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Datasheet

Lissachatina fulica (giant African land snail)

Summary

  • Last modified
  • 20 February 2014
  • Datasheet Type(s)
  • Invasive Species
  • Pest
  • Natural Enemy
  • Host Animal
  • Preferred Scientific Name
  • Lissachatina fulica
  • Preferred Common Name
  • giant African land snail
  • Taxonomic Tree
  • Domain: Eukaryota
  •     Kingdom: Metazoa
  •         Phylum: Mollusca
  •             Class: Gastropoda
  •                 Subclass: Pulmonata
  • Principal Source
  • Datasheet under review

    ...
  • Summary of Invasiveness
  • The giant African land snail L. fulica is a fast-growing polyphagous plant pest that has been introduced from its native range in East Africa to many parts of the world as a commercial food source (for humans, fish and livestock) and as a novelty ...

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Pictures

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PictureTitleCaptionCopyright
Lissachatina fulica (giant African land snail); various specimens, collected nr. Corrientes, Argentina. May, 2013.
TitleAdults
CaptionLissachatina fulica (giant African land snail); various specimens, collected nr. Corrientes, Argentina. May, 2013.
Copyright©Roberto E. Vogler & Ariel A. Beltramino-2013
Lissachatina fulica (giant African land snail); various specimens, collected nr. Corrientes, Argentina. May, 2013.
AdultsLissachatina fulica (giant African land snail); various specimens, collected nr. Corrientes, Argentina. May, 2013.©Roberto E. Vogler & Ariel A. Beltramino-2013
Lissachatina fulica (giant African land snail); adults, collected nr. Corrientes, Argentina. May, 2013.
TitleAdults
CaptionLissachatina fulica (giant African land snail); adults, collected nr. Corrientes, Argentina. May, 2013.
Copyright©Roberto E. Vogler & Ariel A. Beltramino-2013
Lissachatina fulica (giant African land snail); adults, collected nr. Corrientes, Argentina. May, 2013.
AdultsLissachatina fulica (giant African land snail); adults, collected nr. Corrientes, Argentina. May, 2013.©Roberto E. Vogler & Ariel A. Beltramino-2013
A. fulica on vegetation on Martinique, shortly after its discovery on the island in 1989.
TitleAdult
CaptionA. fulica on vegetation on Martinique, shortly after its discovery on the island in 1989.
Copyright©Charles Schotman
A. fulica on vegetation on Martinique, shortly after its discovery on the island in 1989.
AdultA. fulica on vegetation on Martinique, shortly after its discovery on the island in 1989.©Charles Schotman
A. fulica on Martinique.
TitleAdult
CaptionA. fulica on Martinique.
Copyright©Charles Schotman
A. fulica on Martinique.
AdultA. fulica on Martinique.©Charles Schotman
A. fulica on Martinique.
TitleAdults
CaptionA. fulica on Martinique.
Copyright©Charles Schotman
A. fulica on Martinique.
AdultsA. fulica on Martinique.©Charles Schotman
Lissachatina fulica (giant African land snail); eggs. Note scale. nr. Corrientes, Argentina. May, 2013.
TitleEggs
CaptionLissachatina fulica (giant African land snail); eggs. Note scale. nr. Corrientes, Argentina. May, 2013.
Copyright©Roberto E. Vogler & Ariel A. Beltramino-2013
Lissachatina fulica (giant African land snail); eggs. Note scale. nr. Corrientes, Argentina. May, 2013.
EggsLissachatina fulica (giant African land snail); eggs. Note scale. nr. Corrientes, Argentina. May, 2013.©Roberto E. Vogler & Ariel A. Beltramino-2013

Identity

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Preferred Scientific Name

  • Lissachatina fulica (Bowdich)

Preferred Common Name

  • giant African land snail

Other Scientific Names

  • Achatina fulica Bowdich 1822 formerly Férussac 1821

International Common Names

  • English: African giant snail; giant African snail; kalutara snail
  • Spanish: acatina africana; caracol gigante africano; gran caracol africano
  • French: achatine de Madagascar; achatine foulque; achatine mauritanienne; escargot géant africain

Local Common Names

  • Brazil: caramujo gigante africano
  • Germany: Afrikanische Riesenschnecke; grosse Achatschnecke
  • Italy: acatina africana; acatina dell' Isola Maurizio
  • Netherlands: agaatslak; grote Afrikaanse slak

EPPO code

  • ACHAFU (Achatina fulica)

Summary of Invasiveness

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The giant African land snail L. fulica is a fast-growing polyphagous plant pest that has been introduced from its native range in East Africa to many parts of the world as a commercial food source (for humans, fish and livestock) and as a novelty pet. It easily becomes attached to any means of transport or machinery at any developmental stage, is able to go into a state of aestivation in cooler conditions and so is readily transportable over distances. Once escaped it has managed to establish itself and reproduce prodigiously in every tropical location. As a result, L. fulica has been classified as one of the world's top 100 invasive species by The World Conservation Union, IUCN (ISSG, 2003).

Taxonomic Tree

Top of page
  • Domain: Eukaryota
  •     Kingdom: Metazoa
  •         Phylum: Mollusca
  •             Class: Gastropoda
  •                 Subclass: Pulmonata
  •                     Order: Stylommatophora
  •                         Suborder: Sigmurethra
  •                             Superfamily: Achatinoidea
  •                                 Family: Achatinidae
  •                                     Genus: Achatina
  •                                         Species: Lissachatina fulica

Notes on Taxonomy and Nomenclature

Top of pageSince Férussac in 1821, the giant African land snail Achatina fulica has been recognized as a distinct species of pulmonate gastropod belonging to the family Achatinidae. Originally, three races based on colour variation were described from the coastal regions of Kenya and Tanzania. Two of these, A. fulica rodatzi from Zanzibar and the wider ranging A. fulica lamillei have proved to be otherwise morphologically identical. The third, A. fulica bloyeti, is now considered a different species, A. bloyeti. A. fulica has become long-established in many areas of the world largely as a result of passive dispersal, especially in the Indo-Pacific region where it has now become sufficiently distinct to be designated A. fulica fulica. A. fulica was recently redesignated Lissachatina fulica, largely on the basis of Mead's (1961) observations (Naggs, 2002).

Description

Top of pageL. fulica is distinctive in appearance and is readily identified by its large size and relatively long, narrow, conical shell. Reaching a length of up to 20 cm, the shell is more commonly in the size range 5 to 10 cm. The colour can be variable but is most commonly light brown, with alternating brown and cream bands on young snails and the upper whorls of larger specimens. The coloration becomes lighter towards the tip of the shell, which is almost white. There are from six to nine spirally striate whorls with moderately impressed sutures. The shell aperture is ovate-lunate to round-lunate with a sharp, unreflected outer lip. The mantle is dark brown with rubbery skin. There are two pairs of tentacles on the head: a short lower pair and a large upper pair with round eyes situated at the tip. The mouth has a horned mandible containing some 80,000 teeth (Schotman, 1989). Eggs are spherical to ellipsoidal in shape (4.5-5.5 mm in diameter) and are yellow to cream in colour.

Distribution

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L. fulica was originally a native of East African coastal regions, though apart from Tanzania, there is no recent literature on its distribution and abundance in these countries. Its dispersal from Kenya and Tanzania has probably been checked by natural enemies, though it has most likely been long established in the following additional countries: Mozambique, Somalia, Ethiopia, Eritrea, Uganda, Burundi, Rwanda, Democratic Republic of Congo, Malawi, Zambia, Zimbabwe, South Africa and Madagascar.

However, L. fulica owes most of its current wide distribution to human activity (Dharmaraja, 1984). It is now present everywhere in the Indo-Pacific except Banaba Island, Cook Islands, Lord Howe Island, Nauru, Niue, Norfolk Island, Pitcairn Island, Tokelau, Australia and New Zealand, and has been eradicated from Tuvalu (Cowie, 2000). In 1989 it was recorded on both Martinique and Guadeloupe in the Caribbean (Schotman, 1989). It was introduced to South America (Brazil) in the late 1980s as a commercial species, is now present in 25 out of 26 Brazilian states and has been found to be spreading in other countries on the continent (Vogler et al., 2013). It is also likely to have become an established part of the snail fauna of West Africa following reports from Côte d'Ivoire, Togo and Nigeria, and a shell has been identified in Morocco (van Bruggen, 1987), the first discovery of this species from anywhere in the Palearctic.

