Melanoides tuberculata

Invasion History

First Non-native North American Tidal Record: 1966
First Non-native West Coast Tidal Record: 1988
First Non-native East/Gulf Coast Tidal Record: 1966

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General Invasion History:

Melanoides tuberculata has a broad reported native range in subtropical-tropical regions of the Old World, from Africa, the Middle East, East Asia, and Australia. Museum collections include specimens from many Pacific islands, including Fiji, Vanuatu, and Moorea (Academy of Natural Sciences Philadelphia 2013; Museum of Comparative Zoology 2013), where prehistoric or undocumented introductions are possible. Melanoides tuberculata has been introduced to inland and estuarine waters around the world, mostly in warm-temperate to tropical regions, but some populations have become established, at least temporarily, in warm springs and thermal effluents in cooler regions. In North America, this snail has been found in fresh waters in at least 11 continental US states. It was introduced to springs and streams in inland central Texas before 1964, and is now found in 34 water bodies in 15 counties (Karatayev et al. 2009). Inland populations have been found in California, Arizona, Colorado, Utah, Nevada, Wyoming, Idaho and Montana, often in thermal springs (USGS Nonindigenous Aquatic Species Program 2013). It is predominantly a freshwater snail, but can tolerate salinities as high as 34 PSU. It has been collected at marine salinities in Biscayne Bay, Florida (Roessler et al. 1977; Wingard et al. 2007); Oahu, Hawaii (Englund et al. 2000); and the Ceará River estuary, Brazil (in 2005, Barroso and Matthews-Cascon 2009). In Florida and Brazil, this snail reached high population densities in mangrove swamps (Roessler et al. 1977; Wingard et al. 2007; Barroso and Matthews-Cascon 2009).

North American Invasion History:

Invasion History on the West Coast:

Melanoides tuberculata was first reported in California in 1972, from a ditch in Riverside County, California. In 1988, it was collected in the Sacramento-San Joaquin Delta. It has been found at several sites in the inner Delta, in fresh to oligohaline water, at abundances of up to 754 snails m^-3 (Cohen and Carlton 1995; Peterson and Vayssieres 2010; USGS Nonindigenous Aquatic Species Program 2013). Another California population is established in Los Angeles, where the San Gabriel River becomes estuarine (in 2011, USGS Nonindigenous Aquatic Species Program 2013).

Invasion History on the East Coast:

Melanoides tuberculata was first collected on the East Coast of Florida (FL) in Lake Osceola on the University of Miami campus in 1966 (Clench 1970), and by 1973 it was found in brackish water in the Pompano Canal, Fort Lauderdale (Russo 1974, cited by Roessler 1977). By 1975, this snail was abundant in mangrove areas of the Matheson Hammock-Snapper Creek area of Coral Gables, FL adjacent to Biscayne Bay. Melanoides tuberculata reached abundances of up to 23,000 m^-3 (Roessler et al. 1977). In surveys between 2004 and 2007, the snail was abundant along the shore of Biscayne Bay and adjacent canals, and at one location reached 60,000 animals m^-3, higher density than any native snail (Wingard et al. 2007). The Red-Rim Melania reached the upper St. Johns River, FL by 1975 (USGS Nonindigenous Aquatic Species Program 2003; Florida Museum of Natural History 2013; Lee 2013). It was present in the Indian River Lagoon drainage, in the St. Lucie River and Sebastian Creek drainages (Harbor Branch Oceanographic Institution Collections).

Invasion History on the Gulf Coast:

Melanoides tuberculata was first found on the Gulf Coast of Florida in 1969 in the Hillsborough River in Hillsborough State Park (Clench 1970). This snail is now abundant in the tidal tributaries of Tampa Bay (Baker et al. 2004). In 1975, it was found to be abudant in several brackish (2.5-3.0 PSU) canals flowing into Lake Pontchartrain, in New Orleans, Louisiana (Dundee and Paine 1977).

Invasion History in Hawaii:

Melanoides tuberculata was reported from the Hawaiian Islands at least as early as 1915 (Academy of Natural Sciences of Philadelphia 2013), but it may have been a prehistoric introduction to the islands. It is now found in fresh waters on all the main islands, and in streams on Oahu, where it occurs at salinities of 0-34 PSU (USGS Nonindigenous Aquatic Species Program 2013).

Invasion History Elsewhere in the World:

Melanoides tuberculata has been widely introduced in tropical America and the Caribbean, both as an aquarium release and as biocontrol for native snails hosting disease-causing trematodes. It was introduced to Puerto Rico soon after 1963 (Abbott 1973), to Martinique in 1985 (Pointier et al. 2005), Dominica by 1975 (Reeves et al. 2008), Guadeloupe in 1979 (Pointier et al. 1993b), the US Virgin Islands in 1987 (USGS Nonindigenous Aquatic Species Program 2013). It was present and used to make ornaments in Veracruz, Mexico (Abbott 1973) and is now established in at least 12 Mexican states (Conteras-Arquieta 1998). Melanoides tuberculatus was collected in Panama by 1971, and in 1972, was collected from the bottom of the Miraflores Locks of the Panama Canal (USNM 734154, U.S. National Museum of Natural History 2013).

