Linepithema humile (Argentine ant)
- Summary of Invasiveness
- Taxonomic Tree
- Notes on Taxonomy and Nomenclature
- Distribution Table
- History of Introduction and Spread
- Risk of Introduction
- Habitat List
- Hosts/Species Affected
- Biology and Ecology
- Notes on Natural Enemies
- Means of Movement and Dispersal
- Pathway Vectors
- Impact Summary
- Economic Impact
- Environmental Impact
- Threatened Species
- Social Impact
- Risk and Impact Factors
- Detection and Inspection
- Similarities to Other Species/Conditions
- Prevention and Control
- Distribution Maps
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PicturesTop of page
IdentityTop of page
Preferred Scientific Name
- Linepithema humile (Mayr, 1868)
Preferred Common Name
- Argentine ant
Other Scientific Names
- Iridomyrmex humilis (Mayr, 1868)
International Common Names
- Spanish: hormiga Argentina; hormiga invasora (Argentina)
- French: fourmi d'Argentine
- Portuguese: formiga Argentina; formiga-Argentina
Local Common Names
- Argentina: hormiga invasora
- Denmark: argentinsk myre
- Germany: Argentinische Ameise
- Sweden: argentinsk myra
- IRIDHU (Iridomyrmex humilis)
Summary of InvasivenessTop of page
L. humile is one of the most invasive and problematic ant species in the world according to the Invasive Species Specialist Group (www.issg.org/database). Native to South America, it has spread worldwide in areas with Mediterranean-type climates usually associated with disturbed habitats as a result of human commercial activities. Its capacity to tolerate a wide range of abiotic conditions, its generalist dietary requirements and its unicolonial organization, which lead to the formation of large supercolonies, allow the species to spread easily and occupy a wide range of areas. In these areas, L. humile has impacted native ant faunas leading to changes in arthropod communities, ant-vertebrate interactions and ant-plant relationships. Its presence has also had economic effects, such as damage to infrastructure, and negative effects on crops and plantations due to its mutualistic interactions with hemipterans, which can affect the growth and production of the host plant.
Taxonomic TreeTop of page
- Domain: Eukaryota
- Kingdom: Metazoa
- Phylum: Arthropoda
- Subphylum: Uniramia
- Class: Insecta
- Order: Hymenoptera
- Family: Formicidae
- Genus: Linepithema
- Species: Linepithema humile
Notes on Taxonomy and NomenclatureTop of page
Prior to Shattuck’s revision in 1992 (Shattuck, 1992), the species was included in the genus Iridomyrmex under the Latin name of Iridomyrmex humilis.
DescriptionTop of page
The workers of L. humile are small (2-3 mm long), brown and monomorphic. Their body surface is free of erect hairs on the dorsum, thorax and head. Their petiole is small and composed of a single segment (Newell and Barber, 1913).
DistributionTop of page
L. humile is native to the Paraná River drainage basin in subtropical South America (between northern Argentina, southern Brazil, Uruguay and Paraguay) (Suarez et al., 2001; Tsutsui and Suarez, 2003; Wild, 2004). In Argentina both native and invasive colonies have been reported.
The distribution map includes records based on specimens of L. humile from collections in the Natural History Museum of Los Angeles County (LACM), California, USA; Museo Argentina de Ciencias Naturales (MACN), Buenos Aires, Argentina; Museu de Zoologia da Universidade de São Paulo (MZSP), São Paulo, Brazil; Florida Department of Agriculture and Consumer Services, Tallahassee, Florida; Entomology Museum, Oregon Department of Agriculture; and The Field Museum, Chicago, Illinois, USA.