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.

CountryDistributionLast ReportedOriginFirst ReportedInvasiveReferencesNotes

ASIA

BangladeshPresentIntroducedInvasiveMead, 1979
Brunei DarussalamPresentEPPO, 2013
CambodiaPresentIntroducedInvasiveMead, 1979
ChinaRestricted distributionIntroducedInvasiveSchotman, 1989; EPPO, 2013
-FujianPresentLi et al., 2006
-GuangdongPresentMeng et al., 2007
-GuizhouPresentTang et al., 2008
-HainanPresentHu et al., 2007
-Hong KongWidespreadIntroduced1937Schotman, 1989; Mead, 1961; EPPO, 2013
-YunnanPresentChen et al., 2008
Christmas Island (Indian Ocean)PresentIntroducedInvasiveSproul, 1983; Lake & O'Dowd, 1991
IndiaWidespreadIntroduced1847InvasiveRaut & Ghose, 1984; Srivastava et al., 1985; Mead, 1961; EPPO, 2013
-Andaman and Nicobar IslandsWidespreadIntroducedInvasiveShah, 1992
-Andhra PradeshPresentReddy & Sreedharan, 2006
-Arunachal PradeshPresentRamakrishna et al., 2005
-BiharWidespreadIntroducedInvasiveThakur, 1998
-KarnatakaWidespreadIntroducedInvasiveShree & Kumar, 2002; Basavaraju et al., 2000
-Madhya PradeshWidespreadIntroducedInvasiveThakur & Kumari, 1998
-OdishaPresentIntroducedInvasiveMead, 1961
-RajasthanPresentIntroduced1996InvasiveTehsin & Sharma, 2000
-Tamil NaduPresentIntroducedInvasivePadmanaban et al., 2000
-Uttar PradeshPresentIntroducedInvasiveSaxena & Mahendru, 2000
-West BengalWidespreadIntroducedInvasiveRaut & Ghara, 1990; Raut, 1991
IndonesiaRestricted distributionIntroducedInvasiveMead, 1979; EPPO, 2013
-Irian JayaPresentIntroducedInvasiveMead, 1979
-JavaPresentIntroducedInvasiveSchneider et al., 1998; Mead, 1961
-KalimantanPresentEPPO, 2013
JapanRestricted distributionIntroduced1935InvasiveTakeuchi et al., 1991; Tomiyama, 2002; Mead, 1961; EPPO, 2013
-Ryukyu ArchipelagoPresentIntroduced1935InvasiveMead, 1961; EPPO, 2013
MalaysiaRestricted distributionMead, 1979; EPPO, 2013
-Peninsular MalaysiaPresentIntroduced1911InvasiveMead, 1979; Mead, 1961; EPPO, 2013
-SarawakRestricted distributionIntroduced1928InvasiveMead, 1979; Mead, 1961; EPPO, 2013
MaldivesWidespreadIntroduced1957InvasiveMuniappan, 1987; Mead, 1961; EPPO, 2013
MyanmarRestricted distributionIntroducedInvasiveSchotman, 1989; Mead, 1961; EPPO, 2013
PhilippinesWidespreadIntroduced1949InvasiveMead, 1961; EPPO, 2013
SingaporeWidespreadIntroduced1917InvasiveMead, 1979; Mead, 1961; EPPO, 2013
Sri LankaRestricted distributionIntroduced1900InvasiveMead, 1961; EPPO, 2013
TaiwanRestricted distributionIntroduced1932InvasiveMead, 1961; EPPO, 2013
ThailandRestricted distributionMead, 1979; EPPO, 2013
VietnamRestricted distributionIntroduced1937InvasiveMead, 1979; Mead, 1961; EPPO, 2013

AFRICA

Central African RepublicPresentMbétid-Bessane, 2006
Côte d'IvoirePresentIntroducedInvasivede Winter, 1989
KenyaWidespreadIntroducedInvasiveMead, 1979
MadagascarPresentEPPO, 2013
MauritiusWidespreadIntroduced1803InvasiveMead, 1961; Facknath et al., 1999; EPPO, 2013
MoroccoAbsent, intercepted onlyvan Bruggen, 1987
NigeriaPresentAshaye et al., 2001
RéunionWidespreadIntroduced1821InvasiveSchotman, 1989; Graber et al., 1997; Mead, 1961; EPPO, 2013
SeychellesWidespreadIntroduced1840InvasiveSchotman, 1989; Mead, 1961; EPPO, 2013
TanzaniaWidespreadNativeNot invasiveEmberton et al., 1997
TogoPresentIntroducedInvasiveEkoue & Kuevi-Akue, 2002

NORTH AMERICA

USARestricted distributionEPPO, 2013
-FloridaRestricted distributionEPPO, 2013; Mead, 1979; NAPPO, 2011; NAPPO, 2013; NAPPO, 2013
-HawaiiWidespreadIntroduced1936InvasiveCowie, 1998; Mead, 1961; EPPO, 2013

CENTRAL AMERICA AND CARIBBEAN

AnguillaPresentEPPO, 2013
Antigua and BarbudaTransient: actionable, under eradicationEPPO, 2013
BarbadosPresentEPPO, 2013
DominicaPresentEPPO, 2013
GuadeloupePresentIntroduced198*InvasiveAnon., 1989; EPPO, 2013
MartiniqueRestricted distributionIntroduced198*InvasiveAnon., 1989; EPPO, 2013; Mead & Paley, 1992
Netherlands AntillesPresentEPPO, 2013
Saint LuciaRestricted distributionEPPO, 2013; Jn Pierre, 2008
Trinidad and TobagoRestricted distributionIntroducedIPPC, 2009; EPPO, 2013Found at Alyce Glen Gardens Petit Valley, North West Trinidad in Oct. 2008; restricted to an area 1.0 km²
United States Virgin IslandsPresentIntroducedInvasiveMiller, 1994

SOUTH AMERICA

ArgentinaRestricted distributionIntroducedInvasiveVogler et al., 2013; Gutiérrez & Gregoric Núñez, 2012; Gutiérrez Gregoric et al., 2011First detected in 2010 in Puerto Iguazú City (25°36S, 54°35W)
BoliviaPresent only in captivity/cultivationIntroducedVogler et al., 2013Commericalized in La Paz, not reported from the wild
BrazilWidespreadIntroduced1980sInvasivePaiva, 1999; Vogler et al., 2013; EPPO, 2013
-BahiaPresentIntroducedInvasivePaiva, 1999
-Espirito SantoPresentIntroducedInvasivePaiva, 1999
-GoiasPresentPaiva, 1999; do Amaral, 2002
-MaranhaoPresentIntroducedInvasivedo Amaral, 2002
-Minas GeraisWidespreadIntroducedInvasivePaiva, 1999
-ParanaWidespreadIntroducedInvasivePaiva, 1999
-PernambucoPresentIntroducedInvasivePaiva, 1999
-Rio de JaneiroWidespreadIntroducedInvasiveDos Santos et al., 2002; Paiva, 1999; de Vasconcellos & Pile, 2001
-RondoniaPresentIntroducedInvasivePaiva, 1999
-Santa CatarinaWidespreadIntroducedInvasivePaiva, 1999; Magenta Magalhães et al., 2001
-Sao PauloWidespreadIntroducedInvasivePaiva, 1999
ColombiaPresentIntroducedInvasiveVogler et al., 2013; EPPO, 2013
EcuadorPresentIntroducedInvasiveVogler et al., 2013; EPPO, 2013
ParaguayPresentIntroducedInvasiveVogler et al., 2013
PeruPresentIntroducedInvasiveVogler et al., 2013; EPPO, 2013
VenezuelaPresentIntroduced1996InvasiveVogler et al., 2013; EPPO, 2013; Martinez-Escarbassiere & Martinez-Moreno, 1997

