USDA APHIS National Malacology Laboratory The Academy of Natural Sciences Philadelphia, PA USA

The Giant African snail (Lissachatina fulica); its history and reported biology David G. Robinson Ph.D. USDA APHIS National Malacology Laboratory The Academy of Natural Sciences Philadelphia, PA 19103 USA

I. Introduction: Four African snails of the family Achatinidae II. The Giant African snail (GAS) (Lissachatina fulica) serious pest in much of the tropics and subtropics of the world III. The Banana rasp snail (Archachatina marginata) a popular food item in West Africa, but not believed to represent major pest potential IV. Limicolaria aurora potentially a serious pest in the West Indies V. Achatina achatina a popular food item in West Africa, but not believed to represent major pest potential

I. Four species of concern Four species of Achatinidae, all members of the Giant African snail family, are frequently found in passenger baggage arriving in US ports and occasionally in containers Lissachatina fulica (GAS) from East Africa; established throughout the Indian and Pacific Basins, including Hawaii; now in South America and the West Indies Achatina achatina from West Africa Archachatina marginata from West Africa; introduced to Martinique and confirmed as still established in November 2009 Limicolaria aurora from West Africa; now established in Martinique and Singapore

The GIANT AFRICAN SNAIL The first is Lissachatina fulica (Bowdich), generally known as the Giant African snail (GAS), is widely believed to be one of the most serious pests in the world one of the 100 worst pests Achatina fulica

The GIANT AFRICAN SNAIL Summary of its Biology Snails are primarily nocturnal during the day they are generally found under protective cover (for moisture maintenance). Only where populations are very high, will individuals climb trees and walls Hatchlings and adults tend to display homing behaviour Intermediate-size snails (immatures) tend to be more active and disperse more easily GAS tends to aestivate during the dry season(s) in aggregations of up to 100 individuals, and hibernate during winter in the higher latitudes (and higher altitudes) GAS GAS will come out of aestivation/hibernation as the environment becomes wetter/warmer Hatchlings are primarily detritivorous, but can also feed on some preferred plants Intermediate-size t i snails (immatures) are phytophagous h and are serious pests Adults are detritivorous, carpophagous and necropagous As in many pulmonate snails, GAS are protandrous hermaphrodites; their male sexual organs develop first, and therefore the smaller adults behave as males The female organs develop as the snails reach their maximum size, and these snails may behave as females, or can copulate reciprocally and simultaneously Viable allosperm can be stored for more than a year after a single mating Therefore a single, fertilised adult can found an entire colony

The GIANT AFRICAN SNAIL Summary of its Biology (continued continued) Eggs are laid with 8-20 days of copulation Egg clutches (depending on maturity and environmental conditions) can reach 500 or more eggs Eggs hatch within 1-17 days Individual snails can live for 4-5 years (up to 9 years in captivity) Older individuals are more likely to vector Angiostrongylus cantonensis the Rat lungworm For more detailed information on GAS, see Raut and Barker, 2002, p. 55-114 in Molluscs as Crop Pests, CABI Publishing, 468 p. or, download the.pdf: http://www.petsnails.co.uk/docs/afulica_raut_and_barker.pdf

SPREAD OF GIANT AFRICAN SNAIL IN AFRICA, ASIA AND THE PACIFIC A. Distribution- Original distribution: 1. East Africa (coastal Kenya and Tanzania, etc.) - multiple haplotypes 2. Movement out of Africa in the 19 th century a single haplotype B. Early spread in the 1800 s was primarily intentional: 1. To Madagascar prior to 1800 2. To Mauritius and other Mascarene Islands ca. 1800 (for medicinal purposes ). Brews of the snail meat was used for respiratory ailments 3. To eastern India in 1847 by a naturalist (a British sea captain), spreading throughout southern India throughout the rest of the century 4. Introduced into Ceylon (now Sri Lanka) in 1900, probably intentionally

SPREAD OF GIANT AFRICAN SNAIL IN AFRICA, ASIA AND THE PACIFIC C. The spread continued in the 20 th century intentionally and accidentally: 1. Spread in the 20 th Century was Spread from Ceylon in the early 1900 s to southeast Asia (Malaya, Sumatra, Java, Borneo, etc.- probably accidentally 2. Introduced into Japan by 1935- most believe intentionally 3. By the beginning of World War II (1939), the Giant African Snail was intentionally spread as a potential food source, primarily for Japanese prisoner-of-war camps 4. Post-war commerce spread the snail across the Pacific Basin (accidentally as hitch-hikers ), reaching Samoa in 1990 and Micronesia in 1998 5. Some intentional introductions (for escargot -farming, e.g. Tahiti in 1967)