Distribution TableTop of page
The distribution in this summary table is based on all the information available. When several references are cited, they may give conflicting information on the status. Further details may be available for individual references in the Distribution Table Details section which can be selected by going to Generate Report.
|Continent/Country/Region||Distribution||Last Reported||Origin||First Reported||Invasive||Reference||Notes|
|Japan||Present||Introduced||Invasive||Kameyama, 2001; ISSG, 2011|
|-Honshu||Widespread||Introduced||Invasive||Okaue et al., 2007|
|United Arab Emirates||Present||Introduced||Invasive||Collingwood et al., 1997; ISSG, 2011|
|Namibia||Present||Introduced||Espadaler et al., 2004|
|South Africa||Present||Introduced||1901||Invasive||Bond and Slingsby, 1984; ISSG, 2011|
|-Canary Islands||Present||Introduced||1965||Invasive||ISSG, 2011|
|Bermuda||Present||Introduced||Invasive||Haskins and Haskins, 1988; ISSG, 2011|
|Mexico||Present||Introduced||Invasive||Natural History Museum of Los Angeles County; ISSG, 2011|
|-Alabama||Present||Introduced||Invasive||Newell and Barber, 1913|
|-Arizona||Present||Introduced||Invasive||Suarez et al., 2001; ISSG, 2011|
|-Arkansas||Present||Introduced||Invasive||Barber, 1916; ISSG, 2011|
|-California||Present||Introduced||1907||Invasive||Woodworth, 1908; ISSG, 2011|
|-Florida||Present||Introduced||Invasive||Florida Department of Agriculture and Consumer Services, Tallahassee, Florida; ISSG, 2011|
|-Georgia||Present||Introduced||Invasive||Barber, 1916; ISSG, 2011|
|-Hawaii||Present||Introduced||1940||Invasive||Wilson and Taylor, 1967|
|-Louisiana||Present||Introduced||1891||Invasive||Foster, 1908; ISSG, 2011|
|-Minnesota||Present||Introduced||Forshcler and Evans, 1994|
|-Mississippi||Present||Introduced||Invasive||Harned and Smith, 1922; ISSG, 2011|
|-Missouri||Present||Introduced||Invasive||Center for Urban Structural Entomology, 2008|
|-Nevada||Present||Introduced||Invasive||Natural History Museum of Los Angeles County; ISSG, 2011|
|-North Carolina||Present||Introduced||Invasive||Smith, 1936; ISSG, 2011|
|-Oregon||Present||Introduced||Invasive||Entomology Museum, Oregon Department of Agriculture; ISSG, 2011|
|-South Carolina||Present||Introduced||Invasive||Barber, 1916; ISSG, 2011|
|-Tennessee||Present||Introduced||Invasive||Barber, 1916; ISSG, 2011|
|-Texas||Present||Introduced||Invasive||Suarez et al., 2001; ISSG, 2011|
|-Washington||Present||Introduced||Invasive||Suarez et al., 2001; ISSG, 2011|
Central America and Caribbean
|El Salvador||Present||Introduced||Invasive||Natural History Museum of Los Angeles County|
|Argentina||Present||Native||Not invasive||Museo Argentina de Ciencias Naturales, Buenos Aires, Argentina; ISSG, 2011|
|Brazil||Present||Native||Not invasive||ISSG, 2011|
|-Goias||Present||Introduced||Invasive||Museu de Zoologia da Universidade de São Paulo|
|-Minas Gerais||Present||Introduced||Invasive||Museu de Zoologia da Universidade de São Paulo|
|-Rio de Janeiro||Present||Introduced||Invasive||Museu de Zoologia da Universidade de São Paulo|
|Chile||Present||Introduced||Invasive||Natural History Museum of Los Angeles County; ISSG, 2011|
|-Easter Island||Present||Introduced||Invasive||Natural History Museum of Los Angeles County; ISSG, 2011|
|Colombia||Present||Introduced||Invasive||Natural History Museum of Los Angeles County|
|Ecuador||Present||Introduced||Invasive||Natural History Museum of Los Angeles County|
|Paraguay||Present||Native||Not invasive||ISSG, 2011|
|Peru||Present||Introduced||Invasive||Dale, 1974; ISSG, 2011|
|Uruguay||Present||Introduced||Invasive||Newell and Barber, 1913; ISSG, 2011|
|Belgium||Present||Introduced||Invasive||Muséum d'Histoire Naturelle, Geneva, Switzerland|