OCEANIA

American SamoaWidespreadIntroduced1977InvasiveTillier et al., 1993; Cowie, 2000; EPPO, 2013; Tauili'ili & Vargo, 1993
FijiAbsent, formerly presentSouth Pacific Commission, 1979; AQIS, 2003
French PolynesiaWidespreadIntroduced1967InvasiveMurray et al., 1988; Tillier et al., 1993; Cowie, 2000; EPPO, 2013
GuamWidespreadIntroduced1943Tillier et al., 1993; Cowie, 2000; EPPO, 2013
KiribatiAbsent, formerly presentSouth Pacific Commission, 1979; AQIS, 2003
Marshall IslandsPresentIntroducedInvasiveSouth Pacific Commission, 1996; Cowie, 2000; EPPO, 2013
Micronesia, Federated states ofWidespreadIntroduced1938InvasiveTillier et al., 1993; Cowie, 2000; EPPO, 2013
New CaledoniaWidespreadIntroduced1972InvasiveSouth Pacific Commission, 1979; Tillier et al., 1993; Cowie, 2000; EPPO, 2013
New ZealandAbsent, invalid recordEPPO, 2013
NiueAbsent, formerly presentSouth Pacific Commission, 1979; AQIS, 2003
Northern Mariana IslandsWidespreadIntroduced1936InvasiveTillier et al., 1993; Cowie, 2000; EPPO, 2013
PalauWidespreadIntroduced1938InvasiveTillier et al., 1993; Cowie, 2000; EPPO, 2013
Papua New GuineaWidespreadIntroducedInvasiveTillier et al., 1993; Mead, 1961; Cowie, 2000; EPPO, 2013
SamoaPresentIntroduced1990InvasiveSouth Pacific Commission, 1979; Cowie, 2000; EPPO, 2013
Solomon IslandsAbsent, formerly presentSouth Pacific Commission, 1979; Cowie, 2000; EPPO, 2013; AQIS, 2003
TongaAbsent, formerly presentSouth Pacific Commission, 1979; AQIS, 2003
TuvaluEradicatedSouth Pacific Commission, 1979; Cowie, 2000
US Minor Outlying IslandsWidespreadTillier et al., 1993
VanuatuWidespreadTillier et al., 1993; EPPO, 2013
Wallis and Futuna IslandsWidespreadIntroducedInvasiveTillier et al., 1993; Cowie, 2000; EPPO, 2013

History of Introduction and Spread

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L. fulica has been introduced to new regions for farming as a new commercial protein source. Definitive and detailed histories of the spread of L. fulica can be found in Rees (1950) and Mead (1961); and for the more recent spread across South America, see Paiva (1999) and Vogler et al. (2013). Escapes from abandoned breeding facilities, accidental transport with building materials, waste and plants, and use as fishing bait have contributed to the local spread.

Risk of Introduction

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L. fulica has demonstrated its success as an invader in the tropics. It was not introduced to South America until the late 1980s and is now present in almost all states of Brazil and is spreading in several other countries. Bioclimatic models have been used to identify further South American areas that are susceptible to invasion (Vogler et al., 2013).

Habitat List

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CategoryHabitatPresenceStatus
Littoral
Coastal areasPresent, no further detailsHarmful (pest or invasive)
Terrestrial-managed
Cultivated / agricultural landPresent, no further detailsHarmful (pest or invasive)
Disturbed areasPresent, no further detailsHarmful (pest or invasive)
Managed forests, plantations and orchardsPresent, no further detailsHarmful (pest or invasive)
Urban / peri-urban areasPresent, no further detailsHarmful (pest or invasive)
Terrestrial-natural/semi-natural
Natural forestsPresent, no further detailsHarmful (pest or invasive)
RiverbanksPresent, no further details
Scrub / shrublandsPresent, no further detailsHarmful (pest or invasive)
WetlandsPresent, no further details

Hosts/Species Affected

Top of pageL. fulica is a polyphagous pest. Its preferred food is decayed vegetation and animal matter, lichens, algae and fungi. However, the potential of the snail as a pest only became apparent after having been introduced around the world into new environments (Rees, 1950). It has been recorded on a large number of plants including most ornamentals, and vegetables and leguminous cover crops may also suffer extensively. The bark of relatively large trees such as citrus, papaya, rubber and cacao is subject to attack. Poaceous crops (sugarcane, maize, rice) suffer little or no damage from L. fulica. There are reports of L. fulica feeding on hundreds of species of plants (Raut and Ghose, 1984). Thakur (1998) found that vegetables of the genus Brassica were the most preferred food item from a range of various food plants tested.

Host Plants/Plants Affected

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Plant nameFamilyContext
Arachis hypogaea (groundnut)FabaceaeMain
Artocarpus (breadfruit trees)MoraceaeMain
BrassicaBrassicaceaeMain
Carica papaya (pawpaw)CaricaceaeMain
Cucumis melo (melon)CucurbitaceaeMain
Cucurbita pepo (marrow)CucurbitaceaeMain
Daucus carota (carrot)ApiaceaeOther
Dioscorea alata (white yam)DioscoreaceaeMain
Hevea brasiliensis (rubber)EuphorbiaceaeOther
Musa (banana)MusaceaeMain
Polyphagous (polyphagous)Main
Theobroma cacao (cocoa)SterculiaceaeOther

Growth Stages

Top of pageFruiting stage, Seedling stage, Vegetative growing stage

Symptoms

Top of pageIn garden plants and ornamentals of a number of varieties, and vegetables, all stages of development are eaten, leading to severe damage in those species that are most often attacked. However, cuttings and seedlings are the preferred food items, even of plants such as Artocarpus which are not attacked in the mature state. In these plants damage is caused by complete consumption or removal of bark. Young snails up to about 4 months feed almost exclusively on young shoots and succulent leaves. The papaya appears to be the only fruit which is seriously damaged by L. fulica, largely as a result of its preference for fallen and decaying fruit.

In plants such as rice, which are not targets of L. fulica, sometimes sheer weight of numbers can result in broken stems. In general, physical destruction to the cover crop results in secondary damage to the main crop, which relies on the cover crop for manure, shade, soil and moisture retention and/or nitrogen restoration. This in turn can result in a reduction in the available nitrogen in the soil and consequently marked erosion in steeper areas.

Symptoms List

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SignLife StagesType

Fruit

external feeding
lesions: scab or pitting

Growing point

external feeding

Leaves

external feeding
necrotic areas

Roots

external feeding

Stems

discoloration of bark

Whole plant

early senescence
external feeding
uprooted or toppled

Biology and Ecology

Top of pageBequaert (1951) described the biology of Achatininae in general. Mead (1961) considered the biology and ecology of L. fulica in terms of economic damage and pest control.

Genetics

Vercourt (1966) recognized two colour morphs in the East African coastal race: radatzi (unstreaked) and hamillei (streaked), which were otherwise identical in morphology. Mead (1961) commented on the noticeable variability of L. fulica in both shell size and shape, with these factors varying with locality. He reported snails on Guam commonly close to 6 inches, but on Micronesia 3.5 to 4.5 inches being closer to the average. Peterson (in Mead, 1961) recorded one specimen of just under 8 inches. Shells from Anguar in the Pelaus had a deep, rich, contrasting colour pattern, with a thick periostracum and a nacreous gloss, whereas those from Koror Island were almost white and devoid of a periostracum. White snails have been reported from Africa (Owen and Reid, 1986). Specimens from some populations on Sri Lanka and Hawaii are darker in colour with a long axis and reduced body whorl. Such variability in snails is often the result of environmental factors such as varying levels in the supply of calcium carbonate, but the genetic effect resulting from small founding populations may be responsible for much inter-island variation.

The inheritance of shell colour polymorphism in L. fulica has been studied by Allen (1983).