In South America, M. tuberculata was first reported from Venezuela in 1972 (Pointier et al. 1999), from Brazil in 1967 in Sao Paulo (Vaz et al. 1986, cited by Barroso and Matthews-Cascon 2009), and Argentina in 1999 (Peso et al. 2011). This snail is known from 19 of 26 Brazilian states (da Silva and Barros 2011). In Ceara State, in northeastern Brazil, M. tuberculatus was abundant in mangroves along the Ceara River, at salinities of 0-30 PSU (Barroso et al. 2009). Introductions are still occurring in the Eastern Hemisphere around the edges of its native range, including Spain (Oscoz et al. 2009), Malta (Cilia et al. 2013) and New Zealand (Duggan 2002).

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Description

Melanoides tuberculata is a medium-sized, predominantly freshwater snail of warm-temperate to subtropical regions, but it has colonized estuaries with marine salinities (Englund et al. 2000; Wingard et al. 2007; Barroso and Matthews-Cascon 2009). It has an elongate-conical dextrally coiled shell, of 8-12 whorls with a strongly pointed spire, though the tip is usually eroded. The shell is sculptured with fine spiral striations and vertical weakly curved ribs. The middle and upper whorls have more strongly-defined vertical ribs. The aperture has a tear-drop shape and the operculum is horn-colored. The shell reaches a length of about 30-36 mm. The color is light brown, with a dark reddish spiral band formed of transverse streaks and spots (Thompson 2004).

Melanoides tuberculata has a large native and introduced range and displays a wide degree of morphological variation, which is largely genetic in origin. This snail is capable of both sexual and parthenogenetic reproduction and is often patchily distributed. These scattered populations are prone to divergence both through local selection and genetic drift (Samadi et al. 1999). On the Caribbean island of Martinique, multiple introductions and occasional hybridization have led to dramatic genetic variation in morphology and life history among populations in different streams (Samadi et al. 1999; Facon et al. 2008). This degree of local variation is likely seen elsewhere in this species' introduced and native ranges. According to one online database (Galli 2013), there are 47 named species of Melanoides, excluding synonyms, so it is possible that multiple named and unnamed cryptic species have been introduced and identified as M. tuberculata.

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Taxonomy

Ecology

General:

Melanoides tuberculata is a predominantly freshwater snail, found in springs, streams, lakes swamps, etc. (Pointier et al. 1993b; Pointier et al. 1993c; Thompson 2004). It occasionally invades brackish and marine habitats, especially mangrove swamps (Roessler et al. 1977; Wingard et al. 2007; Barroso and Matthews-Cascon 2009). This snail has separate sexes, but reproduces primarily by parthenogenetic reproduction (without fertilization), with only occasional sexual reproduction, resulting in populations that are largely clonal (Samadi et al. 1999; Facon et al. 2008). Males were 20-33% of the populations in Israeli streams (Livshits and Fishelson 1983), but in other populations, males are rare or absent. Reproduction is ovoviviparous, with eggs being incubated in the female's brood pouch in the last (largest) whorl of the shell, and hatching out as small snails, 1.0 - 4.5 mm in length, with 3-6 shell whorls. The time of brooding and the size of the newborn snails decreased as the number of snails in aquaria increased. Annual fecundity is estimated at 365 embryos per year (Keller et al. 2007). The snails reach maturity at about 10-16 mm shell length, in about 3-7 months. Adult snails range about 20-40 mm length and live for about 2 years (Livshits and Fishelson 1983; Rader et al. 2003; Work and Mills 2013; Bolaji et al. 2011). However, populations, even on small islands, such as Martinique, may vary considerably in life history patterns, as a result of multiple introductions, and occasional hybridization of mostly clonal lineages (Samadi et al. 1999; Facon et al. 2008).

Melanoides tuberculata is a snail of warm-temperate to tropical distribution. Snails from the San Marco River, Texas tolerated temperatures of 17 to 32.5°C with little mortality, but a temperature of 13°C killed all the snails within 12 days (Mitchell and Brandt 2005). Many scattered populations in the western US, Europe, and New Zealand are confined to warm springs or thermal effluents (Duggan 2002; Rader et al. 2003; Oscoz et al. 2009). In Biscayne Bay, Florida M. tuberculata occurred at 0-33 PSU and tolerated salinities up to 45 PSU, although reproduction ceased at ~18 PSU (Wingard et al. 2007). In estuaries on Oahu, Hawaii and Ceara, Brazil, the snail occurred from freshwater to 34 and 30 PSU, respectively (Englund et al. 2000; Barroso and Matthews-Cascon 2009). However, another population, in Lagos Lagoon, Nigeria, did not tolerate salinities over 25 PSU (Bolaji et al. 2011). This snail, though aquatic, is quite tolerant of desiccation – individuals survived 80-140 hours of air exposure at 25°C and 20% humidity (Weir and Salice 2012). Given the genetic diversity reported in this species (Samadi et al. 1999; Facon et al. 2008), it is likely that local populations will vary in environmental tolerances.

