Potamopyrgus antipodarum

Invasion History

First Non-native North American Tidal Record: 1997
First Non-native West Coast Tidal Record: 1997
First Non-native East/Gulf Coast Tidal Record: 2021

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General Invasion History:

Potamopyrgus antipodarum is native to fresh and brackish waters of the North and South Islands of New Zealand. In the 18th and 19th centuries, it was introduced to Australia (first reported 1870) and Europe (England- 1859) and spread rapidly in both continents (Winterbourn 1970; Ponder 1988). Possible vectors include dry ballast and the drinking water casks of sailing ships. In Europe, its range runs from the British Isles and Spain, east through the Baltic and Black Seas in interior fresh and coastal brackish waters (Nikolaev 1951; Ponder 1988; Leppakoski and Olenin 2000; Gomiou et al. 2002; Radea et al. 2008). Along the marine coasts of Europe, it prefers the fresher portions of estuaries but does tolerate salinities above 18 PSU (Gerard et al. 2003). This snail has also been introduced to Iraq (Naser and Son 2009) and Japan (Shimada and Urabe 2003).

The New Zealand Mud Snail has been transported by a wide range of vectors. While ships' drinking water barrels were a likely historical vector for introduction to Europe (Ponder 1988), subsequent dispersal modes probably included hull fouling on boats and ships, transport through canals, dry ballast, ballast water, transport on fishing gear, and with stocked fishes or ornamental aquatic plants (Eno et al. 1997; New Zealand Mudsnail Management and Control Plan Working Group 2007; Davidson et al. 2008). Natural dispersal on birds' feet and within guts is also likely (van Leeuwen 2012). The snail's small size, salinity tolerance, and parthenogenetic reproduction have enabled widespread transport (Alonso and Castro-Díez 2012). Multiple parthenogenetic clones exist in New Zealand and Australia, and many have been introduced. At least four clones have been introduced to North America, three probably directly from New Zealand. In the Great Lakes at least one clone arrived via Europe (Dybdahl and Drown 2011).

North American Invasion History:

Invasion History on the West Coast:

In 1987, Potamopyrgus antipodarum was first collected in North America in the headwaters of the Snake River, Idaho and subsequently spread to the headwaters of the Missouri River in Montana. At least three clones exist in Western North America, a widespread form (US 1) derived from New Zealand, and two with restricted ranges in the Snake River (US 1a and US3; Dybdahl and Drown 2011; Hershler et al. 2012). Its spread to other parts of North America has been rapid, but spotty, on boats, fishing gear, boots and waders, with stocked fish, etc. It was collected in the Columbia River estuary near Astoria in 1996 by James Carlton (personal communication, Davidson et al. 2008) and has subsequently spread into fresh and brackish waters up and down the West Coast, including the Rogue River, Oregon (OR) in 1999; Coos Bay, OR in 2005; Alsea Bay, OR in 2007; Yaquina Bay, OR in 2008; Tillamook Bay, OR in 2007; Long Beach, Washington (WA) in 2002; Willapa Bay tributaries, WA (Davidson et al. 2008); and Grays Harbor, WA (USGS Nonindigenous Aquatic Species Program 2013). In 2007, it was collected in Port Alberni Inlet, on the west side of Vancouver Island, British Columbia, at a salinity of 5 PSU (Davidson et al. 2008). By 2009, a population became established in Capitol Lake, Olympia, WA, an artificial lake created by damming an inlet of Puget Sound. An attempt was made to control the population by letting seawater flow into the lake at high tides, raising the salinity to 7.5-27 PSU. The backflush treatment greatly reduced P. antipodarum populations, but survivors showed increased salinity tolerances (Leclair and Cheng 2011).

In California, P. antipodarum was first collected in the Sacramento-San Joaquin Delta in 2003, on the Mokelumne River (USGS Nonindigenous Aquatic Species Program 2008). It has subsequently been collected in at least four other San Francisco Bay tributaries: Calaveras River (in 2004), Napa River, West Antioch Creek, and Alameda Creek (in 2008, USGS Nonindigenous Aquatic Species Program 2008). The New Zealand Mud Snail was also collected in other coastal watersheds from Ventura County to Santa Cruz (in 2008, USGS Nonindigenous Aquatic Species Program 2013).