Reproductive Biology

Like most snails, L. fulica is hermaphroditic and, after a single mating, can produce a number of batches of fertile eggs over a period of months. Early reports of self-fertilization (van der Meer Mohr, 1949) have since been discounted. It lays eggs in batches of 100 to 400, with up to 1200 being laid in a year. These hatch after about 8-21 days under tropical conditions. They are laid on the ground, often in the base of plants. The prodigality of L. fulica is renowned and the literature contains many anecdotes and astonishing estimates of the potential number of progeny if all were to survive (see Mead, 1961, 1979). For example, 20 tonnes of snails were collected on one day in Fiji just 4 years after its introduction.

Courtship behaviour was studied in the field by Tomiyama (1994). This was exclusively a nocturnal activity. Duration in copulation was shorter in young adults than in older adults but occurred during most periods of the night. Older adults were found to copulate only in the middle of the night. Copulation time ranged from 1.5 to 7.5 hours with an average time of 4.6 hours. Courtship behaviour of L. fulica follows a fixed pattern, with different behaviour occurring between the courtship initiator and the acceptor, in contrast to many other species of land snails where it is immediately reciprocal.

Tomiyama (1993) also investigated the correlation between shell growth and the maturation of the reproductive system. The reproductive gland was formed by the end of the year after hatching. Shell growth continued for some months after, but no reflection of the lip was observed - an event commonly considered to indicate sexual maturity in most land snails. Only sperm were produced while shell growth continued, but both sperm and eggs were produced when shell growth stopped. The snails had then become completely hermaphroditic.

Although the adult has an average life span of 5-6 years it may live for as long as 9 years. It will readily enter a state of aestivation and can survive for years in this state. The tendency for a number of aestivating snails to be present in an area at any time, particularly when conditions are optimum for activity, can make control measures difficult.

The life history of Achatina snails was described by Williams (1951). More recently, investigations carried out by Raut (1991) in West Bengal, India, found that L. fulica became sexually mature in their second year and produced 2-13 egg clutches overall up to their fifth year. The number of eggs per clutch ranged from around 30 to 300, with a gradual increase occurring in line with the age of the snails. The mean fertility rate was 94.6%, but again was higher in clutches laid by older snails than in those laid by younger snails. Birth rate varied with the age of the individuals. There was no recruitment up to the age of 422 days (including the aestivation period). The period between ovipositions (21-76 days) was lower during active periods when both active and aestivation periods were considered. The snails survived for between 1 and 1562 days. Mortality rate was higher in young snails, especially during aestivation. Overall, from the time of hatching to sexual maturity, mortality was found to be 49%. The life table for L. fulica indicated that 0-day-old individuals would survive for 638.5 days, whereas individuals that were 1550 days old were expected to survive for another 25 days. The mean total output of eggs per season varied considerably from 100+ to 700, with an average lifetime output of 1300. Raut's findings in general concur with those of other studies.

Environmental Requirements

Although L. fulica is a tropical snail, it can survive cold conditions, even snow, by aestivating, though it is unable to establish itself in temperate regions. It is normally nocturnal and crepuscular in its habits, though it will become active in the daytime during rainy or overcast periods. This indicates that light, temperature, moisture and food are all vital factors in snail activity. There is evidence that growth rate may be density dependent (Sidelnikov and Stepanov, 2000).

Natural Enemies

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Natural enemyTypeLife stagesSpecificityReferencesBiological control inBiological control on
Centropus sinensisPredatorTehsin & Sharma, 2000
Euglandina roseaPredatorAdultsCaribbean
Gecarcoidea natalisPredatorAdults
Gonaxis quadrilateralisPredatorAdults/Eggs/Larvae/Nymphs/Pupae
LamprophorusPredatorAdults
Platydemus manokwariPredatorAdultsMaldives; Philippines
Solenopsis geminataPredatorAdults

Notes on Natural Enemies

Top of pageMead (1961) commented on the paucity of natural enemies of L. fulica in its native East Africa. Although animals from many groups, including birds, mammals and reptiles, will feed on achatinids, he was reviewing the topic in terms of potential natural control in regions where they had been introduced. He noted that those that it does have are of questionable value in effecting any real control. This is the major reason why foreign predators, notably Euglandina rosea and Platydemus manokwari, have been sought as agents of biological control, having been introduced into Pacific islands from Florida, USA, and New Guinea respectively. Two species of East African carnivorous snails of the genus Gonaxis have also been established in many areas, though accounts vary as to their effectiveness on islands of introduction, where their impact on crop protection has been minimal. In addition, two East African species of predatory carabid beetles and five species of lampyrid beetle from South-East Asia are known to prey on L. fulica.

On Christmas Island (Kiritimati), there is good evidence that the endemic red crab, Gecarcoidea natalis, restricts the distribution of L. fulica, especially in undisturbed rain forest (Lake and O'Dowd, 1991). Predatory hermit crabs, primarily Coenibitus perlatus, and robber crabs, Birgus latro, have made it difficult for L. fulica to become established on coral islands. Rats, centipedes, millipedes and fire ants are all known to prey on the giant African snail on Indo-Pacific Islands. In Rajasthan in India, L. fulica was observed by Tehsin and Sharma (2000) to be eaten by the coucal (Centropus sinensis), a small bird related to the cuckoo.

Means of Movement and Dispersal

Top of pageMost dispersal of L. fulica has occurred accidentally, with all developmental stages becoming attached to machinery (e.g., road construction, landscaping) unobserved.

Natural Dispersal

L. achatina is self-propelled.

Agricultural Practices

Eggs and snails accidentally become attached to agricultural machinery and vehicles, and are readily transported in garden waste.

Pathway Causes

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CauseNotesLong DistanceLocalReferences
AgricultureAccidentalYesYes
Escape from confinement/ garden escapeYes
FoodIntroduced as a foodYes
Garden waste disposalAccidentalYes
HitchhikerYesYes
HorticultureAccidentalYesYes
Live food/feed tradeYes
Pet/aquarium tradeYesYes
ResearchYes
Self-propelledYes

Pathway Vectors

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VectorNotesLong DistanceLocalReferences
AircraftYes
BaitYes
Clothing/footwear and possessionsYes
Land vehiclesYes
Plants or parts of plantsNursery tradeYes
Ship/boat structures above the water line/ holds aYes
Soil, sand, gravel etc.Yes

Plant Trade

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Plant parts liable to carry the pest in trade/transportPest stagesBorne internallyBorne externallyVisibility of pest or symptoms
Bulbs, Tubers, Corms, Rhizomeseggs; juvenilesNoYesPest or symptoms usually visible to the naked eye
Leaveseggs; juvenilesNoYesPest or symptoms usually visible to the naked eye
Rootseggs; juvenilesNoYesPest or symptoms usually visible to the naked eye
Stems (above ground), Shoots, Trunks, Brancheseggs; juvenilesNoYesPest or symptoms usually visible to the naked eye
Plant parts not known to carry the pest in trade/transport
Flowers, Inflorescences, Cones, Calyx
Growing medium accompanying plants
Seedlings, Micropropagated plants
True seeds (inc. grain)
Wood

Economic Impact

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L. fulica has a voracious appetite and has been recorded as attacking over 50 different kinds of plants although it has a preference for breadfruit, cassava, cocoa, papaya, peanut, rubber and most species of legumes and cucurbits. The economic impact of L. fulica was considered to be so profound that the new discipline of economic malacology was formulated by zoologist Albert Mead (Mead, 1961, 1979) to take account of a pest species which appeared to be threatening already inadequate food supplies in poor regions of the world. Mead devoted his book to the economic impact of L. fulica. There had been reports that the species would devour virtually anything found in the garden (Jarrett, 1923, 1931) and Mead evaluated L. fulica as a major horticultural and agricultural pest. However, many reports of widespread damage by L. fulica have been anecdotal and more recently Civeyrel and Simberloff (1996) suggest that the lasting impact of L. fulica on agriculture may not be severe, and the human health risk is probably minor.

