferencesNotes
Natural reproductionContinuous restocking
Tasmania New Zealand 1872-1879 Yes Ponder (1988) Other cause - (e.g. drinking water supplies on ships)
UK New Zealand 1859 Yes Ponder (1988) Other cause - (e.g. drinking water supplies on ships)

Risk of Introduction

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The wide range of active and passive methods for dispersal of P. antipodarum makes the control of this species very difficult. Anglers and fish farms may spread it to new waterways (Loo et al., 2007). This species is able to move upstream or float downstream (Haynes et al., 1985). As P. antipodarum is parthenogenetic and has a high reproductive capacity, in theory only one female need arrive at a new ecosystem for a new population to establish.


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Within its native range P. antipodarum lives in freshwater ecosystems, except temporary ponds, as well as brackish waters (Winterbourn, 1973). However, in its non-native range, it can be found in either in freshwater, brackish and even salty water, and has been recorded in streams, rives, lakes, reservoirs, channels, isolated coastal lakes, shallow lakes, estuaries and open seas (Alonso and Castro-Díez, 2008; 2012a).

Habitat List

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Inland saline areas Present, no further details Natural
Estuaries Present, no further details Natural
Lagoons Present, no further details Natural
Coastal areas Present, no further details Natural
Mud flats Present, no further details Natural
Intertidal zone Present, no further details Natural
Salt marshes Present, no further details Natural
  Present, no further details Harmful (pest or invasive)
  Present, no further details Natural
Irrigation channels Present, no further details Natural
Lakes Present, no further details Harmful (pest or invasive)
Lakes Present, no further details Natural
Reservoirs Present, no further details Natural
Rivers / streams Present, no further details Harmful (pest or invasive)
Rivers / streams Present, no further details Natural
Ponds Present, no further details Harmful (pest or invasive)
Ponds Present, no further details Natural

Biology and Ecology

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In its native (New Zealand) range, P. antipodarum is thought to be composed of sexually reproducing diploids and parthenogenetic (apomictic) triploid females (Wallace, 1992). In introduced populations, most individuals are triploid clonal females (Hauser et al., 1992; Hughes, 1996).

Reproductive biology

Although, in its natural range, both sexual and asexual reproduction coexists, non-native populations are parthenogenetic and consist almost exclusively of females (Jokela et al., 1997; Alonso and Castro-Díez, 2008). P. antipodarum is ovoviviparous, and females brood their offspring in a brood pouch until they reach the 'crawl-away' developmental stage (Jokela et al., 1997). Individuals reach sexual maturity at a shell length of 3-3.5 mm of shell length (Moller et al., 1994; Richards, 2002). There are between 1 and 6 generations per year, and one adult can produce an average of 230 juveniles per year (Moller et al., 1994; Richards, 2002).

Physiology and phenology

P. antipodarum has gills for aquatic respiration. It has a wide range of tolerance to different environmental parameters (such as salinity and water temperature). A study by Hoy et al. (2012) suggested that P. antipodarum may adapt to salty water. Alonso and Camargo (2003) showed it has a high tolerance to the toxicity of nitrogen compounds (ammonia, nitrate and nitrite). P. antipodarum also possess a high tolerance to air exposure (Alonso and Castro-Díez, 2012b). Given that most physiological studies have been focused on exotic populations, comparison of tolerances between native and exotic populations are not available. Kistner and Dybdahl (2013) suggested that phenotypic plasticity in conjunction with evolution may be driving variation of shell morphology.

Activity patterns

P. antipodarum can bury itself into the sediment during dry or cold periods (Duft et al., 2003).

Population size and structure

P. antipodarum presents contrasting densities and population sizes within its invaded range. Several studies (e.g. Richards et al., 2001; Hall et al., 2003) have shown high densities of up to 500,000 snails/m2 in some streams in the USA, or even up to 800,000 in a lake in Switzerland (Dorgelo, 1987). Lower densities do occur, however (Brzezinski and Kolodziejczyk, 2001; Lewin and Smolinski, 2006). Additionally, population sizes change throughout the year, with higher densities in summer and very low densities in winter, especially at temperatures near freezing (Moffitt and James, 2012).


P. antipodarum has a broad diet that includes organic matter (detritus), living plants (macrophytes) and micreoorganisms (periphyton) (Alonso and Castro-Díez, 2008).

Environmental requirements

This species can tolerate water temperature from 31ºC to near 0ºC, but cannot tolerate freezing or sub-zero temperatures (Quinn et al., 1994; Moffitt and James, 2012; Hamada et al., 2013b).

P. antipodarum can tolerate salinity up to 32‰, but prefers 15‰ (Jacobsen and Forbes, 1997; Costil et al., 2001; Gerard et al., 2003).