Potamopyrgus antipodarum spread in a spotty fashion in the Intermountain West of North America, from the Snake River, Idaho in 1997, appearing in the Madison River, Montana-Wyoming in 1994-1995, and in the Green-Colorado system and the Great Salt Lake basin in Utah in 2001. It was found in the Colorado River in Arizona in 1996, and in southeastern California, in the Owens River Valley. In 2004-2005, populations were found in the South Platte River drainage, east of the Rockies in Colorado (USGS Nonindigenous Aquatic Species Program 2013; New Zealand Mudsnail Management and Control Plan Working Group 2007). This pattern of widely scattered dispersal suggests a diverse mixture of anthropogenic vectors, and possibly natural transport by birds, as well.

Invasion History on the East Coast:

In 1991, Potamopyrgus antipodarum was found at Wilson, New York on Lake Ontario (Zaranko et al. 1997). It spread in the Great Lakes - St. Lawrence system to Prescott, Ontario on the St. Lawrence River in 2004 (USGS Nonindigenous Aquatic Species Program 2013); Lake Erie in 2005; Thunder Bay, Ontario in 2005; the St. Louis River estuary, Minnesota in 2005; and Lake Michigan in 2006 (Trebitz et al. 2010; USGS Nonindigenous Aquatic Species Program 2013). The Great Lakes population belongs to a clone (US2) which has a mitochondrial DNA profile identical to the clone (EU A), which is widespread in Europe (Dybdahl and Drown 2011). This snail was probably transported to the Great Lakes in ballast water of transatlantic ships (Zaranko et al. 1997).

In 2013, it was found to be abundant in Spring Creek, in the Bald Eagle Creek (upper Susquehanna) watershed in central Pennsylvania (USGS Nonindigenous Aquatic Species Program 2013). This is its first occurrence in an Atlantic drainage outside the Great Lakes - St. Lawrence system. In September, 2017, P. antipodarum was found closer to Chesapeake Bay, in the Big Gunpowder River, about 50 km from tidal waters (Dance 2017). GARP modeling (Genetic Algorithm for Rule-set Production) predicts that this snail could colonize coastal drainages from Chesapeake Bay to Nova Scotia, and the Mississippi Basin to the edge of the Great Plains (Loo et al. 2007).

Invasion History Elsewhere in the World:

Potamopyrgus antipodarum is likely introduced in Australia. It belongs to a genus that has several species in New Zealand, but has no fossil record and is the only Potamopyrgus species in Australia. It was first reported in Hobart, on the south coast of Tasmania, in 1872 (as Paludestrina wisemaniana and Bythinella legrandiana) and was subsequently found in Launcestown, Tasmania in 1879; and Melbourne, Victoria in 1895. It was first collected in Adelaide, South Australia in 1926 and near Sydney, New South Wales in 1963 (Ponder 1988). It is now widespread in coastal drainages of Victoria and also in streams around Adelaide and Sydney. It is predicted to be capable of colonizing most of the southeastern and some of the southwestern (Perth area) coastal region of Australia (Loo et al. 2007).