It is difficult to quantify the damage wrought by L. fulica to gardens and crops, but suffice to say that it is considered by most authorities to be the most damaging land snail in the world. However, Civeyrel and Simberloff (1996) believe that the damage done to endemic species of snail by ill-judged biological control programmes outweighs the impact of the pest species. The dramatic population crashes commonly observed in populations of L. fulica which had increased rapidly in size following introduction into new environments, may well lessen the deleterious long-term economic impact of the species, though it remains a serious pest in many areas.

Indirectly, L. fulica may have an impact as a vector of plant diseases as it has been implicated in the transmission of Phytophthora palmivora (Schotman, 1989).

Environmental Impact

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Cowie (ISSG, 2003) believes the agricultural impacts of L. fulica may have been exaggerated, the nuisance factor perhaps being more important. By reaching such enormous numbers and invading native ecosystems they also pose a serious conservation problem by eating native plants, modifying habitat, and probably out-competing native snails.

Social Impact

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Perhaps the biggest social impact wrought by L. fulica is its nuisance value as large numbers of snails build up. Not only are they unsightly but cadavers make smell and mess, especially where they are run over by traffic, which invariably happens during rapid growth of numbers.

L. fulica can act as a vector of the human disease, eosinophilic meningitis, which is caused by the rat lungworm parasite, Angiostrongylus cantonensis. The parasite is passed to humans through eating raw or improperly cooked snails or freshwater prawns. It is therefore advisable to wash one's hands after handling the snail. However, Cowie (2000) states that many other introduced snails in the tropics are vectors of this parasite and the spread of the disease has not definitively been related to the spread of L. fulica.

Uses

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In its native East Africa, L. fulica is a source of protein for some local people. Tillier et al. (1993) stated that although L. fulica is a vector of rat lungworm which causes eosinophilic meningitis in humans, it is nevertheless suitable for human consumption if properly prepared and cooked, and has not been in contact with poisoned bait. Instructions for that process were included in the Pest Advisory Leaflet of the South Pacific Commission (Lambert, 1999). It is widely advertised as food on Chinese Internet sites. L. fulica can also be useful in making fertilizer and chicken feed. It has been tested as a cheap alternative source of feed in fish farming in Sri Lanka. In Java, although consumption depends on the consumer's ethnic background, the breeding of L. fulica can benefit disadvantaged groups and help to conserve the natural snail population (Schneider et al., 1998).

In common with a number of other molluscan species, L. fulica has been used as a source of biological compound in clinical and experimental laboratories. It has been widely utilised in neurobiology and electrophysiology (Tamamaki, 1989; Zhang et al., 1996), endocrinology (Takeda and Ohtake, 1994; Bose et al., 1997), comparative biochemistry and physiology (Kalyani, 1990; Misra and Shrivastava, 1994; Indra and Ramalingam, 1996), reproductive biology (Sretarugsa et al., 1991), parasitology (Utomo et al., (1991), immunology (Harris et al., 1992) and molecular biology (Obara et al., 1992).

In the UK and in a few US states, where it is permissable to do so, L. fulica (along with the two main West African species) is sometimes kept as a pet.

Uses List

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Animal feed, fodder, forage

  • Bait/attractant
  • Fodder/animal feed

General

  • Botanical garden/zoo
  • Laboratory use
  • Pet/aquarium trade

Human food and beverage

  • Meat/fat/offal/blood/bone (whole, cut, fresh, frozen, canned, cured, processed or smoked)

Medicinal, pharmaceutical

  • Source of medicine/pharmaceutical

Detection and Inspection

Top of pageL. fulica is a large and conspicuous crop pest which hides during the day. Surveys are best carried out at night using a flashlight. It is easily seen, and attacked plants exhibit extensive rasping and defoliation. Weight of numbers can break the stems of some species. Its presence can also be detected by signs of ribbon-like excrement, and slime trails on plants and buildings.

Similarities to Other Species/Conditions

Top of pageThere are a number of species of giant achatinids distributed across sub-Saharan Africa from Zanzibar to Freetown, though three are most often encountered. L. fulica from East Africa is somewhat smaller than its similar counterpart in West Africa, Achatina marginata, and has a more pointed apex to its shell. Another similar species, Achatina achatina, also from West Africa, is the world's largest gastropod with a maximum recorded shell length of 27 cm and a weight of almost 1 kg. A. achatina lays around six large (15 mm) eggs at a time compared to several hundred smaller (3-4 mm) eggs laid by L. fulica. All three species form part of the diet of local people.

Prevention and Control

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A vast body of literature has accrued on the various methods of control available for L. fulica. Half of Mead's 1961 book on the economic impact of the giant African snail was devoted to control. Perhaps the major problem with testing the efficacy of control measures in experiments is the ability to distinguish the level of success of the method under test and the natural demographic crashes that are a renowned feature of the population dynamics of this species. There is only one officially documented example of eradication, in Florida, USA, in 1969 (Schotman, 1989).

Biological Control

Again, Mead (1961) reviewed in considerable depth all the species of animals, from microbes to mammals, that either prey on L. fulica or may do so given the opportunity, and assessed their potential as biological control agents. Those species that offered the best potential prospects, namely predatory insects and molluscs, had a major drawback: they either failed to survive or become established on introduction, or they were likely to pose a serious potential threat to native species if they did. It is interesting to note that even 40 years ago "the predatory snails Gonaxis kibwiensis and Euglandina rosea have taken the limelight almost entirely in the program of biological control of A. fulica" (Mead 1961).

It is unfortunate for many species of terrestrial gastropods on the islands of the Indian and Pacific Oceans that the introductions of the alien predatory molluscs as biological control agents against L. fulica were often conducted in haste, contrary to expert advice and without satisfactory field trials. This has led to widespread extinctions of endemic snail species without evidence that the introductions were effective against the target species (Murray et al., 1988). The principal agent of this ecological disaster was, and is, Euglandina rosea, a carnivorous snail species from Florida, USA. It has been introduced into many islands of the Indo-Pacific in an attempt to eradicate L. fulica but has had a catastrophic effect on endemic gastropods, most notably in French Polynesia (Cowie, 1992) and Hawaii (Hadfield et al., 1993). In French Polynesia population crashes of L. fulica were observed both on Moorea, after the introduction of E. rosea, and on Huahine in the absence of the predator (Murray et al., 1988).

E. rosea and a natural predator of L. fulica, Gonaxis quadrilateralis, both failed to control the pest in India (Srivastava, 1985). Further, the local carnivorous snail, Gulella indoennea bicolor, was killed by chemicals used against L. fulica. Gonaxis kibweziensis has failed to make an impression on populations of L. fulica on Mauritius, as has Edentulina ovoidea on Réunion Island.

Good control has been reported from Guam, the Marianas Islands and in the Maldives in the 1980s with the use of a non-specific planarian worm Platydemus manokwari (Muniappan, 1990). The same researchers describe similar success on Bugsuk Island in the Philippines (Muniappan et al., 1986) by distributing the flatworm in piles of coconut husks. P. manokwari is still seen in some quarters to be a potential control agent for L. fulica (Kaneda et al., 1990) but there are dangers in introducing a species where little is known of its ecology and life history. Invertebrate researchers and conservationists are disturbed by the non-specificity of P. manokwari and its targets and the lack of control over its dispersal, and strongly recommend that no further introductions of this species take place (Pearce Kelly et al., 1994). Another flatworm, Geoplana sp., was reported to have made an impact on populations of L. fulica in Guam (Schotman, 1989) but it should have no future as a biological control agent because it is a vector of human disease.

Kurozumi (1985) has discussed the possibility of using the slug, Incillaria sp., a common predator of eggs and adults of several species of land snail in Japan, as a potential biological control agent.

Chemical Control

The most common means of chemical control against L. fulica has been the use of metaldehyde, though chemical molluscicides are no longer favoured and have proved ineffective in the control of this species. Both the authorities and local farmers on Samoa oppose the use of metaldehyde. Mead (1979) and Raut and Ghose (1984) have stated that all chemical means of control against L. fulica have so far failed. The FAO (Schotman, 1989) suggest that metaldehyde poison baits can be effective in small scale cultivation, but are not practicable elsewhere. Sharma and Agarwal (1989) recommended the use of 5% metaldehyde pellets, but only in conjunction with wider sanitation and physical control measures. However, Salmijah et al. (2000) record the development of resistance by L. fulica to metaldehyde.

