Linepithema humile

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Linepithema humile
Scientific classification
Kingdom: Animalia
Phylum: Arthropoda
Class: Insecta
Order: Hymenoptera
Family: Formicidae
Subfamily: Dolichoderinae
Tribe: Leptomyrmecini
Genus: Linepithema
Species: L. humile
Binomial name
Linepithema humile
(Mayr, 1868)

Linepithema humile casent0005323 profile 1.jpg

Linepithema humile casent0005323 dorsal 1.jpg

Specimen labels


Common Name
Argentine Ant
Language: English
Language: Japanese

This is the notoriously pestiferous Argentine ant, one of the 100 worst invasive species in the world (IUCN/SSC Invasive Species Specialist Group). Its colonies are large and polygynous, sometimes comprising hundreds of queens and many thousands of workers. New nests are founded by migration or budding. The Argentine ant commonly infests houses and other premises, contaminating and spoiling stored food and other products. It tends homopterous agricultural and horticultural insect pests, and severely damages and depletes populations of native ant species in infested areas. L. humile is native to parts of South America (Brazil and Argentine) and has been distributed throughout much of the world by human commerce.

At a Glance • Highly invasive  • Supercolonies  • Polygynous  

Photo Gallery

  • The odd insect at right is a male ant. He looks nothing like his sisters, bearing wings and large eyes to better serve his role of dispersal and mating. Davis, California, USA. Photo by Alex Wild.


Worker Eyes large (OI > 30); antennal scapes long (SI > 105); pronotum and first two gastric tergites lacking erect setae; mesopleura and metapleura densely pubescent.

Workers of the sister species Linepithema oblongum, from the high Andes of Bolivia and northern Argentina, normally have at least some members of each series with dilute pubescence on gastric tergites 2–4. These ants also have, on average, smaller eyes (OI 28–38) and longer antennal scapes (SI 120–139) than L. humile. Workers of Linepithema anathema, a rarely-collected Brazilian species, have a more produced propodeum, a narrow head (CI < 86), and usually bear short standing setae on gastric tergites 1–2. Workers of other Humile-group species have shorter antennal scapes and often bear erect setae on the pronotum and basal gastric tergites. Males of related species are much smaller than L. humile and lack the greatly swollen mesosoma.

Male Forewing with single submarginal cell; mesosoma robust (MML > 1.3), mesoscutum greatly enlarged and overhanging pronotum; wings short relative to mesosomal length (WI < 21).

Keys including this Species


Native to the Paraná river drainage of Brazil, Paraguay, Argentina, and Uruguay. Introduced worldwide.

This introduced species occurs sporadically throughout Florida, in places forming massive populations. It occurs in both moist and dry open habitats, usually in heavily disturbed sites. Pest status: can become a nuisance by sheer numbers, trailing long distances to outdoor eating areas and into buildings. First published Florida record: Wheeler 1932; earlier specimens: 1914. (Deyrup, Davis & Cover, 2000.)

Linepithema humile was previously reported in Colombia (Sanabria & Chacón de Ulloa 2009; Wild 2004, 2007). However, we found that the specimens identified as L. humile in Sanabria & Chacón de Ulloa (2009), and many other “Linepithema humile” specimens in Colombian entomological collections, were misidentified. In most cases these specimens were Linepithema piliferum or Linepithema neotropicum, and in no cases were L. humile. In order to confirm the occurrence of L. humile in Colombia (Wild 2007), we studied the L. humile specimens reported in Wild (2007) and deposited in WPMC. We confirm that the specimens are L. humile, and this remains the only known collection of the species in Colombia. These ants were collected in the Colombian coffee zone (Armenia, Quindio) in 1973, but despite subsequent intensive sampling done in that area by the program Paisajes Rurales (Instituto Alexander von Humboldt), there are no more L. humile records from Armenia or elsewhere. Thus it is possible that this was an introduction that did not persist and the species no longer occurs in Colombia. (Escarraga & Guerrero, 2016)

Latitudinal Distribution Pattern

Latitudinal Range: 25.68015° to -37.533333°.