|France||Present||Introduced||Invasive||Benois, 1973; ISSG, 2011|
|Italy||Present||Collingwood et al., 1997|
|Netherlands||Present||Introduced||Boer and Vierbergen, 2008|
|Norway||Present only under cover/indoors||Introduced||Not invasive||Gómez et al., 2005|
|Portugal||Present||Introduced||1900||Invasive||Silva Dias, 1955; ISSG, 2011|
|Spain||Present||Introduced||Invasive||Espadaler and Gómez, 2003; ISSG, 2011|
|UK||Present||Introduced||Invasive||Cornwell, 1978; ISSG, 2011|
|Australia||Restricted distribution||Introduced||1939||Invasive||ISSG, 2011|
|-New South Wales||Present, few occurrences||Introduced||Invasive||The Field Museum, Chicago, Illinois|
|-Tasmania||Present||Introduced||Invasive||Harris, 2002; ISSG, 2011|
|-Western Australia||Present||Introduced||Invasive||Schagen et al., 1994|
|New Zealand||Restricted distribution||Introduced||March 2000||Invasive||Harris, 2002; ISSG, 2011|
History of Introduction and SpreadTop of page
In Hawaii, L. humile has frequently been intercepted in goods shipped from California. In Honolulu its establishment was first reported in 1940 (Zimmerman, 1941). It reached Maui by 1950 (Wilson and Taylor, 1967) and Haleakala National Park by 1967 (Huddleston and Fluker, 1968). The date when L. humile reached the island of Hawaii remains unknown.
According to Newell and Barber (1913), L. humile is supposed to have been introduced into New Orleans by means of the coffee ships passing back and forth between that city and Brazilian ports, as this was the only way of communication between New Orleans and the areas native to the species, and because the ants were first noticed near the areas where these ships unloaded their cargoes.
The date at which L. humile reached the Iberian Peninsula also remains unknown, but the first reported observations of the species in this area were in Oporto (the western part of the Peninsula) and date back to 1894 (Martins, 1907). This means that the first introduction of the ant would probably have been much earlier, perhaps in the 1800s, as a result of the intense commercial shipping between the Iberian Peninsula and the Río de la Plata region, the native area of this species (Tsutsui et al., 2001).
IntroductionsTop of page
|Introduced to||Introduced from||Year||Reason||Introduced by||Established in wild through||References||Notes|
|Natural reproduction||Continuous restocking|
|New Zealand||1990||Yes||No||Green (1990)||Accidental introduction|
Risk of IntroductionTop of page
L. humile has two main dispersion processes: diffusion dispersal through budding and long-distance jump dispersion through human-mediated transport (Suarez et al., 2001). The former is the mechanism used by the ant to extend the invasion to contiguous habitats and is characterized by a low rate of spread (Suarez et al., 2001). The latter is the process responsible for its worldwide dispersion outside its natural range and may involve anything from a few to a thousand kilometres. Jump-dispersal linked to human-mediated transport is nowadays the main factor behind the expansion of the invasion to new areas. It means that, in global terms, the risk of the introduction of the species into new areas is extremely high, since preventing its accidental transport is almost impossible.
HabitatTop of page
L. humile appears to be more successful in subtropical and Mediterranean climates than in extremely cold, arid or tropical climates (Hölldobler and Wilson, 1990; Passera, 1994). Where it has been introduced, this species is mainly found in close association with humans, with a preference for anthropogenically disturbed areas, but it also has the capacity to invade natural ecosystems around the world (Holway, 1995; Human and Gordon, 1996; Suarez et al., 2001; Gomez et al., 2003; Carpintero et al., 2005).