Various insecticides and fungicides were tested against L. fulica by Kakoty and Das (1987) who found that only copper sulphate solution produced high mortality rates. An insecticidal bait has also been tested by Sarkar et al. (1997). Rao et al. (2000, 2003) looked at the effect of single and binary treatments of plant-derived molluscicides on reproduction and survival, and on different enzyme activities in the nervous tissue of L. fulica. Although results were positive, it was agreed that the sublethal exposure of these molluscicides on snail reproduction is a complex process. Saxena and Mahendru (2000) evaluated the efficacy of different baits (wet wheat, gram, barley and corn flour) and insecticides (malathion, trichlorfon and mexacarbate) against L. fulica, at different concentrations. The results showed that wet wheat flour was the best bait.

Mechanical Control

As L. fulica is primarily a pest in areas of human habitation, physical control can often be as effective as any other means where they congregate in large numbers. Shah (1992) recommends that snails on the Andaman and Nicobar Islands, where L. fulica is a major crop pest, can be collected and destroyed during aestivation between January and April when they hide under hedges and debris. Potential hiding places should be removed from fields. The snails can then be killed by sprinkling with salt or by exposure to the sun.

Physical control can be efficient by making a strip of 1.5 m wide bare soil around nurseries. Barriers or screens can be constructed using corrugated tin, security wire mesh and ditches dug around fields (Schotman, 1989). Snails can be collected each day and destroyed by crushing or drowning. The public can be involved in such collections by using organised campaigns. In developing countries children are often keen to be involved, treating the collection and destruction of large snails as a game. In some countries they are gathered for food in large quantities. L. fulica is one of four species of giant African snails, 30 tonnes of which are collected each year in Togo (Ekoue and Kuevi-Akue, 2002).

Integrated Pest Management

Ultimately, if control against L. fulica is to be effective it will have to involve some form of integration of the above methods. Srivastava et al. (1985) reviewed the biology and management of L. fulica in India with sections on its cultural, chemical and biological control. They noted the ineffectiveness of predatory molluscs, or their susceptibility to molluscicides used against L. fulica. An integrated approach was recommended using an aqueous extract of diseased L. fulica sprayed onto the snails and their food plants, in conjunction with non-molluscan snail predators such as the millipede, Orthomorha sp., and hermit crabs, Coenobita sp. An integrated approach is also recommended by the South Pacific Commission (Lambert, 1999) using chemical control by metaldehyde, and physical control using barriers of bare land around crops and collection after rain. The commission does not recommend the use of predatory snails as a control agent of L. fulica. Raut and Barker (2002) review the control (physical, chemical and biological) of L. fulica and other Achatinidae pests across Africa and other countries.

Phytosanitary Measures

The FAO International Plant Quarantine Treatment Manual (FAO, 1981) describes protocols for treatments to eliminate infestations of L. fulica on non-plant cargo using hydrogen cyanide and cold treatments (Schotman, 1989). In addition to the plant quarantine inspection of incoming carriers and cargo at international points of entry, thorough predeparture quarantine inspection from countries where L. fulica occurs should also be undertaken.

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Schotman CYL, 1989. Data sheet on the giant African snail Achatina fulica Bowdich (Mollusca: Achatinidae). RLAC-PROVEG, No. 19:16-21.

Shah NK, 1992. Management of the giant African snail. Indian Farming, 41(11):21.

Sharma DD, Agarwal ML, 1989. Save your crops from giant African snail. Indian Farming, 38(12):15, 22.

Sheela Thakur, Rina Kumari, 1998. Seasonal behaviour of giant African snail Achatina fulica in Bihar. Journal of Ecotoxicology & Environmental Monitoring, 8(2):153-160.

Shree MP, Kumar KR, 2002. Effect of gaint African snail-Achatina fulica Bowdich infestation on the nutritional quality of mulberry (Morus sp.) leaves. Bulletin of Indian Academy of Sericulture, 6(1):50-56.

Sidelnikov AP, Stepanov II, 2000. Influence of population density on growth and regenerative capacity of the snail Achatina fulica. Biology Bulletin of the Russian Academy of Sciences, 27(5):438-444; [translated from Izvestiya Akademii Nauk, Seriya Biologicheskaya (2000) No. 5, 525-532 (Ru)].

South Pacific Commission, 1979. Regional workshop on biological control (Noumea, New Caledonia, 6-10 August 1979). Report. Regional workshop on biological control (Noumea, New Caledonia, 6-10 August 1979). Report., 28 pp.

South Pacific Commission, 1996. Introduction of giant African snail to various Pacific islands. Ag-Alert, No. 15.

Sproul AN, 1983. Christmas Island snail search. Journal of Agriculture - Western Australia, 24(3):94-96

Sretarugsa P, Ngowsiri U, Kruatrachue M, Sobhon P, Chavadej J, Upatham ES, 1991. Spermiogenesis in Achatina fulica as revealed by electron microscopy. Journal of Medical and Applied Malacology, 3:7-18.

Srivastava PD, Gupta GP, Doharey KL, 1985. Giant African snail and its management. Non-insect pests and predators. Proceedings of the National Symposium on 'Impact of Non-insect Pests and Predators on Food Production and Environment', held from March 12-24, 1984, 72-78, 261-262.

Takeda N, Ohtake SI, 1994. Cilia found in the endocrine bodies of the giant African snail, Achatina fulica. Journal of Biological Chemistry, 1996, 271(20):11750-11755.

Takeuchi K, Koyano S, Numazawa K, 1991. Occurrence of the giant African snail in the Ogasaware (Bonin) Islands, Japan. Micronesica, No. 3, Supplement:109-116.

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Principal Source

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Datasheet under review

Contributors

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16/10/13 Updated by:

Roberto E Vogler, Universidad Nacional de Misiones, Facultad de Ciencias Exactas Químicas y Naturales, Departamento de Biología, Rivadavia 2370, N3300LDX, Posadas, Argentina

Ariel A Beltramino, Universidad Nacional de La Plata, Facultad de Ciencias Naturales y Museo, División Zoología Invertebrados, Paseo del Bosque S/N, B1900FWA, La Plata, Argentina