Tropical South

Distribution based on Regional Taxon Lists

Afrotropical Region: Lesotho, Mozambique, Namibia, Saint Helena, United Arab Emirates.
Australasian Region: Australia, New Zealand, Norfolk Island.
Indo-Australian Region: Hawaii, Vanuatu.
Nearctic Region: United States.
Neotropical Region: Argentina (type locality), Bermuda, Brazil, Ecuador, French Guiana, Mexico, Paraguay, Uruguay.
Palaearctic Region: Balearic Islands, Belgium, Bulgaria, Canary Islands, Channel Islands, Democratic Peoples Republic of Korea, France, Germany, Gibraltar, Greece, Iberian Peninsula, Iran, Italy, Japan, Malta, Monaco, Montenegro, Morocco, Poland, Portugal, Spain, Türkiye, United Kingdom of Great Britain and Northern Ireland.

Distribution based on AntMaps


Distribution based on AntWeb specimens

Check data from AntWeb

Countries Occupied

Number of countries occupied by this species based on AntWiki Regional Taxon Lists. In general, fewer countries occupied indicates a narrower range, while more countries indicates a more widespread species.

Estimated Abundance

Relative abundance based on number of AntMaps records per species (this species within the purple bar). Fewer records (to the left) indicates a less abundant/encountered species while more records (to the right) indicates more abundant/encountered species.


There is an Antwiki webpage with a list of some recent studies of the Argentine ant.

Wild (2007) - This important pest species has a literature too extensive to be covered in depth here. An early general review of the biology of this ant is given by Newell and Barber (1913). The spread of Argentine ants around the world is documented by Roura-Pascual et al. (2004), Wild (2004), Giraud et al. (2002), and Suarez et al. (2001). Ecological impacts of Argentine ant invasion have been detailed in numerous studies, including Suarez and Case (2003), Touyama et al. (2003), Christian (2001), and Human and Gordon (1997). Colony structure has also received considerable attention, and relevant papers include Holway and Suarez (2004), Tsutsui and Case (2001), Reuter et al. (2001), and Kreiger and Keller (2000). A series of studies by Cavill and colleagues (Cavill and Houghton 1973, Cavill and Houghton 1974, Cavill et al. 1980) describe some of the glandular and cuticular chemistry of L. humile. Chemical and biological control options are reviewed by Harris (2002).

Of the L. humile material examined, more than 90% of native range records are within 10 kilometers of a major river in the Paraná drainage. Contrary to some reports (Buczkowski et al. 2004), L. humile can reach high densities in urban areas in Argentina and Paraguay (Wild 2004) as well as in less disturbed habitats (Heller 2004). Where nest information was recorded in the native range, 24 nests are from soil, five from under covering objects such as stones or garbage, one from an old termite mound, and one from under bark. This species is polygynous and polydomous, and many nests are recorded as having numerous dealate queens. In contrast to introduced populations, alate queens are not uncommon in nests in Argentina (Wild 2004). One observation in Victoria, Argentina, notes a live lycaenid larva in the brood nest (Wild, pers. obs.).

This ant forms what has been called supercolonies: a large number of nests spread over large areas where the individuals from one nest can be brought to any other nest and accepted as nestmates (Corin et al., 2007). Linepithema humile do not have nuptial flights (Passera and Keller 1990). Queens mate with related males in their natal nests (Markin 1970).

Inou et al. (2015) suggested the results of a genetic study of four supercolonies in Kobe Japan showed that these colonies replaced their queens during the reproductive season. Genetic differentiation among workers varied significantly in comparing May samples to September samples. They feel this provides evidence for queen execution, which has been reported in two introduced populations (USA, Markin 1970 and France Keller et al. 1989).

Bertelsmeier et al. (2015a, b) examined elements of interspecific aggression, and food resource discovery and dominance, between this species and several other highly invasive ants. In laboratory assays Linepithema humile was highly aggressive when confronted with workers of other invasive ants. Of the group of four species that were found to be aggressive, L. humile was found to be fairly adept at finding and recruiting to food in a laboratory arena experiment.