Melanoides tuberculata grazes on microalgae and detritus, but does not feed on the leaves of larger plants (Miranda et al. 2012). It is eaten by crabs, fishes, and birds, as indicated by a diversity of parasites which use it as an intermediate host. At least 37 described trematode species and many more undescribed larval forms have been reported from this snail (Alves-Pinto and de Melo 2011).

Food:

Detritus, microalgae, aquatic macrophytes

Consumers:

Parasites, Birds, fishes

Trophic Status:

Deposit Feeder

DepFed

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Habitats

General Habitat	Fresh (nontidal) Marsh	None
General Habitat	Grass Bed	None
General Habitat	Coarse Woody Debris	None
General Habitat	Swamp	None
General Habitat	Nontidal Freshwater	None
General Habitat	Tidal Fresh Marsh	None
General Habitat	Canals	None
General Habitat	Mangroves	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	Endobenthic	None
Vertical Habitat	Epibenthic	None

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Tolerances and Life History Parameters

Minimum Temperature (ºC)	18	10% mortality over 27 days at 17 C (Mitchell and Brandt 2005)
Maximum Temperature (ºC)	32.5	10% mortality over 27 days at 17 C (Mitchell and Brandt 2005)
Minimum Salinity (‰)	0	Widespread in freshwater
Maximum Salinity (‰)	45	Experimental (Wingard et al. 2007); Up to 34 in field observations, Biscayne Bay FL mangrove areas in the Matheson Hammock-Snapper Creek area of Coral Gables, FL/adjacent to Biscayne Bay (Roesseler et al. 1977); Up to 34 PSU in coast streams, Oahu, Hawaii (Englund et al. 2000)
Minimum Reproductive Salinity	0	This is a freshwater species
Maximum Reproductive Salinity	18	Experimental, in tanks with gradually increasing salinity (Wingard et al. 2007)
Minimum Length (mm)	10	Minimum length at maturity (Rader et al. 2003)
Maximum Length (mm)	36	Thompson (2004)
Broad Temperature Range	None	Subtropical-tropical
Broad Salinity Range	None	Nontidal Limnetic-Euhaline

General Impacts

Melanoides tuberculata (Red-Rim Melania) has had ecological impacts as a competitor with native snails and as an intermediate host of many parasites. In many cases, its competition has had positive impacts on human health, as it has displaced some native snails which carry trematodes causing human disease, leading to the use of this snail as a biocontrol agent (Pointier et al. 1993; Pointier et al. 1994; Guimarães et al. 2001; Pointier 2001). However, it is also a host for a number of disease-causing trematodes itself (Alves-Pinto et al. 2011). Aside from disease, the attractive appearance of this snail, and its ability to consume algae and detritus, without damaging plants, has led to its widespread use and dispersal in the aquarium trade (Clench 1970; McCann et al. 1996; USGS Nonindigenous Aquatic Species Program 2013).

Economic Impacts

Human Health: Melanoides tuberculata has been seen in different parts of the world, as having both negative and positive impacts on human health. On one hand, it is an intermediate host to many (37+) trematode species, and has been a vector for the introduction of at least three species to the US. These parasites develop in snails, then fishes, and have birds and mammals as their final hosts. One of these species, a liver fluke, Haplorchis pumilio, does infect humans who eat fish (Alves-Pinto et al. 2011). Melanoides tuberculata populations in the Western Hemisphere are potential hosts for trematodes causing more serious human disease, such as the Asian blood-fluke Paragonimus westermani, whose other intermediate host, the Chinese Mitten Crab (Eriocheir sinensis) is also established in the San Francisco estuary. However, the parasite is not established in the estuary, or elsewhere in North America (Chinese Mitten Crab Working Group 2003). With increased immigration from Asia, differing diets and methods of food preparation, together with the spread of potential animal hosts, the potential for establishment of new human parasites increases. However, a recent survey in Biscayne Bay found no harmful parasites (Wingard et al. 2007).

In some parts of the world, M. tuberculata has been introduced as a biocontrol agent to control native snails (Biomphalaria spp.) hosting trematodes causing schistosomiasis in humans. In Martinique, deliberate introductions of this snail (starting in 1973) resulted in the local extinction of B. glabrata and great reductions of B. straminea (Pointier 2001). Similar introduction programs have been successful in St. Lucia (Prentice 1983, cited by Pointier et al. 1994), Guadeloupe (Pointier et al. 1993), Venezuela (Pointier et al. 1994), and Minas Gerais, Brazil (Guimarães et al. 2001).