Distribution Maps

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Distribution map Antigua and Barbuda: Transient: actionable, under eradication
EPPO, 2013Anguilla: Present
EPPO, 2013Netherlands Antilles: Present
EPPO, 2013Argentina: Restricted distribution, introduced, invasive
Vogler et al., 2013; Gutiérrez & Gregoric Núñez, 2012; Gutiérrez Gregoric et al., 2011American Samoa: Widespread, introduced, invasive
Tillier et al., 1993; Cowie, 2000; EPPO, 2013Barbados: Present
EPPO, 2013Barbados: Present
EPPO, 2013Bangladesh: Present, introduced, invasive
Mead, 1979Brunei Darussalam: Present
EPPO, 2013Bolivia: Present only in captivity/cultivation, introduced
Vogler et al., 2013Brazil: Widespread, introduced, invasive
Paiva, 1999; Vogler et al., 2013; EPPO, 2013Brazil
See regional map for distribution within the countryBrazil
See regional map for distribution within the countryBrazil
See regional map for distribution within the countryBrazil
See regional map for distribution within the countryBrazil
See regional map for distribution within the countryBrazil
See regional map for distribution within the countryBrazil
See regional map for distribution within the countryBrazil
See regional map for distribution within the countryBrazil
See regional map for distribution within the countryBrazil
See regional map for distribution within the countryBrazil
See regional map for distribution within the countryCentral African Republic: Present
Mbétid-Bessane, 2006Côte d'Ivoire: Present, introduced, invasive
de Winter, 1989China: Restricted distribution, introduced, invasive
Schotman, 1989; EPPO, 2013China: Restricted distribution, introduced, invasive
Schotman, 1989; EPPO, 2013China
See regional map for distribution within the countryChina
See regional map for distribution within the countryChina
See regional map for distribution within the countryChina
See regional map for distribution within the countryChina
See regional map for distribution within the countryChina
See regional map for distribution within the countryColombia: Present, introduced, invasive
Vogler et al., 2013; EPPO, 2013Colombia: Present, introduced, invasive
Vogler et al., 2013; EPPO, 2013Christmas Island (Indian Ocean): Present, introduced, invasive
Sproul, 1983; Lake & O'Dowd, 1991Dominica: Present
EPPO, 2013Ecuador: Present, introduced, invasive
Vogler et al., 2013; EPPO, 2013Micronesia, Federated states of: Widespread, introduced, invasive
Tillier et al., 1993; Cowie, 2000; EPPO, 2013Guadeloupe: Present, introduced, invasive
Anon., 1989; EPPO, 2013Guam: Widespread, introduced
Tillier et al., 1993; Cowie, 2000; EPPO, 2013Indonesia: Restricted distribution, introduced, invasive
Mead, 1979; EPPO, 2013Indonesia: Restricted distribution, introduced, invasive
Mead, 1979; EPPO, 2013Indonesia
See regional map for distribution within the countryIndonesia
See regional map for distribution within the countryIndonesia
See regional map for distribution within the countryIndonesia
See regional map for distribution within the countryIndonesia
See regional map for distribution within the countryIndia: Widespread, introduced, invasive
Raut & Ghose, 1984; Srivastava et al., 1985; Mead, 1961; EPPO, 2013India
See regional map for distribution within the countryIndia
See regional map for distribution within the countryIndia
See regional map for distribution within the countryIndia
See regional map for distribution within the countryIndia
See regional map for distribution within the countryIndia
See regional map for distribution within the countryIndia
See regional map for distribution within the countryIndia
See regional map for distribution within the countryIndia
See regional map for distribution within the countryIndia
See regional map for distribution within the countryIndia
See regional map for distribution within the countryJapan: Restricted distribution, introduced, invasive
Takeuchi et al., 1991; Tomiyama, 2002; Mead, 1961; EPPO, 2013Japan
See regional map for distribution within the countryKenya: Widespread, introduced, invasive
Mead, 1979Cambodia: Present, introduced, invasive
Mead, 1979Saint Lucia: Restricted distribution
EPPO, 2013; Jn Pierre, 2008Sri Lanka: Restricted distribution, introduced, invasive
Mead, 1961; EPPO, 2013Madagascar: Present
EPPO, 2013Marshall Islands: Present, introduced, invasive
South Pacific Commission, 1996; Cowie, 2000; EPPO, 2013Myanmar: Restricted distribution, introduced, invasive
Schotman, 1989; Mead, 1961; EPPO, 2013Northern Mariana Islands: Widespread, introduced, invasive
Tillier et al., 1993; Cowie, 2000; EPPO, 2013Martinique: Restricted distribution, introduced, invasive
Anon., 1989; EPPO, 2013Mauritius: Widespread, introduced, invasive
Mead, 1961; Facknath et al., 1999; EPPO, 2013Maldives: Widespread, introduced, invasive
Muniappan, 1987; Mead, 1961; EPPO, 2013Malaysia: Restricted distribution
Mead, 1979; EPPO, 2013Malaysia
See regional map for distribution within the countryMalaysia
See regional map for distribution within the countryNew Caledonia: Widespread, introduced, invasive
South Pacific Commission, 1979; Tillier et al., 1993; Cowie, 2000; EPPO, 2013Nigeria: Present
Ashaye et al., 2001Peru: Present, introduced, invasive
Vogler et al., 2013; EPPO, 2013French Polynesia: Widespread, introduced, invasive
Murray et al., 1988; Tillier et al., 1993; Cowie, 2000; EPPO, 2013Papua New Guinea: Widespread, introduced, invasive
Tillier et al., 1993; Mead, 1961; Cowie, 2000; EPPO, 2013Papua New Guinea: Widespread, introduced, invasive
Tillier et al., 1993; Mead, 1961; Cowie, 2000; EPPO, 2013Philippines: Widespread, introduced, invasive
Mead, 1961; EPPO, 2013Philippines: Widespread, introduced, invasive
Mead, 1961; EPPO, 2013Palau: Widespread, introduced, invasive
Tillier et al., 1993; Cowie, 2000; EPPO, 2013Palau: Widespread, introduced, invasive
Tillier et al., 1993; Cowie, 2000; EPPO, 2013Paraguay: Present, introduced, invasive
Vogler et al., 2013Réunion: Widespread, introduced, invasive
Schotman, 1989; Graber et al., 1997; Mead, 1961; EPPO, 2013Seychelles: Widespread, introduced, invasive
Schotman, 1989; Mead, 1961; EPPO, 2013Singapore: Widespread, introduced, invasive
Mead, 1979; Mead, 1961; EPPO, 2013Togo: Present, introduced, invasiveThailand: Restricted distribution
Mead, 1979; EPPO, 2013Trinidad and Tobago: Restricted distribution, introduced
IPPC, 2009; EPPO, 2013Trinidad and Tobago: Restricted distribution, introduced
IPPC, 2009; EPPO, 2013Taiwan: Restricted distribution, introduced, invasive
Mead, 1961; EPPO, 2013Taiwan: Restricted distribution, introduced, invasive
Mead, 1961; EPPO, 2013Tanzania: Widespread, native, not invasive
Emberton et al., 1997US Minor Outlying Islands: Widespread
Tillier et al., 1993USA: Restricted distribution
EPPO, 2013USA: Restricted distribution
EPPO, 2013USA
See regional map for distribution within the countryUSA
See regional map for distribution within the countryUSA
See regional map for distribution within the countryVenezuela: Present, introduced, invasive
Vogler et al., 2013; EPPO, 2013Venezuela: Present, introduced, invasive
Vogler et al., 2013; EPPO, 2013United States Virgin Islands: Present, introduced, invasive
Miller, 1994Vietnam: Restricted distribution, introduced, invasive
Mead, 1979; Mead, 1961; EPPO, 2013Vanuatu: Widespread
Tillier et al., 1993; EPPO, 2013Wallis and Futuna Islands: Widespread, introduced, invasive
Tillier et al., 1993; Cowie, 2000; EPPO, 2013Samoa: Present, introduced, invasive
South Pacific Commission, 1979; Cowie, 2000; EPPO, 2013
  • = Present, no further details
  • = Evidence of pathogen
  • = Widespread
  • = Last reported
  • = Localised
  • = Presence unconfirmed
  • = Confined and subject to quarantine