Regional Notes

Brazil - DaRocha et al. (2015) studied the diversity of ants found in bromeliads of a single large tree of Erythrina, a common cocoa shade tree, at an agricultural research center in Ilhéus, Brazil. Forty-seven species of ants were found in 36 of 52 the bromeliads examined. Bromeliads with suspended soil and those that were larger had higher ant diversity. Linepithema humile was found in a single bromeliad and was associated with the suspended soil and litter of the plant.

Canary Islands - Espadaler (2007): The Argentine ant is known from all the Canary Islands (Espadaler & Bernal, 2003). At El Hierro it occupies habitats from next to sea level to one thousand meters, in pine forests. Confronted with the two populations known to exist in North Mediterranean Europe (Giraud et al., 2002), the Argentine ants from El Hierro showed aggressiveness towards the “Catalan” population and reacted peacefully towards the “Main” population from mainland Europe. Aggression tests (one to one worker; five replicates) were run with two samples from El Hierro (La Frontera; Mirador de las Playas). I conclude that both samples from El Hierro belong to the genotypic profile of the “Main” population, the more abundant in Western Mediterranean Europe.

Europe - Collingwood (1979): This species was introduced into Europe from South America and has become an established and notorious pest in the Mediterranean area, developing very populous multi-queened colonies along the coast. It is sometimes brought into North Europe with plant materials and occasionally colonises heated premises. It does not appear to be able to establish outside in northern latitudes but is present and said to be increasing in the Channel Islands.

Greece Borowiec and Salata (2022) - Noted only from tourist resorts. Ants were collected on ground in botanical garden, grassland and small garden in urban area.

New Zealand: Corin, Abbott et al. (2007) found that the introduced population of Linepithema humile in New Zealand effectively forms a single unicolonial population, a supercolony, which is likely the result of colonization from a single source population from Australia (Corin, Lester et al., 2007).

Flight Period

Jan Feb Mar Apr May Jun Jul Aug Sep Oct Nov Dec


Association with Other Organisms

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  • This species is a host for the phorid fly Apocephalus silvestrii (a parasitoid) (Quevillon, 2018) (encounter mode primary; direct transmission; transmission outside nest).
  • This species is a host for the phorid fly Ceratoconus setipennis (a parasitoid) (Quevillon, 2018) (encounter mode primary; direct transmission; transmission outside nest).
  • This species is a host for the phorid fly Pseudacteon pusillus (a parasitoid) (Quevillon, 2018) (encounter mode primary; direct transmission; transmission outside nest).
  • This species is a prey for the syrphid fly Mixogaster lanei (a predator) (Quevillon, 2018).


  • This species is a mutualist for the aphid Aphis coreopsidis (a trophobiont) (Altfeld and Stiling, 2009; Saddiqui et al., 2019).
  • This species is a mutualist for the aphid Aphis gossypii (a trophobiont) (Powell and Silverman, 2010; Tena et al., 2013; LeVan and Holway, 2015; Saddiqui et al., 2019).
  • This species is a mutualist for the aphid Aphis nerii (a trophobiont) (Bristow, 1991; Pringle et al., 2014; Saddiqui et al., 2019).
  • This species is a mutualist for the aphid Aphis spiraecola (a trophobiont) (Tena et al., 2013; Saddiqui et al., 2019).
  • This species is a mutualist for the aphid Chaitophorus populicola (a trophobiont) (Mondor and Addicott, 2007; Saddiqui et al., 2019).
  • This species is a mutualist for the aphid Myzus persicae (a trophobiont) (Powell and Silverman, 2010; Saddiqui et al., 2019).