Habitat ListTop of page
|Buildings||Principal habitat||Harmful (pest or invasive)|
|Cultivated / agricultural land||Present, no further details||Harmful (pest or invasive)|
|Disturbed areas||Principal habitat||Harmful (pest or invasive)|
|Protected agriculture (e.g. glasshouse production)||Present, no further details||Harmful (pest or invasive)|
|Rail / roadsides||Present, no further details||Harmful (pest or invasive)|
|Urban / peri-urban areas||Principal habitat||Harmful (pest or invasive)|
|Natural forests||Present, no further details||Harmful (pest or invasive)|
|Scrub / shrublands||Present, no further details||Harmful (pest or invasive)|
Hosts/Species AffectedTop of page
Nearly all of the effects of L. humile on crops or plantations are due to its mutualistic interactions with hemipterans, which are protected from their natural enemies by the ant, thereby producing a large increase in their populations and so affecting the host plant (Newell and Barber, 1913; Ness and Bronstein, 2004).
Biology and EcologyTop of page
In native populations of L. humile, workers from different nests compete aggressively for resources. The introduced populations of the species are characterized by a lack of aggressive behaviour between workers from different nests, leading to the formation of unicolonies. These unicolonies can consist of cooperative interconnected nests ranging over thousands of kilometres, as is the case of the so-called European supercolonies (Giraud et al., 2002), with high densities of workers leading to a numerical dominance over native ant species (Holway et al., 2002a). This lack of a capacity of workers in introduced populations to distinguish between workers from other nests seems to be related to a lower level of genetic diversity compared with the native populations. This has resulted in the elimination of the mechanisms required for workers to recognize workers from a different nest (Tsutsui et al., 2000).
Once virgin queens are fertilized, workers cut off their wings. This allows ovarian maturation and consequent production of eggs (Passera and Aron, 1993). Fertilized eggs (also known as diploid eggs) will develop into females (workers and queens), and the non-fertilized ones (also known as haploid eggs) will develop into males.
Queens lay eggs during most of the year except in winter, due to a physiological rest period. There are two periods during the remainder of the year when egg laying is at its maximum: one in spring and the other, to a lesser extent, in autumn (Benois, 1973). In June and July there is a strong demographic increase in males and workers, respectively, and virgin queens appear in the nest from June to August (Benois, 1973).
L. humile is an omnivorous species. Its diet is basically composed of liquid food (Markin, 1970b; Abril et al., 2007), but it also contains a small percentage of solid food, mainly insects (Abril et al., 2007). The collection of liquid food is related to the feeding of males and workers which feed mainly on carbohydrates, the principal nutrient of the liquid food collected. This, for the most part, consists of honeydew or nectar. The collection of solid food is linked to the feeding of larvae and queens. Due to its high protein content, this is the main nutrient consumed by these individuals for their development and egg-laying, respectively (Markin, 1970a; Abril et al., 2007).
ClimateTop of page
|C - Temperate/Mesothermal climate||Preferred||Average temp. of coldest month > 0°C and < 18°C, mean warmest month > 10°C|
Notes on Natural EnemiesTop of page
There are no recorded cases of natural enemies of L. humile in its introduced range (Orr et al., 2001).
Means of Movement and DispersalTop of page
Natural Dispersal (Non-Biotic)
The dispersion of L. humile worldwide is produced via accidental transportation by means of human commercial activities. This means of dispersion is commonly known as long-distance jump dispersion (Suarez et al., 2001) and allows dispersion from a few to thousands of kilometres. There is also evidence of accidental dispersion of small fragments of colonies by means of road vehicles (Hee et al., 2000).
There are no recorded instances of the intentional introduction of L. humile.
Pathway VectorsTop of page
Impact SummaryTop of page
Economic ImpactTop of page
L. humile produces negative economic effects when invading crops and plantations because of its mutualistic interactions with hemipterans, which can affect the growth and production of the host plant (Buckley, 1987; Ness and Bronstein, 2004).
There is also evidence of its capacity to transfer the pathogenic fungus Phytophthora citricola to Persea americana plantations, with a resulting serious economic impact (El-Hamalawi and Menge, 1996).