  • = See regional map for distribution within the country
  • = Occasional or few reports
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Distribution map (asia) Bangladesh: Present, introduced, invasive
Mead, 1979Brunei Darussalam: Present
EPPO, 2013China: Restricted distribution, introduced, invasive
Schotman, 1989; EPPO, 2013Fujian: Present
Li et al., 2006Guangdong: Present
Meng et al., 2007Guizhou: Present
Tang et al., 2008Hainan: Present
Hu et al., 2007Hong Kong: Widespread, introduced
Schotman, 1989; Mead, 1961; EPPO, 2013Yunnan: Present
Chen et al., 2008Christmas Island (Indian Ocean): Present, introduced, invasive
Sproul, 1983; Lake & O'Dowd, 1991Indonesia: Restricted distribution, introduced, invasive
Mead, 1979; EPPO, 2013Irian Jaya: Present, introduced, invasive
Mead, 1979Java: Present, introduced, invasive
Schneider et al., 1998; Mead, 1961Kalimantan: Present
EPPO, 2013India: Widespread, introduced, invasive
Raut & Ghose, 1984; Srivastava et al., 1985; Mead, 1961; EPPO, 2013Andaman and Nicobar Islands: Widespread, introduced, invasive
Shah, 1992Andhra Pradesh: Present
Reddy & Sreedharan, 2006Arunachal Pradesh: Present
Ramakrishna et al., 2005Bihar: Widespread, introduced, invasive
Thakur, 1998Karnataka: Widespread, introduced, invasive
Shree & Kumar, 2002; Basavaraju et al., 2000Madhya Pradesh: Widespread, introduced, invasive
Thakur & Kumari, 1998Odisha: Present, introduced, invasive
Mead, 1961Rajasthan: Present, introduced, invasive
Tehsin & Sharma, 2000Tamil Nadu: Present, introduced, invasive
Padmanaban et al., 2000Uttar Pradesh: Present, introduced, invasive
Saxena & Mahendru, 2000West Bengal: Widespread, introduced, invasive
Raut & Ghara, 1990; Raut, 1991Japan: Restricted distribution, introduced, invasive
Takeuchi et al., 1991; Tomiyama, 2002; Mead, 1961; EPPO, 2013Ryukyu Archipelago: Present, introduced, invasive
Mead, 1961; EPPO, 2013Cambodia: Present, introduced, invasive
Mead, 1979Sri Lanka: Restricted distribution, introduced, invasive
Mead, 1961; EPPO, 2013Myanmar: Restricted distribution, introduced, invasive
Schotman, 1989; Mead, 1961; EPPO, 2013Maldives: Widespread, introduced, invasive
Muniappan, 1987; Mead, 1961; EPPO, 2013Malaysia: Restricted distribution
Mead, 1979; EPPO, 2013Peninsular Malaysia: Present, introduced, invasive
Mead, 1979; Mead, 1961; EPPO, 2013Sarawak: Restricted distribution, introduced, invasive
Mead, 1979; Mead, 1961; EPPO, 2013Papua New Guinea: Widespread, introduced, invasive
Tillier et al., 1993; Mead, 1961; Cowie, 2000; EPPO, 2013Philippines: Widespread, introduced, invasive
Mead, 1961; EPPO, 2013Palau: Widespread, introduced, invasive
Tillier et al., 1993; Cowie, 2000; EPPO, 2013Singapore: Widespread, introduced, invasive
Mead, 1979; Mead, 1961; EPPO, 2013Thailand: Restricted distribution
Mead, 1979; EPPO, 2013Taiwan: Restricted distribution, introduced, invasive
Mead, 1961; EPPO, 2013Vietnam: Restricted distribution, introduced, invasive
Mead, 1979; Mead, 1961; EPPO, 2013
Distribution map (europe)
Distribution map (africa) Central African Republic: Present
Mbétid-Bessane, 2006Côte d'Ivoire: Present, introduced, invasive
de Winter, 1989Kenya: Widespread, introduced, invasive
Mead, 1979Madagascar: Present
EPPO, 2013Mauritius: Widespread, introduced, invasive
Mead, 1961; Facknath et al., 1999; EPPO, 2013Nigeria: Present
Ashaye et al., 2001Réunion: Widespread, introduced, invasive
Schotman, 1989; Graber et al., 1997; Mead, 1961; EPPO, 2013Seychelles: Widespread, introduced, invasive
Schotman, 1989; Mead, 1961; EPPO, 2013Togo: Present, introduced, invasiveTanzania: Widespread, native, not invasive
Emberton et al., 1997
Distribution map (north america) USA: Restricted distribution
EPPO, 2013Florida: Restricted distribution
EPPO, 2013; Mead, 1979; NAPPO, 2011; NAPPO, 2013; NAPPO, 2013Hawaii: Widespread, introduced, invasive
Cowie, 1998; Mead, 1961; EPPO, 2013
Distribution map (central america) Antigua and Barbuda: Transient: actionable, under eradication
EPPO, 2013Anguilla: Present
EPPO, 2013Netherlands Antilles: Present
EPPO, 2013Barbados: Present
EPPO, 2013Colombia: Present, introduced, invasive
Vogler et al., 2013; EPPO, 2013Dominica: Present
EPPO, 2013Guadeloupe: Present, introduced, invasive
Anon., 1989; EPPO, 2013Saint Lucia: Restricted distribution
EPPO, 2013; Jn Pierre, 2008Martinique: Restricted distribution, introduced, invasive
Anon., 1989; EPPO, 2013Trinidad and Tobago: Restricted distribution, introduced
IPPC, 2009; EPPO, 2013USA: Restricted distribution
EPPO, 2013Florida: Restricted distribution
EPPO, 2013; Mead, 1979; NAPPO, 2011; NAPPO, 2013; NAPPO, 2013Venezuela: Present, introduced, invasive
Vogler et al., 2013; EPPO, 2013United States Virgin Islands: Present, introduced, invasive
Miller, 1994
Distribution map (south america) Argentina: Restricted distribution, introduced, invasive
Vogler et al., 2013; Gutiérrez & Gregoric Núñez, 2012; Gutiérrez Gregoric et al., 2011Barbados: Present
EPPO, 2013Bolivia: Present only in captivity/cultivation, introduced
Vogler et al., 2013Brazil: Widespread, introduced, invasive
Paiva, 1999; Vogler et al., 2013; EPPO, 2013Bahia: Present, introduced, invasive
Paiva, 1999Espirito Santo: Present, introduced, invasive
Paiva, 1999Goias: Present
Paiva, 1999Maranhao: Present, introduced, invasiveMinas Gerais: Widespread, introduced, invasive
Paiva, 1999Pernambuco: Present, introduced, invasive
Paiva, 1999Parana: Widespread, introduced, invasive
Paiva, 1999Rio de Janeiro: Widespread, introduced, invasive
Dos Santos et al., 2002; Paiva, 1999Rondonia: Present, introduced, invasive
Paiva, 1999Santa Catarina: Widespread, introduced, invasive
Paiva, 1999Sao Paulo: Widespread, introduced, invasive
Paiva, 1999Colombia: Present, introduced, invasive
Vogler et al., 2013; EPPO, 2013Ecuador: Present, introduced, invasive
Vogler et al., 2013; EPPO, 2013Peru: Present, introduced, invasive
Vogler et al., 2013; EPPO, 2013Paraguay: Present, introduced, invasive
Vogler et al., 2013Trinidad and Tobago: Restricted distribution, introduced
IPPC, 2009; EPPO, 2013Venezuela: Present, introduced, invasive
Vogler et al., 2013; EPPO, 2013
Distribution map (pacific) American Samoa: Widespread, introduced, invasive
Tillier et al., 1993; Cowie, 2000; EPPO, 2013China: Restricted distribution, introduced, invasive
Schotman, 1989; EPPO, 2013Micronesia, Federated states of: Widespread, introduced, invasive
Tillier et al., 1993; Cowie, 2000; EPPO, 2013Guam: Widespread, introduced
Tillier et al., 1993; Cowie, 2000; EPPO, 2013Indonesia: Restricted distribution, introduced, invasive
Mead, 1979; EPPO, 2013Irian Jaya: Present, introduced, invasive
Mead, 1979Kalimantan: Present
EPPO, 2013Marshall Islands: Present, introduced, invasive
South Pacific Commission, 1996; Cowie, 2000; EPPO, 2013Northern Mariana Islands: Widespread, introduced, invasive
Tillier et al., 1993; Cowie, 2000; EPPO, 2013New Caledonia: Widespread, introduced, invasive
South Pacific Commission, 1979; Tillier et al., 1993; Cowie, 2000; EPPO, 2013French Polynesia: Widespread, introduced, invasive
Murray et al., 1988; Tillier et al., 1993; Cowie, 2000; EPPO, 2013Papua New Guinea: Widespread, introduced, invasive
Tillier et al., 1993; Mead, 1961; Cowie, 2000; EPPO, 2013Philippines: Widespread, introduced, invasive
Mead, 1961; EPPO, 2013Palau: Widespread, introduced, invasive
Tillier et al., 1993; Cowie, 2000; EPPO, 2013Taiwan: Restricted distribution, introduced, invasive
Mead, 1961; EPPO, 2013US Minor Outlying Islands: Widespread
Tillier et al., 1993Vanuatu: Widespread
Tillier et al., 1993; EPPO, 2013Wallis and Futuna Islands: Widespread, introduced, invasive
Tillier et al., 1993; Cowie, 2000; EPPO, 2013Samoa: Present, introduced, invasive
South Pacific Commission, 1979; Cowie, 2000; EPPO, 2013