Neoponera villosa

A common species that is found in a variety of habitats.

Identification
From Mackay and Mackay (2010): Neoponera villosa vies with Pachycondyla harpax for being the most common and widely distributed member of ponerine in the New World. Neoponera villosa is much larger than P. harpax (total length of  the worker, female and male of P. harpax < 10 mm). Workers of N. villosa can be easily confused with Neoponera holcotyle, Neoponera  theresiae,  and Neoponera foetida. The side of the petiole of N. villosa is without any evidence of striae, which are present in the other three species. The first two of these three species are also smaller (total length < 11 mm), N. foetida is approximately the same size. Neoponera foetida also has horizontal striae on the posterior face of the petiole, which are lacking in N. villosa. Neoponera villosa could also misidentified as Neoponera bugabensis, as both lack striae on the sides and posterior face of the petiole. It is much larger (total length of workers of N. bugabensis is < 10 mm) and the legs are dark red (ferrugineous red in color in N. bugabensis).

It is easy to confuse this common species with several other species. For example, smaller specimens with striae on the dorsum of the head between the eyes are Neoponera lineaticeps, specimens with a rectangular shaped petiole, a concave pygidium with horizontal lateral striae are Pachycondyla impressa,  specimens with two lateral angulate processes on the pygidium are Pachycondyla crassinoda. It is mostly dull in color, not shiny black as in the relatively large Neoponera commutata, Neoponera laevigata  and Neoponera marginata. Neoponera fisheri is a similar size, but lacks the preocular and pronotal carinae.

It is similar to the closely related Neoponera curvinodis and Neoponera inversa, but can be separated from both species by the straight anterior face of the petiole (concave in the other two species). Mariano et al. (2000) compare the cytogenetics of N. villosa and N. inversa.

The males of three common species are very similar: N. villosa, Pachycondyla striata and P. impressa. Neoponera villosa males can be separated from the males of the other two species by the presence of the relatively large sharp subpetiolar process, which is blunt and rounded or only angulate anteriorly in the other two species. The erect hairs of N. villosa are not as abundant as they are in P. striata and the petiole is not covered with rugae as it is in P. impressa. The males of the closely related N. curvinodis and N. inversa are unknown, but it could be expected that the anterior faces of the petioles of these two species would be concave.

Fernandes et al. (2014) - Worker: Anterior border of clypeus concave medially; anterior face of petiole vertical and broadly convex on dorsum. Male: Posteropropodeum strongly striate; anterior face of petiole vertical, petiole robust with posterolateral carina.

Neoponera villosa can be differentiated from other species of the Neoponera foetida species complex by the anterior margin of its clypeus, which is concave medially without striae, and by the anterior face of the petiole, which is vertical and convex with a broadly rounded posterior face, characters absent in the other species. Neoponera curvinodis can be differentiated by the shape of the petiole, which has a concave anterior face, while that in N. villosa is vertical. Neoponera theresiae can be differentiated by striae on the side and dorsum of the petiole, characters not observed in N. villosa.

Neoponera villosa has a wide distribution in the New World, occurring from the southern United States to Argentina. The species shows little variation in color, although specimens collected nearer to the equator circle have the petiole, mesosoma, gaster, and legs darker brown. Body size is another variable character, but is not correlated with any changes in morphological characters. Diet and age of nest are known to influence body size in many ant species.

Distribution
Southern United States through South America, Trinidad, Puerto Rico (Mackay and Mackay 2010).

Distribution based on Regional Taxon Lists
Nearctic Region: United States. Neotropical Region: Argentina, Belize, Bolivia, Brazil, Colombia, Costa Rica, Ecuador, El Salvador, French Guiana, Guatemala, Guyana, Honduras, Mexico, Nicaragua, Panama, Paraguay, Peru, Puerto Rico, Suriname, Trinidad and Tobago, Venezuela.