  • This species is a host for the diapriid wasp Basalys sp. (a parasite) in Argentina (Loiacono, 2013; Gonzalez et al., 2016).
  • This species is a host for the diapriid wasp Trichopria sp. (a parasite) in Argentina (Loiacono, 2013; Gonzalez et al., 2016).
  • This species is a host for the aphelinid wasp Aphytis melinus (a parasite) (Universal Chalcidoidea Database) (associate).
  • This species is a host for the aphelinid wasp Cales noacki (a parasite) (Universal Chalcidoidea Database) (associate).
  • This species is a host for the aphelinid wasp Eretmocerus sp. (a parasite) (Universal Chalcidoidea Database) (associate).
  • This species is a host for the encyrtid wasp Comperiella bifasciata (a parasite) (Universal Chalcidoidea Database) (associate).
  • This species is a host for the encyrtid wasp Leptomastix dactylopii (a parasite) (Universal Chalcidoidea Database) (associate).
  • This species is a host for the encyrtid wasp Metaphycus anneckei (a parasite) (Universal Chalcidoidea Database) (associate).
  • This species is a host for the encyrtid wasp Metaphycus hageni (a parasite) (Universal Chalcidoidea Database) (associate).
  • This species is a host for the encyrtid wasp Metaphycus lounsburyi (a parasite) (Universal Chalcidoidea Database) (associate).
  • This species is a associate (details unknown) for the encyrtid wasp Ananusia longiscapus (a associate (details unknown)) (Quevillon, 2018) (as Iridomyrmex domestica).
  • This species is a associate (details unknown) for the encyrtid wasp Comperiella bifasciata (a associate (details unknown)) (Quevillon, 2018).


  • This ant has been observed tending larvae of Lampides boeticus (Obregon et al. 2015).


  • Pekár et al. (2018) - In the Iberian Penisula, this ant is preyed upon by a spider species in the genus Zodarion sp. (Araneae: Zodariidae). All members of this genus are specialized ant predators that exclusively prey on ants.


  • This species is a host for the nematode Diploscapter lycostoma (a parasite) (Markin & McCoy, 1968).


  • This species is a host for the virus Aparavirus: Kashmir bee virus (a parasite) (Quevillon, 2018) (encounter mode primary; direct transmission; transmission within nest).
  • This species is a host for the virus Iflavirus: Deformed wing virus (a parasite) (Quevillon, 2018) (encounter mode primary; direct transmission; transmission within nest).
  • This species is a host for the virus Linepithema humile virus-1 (a parasite) (Quevillon, 2018) (encounter mode primary; direct transmission; transmission within nest).
  • This species is a host for the virus Triatovirus: Black queen cell virus (a parasite) (Quevillon, 2018) (encounter mode primary; direct transmission; transmission within nest).


Linepithema humile has had their entire genome sequenced.

Palomeque et al. (2015) found class II mariner elements, a form of transposable elements, in the genome of this ant.

Life History Traits

  • Queen number: polygynous (Bartels, 1983; Newell & Barber, 1913; Frumhoff & Ward, 1992)
  • Colony type: supercolony
  • Foraging behaviour: mass recruiter


Queens differ in their Cuticular Hydrocarbons according to ovarian activity. Whereas the cuticular profile of non-laying queens is similar to that of sterile workers, it gradually changes both qualitatively and quantitatively once queens start to lay eggs. These changes are independent of mating status, since virgin egg-laying queens show a CH profile similar to that of mated egg-laying queens (de Biseau et al. 2004).

Images from AntWeb


Linepithema humile casent0006019 head 1.jpgLinepithema humile casent0006019 profile 1.jpgLinepithema humile casent0006019 dorsal 1.jpgLinepithema humile casent0006019 label 1.jpg
Worker. Specimen code casent0006019. Photographer April Nobile, uploaded by California Academy of Sciences. Owned by CAS, San Francisco, CA, USA.
Linepithema humile casent0006020 head 1.jpgLinepithema humile casent0006020 profile 1.jpgLinepithema humile casent0006020 dorsal 1.jpgLinepithema humile casent0006020 label 1.jpg
Worker. Specimen code casent0006020. Photographer April Nobile, uploaded by California Academy of Sciences. Owned by CAS, San Francisco, CA, USA.
Linepithema humile casent0104147 head 1.jpgLinepithema humile casent0104147 profile 1.jpgLinepithema humile casent0104147 dorsal 1.jpgLinepithema humile casent0104147 label 1.jpg
Worker. Specimen code casent0104147. Photographer April Nobile, uploaded by California Academy of Sciences. Owned by ABS, Lake Placid, FL, USA.
Linepithema humile casent0106983 head 1.jpgLinepithema humile casent0106983 profile 1.jpgLinepithema humile casent0106983 dorsal 1.jpgLinepithema humile casent0106983 label 1.jpg
Worker. Specimen code casent0106983. Photographer Alexander Wild, uploaded by California Academy of Sciences. Owned by ALWC, Alex L. Wild Collection.