Water Tolerances

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ParameterMinimum ValueMaximum ValueTypical ValueStatusLife StageNotes
Ammonia [unionised] (mg/l) 0.02 Harmful
Ammonium [ionised] (mg/l) Harmful High levels tolerated if water temperature and pH are low
Conductivity (µmhos/cm) Optimum This species is limited by low conductivity
Dissolved oxygen (mg/l) Harmful Wide range of tolerances
Salinity (part per thousand) 15 Optimum
Salinity (part per thousand) 32 Harmful
Turbidity (JTU turbidity) Harmful Wide range of tolerances
Velocity (cm/h) 86400 Harmful it prefers low water velocities
Water temperature (ºC temperature) 18 Optimum
Water temperature (ºC temperature) 0 31 Harmful

Notes on Natural Enemies

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Some predators (such as rainbow trout Oncorhynchus mykiss) and parasites (such as the digenetic trematode Microphallus sp.) have been reported for P. antipodarum (Kopp and Jokela, 2007; Vinson and Baker, 2008). A high proportion of P. antipodarum are able to survive the passage through the digestive tracts of fish such as rainbow trout, although Hellmair et al. (2011) reported that the endangered tidewater goby (Eucyclogobius newberryi) may digest it more effectively. In the case of parasites, the number of parasite species and their incidence on P. antipodarum populations in invaded ecosystems has been reported to be very low (Zbikowski and Zbikowska, 2009).

Means of Movement and Dispersal

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Natural dispersal

P. antipodarum is able to spread upstream up to 60 meters in three months (Adam, 1942). It can also move downstream by drifting in the current, or on floating aquatic plants (Alonso and Castro-Díez, 2008).

Vector transmission (biotic)

P. antipodarum can be transported in the gut of fish. A high proportion of mud snails are able to survive the pass through the digestive tracts of different fish species; up to 50% in the case of rainbow trout (Oncorhynchus mykiss) (Vinson and Baker, 2008). P. antipodarum can be also carried in the digestive tract of birds, or attached to their legs or feathers.

Accidental introduction

Most accidental introductions are caused by ballast waters and aquatic movements of boats, or by recreational vessels such as rafts and kayaks (Zaranko et al., 1997; Hosea and Finlayson, 2005). It can also be introduced in water tanks or via trade of aquatic plants, or via fish farms (Bowler, 1991). Anglers may introduce P. antipodarum to new waterways.

Intentional introduction

There is no commercial interest in P. antipodarum as food nor as a pet and so most or all introductions are thought to be accidental. There are no documented intentional introductions.

Pathway Causes

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CauseNotesLong DistanceLocalReferences
AquacultureTransport of aquaculture products (trout eggs, live fish); in the guts of introduced fish farm fish Yes Zaranko et al., 1997
Digestion and excretionMudsnails are able to survive the pass through the digestive tracts of different fish species Yes Vinson and Baker, 2008
Hunting, angling, sport or racingMovement of anglers between different aquatic ecosystems Yes Alonso and Castro-Díez, 2008

Pathway Vectors

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VectorNotesLong DistanceLocalReferences
Aquaculture stockTransport of aquaculture products (trout eggs, live fish); in the guts of introduced fish farm fish Yes Zaranko et al., 1997
Floating vegetation and debrisDownstream movements Yes Alonso and Castro-Díez, 2008
Ship ballast water and sedimentOne of the main causes of long distance transport of mudsnail Yes Yes Alonso and Castro-Díez, 2008

Impact Summary

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Environment (generally) Positive and negative

Environmental Impact

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At high densities, P. antipodarum it can dominate secondary production and is capable of increasing it to some of the highest values ever reported among stream invertebrates (194 g of ash free dry mass/m2/year) (Hall et al., 2006). This allows P. antipodarum to alter the overall nitrogen fixation rate of an ecosystem by consuming a high proportion of green algae, which causes an increase of nitrogen-fixing diatoms (Arango et al., 2009). Some studies show domination of mollusc communities by this species (Gerard et al., 2003; Lewin and Smolinski, 2006) and also a reduction in the growth of native molluscs (Riley et al., 2008) due to competition for space and food. Because P. antipodarum can survive travelling through the digestive tract of fish, fish that eat lots of P. antipodarum tend to lose weight compared to those which do not (Vinson and Baker, 2008).