Ornamental: Melanoides tuberculata is a popular aquarium snail, often sold under the name 'Malaysian Trumpet Snail'. Some of the escapes in Florida and elsewhere, probably came from commercial tropical fish culture operations, in addition to aquarium discards (Clench 1970; Abbott 1973; McCann et al. 1996).

Ecological Impacts

Competition: Melanoides tuberculata has a proven history of replacing some native snail species in tropical and subtropical environments, an ability that has been exploited in biocontrol programs to control snails which are hosts for schistosomiasis (Pointier et al. 1994; Guimarães et al. 2001; Pointier 2001). This competitive ability is a concern, when this snail invades water bodies which contain rare native species. So far, impacts on native marine snails in Biscayne Bay are limited, in part because this snail is concentrated in sites near the mouths of canals and other disturbed sites (Wingard et al. 2007). The central Texas springs invaded by M. tuberculata are habitat for a number of endemic and highly localized snail species, including Elimia comalensis, Tryonia circumstriata, Pseudotryonia adamantia, Assiminea pecos, and Physella spp. However, in some springs, high densities of native snails coexist with M. tuberculata (Karatayev et al. 2009; Ladd and Rogowski 2012). The exact mechanisms by which M. tuberculata displace other snail species has not been studied in detail; however, behavioral interference, food competition, and egg predation are possibilities (Ladd and Rogowski 2012).

Predation: Predation on egg masses is one possible means by which M. tuberculata displaces native snails. However, in experiments using eggs of Physella spp. from a Texas spring, predation rates were quite low (Ladd and Rogowski 2012). This snail did eat the egg masses of an endangered central Texas fish, Etheostoma fonticola (Fountain Darter), but at rates lower than those of most other native and introduced snails (Phillips et al. 2010).

Habitat Change: A novel effect of a freshwater snail invasion was observed in Tobago, where large shells of M. tuberculata and Tarebia granifera (Quilted Melania) were washed down streams into coastal waters during storms, providing a new empty-shell resource for hermit crabs (Clibanarius tricolor and C. vittatus) and drastically changing the patterns of shell use (van Oosterhout et al. 2013).

Parasite-Predator Vector: Melanoides tuberculata is a host to 37+ parasites in its native range and has been a vector for at least 3 trematodes (Centrocestus formosanus; Haplorchis pumilio; Philophthalmus gralli) which have been introduced to the Western Hemisphere, and the US, including central Texas. These parasites develop in snails, then fishes, with birds and mammals as their final hosts (Karatyev et al. 2009; Alves-Pinto et al. 2011). No harmful parasites were found in a survey of M. tuberculata in Biscayne Bay, Florida. However, the 3 introduced Asian trematodes, C. formosanus and P. gralli are found in M. tuberculata in central Texas springs and streams. Centrocestus formosanus infects the gills of fishes and can cause extensive fish mortality. This trematode is a threat to several local endemic fishes, including the endangered Fountain Darter (Karatyev et al. 2009; Ladd and Rogowski 2012).

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Regional Impacts

CAR-I	Northern Yucatan, Gulf of Mexico, Florida Straits, to Middle Eastern Florida		Ecological Impact	Competition
Melanoides tuberculata is suspected to have competitive impacts on native snails on the shoreline of Biscayne Bay, because it is present in abundances which greatly exceed those of native snails (Wingard et al. 2007). The likeliest competitor is Neritina virginea (Virgin Nerite), which also grazes microalgae. However, impacts are likely to be localized, as M. tuberculata is strongly concentrated near the mouths of canals (Roessler et al. 1977; Wingard et al. 2007).
S200	Biscayne Bay		Ecological Impact	Competition
Melanoides tuberculata is suspected to have competitive impacts on native snails on the shoreline of Biscayne Bay, because it is present in abundances which greatly exceed those of native snails (Wingard et al. 2007). The likeliest competitor is Neritina virginea (Virgin Nerite), which also grazes microalgae. However, impacts are likely to be localized, as M. tuberculata is strongly concentrated near the mouths of canals (Roessler et al. 1977; Wingard et al. 2007).
FL	Florida		Ecological Impact	Competition
Melanoides tuberculata is suspected to have competitive impacts on native snails on the shoreline of Biscayne Bay, because it is present in abundances which greatly exceed those of native snails (Wingard et al. 2007). The likeliest competitor is Neritina virginea (Virgin Nerite), which also grazes microalgae. However, impacts are likely to be localized, as M. tuberculata is strongly concentrated near the mouths of canals (Roessler et al. 1977; Wingard et al. 2007).

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