Linepithema humile casent0104070 head 1.jpgLinepithema humile casent0104070 profile 1.jpgLinepithema humile casent0104070 dorsal 1.jpgLinepithema humile casent0104070 label 1.jpg
Queen (alate/dealate). Specimen code casent0104070. Photographer April Nobile, uploaded by California Academy of Sciences. Owned by ABS, Lake Placid, FL, USA.


The following information is derived from Barry Bolton's Online Catalogue of the Ants of the World.

  • humile. Hypoclinea humilis Mayr, 1868b: 164 (w.) ARGENTINA. Forel, 1908c: 395 (m.); Newell, 1908: 28 (q.); Wheeler, G.C. & Wheeler, J. 1951: 186 (l.); Crozier, 1969: 250 (k.). Combination in H. (Iridomyrmex): Mayr, 1870b: 959; in Iridomyrmex: Emery, 1888d: 386; in Linepithema: Shattuck, 1992a: 16. Senior synonym of arrogans, riograndensis: Wild, 2004: 1207. See also: Gallardo, 1916a: 97; Bernard, 1967: 251; Collingwood, 1979: 33; Smith, D.R. 1979: 1418; Ward, 1987: 1; Wheeler, G.C. & Wheeler, J. 1990a: 465; Shattuck, 1994: 123; Wild, 2007a: 61; Solis, Fox, Rossi & Bueno, 2010: 19.
  • arrogans. Iridomyrmex humilis var. arrogans Chopard, 1921: 237 (footnote),:241, figs. 1-31 (w.q.m.l.) FRANCE. [Also described as new by Santschi, 1929d: 306 (in key).] Combination in Linepithema: Shattuck, 1992a: 16. Junior synonym of humile: Bernard, 1967: 251. Revived from synonymy as subspecies of humile: Shattuck, 1992a: 16. Junior synonym of humile: Wild, 2004: 1207.
  • riograndensis. Iridomyrmex riograndensis Borgmeier, 1928b: 64 (w.) BRAZIL. Combination in Linepithema: Shattuck, 1992a: 16. Junior synonym of humile: Wild, 2004: 1207.

Type Material

Taxonomic Notes

The taxonomy and distribution of L. humile was reviewed in depth by Wild (2004). The following description is from Wild (2007):



Holotype: HL 0.74, HW 0.66, MFC 0.16, SL 0.76, FL 0.65, LHT 0.68, PW 0.45, ES 2.93, SI 115, CI 89, CDI 24, OI 40.

Worker: (n = 81) HL 0.62–0.78, HW 0.53–0.72, MFC 0.14–0.18, SL 0.62–0.80, FL 0.52–0.68, LHT 0.57–0.76, PW 0.35–0.47, ES 1.98–3.82, SI 108–126, CI 84–93, CDI 23–28, OI 32–49.

Head in full face view longer than broad (CI 84–93), narrowed anteriorly and reaching its widest point just posterior to compound eyes. Lateral margins broadly convex, grading smoothly into posterior margin. Posterior margin straight in smaller workers to weakly concave in larger workers. Compound eyes large (OI 32–49), comprising 82–110 ommatidia (normally around 100). Antennal scapes long (SI 108–126), as long or slightly longer than HL and easily surpassing posterior margin of the head in full face view. Frontal carinae narrowly to moderately spaced (CDI 23–28). Maxillary palps relatively short, shorter than ½ HL, ultimate segment (segment six) noticeably shorter than segment 2.

Pronotum and mesonotum forming a continuous convexity in lateral view, mesonotal dorsum nearly straight, not angular or strongly impressed, although sometimes with a slight impression in anterior portion. Metanotal groove moderately impressed. Propodeum in lateral view inclined anteriad. In lateral view, dorsal propodeal face meeting declivity in a distinct though obtuse angle, from which the declivity descends in a straight line to the level of the propodeal spiracle.

