Linepithema humile

This is the notoriously pestiferous Argentine ant. 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.

Identification
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).

Distribution
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)

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, 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, Japan, Malta, Monaco, Montenegro, Morocco, Poland, Portugal, Spain, United Kingdom of Great Britain and Northern Ireland.

Biology
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.

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).

Lepidoptera

 * This ant has been observed tending larvae of (Obregon et al. 2015).

Predators

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

Genetics
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.

Castes
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).

Nomenclature

 *  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.

Taxonomic Notes
The taxonomy and distribution of L. humile was reviewed in depth by Wild (2004).

The following described based on Wild (2007):

Worker
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.

Queen
(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.

Male
(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.

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 fi scheriana (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.
 * Heller, N.E. 2004. Colony structure in introduced and native populations of the invasive Argentine ant, Linepithema humile. Insectes Sociaux 51(4): 378-386.
 * Ilha C., J. A. Lutinski, D. Von Muller Pereira, F. R. Mello Garcia. 2009. Riqueza de formigas (Hymenoptera: Formicidae) de Bacia da Sanga Caramuru, municipio de Chapeco-SC. Biotemas 22(4): 95-105.
 * Iop S., V. M. Caldart, J. A. Lutinski, and F. R. Mello Garcia. 2009. Formigas urbanas da cidade de Xanxerê, Santa Catarina, Brasil. Biotemas 22(2): 55-64.
 * Kamura, C.M., M.S.C. Morini, C.J. Figueiredo, O.C. Bueno, and A.E.C. Campos-Farinha. 2007. Comunidades de formigas (Hymenoptera: Formicidae) em um ecossistema urbano próximo à Mata Atlântica. Brazilian Journal of Biology 67(4): 635-641
 * Kempf, W.W. 1972. Catalago abreviado das formigas da regiao Neotropical (Hym. Formicidae) Studia Entomologica 15(1-4).
 * Kusnezov N. 1953. La fauna mirmecológica de Bolivia. Folia Universitaria. Cochabamba 6: 211-229.
 * Lapola D. M., and H. G. Fowler. 2008. Questioning the implementation of habitat corridors: a case study in interior São Paulo using ants as bioindicators. Braz. J. Biol., 68(1): 11-20.
 * LeBrun, E.G., C. V. Tillberg, A. V. Suarez, P. J. Folgarait, C. R. Smith and D. A. Holway. 2007. An Experimental Study of Competition between Fire Ants and Argentine Ants in Their Native Range. Ecology 88(1):63-75
 * Luederwaldt H. 1918. Notas myrmecologicas. Rev. Mus. Paul. 10: 29-64.
 * Lutinski J. A., B. C. Lopes, and A. B. B.de Morais. 2013. Diversidade de formigas urbanas (Hymenoptera: Formicidae) de dez cidades do sul do Brasil. Biota Neotrop. 13(3): 332-342.
 * Lutinski J. A., F. R. Mello Garcia, C. J. Lutinska, and S. Iop. 2008. Ants diversity in Floresta Nacional de Chapecó in Santa Catarina State, Brazil. Ciência Rural, Santa Maria 38(7): 1810-1816.
 * Marinho C. G. S., R. Zanetti, J. H. C. Delabie, M. N. Schlindwein, and L. de S. Ramos. 2002. Ant (Hymenoptera: Formicidae) Diversity in Eucalyptus (Myrtaceae) Plantations and Cerrado Litter in Minas Gerais, Brazil. Neotropical Entomology 31(2): 187-195.
 * Marques G. D. V., and K. Del-Claro. 2006. The Ant Fauna in a Cerrado area: The Influence of Vegetation Structure and Seasonality (Hymenoptera: Formicidae). Sociobiology 47(1): 1-18.
 * Martinez M.J. and Weis E.M. 2011. field observations of two species of invasive ants, Linepithema humile Mayr, 1868 and Tetramoirum bicarinatum Nylander, 1846 (Hymenoptera: Formicidae), at a suburban park in Southern California. The Pan-Pacific Entomologist. 87: 57-61