Environmental ImpactTop of page
Impact on Biodiversity
Threatened SpeciesTop of page
|Threatened Species||Conservation Status||Where Threatened||Mechanism||References||Notes|
|Charadrius alexandrinus nivosus||USA ESA listing as threatened species USA ESA listing as threatened species||California; Oregon; Washington||Predation||US Fish and Wildlife Service, 2007|
|Desmocerus californicus dimorphus (valley elderberry longhorn beetle)||USA ESA listing as threatened species USA ESA listing as threatened species||California||Predation||US Fish and Wildlife Service, 2012|
|Perognathus longimembris pacificus (Pacific pocket mouse)||USA ESA listing as endangered species USA ESA listing as endangered species||California||Ecosystem change / habitat alteration; Predation||US Fish and Wildlife Service, 2010b|
|Polioptila californica californica (coastal California gnatcatcher)||USA ESA listing as threatened species USA ESA listing as threatened species||California||Predation||US Fish and Wildlife Service, 2010a|
|Schiedea haleakalensis (Haleakala schiedea)||CR (IUCN red list: Critically endangered) CR (IUCN red list: Critically endangered); USA ESA listing as endangered species USA ESA listing as endangered species||Hawaii||Ecosystem change / habitat alteration||US Fish and Wildlife Service, 2011|
|Silene lanceolata (Kauai catchfly)||USA ESA listing as endangered species USA ESA listing as endangered species||Hawaii||Herbivory/grazing/browsing||US Fish and Wildlife Service, 2010c|
|Sterna antillarum browni (California least tern)||USA ESA listing as endangered species USA ESA listing as endangered species||California||Predation||US Fish and Wildlife Service, 2006|
Social ImpactTop of page
There is no relevant information on any negative social impact of this ant species.
Risk and Impact FactorsTop of page Invasiveness
- Proved invasive outside its native range
- Has a broad native range
- Is a habitat generalist
- Tolerates, or benefits from, cultivation, browsing pressure, mutilation, fire etc
- Pioneering in disturbed areas
- Capable of securing and ingesting a wide range of food
- Highly mobile locally
- Benefits from human association (i.e. it is a human commensal)
- Has high reproductive potential
- Altered trophic level
- Damaged ecosystem services
- Ecosystem change/ habitat alteration
- Infrastructure damage
- Negatively impacts agriculture
- Negatively impacts forestry
- Reduced native biodiversity
- Threat to/ loss of endangered species
- Threat to/ loss of native species
- Competition - monopolizing resources
- Highly likely to be transported internationally accidentally
- Difficult to identify/detect as a commodity contaminant
- Difficult/costly to control
Detection and InspectionTop of page
The easiest way to detect L. humile on the crop is by searching for foraging workers on the trunk or the stem of the plant, because they associate closely with honeydew-producing hemipterans while searching out honeydew.
Similarities to Other Species/ConditionsTop of page
L. humile is easily distinguishable from other species of ants by its extremely fast movements and high number of recruitment workers. The workers, when crushed between the fingers, give off no perceptible odour, and this characteristic allows the species to be distinguished from other similar ants like Iridomirmex analis, as well as from other similar ant species from the genus Tapinoma.
Prevention and ControlTop of page
Cultural control and sanitary measures
Baits can be used to control L. humile invasions by using workers as a vehicle for transmitting toxins through tropholaxis to larvae and queens. Baits consist of an active toxic ingredient and an attractant in the form of a food resource (Rust, 2001). This method has resulted in significant reductions in L. humile numbers in infected areas (Forschler, 1997; Vega and Rust, 2003). One example of this is the reduction of the invasion in the Haleakala National Park, Hawaii, by using granular baits (Krushelnycky et al., 2004). However, this method is only successful temporarily, as suspension of the treatment leads to regeneration of the ant population and renewed expansion of the invasion (Krushelnycky et al., 2004).
BibliographyTop of page
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ReferencesTop of page
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ContributorsTop of page
29/04/08 Original text by:
Crisanto Gómez, Universitat de Girona, Dept Ciències Ambientals, Facultat de Ciències, Campus de Montilivi7071- Girona, Spain
Silvia Abril, Universitat de Girona, Dept Ciències Ambientals, Facultat de Ciències7071- Girona, Spain
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