Potamopyrgus antipodarum was collected in Europe before it was found in Tasmania. It was found in the Thames estuary as early as 1859, and was formally named as Hydrobia jenkinsi Smith in 1889. It spread slowly in coastal areas at first and then rapidly along canals and coastal rivers. It was widespread by 1920 and now occurs from the Scilly Islands in the south, to the Shetland Islands in the north, although in Scotland it is largely confined to coastal areas (Ponder 1988; Eno et al. 1997). By 1887, this snail was collected in Germany in the Western Baltic Sea (Olenin and Leppakoski 2000), and within two decades was found in Copenhagen and the Oder-Odra lagoon (Gruszka 1999; Jensen and Knudsen 2005). The New Zealand Mudsnail spread further into the Baltic, reaching the mid-Baltic (Gotland, Sweden; Curonian Lagoon, Lithuania) by 1920, the Gulf of Finland by 1926, and the Gulf of Bothnia by 1945 (Nikolaev 1951; Leppakoski and Olenin 2000). It reached southern Norway by 1952 (Hopkins 2002), and was introduced to Galicia, Spain in the late 19th century, and Catalonia, on the Mediterranean by 1936 (Altimira 1969, cited by Múrria et al. 2008). Potamopyrgus antipodarum reached the Black Sea in Romania by 1952 (Gomoiu et al. 2002) and reached the region around Odessa, Ukraine by 2005 (Son 2008). The New Zealand Mud Snail has colonized the interior of Europe as well as the coast, reaching such landlocked countries as Switzerland and Austria (Alonso and Castro-Diez 2012).  In 2017, it was found in freshwater the upper and lower regions streams on the Portuguese island of Madeira, and is assumed to occur in their estuaries (Órfão et al. 2024).

Potamopyrgus antipodarum has had two widely separated introductions in Asia, in Japan and Iraq. It was first found in Japan, in Mie prefecture, Honshu in 1990, and is now found on all of Japan's main islands (National Institute for Environmental Studies 2013). In 2008, this snail was found in the Garmat Ali River, Basra, Iraq, part of Shat-al-Arab estuary (Naser and Son 2009). The authors consider bird dispersal to be the likeliest vector, but given the military and oil industry activity in the region, shipping cannot be ruled out.

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Description

Potamopyrgus antipodarum is a small snail, primarily occurring in freshwater, but able to tolerate brackish to near-marine salinities in estuaries (Leclair and Cheng 2011; Hoy et al. 2012). Its overall body shape is conical-oval, with a sharp spire. Its shell is solid, but not thick-walled, and is dextrally coiled, usually with 5-6 and sometimes 7-8 whorls. The aperture is oval, and the inner lip extends completely along the parietal wall. The first whorl is minute and raised, and the subsequent whorls are evenly rounded and separated by a deep suture. The body whorl is bluntly angular. The umbilicus is a narrow slit. The operculum is oval, with the nucleus off-center and a calcium deposit on the inner surface. Specimens in New Zealand may reach 13 mm in shell length, with 7-8 whorls, but a more typical size in introduced populations is 4-6 mm with 5-6 whorls. The color of the shell ranges from light to dark brown. Description from: Winterbourn 1970, Zaranko et al. 1997, New Zealand Mudsnail Management and Control Plan Working Group 2007, and Loo 2012.

All introduced populations are clonal, reproducing parthenogenetically. Individuals can vary considerably in shape, size, color, and shell ornamentation, even within one population, as well as among clones. Some individuals and clones show a sharp, bristly keel on the whorls (Winterbourn 1970; Ponder 1988; New Zealand Mudsnail Management and Control Working Group 2007; Butkus et al. 2012; Loo 2012). At least four clones are known from North America, three from Western North America, and one in the Great Lakes. The clone found in the Great Lakes is also widespread in Europe (Dybdahl and Drown 2011). Multiple lineages havebeeen involved in invasions in Europe. A largely freshwater Lineage T is widespread in Central Europe, but a more salinity tolerant Lineage Z is establoished around the Black Sea, while both lineages occur in Baltic, though usually in different habitats (Butkus et al. 2020).

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Taxonomy

Ecology

General:

Potamopyrgus antipodarum is a euryhaline freshwater snail, capable of surviving near-marine salinities (Gerard et al. 2003; Leclair and Cheng 2011; Hoy et al. 2012). In its native New Zealand, this snail has both sexually reproducing and clonal populations. Introduced populations are triploid parthenogenetic, consisting of females reproducing without fertilization, though they occasionally produce males (Ponder 1988; New Zealand Mudsnail Management and Control Plan Working Group 2007; Loo 2012; Liu et al. 2012). Some European populations have up to 20% males, although the usual range is 0 to 5%. Males do not seem to contribute to population growth or invasion success (Neiman et al. 2012). The snails are oviviparous, brooding eggs, and producing an average of 50 live young per brood and an average of 230 young per year. Females reach breeding size at about 3.5 mm and 3-6 months of age, and their life cycle is annual (Winterbourn 1970; Lassen 1979; Crosier et al. 2006). Although invading populations are parthenogenetic, there seems to be sufficient genetic diversity for adaptation to a wide range of habitats and environmental conditions. The concept of a small number of clones, as defined by mitochondrial DNA (Dybdahl and Drown 2011) may underestimate the genetic diversity of the introduced populations (Liu et al. 2012; Hershler et al. 2012).