 * Menozzi C. 1926. Neue Ameisen aus Brasilien. Zoologischer Anzeiger. 69: 68-72.
 * Morini M. S. de C., C. de B. Munhae, R. Leung, D. F. Candiani, and J. C. Voltolini. 2007. Comunidades de formigas (Hymenoptera, Formicidae) em fragmentos de Mata Atlântica situados em áreas urbanizadas. Iheringia, Sér. Zool., Porto Alegre, 97(3): 246-252.
 * Oliveira R. F., L. C. de Almeida, D. R. de Suza, C. Bortoli Munhae, O. C. Bueno, and M. S. de Castro Morini. 2012. Ant diversity (Hymenoptera: Formicidae) and predation by ants on the different stages of the sugarcane borer life cycle Diatraea saccharalis (Lepidoptera: Crambidae). Eur. J. Entomol. 109: 381387.
 * Orr M. R., and S. H. Seike. 1998. Parasitoids deter foraging by Argentine ants (Linepithema humile) in their native habitat in Brazil. Oecologia 117: 420-425.
 * Pacheco R., and H. L. Vasconcelos. 2007. Invertebrate conservation in urban areas: ants in the Brazilian Cerrado. Landscape and Urban Planning 81: 193199.
 * Pacheco, R., R.R. Silva, M.S. de C. Morini, C.R.F. Brandao. 2009. A Comparison of the Leaf-Litter Ant Fauna in a Secondary Atlantic Forest with an Adjacent Pine Plantation in Southeastern Brazil. Neotropical Entomology 38(1):055-065
 * Pedersen, J.S., M.J.B. Krieger, V. Vogel, T. Giraud and L. Keller 2006. Native Supercolonies of Unrelated Individuals in the Invasive Argentine Ant. Evolution 60(4):782-791
 * Pereira M. C., J. H. C. Delabie, Y. R. Suarez, and W. F. Antonialli Junior. 2013. Spatial connectivity of aquatic macrophytes and flood cycle influence species richness of an ant community of a Brazilian floodplain. Sociobiology 60(1): 41-49.
 * Pignalberi C. T. 1961. Contribución al conocimiento de los formícidos de la provincia de Santa Fé. Pp. 165-173 in: Comisión Investigación Científica; Consejo Nacional de Investigaciones Científicas y Técnicas (Argentina) 1961. Actas y trabajos del primer Congreso Sudamericano de Zoología (La Plata, 12-24 octubre 1959). Tomo III. Buenos Aires: Librart, 276 pp.
 * Piva A., and A. E. de C. Campos. 2012. Ant Community Structure (Hymenoptera: Formicidae) in Two Neighborhoods with Different Urban Profiles in the City of Sao Paulo, Brazil. Psyche 2012 (390748): 1-8
 * Ramos L. S., R. Z. B. Filho, J. H. C. Delabie, S. Lacau, M. F. S. dos Santos, I. C. do Nascimento, and C. G. S. Marinho. 2003. Ant communities (Hymenoptera: Formicidae) of the leaf-litter in cerrado stricto sensu areas in Minas Gerais, Brazil. Lundiana 4(2): 95-102.
 * Ramos L. de S., C. G. S. Marinho, R. Zanetti, and J. H. C. Delabie. 2003. Impacto de iscas formicidas granuladas sobre a mirmecofauna não-alvo em eucaliptais segundo duas formas de aplicacação / Impact of formicid granulated baits on non-target ants in eucalyptus plantations according to two forms of application. Neotropical Entomology 32(2): 231-237.
 * Ramos L. de S., R. Zanetti, C. G. S. Marinho, J. H. C. Delabie, M. N. Schlindwein, and R. P. Almado. 2004. Impact of mechanical and chemical weedings of Eucalyptus grandis undergrowth on an ant community (Hymenoptera: Formicidae). Rev. Árvore 28(1): 139-146.
 * Ribas C. R., J. H. Schoereder, M. Pic, and S. M. Soares. 2003. Tree heterogeneity, resource availability, and larger scale processes regulating arboreal ant species richness. Austral Ecology 28(3): 305-314.
 * Rosa da Silva R. 1999. Formigas (Hymenoptera: Formicidae) do oeste de Santa Catarina: historico das coletas e lista atualizada das especies do Estado de Santa Catarina. Biotemas 12(2): 75-100.
 * Santoandre S., J. Filloy, G. A. Zurita, and M. I. Bellocq. 2019. Ant taxonomic and functional diversity show differential response to plantation age in two contrasting biomes. Forest Ecology and Management 437: 304-313.