Petiolar scale sharp and inclined anteriorly, in lateral view falling short of the propodeal spiracle.

Dorsum of head (excluding clypeus), mesosoma, petiole, and gastric tergites 1–2 ( = abdominal tergites 3–4) devoid of erect setae (very rarely with a pair of small setae on gastric tergite 2). Gastric tergites 3–4 each bearing a pair of long, erect setae. Venter of metasoma with scattered erect setae.

Integument shagreened and lightly shining. Body and appendages including gula, entire mesopleura, metapleura, and abdominal tergites covered in dense pubescence.

Body and appendages concolorous, most commonly a medium reddish or yellowish brown but ranging in some populations from testaceous to dark brown, never yellow or piceous.


(n = 13) HL 0.83–0.92, HW 0.83–0.93, SL 0.81–0.89, FL 0.78–0.90, LHT 0.88–0.97, EL 0.31–0.36, MML 1.67–2.09, WL 4.42–4.51, CI 93–101, SI 96–102, OI 36–39, WI 24–27, FI 40–48.

Moderately large species (MML 1.67–2.09). Head slightly longer than broad to as broad as long in full face view (CI 93–101), posterior margin slightly concave to slightly convex. Eyes of moderate size (OI 36–39). Ocelli small. Antennal scapes relatively long (SI 96–102), in full face view scapes in repose surpassing posterior margin by a length greater than length of first funicular segment.

Forewings short relative to mesosomal length (WI 24–27). Forewings with Rs+M at least three times longer than M.f2. Legs of moderate length relative to mesosomal length (FI 40–48).

Dorsum of mesosoma and metasoma with scattered standing setae. Mesoscutum bearing 2–11 standing setae. Body color medium reddish brown. Antennal scapes, legs, and mandibles concolorous with body.


(n = 12) HL 0.56–0.70, HW 0.56–0.74, SL 0.13–0.16, FL 0.60–0.77, LHT 0.51–0.66, EL 0.31–0.34, MML 1.40–1.96, WL 2.55–3.26, PH 0.25–0.34, CI 99–106, SI 22–27, OI 51–55, WI 17–20, FI 37–45.

Head about as broad as long in full face view (CI 99–106). Eyes large (OI 51–55), occupying much of anterolateral surface of head and separated from posterolateral clypeal margin by a length less than width of antennal scape. Ocelli large and in full frontal view set above adjoining posterolateral margins. Antennal scape of moderate length (SI 22–27), about 2/3 length of 3rd antennal segment. Anterior clypeal margin straight to broadly convex. Mandibles small, bearing a single apical tooth and 4-8 denticles along masticatory margin and rounding into inner margin. Masticatory margin relatively short, subequal in length to inner margin. Inner margin roughly parallel to, or converging distally with, exterior lateral margin.

Mesosoma unusually well developed, considerably wider than head width, and larger in bulk and in length than metasoma. Mesoscutum greatly enlarged, projecting forward in a convexity overhanging pronotum. Scutellum large, convex, nearly as tall as mesoscutum and projecting well above level of propodeum. Propodeum well developed and overhanging petiolar node, posterior propodeal face strongly concave. Forewings short relative to mesosomal length (WI 17–20) and bearing a single submarginal cell. Wing color whitish or yellowish with dark brown veins and stigma. Legs short relative to mesosoma length (FI 37–45).

Petiolar scale taller than node length and bearing a broad crest. Ventral process well developed. Gaster oval in dorsal view, nearly twice as long as broad. Gonostylus produced as a bluntly rounded pilose lobe. Volsella with cuspis present, digitus short and downturned distally.

Dorsal surfaces of body largely devoid of erect setae, occasionally with a few fine, short setae scattered on mesoscutum, scutellum, and posterior abdominal tergites. Venter of gaster with scattered setae. Pubescence dense on body and appendages, becoming sparse only on medial propodeal dorsum.

Color as for worker.

Worker Morphology

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  • Caste: monomorphic


  • n = 8, 2n = 16 (Australia; Spain) (Crozier, 1968a; Crozier, 1975; Lorite et al., 1996b; Lorite et al., 1998b) (as Iridomyrmex humilis).