SPREAD OF GIANT AFRICAN SNAIL IN AFRICA, ASIA AND THE PACIFIC C. The spread continued in the 20 th century intentionally and accidentally (continued): 6. The Giant African Snails reached the Hawaiian Islands in 1936, intentionally to two islands, and later as accidental hitch-hikers. 7. Introduced into Florida in 1966 by a tourist returning from Hawaii; eradicated by 1972. 8. Accidentally introduced into Queensland, Australia, in 1977; successfully eradicated the same year due to the rapid response by the Australian quarantine authorities

SPREAD OF GIANT AFRICAN SNAIL IN AFRICA, ASIA AND THE PACIFIC (modified from Raut & Barker, 2002)

SPREAD OF GIANT AFRICAN SNAIL IN THE LESSER ANTILLES The intentional introduction of the giant African snail (GAS) into the Lesser Antilles occurred in the Parc National de la Guadeloupe, on the island of Basse-Terre in 1984 (probably from French islands in the Southwestern Pacific) Ground Zero: La Cascades aux Écrevisses, Parc National de la Guadeloupe (photo: M. Ciomperlik)

SPREAD OF GIANT AFRICAN SNAIL (continued) Infestation ti in St. Lucia (since 2000) Two color forms of GAS in Choc (near Castries), St. Lucia GAS in trees in Bisée

SPREAD OF GIANT AFRICAN SNAIL (continued) The intentional introduction of the giant African snail (GAS) into the Lesser Antilles occurred in the Parc National de la Guadeloupe, on the island of Basse-Terre in 1984 In 1987 school-children children transported GAS to Sainte-Anne on Grande-Terre, Guadeloupe, and shortly thereafter the snail spread throughout both principal islands of Guadeloupe By 1988 the island of Martinique became infested By 1995 the islands of Marie-Galante and Saint Martin/Sint Maarten became infested In early 2000, agriculture officials indicated that GAS was introduced to northern Saint Lucia Later the same year (2000) the snail was detected in Barbados, probably introduced on shipments from Martinique to the Mount Gay rum factory

SPREAD OF GIANT AFRICAN SNAIL (continued) Barbados infestation ti (since 2000) GAS on tree trunks in Salisbury Dead shells along the roadside

SPREAD OF GIANT AFRICAN SNAIL (continued) Infestation ti in Barbados (since 2000) Juveniles under a rock, Dairy Meadows, St. James Copulating pair, Rock Hall, St. Thomas

SPREAD OF GIANT AFRICAN SNAIL (continued) The intentional introduction of the giant African snail (GAS) into the Lesser Antilles occurred in the Parc National de la Guadeloupe, on the island of Basse-Terre in 1984 In 1987 school-children children transported GAS to Sainte-Anne on Grande-Terre, Guadeloupe, and shortly thereafter the snail spread throughout both principal islands of Guadeloupe By 1988 the island of Martinique became infested By 1995 the islands of Marie-Galante and Saint Martin/Sint Maarten became infested In early 2000, agriculture officials indicated that GAS was introduced to northern Saint Lucia, and later the same year the snail was detected in Barbados By 2005 GAS was established in Anguilla, and in April of 2007 a population was detected in northern Dominica GAS was also introduced into northern Trinidad in 2008? (not shown on map) GAS is now in Aruba as of 2011

The GIANT AFRICAN SNAIL (continued) The damage caused by GAS (L. fulica) in the Southeast Asia and islands groups in the Pacific is reported as extremely extensive However, in the Lesser Antilles, the damage has been less than anticipated; the reasons why are debatable. Some preliminary genetic research at the University of Cambridge (UK) indicates that most GAS populations worldwide are identical. Nevertheless, Guadeloupe has reported damage to sugar cane seedlings, as well as cucumber, banana, yam, dasheen, and papaya Initially, fruit trees in Saint Lucia were targeted, including papaya, mango, beadrfruit, and some ornamentals In Barbados, there are unconfirmed reports of some damage to cabbage, cucumber, sweet potato, and breadfruit In the Antilles in general, GAS has been more of a garden pest and a nuisance Due to expanding populations of GAS in areas of human habitation, householders complain increasingly about the accumulations of snail faeces, slime trails and decomposing snails on their property Potentially the nuisance aspect of large populations of these snails will start to affect tourism Reports of disease associated with GAS will affect tourism

The GIANT AFRICAN SNAIL (continued) GAS (Achatina fulica) is recognized known vector of eosinophilc meningitis, particularly in the Pacific Basin the distribution of snails mirrors the distribution of the Rat lungworm (Angiostrongylus cantonensis) that affects humans and livestock The Rat lungworm has been reported in Cuba, Jamaica, Hispaniola and Puerto Rico, i.e. where GAS has not yet been introduced, and also in Martinique So far, GAS become established in the Lesser Antilles, where no Rat lungworm has been reported (other than Martinique) GAS is a known vector of Aeromonas hydrophila, a gram-negative bacterium causing a wide range of symptoms, including serious gastroenteritis, cellulitis, myonecrosis, and eczema in children and people with compromised immune systems GAS is a known vector of Salmonella, a gram-negative bacterium causing diarrhea, fever and abdominal cramps GAS is also a vector of plant pathogens, including Phytophthora spp. that cause disease in cacao, coffee, banana, papaya and other crops