 * Santos M. S., J. N. C. Louzada, N. Dias, R. Zanetti, J. H. C. Delabie, and I. C. Nascimento. 2006. Litter ants richness (Hymenoptera, Formicidae) in remnants of a semi-deciduous forest in the Atlantic rain forest, Alto do Rio Grande region, Minas Gerais, Brazil. Iheringia, Sér. Zool., Porto Alegre, 96(1): 95-101.
 * Santschi F. 1912. Quelques fourmis de l'Amérique australe. Revue Suisse de Zoologie 20: 519-534.
 * Santschi F. 1916. Formicides sudaméricains nouveaux ou peu connus. Physis (Buenos Aires). 2: 365-399.
 * Santschi F. 1925. Fourmis des provinces argentines de Santa Fe, Catamarca, Santa Cruz, Córdoba et Los Andes. Comunicaciones del Museo Nacional de Historia Natural "Bernardino Rivadavia" 2: 149-168.
 * Schoereder J. H., T. G. Sobrinho, M. S. Madureira, C. R. Ribas, and P. S. Oliveira. 2010. The arboreal ant community visiting extrafloral nectaries in the Neotropical cerrado savanna. Terrestrial Arthropod Reviews 3: 3-27.
 * Silva F. H. O., J. H. C. Delabie, G. B. dos Santos, E. Meurer, and M. I. Marques. 2013. Mini-Winkler Extractor and Pitfall Trap as Complementary Methods to Sample Formicidae. Neotrop Entomol 42: 351358.
 * Silvestre R., M. F. Demetrio, and J. H. C. Delabie. 2012. Community Structure of Leaf-Litter Ants in a Neotropical Dry Forest: A Biogeographic Approach to Explain Betadiversity. Psyche doi:10.1155/2012/306925
 * Tillberg, C.V., D.P. McCarthy, A.G. Dolezal and A.V. Suarez. 2006. Measuring the trophic ecology of ants using stable isotopes. Insectes Sociaux 53:65-69
 * Ulyssea M.A., C. E. Cereto, F. B. Rosumek, R. R. Silva, and B. C. Lopes. 2011. Updated list of ant species (Hymenoptera, Formicidae) recorded in Santa Catarina State, southern Brazil, with a discussion of research advances and priorities. Revista Brasileira de Entomologia 55(4): 603-611.
 * Vasconcelos, H.L., J.M.S. Vilhena, W.E. Magnusson and A.L.K.M. Albernaz. 2006. Long-term effects of forest fragmentation on Amazonian ant communities. Journal of Biogeography 33:1348-1356
 * Vittar, F. 2008. Hormigas (Hymenoptera: Formicidae) de la Mesopotamia Argentina. INSUGEO Miscelania 17(2):447-466
 * Vittar, F., and F. Cuezzo. "Hormigas (Hymenoptera: Formicidae) de la provincia de Santa Fe, Argentina." Revista de la Sociedad Entomológica Argentina (versión On-line ISSN 1851-7471) 67, no. 1-2 (2008).
 * Wild A. L. 2004. Taxonomy and distribution of the Argentine ant, Linepithema humile (Hymenoptera: Formicidae). Annals of the Entomological Society of America 97: 1204-1215
 * Wild A. L. 2007. Taxonomic revision of the ant genus Linepithema (Hymenoptera: Formicidae). University of California Publications in Entomology 126: 1-151
 * Wild, A. L.. "A catalogue of the ants of Paraguay (Hymenoptera: Formicidae)." Zootaxa 1622 (2007): 1-55.
 * Zoppas de Albuquerque E., and E. Diehl. 2009. Análise faunística das formigas epígeas (Hymenoptera, Formicidae) em campo nativo no Planalto das Araucárias, Rio Grande do Sul. Revista Brasileira de Entomologia 53(3): 398403.
 * da Silva Araujo, M., Castro Della Lucia, T.M., DA VEIGA, Clayton E y CARDOSO DO NASCIMENTO, Ivan. 2004. Efeito da queima da palhada de cana-de-açúcar sobre comunidade de formicídeos. Ecol. austral. 14(2): 191-200.
 * da Silva T. F., D. Russ Solis, T. de Carvalho Moretti, A. Calazans da Silva, M. E. Din Mostafa Habib. 2009. House-infesting ants (Hymenoptera: Formicidae) in a municipality of southeastern Brazil. Sociobiology 54(1): 153-159.
 * de Almeida Soares S., Y. R. Suarez, W. D. Fernandes, P. M. Soares Tenorio, J. H. C. Delabie, and W. F. Antonialli-Junior. 2013. Temporal variation in the composition of ant assemblages (Hymenoptera, Formicidae) on trees in the Pantanal floodplain, Mato Grosso do Sul, Brazil. Rev. Bras. entomol. 57: 84-90
 * de Zolessi, L.C., Y.P. de Abenante and M.E. Philippi. 1987. Lista sistemática de las especies de formícidos del Uruguay. Comunicaciones Zoologicas del Museo de Historia Natural de Montevideo 11(165):1-9