Additional References

See also a list of some recent studies of the Argentine ant.

References based on Global Ant Biodiversity Informatics

  • Andrade T., G. D. V. Marques, K. Del-Claro. 2007. Diversity of ground dwelling ants in Cerrado: an analysis of temporal variations and distinctive physiognomies of vegetation (Hymenoptera: Formicidae). Sociobiology 50(1): 121-134.
  • Borgmeier T. 1923. Catalogo systematico e synonymico das formigas do Brasil. 1 parte. Subfam. Dorylinae, Cerapachyinae, Ponerinae, Dolichoderinae. Archivos do Museu Nacional (Rio de Janeiro) 24: 33-103.
  • Caldart V. M., S. Iop, J. A. Lutinski, and F. R. Mello Garcia. 2012. Ants diversity (Hymenoptera, Formicidae) of the urban perimeter of Chapecó county, Santa Catarina, Brazil. Revista Brasileira de Zoociências 14 (1, 2, 3): 81-94.
  • Clemes Cardoso D., and J. H. Schoereder. 2014. Biotic and abiotic factors shaping ant (Hymenoptera: Formicidae) assemblages in Brazilian coastal sand dunes: the case of restinga in Santa Catarina. Florida Entomologist 97(4): 1443-1450.
  • Clemes Cardoso D., and M. Passos Cristiano. 2010. Myrmecofauna of the Southern Catarinense Restinga sandy coastal plain: new records of species occurrence for the state of Santa Catarina and Brazil. Sociobiology 55(1b): 229-239.
  • Corassa J. N., I. C. Magistrali, J. C. Moreno, E. B. Cantarelli, and A. Corassa. Effect of formicid granulated baits on non-target ants biodiversity in eucalyptus plantations litter. Comunicata Scientiae 4(1): 35-42.
  • Costa-Milanez C. B., G. Lourenco-Silva, P. T. A. Castro, J. D. Majer, and S. P. Ribeiro. 2014. Are ant assemblages of Brazilian veredas characterised by location or habitat type? Braz. J. Biol. 74(1): 89-99.
  • Dias N. S., R. Zanetti, M. S. Santos, J. Louzada, and J. H. C. Delabie. 2008. Interaction between forest fragments and adjacent coffee and pasture agroecosystems: responses of the ant communities (Hymenoptera, Formicidae). Iheringia, Sér. Zool., Porto Alegre, 98(1): 136-142.
  • Fagundes R., G. Terra, S. P. Ribeiro, and J. D. Majer. 2010. The Bamboo Merostachys fischeriana (Bambusoideae: Bambuseae) as a Canopy Habitat for Ants of Neotropical Montane Forest. Neotropical Entomology 39(6):906-911
  • Favretto M. A., E. Bortolon dos Santos, and C. J. Geuster. 2013. Entomofauna from West of Santa Catarina State, South of Brazil. EntomoBrasilis 6 (1): 42-63.
  • Fernández, F. and S. Sendoya. 2004. Lista de las hormigas neotropicales. Biota Colombiana Volume 5, Number 1.
  • Fleck M. D., E. Bisognin Cantarelli, and F. Granzotto. 2015. Register of new species of ants (Hymenoptera: Formicidae) in Rio Grande do Sul state. Ciencia Florestal, Santa Maria 25(2): 491-499.
  • Forel A. 1908. Ameisen aus Sao Paulo (Brasilien), Paraguay etc. gesammelt von Prof. Herm. v. Ihering, Dr. Lutz, Dr. Fiebrig, etc. Verhandlungen der Kaiserlich-Königlichen Zoologisch-Botanischen Gesellschaft in Wien 58: 340-418.
  • Gallardo A. 1916. Las hormigas de la República Argentina. Subfamilia Dolicoderinas. Anales del Museo Nacional de Historia Natural de Buenos Aires 28: 1-130.
  • Gallardo A. 1920. Las hormigas de la República Argentina. Subfamilia Dorilinas. Anales del Museo Nacional de Historia Natural de Buenos Aires 30: 281-410.
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