Habitat
This common species is found in a variety of habitats, ranging from grasslands (Quiroz-Robledo and Valenzuela-González, 1995), primary forest (Roth et al., 1994), secondary rain forest, tropical rain forest, wet tropical rain forest, lowland forest, mangrove, bosque medio, the edge of natural forest, gallery forest, epiphyte forest, open grassy areas with a few trees, area with scattered oaks, to highly disturbed areas such as cacao plantations and even dry scrub forest. Elevations range from 1.5 - 1100 m. Wild (2002) includes undisturbed primary tall forest, gallery forest, scrub forest, campo cerrado [scrub vegetation], pastures, lawns and orchards. This species has been collected in caves near the entrances (Reddell and Cokendolpher, 2001). (Mackay and Mackay 2010)

Biology
Sanz-Veiga et al. (2017) - This species was observed visiting the extrafloral nectaries of Tocoyena formosa plants occurring in a southeastern Brazilian cerrado study-site.

From Mackay and Mackay (2010): This species nests in wood, including dead, hard trees, dead branches and trunks, stumps, under bark and even in dead logs on the ground. It is one of the rare Ponerinae that nests in trees (Dejean and Corbara, 1990b). Dealate queens sometimes establish their first nests in hollow twigs. Nests usually start cooperatively with two or more queens who establish a dominance order with a division of labor (D’Ettorre et al., 2005). One nest was at the base of small tree.

Brood was collected in a nest in July (Ecuador). Males were found in nests in June (Nicaragua), July (Ecuador) and December (Texas). Winged females have been collected in January and March (Costa Rica), March - April (Guatemala), June (USA), July (USA, Panamá), November (British Guyana) and December (USA, Perú, Costa Rica). Males have been collected in May (Costa Rica) and August (Honduras). Dealate females were collected in February (Costa Rica), March (México), May (Costa Rica), July (México, Panamá), August (Brasil) and September (Texas, Brasil).

It is a common arboreal ant in Brasil (De Medeiros et al., 1995) and also forages commonly on the forest floor. Workers are predaceous (Maes, 1989; Dejean et al., 1990) and prey on termites (Wild, 2005). Foragers also feed on nectar (Paul and Roces, 2003) and can carry droplets of liquids between their mandibles (Hölldobler, 1985). They can be found in forest leaf litter, on trees, at baits on the soil surface and in pitfall traps.

They nest almost exclusively in bromeliad epiphytes (Aechmea bracteata) in the state of Quintana Roo, México (Dejean, 1990; Dejean and Olmsted, 1997) and are occasionally found in Cecropia sp., especially in Cecropia hispidissima [Cecropiaceae]. Wheeler (1942) lists this species as nesting in pseudobulbs of Schomburgkia tibicinis [Orchidaceae]. This species lives in the epiphytes Tillandsia bulbosa [Bromeliaceae] and T. streptophylla (Dejean et al., 1995). It nests in peripheral cavities of trees of Bursera simaruba [Burseraceae] and protects it from attack from the leaf cutting ant Atta cephalotes (Dejean et al., 1992). It collects nectar from the mistletoe Phoradendron tomentosum [Loranthaceae] in southern Texas (Whittaker, 1984). Fagundes et al. (2010) report it nesting in bamboo.

Zara et al. (2002) did chemical analysis of the different larval instars and (2003) discussed the cytochemical and chemical analysis of the fat bodies of this species. Caetano (1988) described the digestive and excretory system. Lipids are the first materials to be deposited in the oocytes (Caperucci and Camargo-Mathias, 2006). The size of the corpora allata in females, located dorsolaterally to the esophagus, is inversely related to the development of the ovaries (Camargo and Caetano, 1995b).

Neoponera villosa is eaten by the leptodactylid “veragua robber” frog, Eleutherodactylus biporcatus in Nicaragua (Wild, 2005). It is the prey of the army ant Eciton mexicanum. Neoponera villosa is mimicked by the salticid spider Zuniga magna (Reiskind, 1977), but only the female (McIver and Stonedahl, 1993). The male of the spider mimics the ant Pseudomyrmex gracilis (McIver and Stonedahl, 1993). One colony was nesting together with an ant of the genus Cephalotes in Colombia. It apparently has a positive relationship with the carpenter ant Camponotus atriceps (Majer et al., 1994, listed as C. abdominalis).