SPREAD OF GIANT AFRICAN SNAIL (continued) The intentional introduction of the giant African snail (GAS) into the Lesser Antilles occurred in the Parc National de la Guadeloupe, on the island of Basse-Terre in 1984 In 1987 school-children children transported t GAS to Sainte-Anne on Grande-Terre, Guadeloupe, and shortly thereafter the snail spread throughout both principal islands of Guadeloupe By 1988 the island of Martinique became infested By 1995 the islands of Marie-Galante and Saint Martin/Sint Maarten became infested In early 2000, agriculture officials indicated that GAS was introduced to northern Saint Lucia, and later the same year the snail was detected in Barbados By 2005 GAS was established in Anguilla, and in April of 2007 a population was detected in northern Dominica In March 2008, Antigua reported infestations on the island

Two other species of African snails are introduced in Martinique; Archachatina marginata and Limicolaria aurora How they reached Martinique is uncertain: - possibly intentionally introduced by someone as pets - possibly introduced by workers returning from West Africa

SPREAD OF GIANT AFRICAN SNAIL (continued (continued)) In South America Lissachatina fulica is widely distributed In Brazil, the snail was intentionally introduced to Curitiba, Paraná Currently 25 of the 26 states are reported as infested GAS was introduced into Venezuela for the beautyy cream industry More than 3 years ago, a fisherman in Argentina, introduced the snail from Paraná Paraná, Brazil Detected the snail in Ecuador 2005; now more than half the country is infested G l Galapagos en 2010 (2006?) Colombia? Bolivia - confirmed two years ago Paraguay reported GAS this year

Archachatina marginata (continued) Archachatina marginata (Swainson) is the largest species of the invasive i achatinids id (it can considerably larger than GAS up to 21 cm in length, and is sometimes known as the Banana rasp snail It is believed to have been introduced in Martinique in the early 1990 s Even though it is phytophagous, it is not considered to represent a major pest threat as its reproductive capacity is very low It lays small clutches of pigeon egg-sized eggs It was part of the 2009 Miami infestation, but does not seem to have been established Archachatina marginata (photo: P. Marquez)

Archachatina marginata (continued) Archachatina marginata is a popular food item in West Africa; a recent study (Udofia, 2009) indicates that its meat has more protein and iron, and is better-flavoured than beef In contrast, West Africans refuse to consume GAS as it has poor flavour Archachatina marginata for sale as food near Ibadan, Nigeria (photos: L. Garcia)

Archachatina marginata (continued) Archachatina marginata is frequently intercepted in US. airports in airline passenger baggage from West Africa, both alive and/or already barbecued Archachatina marginata confiscated from passenger baggage in Detroit airport (photos: B. Sullivan)

New Pest Response Guidelines from USDA APHIS PPQ a good resource for preparing for giant African snail infestations Obtain.pdf online at: http://www.aphis.usda.gov/import / po t _export/plants/manuals/emergen cy/downloads/nprg_gas.pdf Should be modified according to the current experience in Miami

The Miami Infestations In 1966 GAS was introduced into Miami, Florida in by young boy returning from a holiday in Hawaii, carrying 3 live snails in his baggage The USDA and Florida Department of Agriculture killed 18,000 snails and enumerable eggs; 1973 declared successfully eradicated; 1975 eradication costs determined to be almost $1 million (approximately $3 million in today s $$ s)

The Miami Infestations Introduction into the USA: Lissachatina fulica

Potential risk for Lissachatina infestation on the mainland (courtesy of G. Fowler & J. Smith (CPHST PERAL)

The Miami Infestations (continued) On 8 September 2011, the USDA was notified by Florida Department of Agriculture of possible Giant African snail (Lissachatina fulica) = GAS USDA initiated field operations and confirmed the identification of the GAS on 9 September 2011

Conclusions The giant African snail (GAS )(Lissachatina fulica) will eventually be introduced into every country in the South America, the West Indies, Central America and southern USA; our only realistic goal is to delay its progress for as long as possible. Once introduced, GAS is likely spread rapidly, especially in limestone (high calcium carbonate) environments, areas of human habitation, agricultural areas, and other disturbed habitats GAS has the potential to be a serious agricultural pests and disease carriers; however, in the West Indies, it has been more of a nuisance, and its impact on agriculture (so far) been relatively low (as compared with Southeast Asia and the Pacific). This does not mean that t GAS will never be a serious pest in the New World; in some countries in southern Asia, the snail was well established for decades before changing its feeding preferences to include economically important crops. GAS can be an important vector of diseases: to humans, livestock and plant crops GAS can and will affect tourism

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