Risk and Impact Factors

Top of page Invasiveness
  • Proved invasive outside its native range
  • Highly adaptable to different environments
  • Is a habitat generalist
  • Pioneering in disturbed areas
  • Fast growing
  • Has high reproductive potential
  • Reproduces asexually
Impact outcomes
  • Altered trophic level
  • Damaged ecosystem services
  • Ecosystem change/ habitat alteration
  • Modification of natural benthic communities
  • Modification of nutrient regime
  • Reduced native biodiversity
  • Threat to/ loss of native species
Impact mechanisms
  • Rapid growth
Likelihood of entry/control
  • Highly likely to be transported internationally accidentally
  • Difficult to identify/detect as a commodity contaminant
  • Difficult to identify/detect in the field

Uses List

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  • Research model

Detection and Inspection

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Adult and juvenile P. antipodarum can be seen without equipment, but newborn snails have such a light shell and small size (0.4 mm shell length) that their detection is difficult. DNA detection methods have been developed to detect P. antipodarum at densities as low as one animal per 1.5 liters of water for two days (Goldberg et al., 2013). As P. antipodarum can be confused with many other aquatic snails, reliable identification requires a specialist in the Tateidae family.

Similarities to Other Species/Conditions

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P. antipodarum can be confused with others species belonging to different genera of the Hydrobiidae family (Plesiella, Belgrandia, Mercuria, Pseudamnicola, Colligyrus, Pyrgulopsis, Eremopyrgus) and even with species of the genus Fluminicola (Lithoglyphidae family). Therefore, a reliable identification requires a specialist in the Tateidae and Hydrobiidae families.

Prevention and Control

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SPS measures and Early warning systems

Although no specific methods to prevent the introduction of P. antipodarum have been described, techniques applied to other molluscs may prove effective for this species too. For instance, the treatment of ballast water in local and international trade can be applied to P. antipodarum. Additionally, examination of aquaculture products (live trout, eggs, etc.) is necessary to avoid the new introductions of this species. In the case of transporting live fish a quarantine to avoid the introduction of live P. antipodarum in the digestive tract is recommended.

Rapid response

In the case of recent introductions several steps should be followed: firstly, reports of the presence of P. antipodarum are confirmed by an expert along with the population density and distribution. Once confirmed, the feasibility of eradication/control has to be assessed. These actions have to be communicated to environmental agencies. Additionally, a monitoring program needs to be established to ensure that the action taken is successful.

Public awareness

Public awareness campaigns would have to be aimed at anglers, fish farm owners, hunters, natural resource management personnel, hikers and any people that could come in contact with aquatic ecosystems. Anglers should be encouraged to carefully clean their equipment and waders.


Most of the cited control practices focused on chemical control (Oplinger et al., 2009; Oplinger and Wagner, 2009; 2010; 2011; Myrick and Conlin, 2011; Hoyer and Myrick, 2012) with some cases of physical control by temperature, desiccation and hydrocyclonic separation (Strzelec, 2000; Richards et al., 2004; Nielson et al., 2012).


No information on eradication programs for P. antipodarum is available. Generally speaking, eradication may be possible in isolated ecosystems, such as small lakes or ponds that can be temporally isolated from water sources, whereas in rivers, streams or lakes the chemical or physical eradication is not feasible as the damage to other elements of the ecosystems will be unacceptable. In the case of fish farms this species can be completely eradicated by desiccation or chemical treatments of ponds.


Isolated ecosystems (such as small lakes or ponds) may be contained, but this control method is practically impossible in rivers, streams or big lakes.

Monitoring and surveillance (incl. remote sensing)

Sampling protocols used for invertebrate monitoring (such us Surber and Hess samplers and kick-net samplers) could be used to detect the presence of P. antipodarum in aquatic ecosystems.

Gaps in Knowledge/Research Needs

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There are several topics that need further research:

1. Identifying the properties of ecosystems (such as degree of environmental degradation, or biodiversity) that allow for a rapid spread of P. antipodarum, as this species has not always shown invasive behaviour. Such knowledge would help to develop programs to prevent the introduction of P. antipodarum into vulnerable ecosystems.

2. Assessing the effectiveness of control and eradication methods is necessary. There are several studies focused on control, but most of them are under laboratory conditions. Research in invaded natural ecosystems is necessary to test if these methods are effective in natural scenarios, including biological control by natural parasites.

3. Understanding the long-term dynamic of P. antipodarum populations in invaded ecosystems will help explain whether the high densities that are found in some ecosystems can be maintained or whether these populations may experience a collapse.

4. Regarding P. antipodarum distribution, more information is needed for Central and South America and Africa.


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Brzezinski T; Kolodziejczyk A, 2001. Distribution of Potamopyrgus antipodarum (Gray, 1843) in waters of the Wigry National Park and the effect of selected habitat factors on its occurrence. Folia Malacologica, 9:125-135.

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USA: California Department of Fish and Wildlife (CDFW),

USA: United States Department of Agriculture, 1400 Independence Avenue, Washingon DC 20250,


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08/10/13 Original text by:

Alvaro Alonso, University of Alcala, Spain

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