Potamopyrgus antipodarum inhabits a wide range of freshwater and brackish habitats, including streams, springs, ponds, lakes and estuaries. It is often more abundant in disturbed areas, such as pastures, compared to forests, due to thicker algal growth, with more light and nutrients (Ponder 1988; Loo et al. 2007). In Australia, it has been found in drinking water tanks and pipes (Ponder 1988). It tolerates temperatures as low as 2°C, but cannot withstand freezing (Moffit and James 2012). The New Zealand Mud Snail is also tolerant of varying salinities and can be very abundant in estuarine habitats (Lassen 1979; Gerard et al. 2003; Ezhova et al. 2005; Davidson et al. 2010; Brenneis et al. 2010). Published accounts, using snails from New Zealand, Europe, and the western US give widely varying upper salinity limits for this animal, from 21 to 64 PSU. This may reflect differences in method and acclimation in the wild and laboratory, as well as clonal differences between populations (Leclair and Cheng 2011). Hoy et al. (2012) compared salinity tolerances of a population from the mouth of the Columbia River, where salinities fluctuated between 0 and 32 PSU, with those from freshwater, coastal Devils Lake (Oregon). With gradual acclimation, the Devils Lake population had the same 50% tolerance limit as the Columbia River population – 34 PSU. European estuarine populations occur in salinities at least up to 26 PSU (Gerard et al. 2003). The New Zealand Mud Snail is not normally amphibious, but does occur on exposed water-saturated mud and can survive up to 10 days on a saturated surface. On a dry surface, it can survive 30-50 hours in 'dry' air (Winterbourn 1970) and 68% relative humidity (Alonso and Castor-Diez 2012), respectively. Thus, this snail has a strong potential for transport on fishing gear, boots, boats, and many other vectors.

The food of P. antipodarum consists of detritus, periphyton (attached algae), and benthic diatoms (Brenneis et al. 2010; Loo 2012). The snail is a potential prey for fish and invertebrates, including the Signal Crayfish (Pacifastacus leniusculus) (Brenneis et al. 2010). However, it is poorly digested by most fishes and birds (New Zealand Mudsnail Management and Control Plan Working Group 2007; Bersine et al. 2008; Vinson and Baker 2008; van Leeuwen et al. 2012). New Zealand Mudsnails are hosts to numerous parasites in their native region (14 trematode spp., Winterbourn 1970; Krist and Lively 1998), but very few (0-2) in invaded waters (France, Gerard et al. 2003; Ontario Creek, Karatyev et al. 2012; Wyoming, Adema et al. 2009). Release from parasites and predators is probably one of many factors contributing to this species invasion success (Gerard et al. 2003; Alonso et al. 2012).

Food:

Algae, detritus

Consumers:

Fishes, birds

Competitors:

Trophic Status:

Deposit Feeder

DepFed

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Habitats

General Habitat	Fresh (nontidal) Marsh	None
General Habitat	Tidal Fresh Marsh	None
General Habitat	Unstructured Bottom	None
General Habitat	Salt-brackish marsh	None
General Habitat	Nontidal Freshwater	None
Salinity Range	Limnetic	0-0.5 PSU
Salinity Range	Oligohaline	0.5-5 PSU
Salinity Range	Mesohaline	5-18 PSU
Salinity Range	Polyhaline	18-30 PSU
Tidal Range	Subtidal	None
Tidal Range	Low Intertidal	None
Vertical Habitat	Epibenthic	None

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Life History

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Tolerances and Life History Parameters