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. Neoponera villosa was found in 13 different bromeliads but was associated with twigs and bark cavities, rather than suspended soil or litter, of the plant.

Dejain et al. (2018) found this species nesting in Aechmea bracteata, a large tank bromeliad (leaves ca. 1-m-long; inflorescences up to 1.7-m-long). Each plant forms numerous shoots, with each bearing multiple reservoirs within their leaves and a central dry cavity where ants can nest. Other ants were also found, and both the presence of ants and the ant species present was found to be correlated with changes in the aquatic macroinvertebrate communities within the plants' reservoirs. Neoponera villosa was found in 33 of the 92 plants sampled. The study was carried out in an inundated forest dominated by 10-m-tall Metopium brownei (Anacardiaceae). This forest was located in southern Quintana Roo, Mexico (18.42678 N; 88.80438 W; 120 meters a.s.l).

Behavior
This is a very aggressive ant, with a painful sting (feels like being stabbed with a hot needle). The sharp pain mostly subsides after about a half an hour, but remains tender for a few days. They are avoided by the army ant Eciton burchellii (García-Pérez, 1989).

Forty percent of founding colonies are pleometrotic, with two (24%) or three (16%) nest females and may lead to polygynous nests (Trunzer et al., 1998, probably refers to Neoponera inversa - see Kolmer and Heinze, 2000b). The females leave the nest to forage before the first workers are developed (Trunzer et al., 1998). Workers lay eggs which are utilized as food by the colony (Camargo Mathias and Caetano, 1995a). The queens mate with two or more males and the queens that cooperatively form new nests are not closely related (Kellner et al., 2007).

Workers in queenless colonies form a dominance hierarchy, using “boxing” and biting (Heinze et al., 1996). The top ranking individuals lay the most eggs, but lower ranking individuals also lay eggs, which result in the production of males (Trunzer et al., 1999). The dominant workers eat the eggs of other workers. The presence of larvae reduces the numbers of eggs in the nest, by feeding on the eggs and fewer eggs are laid when larvae are present (Heinze et al., 1996).

Individual foragers show strong fidelity to a particular area, but forage individually, without recruitment, although tandem running has been observed during nest relocations  (Fresneau, 1985). Neoponera villosa responds differently to different prey types and does not always use the sting (Dejean and Corbara, 1990b). Dejean et al. (1990), discuss the behavior of Neoponera villosa when foragers prey on rhinotermitid termites. They move their antennae posteriorly and lift as many legs as possible from the soil surface to avoid contact with the termites. Foragers follow four types of paths including classical exploration, area-concentrated searching, reserve behavior and homing (Dejean and Corbara, 1998).

Workers demonstrate polyethism, with tasks changing during their development (Pérez-Bautista et al., 1985).

Nomenclature

 *  villosa. Formica villosa Fabricius, 1804: 409 (w.) CENTRAL AMERICA. Roger, 1861a: 2 (w.q.); Wheeler, G.C. & Wheeler, J. 1952c: 615 (l.); Petralia & Vinson, 1980: 378 (l.). Combination in Pachycondyla: Mayr, 1862: 720; in Neoponera: Emery, 1901a: 43; in Pachycondyla: Brown, in Bolton, 1995b: 311; in Neoponera: Schmidt & Shattuck, 2014: 152. Senior synonym of bicolor, pedunculata, pilosa: Roger, 1861: 1; Roger, 1862c: 288. See also: Wheeler, W.M. 1908e: 404; Gallardo, 1918b: 56; Mackay & Mackay, 2010: 571.
 * bicolor. Ponera bicolor Guérin-Méneville, 1844a: 424 (w.) MEXICO. Junior synonym of villosa: Roger, 1861a: 1; Roger, 1862c: 288.
 * pedunculata. Ponera pedunculata Smith, F. 1858b: 96, pl. 6, fig. 25 (w.) BRAZIL. Junior synonym of villosa: Roger, 1861a: 1; Roger, 1862c: 288.
 * pilosa. Ponera pilosa Smith, F. 1858b: 95 (m.) BRAZIL. Junior synonym of villosa: Roger, 1861a: 1; Roger, 1862c: 288.