Minimum Temperature (ºC)	2	Little mortality at 2-4 C, but 100% mortality at 0 C (Moffit and James 2012)
Maximum Temperature (ºC)	31	Experimental, Cox and Rutherford 2000.
Minimum Salinity (‰)	0	This is a freshwater species.
Maximum Salinity (‰)	32	Experimental, 9 day LC 50, Hoy et al. 2012, snails from mouth of Columbia River. (72 h LC50, 28 PSU (Piscart et al. 2011), short term survival at 32 PSU (Lucas 1965, cited by Gerard et al. 2003). Reported field upper salinities range from 22-28, while laboratory tolerances range for 15 -64 PSU. Salinity tolerance may vary among genetic strains, and with acclimation (LeClair and Cheng 2011; Hoy et al. 2012).
Minimum Reproductive Salinity	0	This is a freshwater species.
Maximum Reproductive Salinity	15	Experimental, Danish (Jacobsen and Forbes 1997) and Columbia River, and New Zealand clones, as well as New Zealand sexual populations (Drown et al. 2011) showed similar patterns with a peak in reproductive output at 5-10 PSU, and a sharp reduction in reproduction at 15 PSU. Orlova and Kommendatov (2013), using populations from the Gulf of Finland, and different modes of acclimation, found that reproduction decreased at 10 PSU, and ceased at 18 PSU.
Minimum Length (mm)	3.5	Parthenogenetic females begin laying eggs at 3-4 mm (Loo 2011; Zarenko et al.1997)
Maximum Length (mm)	12	Specimens in New Zealand may reach 13 mm in shell length, with 7-8 whorls, but a more typical size in introduced populations is 4-6 mm, with 5-6 whorls (Zaranko et al. 1997; Loo 2011)
Broad Temperature Range	None	Cold temperate-Warm temperate
Broad Salinity Range	None	Nontidal Limnetic-Polyhaline

General Impacts

The rapid spread and high densities (up to 500,000-800,000 m-3) achieved by Potamopyrgus antipodarum in its non-native range have prompted concern about competition with native invertebrates, and adverse effects on freshwater and estuarine food webs, including sport and commercial fishes.

Specific economic impacts of the New Zealand Mud Snail have not been reported in North America. However, it is suspected to have a negative impact on fisheries, because it seems to be poorly digested, and under-consumed relative to its abundance, by many fish species. It is generally regarded as a competitor with native snails and a threat to biodiversity, especially of isolated streams and springs (New Zealand Mudsnail Management and Control Plan Working Group 2007; Bersine et al. 2008; Vinson and Baker 2008; Brenneis et al. 2011). These concerns have prompted research on control and eradication methods, and prompted education programs for fishermen and boaters (Richards et al. 2012; New Zealand Mudsnail Management and Control Plan Working Group 2007; Leclair and Cheng 2011; Alonso and Castro-Díez 2012). In Australia, this species has been known to foul water tanks and clog drinking water pipes (Ponder 1988).

Ecological Impacts

Competition with native snails and other invertebrates, habitat impacts, and food web and nutrient impacts have been reported in freshwater systems, but the nature and magnitude of these impacts have varied greatly (Kerans et al. 2005; Múrria et al. 2008; Alonso and Castro-Diez 2012; Loo 2012; Moore et al. 2012). In estuaries, in France and Denmark, and in portions if the Baltic Sea, very high densities and numerical dominance of benthic communities have been observed, leading to inferences of competition, habitat changes, and alteration of food webs and nutrient cycles (Lassen 1979; Gerard et al. 2003; Thomsen et al. 2009; Zaiko et al. 2011). However, the only experimental evaluations of impacts of P. antipodarum in estuaries have been conducted in the Columbia River estuary (Brenneis et al. 2010; Brenneis et al. 2011). The results of these experiments, so far, suggest that impacts on native food webs are subtle, and do not represent drastic alterations.

Competition: Competition of P. antipodarum with other invertebrates, especially snails, has often been inferred from the very high densities and numerical dominance that this species achieves in estuaries. Lassen (1979) noted its high reproductive output relative to other hydrobiids, while Gerard et al. (2003) noted the near-absence of trematode parasites in this snail. Zaiko et al. (2011) listed this species as having some or moderate 'community impacts', (presumed to be competition with native invertebrates) in 5 of 8 regions of the Baltic Sea, based largely on dominance or historical increases in abundance (e.g. Ezhova et al. 2005). Experimental and field observations in Wyoming, Idaho, and Montana springs and streams provide some evidence for competition with native snails, summarized by Alonso and Castro-Diez (2012). In field experiments, tiles with high densities of New Zealand Mudsnails had reduced settlement of native invertebrates (insects and mollusks, Kerans et al. 2005). In the Owens River, California, a population boom of P. antipodarum was accompanied by a crash of native grazers, which recovered when the snail abundance sharply declined, strongly indicative of competition (Moore et al. 2012). However, in the Columbia River estuary, a competition experiment using P. antipodarum and the native isopod Gnorimosphaeroma insulare found that snail density had no effect on isopod feeding, but isopod density reduced snail feeding (Brenneis et al. 2011).

Habitat Change: Potamopyrgus antipodarum has the potential to alter habitats by reducing algal cover. However, grazing rates have not been measured in estuaries, and observations on changes in algal cover in streams are contradictory (Hall et al. 2003; Riley et al. 2008; Alonso and Castro-Diez 2012). At high snail densities, heavy deposition of shells could have an effect on sediment quality (Schmidlin et al. 2010, cited by Alonso and Castro-Diez 2012). Habitat impacts were suggested for P. antipodarum in the Vistula Lagoon, Baltic Sea (Ezhova et al. 2005; Zaiko et al. 2011).

Food/Prey: Potamopyrgus antipodarum is a potential prey for fishes, birds, and invertebrates, including the Signal Crayfish (Pacifastacus leniusculus) (Brenneis et al. 2011). However, it is poorly digested by most fishes and birds (New Zealand Mudsnail Management and Control Plan Working Group 2007; Bersine et al. 2008; Vinson and Baker 2008). Vinson and Baker (2008) found that Rainbow Trout (Oncorhynchus mykiss) grew poorly on an exclusive diet of P. antipodarum. Rainbow and Brown Trout (Salmo trutta) from the Green River, Utah, with New Zealand Mud Snails in their gut contents, had poorer body condition (Vinson and Baker 2008). Field evidence form the Columbia River estuary suggested that many species of estuarine fishes avoided the snails as prey (Bersine et al. 2008). One exception is the Tidewater Goby (Eucyclogobius newberryi), an endangered species of the California coast, for which this snail has become an important food source (Hellmair et al. 2011).

Trophic Cascade: High population densities of P. antipodarum have the potential to alter food webs and nutrient cycles in aquatic ecosystems. In some inland streams (MT, ID, WY), grazing by P. antipodarum has resulted in consumption of much of the primary production and releases of large quantities of ammonium, stimulating plant and microbial growth (Hall et al. 2003; Alonso and Castro-Diez 2012). In one WY stream, heavy grazing of filamentous green algae, led to its replacement by nitrogen-fixing diatoms, increasing the nitrogen budget in the stream (Arango et al, 2009). These changes in nutrient cycles have been suggested to occur in estuaries in the Baltic (Thomsen et al. 2009; Zaiko et al. 2011). A smaller-scale food web change was seen in experiments with Columbia River estuary species, when high abundances of P. antipodarum led to increased consumption by native fishes and a shift away from the native isopod Gnorimosphaeroma insulare to the native amphipod Americorophium salmomis. This change in diet may have resulted from a change in foraging behavior in the fishes in the presence of the snails (Brenneis et al. 2011).

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Regional Impacts

P135	_CDA_P135 (Mad-Redwood)		Ecological Impact	Food/Prey
Potamopyrgus antipodarum has become a frequent food item for the Tidewater Goby (Eucyclogobius newberryi), an endangered fish of California coastal streams and lagoons (Hellmair et al. 2011).
CA	California		Ecological Impact	Food/Prey
Potamopyrgus antipodarum has become a frequent food item for the Tidewater Goby (Eucyclogobius newberryi), an endangered fish of California coastal streams and lagoons (Hellmair et al. 2011).

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