Type Material
Central America; México; Brasil, Villa Nova; Brasil, Rio Constantia. Worker seen, was compared to type by Brown, MCZC (Mackay and Mackay 2010)

Ponera pedunculata

Holotype worker in. Labelled “Rio. Constancia.”

A specimen in previously indicated as type is labelled “Ega. 58/6” (= Brazil) and therefore is not type-material. A specimen in has “pedunculata” on a label but no other data. All specimens represent a single species.

Ponera pilosa

Holotype male in. Labelled “Brazil. Villa Nova.” There is another male in, mounted as the holotype but without the correct data labels.

Worker
From Mackay and Mackay (2010) - Workers of this species are large (total length about 15 mm) black ants with black or reddish brown tibiae. The anterior border of the clypeus is convex but is concave medially. The malar carina is well developed and sharp, shining and well marked against a background of golden appressed pubescence. The maximum diameter the eye is about one fourth the length of the side of the head and is located more than one maximum diameter from the insertion of the mandible (side view). The scape extends approximately the first two funicular segments past the posterior lateral corner of the head. The carina on the pronotal shoulder is sharp and slightly overhangs the side of the pronotum. The metanotal suture is well marked on the dorsum of the mesosoma and depresses the level slightly below the remainder of the mesosoma. The propodeal spiracle is slit-shaped. The posterior face of the propodeum is separated from the side of the propodeum by a definite carina. The petiole is thick when viewed in profile, with a vertical anterior face and a convex broadly rounded posterior face that meets the anterior face at the anterior point of the apex of the petiole. The posterior lateral edges of the petiole nearly form carinae. The stridulatory file is well developed on the second pretergite.

Erect hairs are abundant on most surfaces, including the dorsal and ventral surfaces of the head, the antennal scapes, the posterior border of the head, the dorsum of the mesosoma, dorsum of the petiole, all surfaces of the gaster, the hairs on the legs are either erect or suberect. Appressed golden pubescence is abundant on most surfaces, especially the mesosoma, petiole and gaster.

The head is covered with coarse punctures, which are somewhat aligned in rows. The dorsum of the mesosoma has finer punctures, and surfaces which are not hidden by appressed pubescence are glossy and shiny. The side of the mesosoma is mostly punctate, but the surfaces are nearly completely hidden by appressed pubescence. The petiole is finely punctate with surfaces not hidden by appressed pubescence, being shining. The dorsum of the gaster is punctate, but most surfaces are densely covered with appressed pubescence.

Fernandes et al. (2014) - (n = 261): SL: 2.60-2.80; HW: 2.67-2.77; HL: 2.89-2.98; PW: 1.79-1.84; WL: 4.50-4.59; NLd: 0.96-1.02; NWd: 1.29-1.36; NHl: 1.01-1.10; NLl: 1.30-1.40; NIl: 127.27-128.71; SI: 97.37-101.08.

Queen
Fernandes et al. (2014) - (n = 94): SL: 2.90-3.10; HW: 3.01-3.09; HL: 3.20-3.34; PW: 2.28-3.38; WL: 5.27-5.70; NLd: 1.22-1.34; NWd: 1.58-1.84; NHl: 1.06-1.19; NLl: 1.20-1.45; NIl: 113.20-121.84; SI: 96.34-100.32.

Redescription of queen type: SL: 3.03; HW: 3.08; HL: 3.29; PW: 2.35; WL: 5.73; NLd: 1.31; NWd: 1.81; NHl: 1.18; NLl: 1.43; NIl: 121.18; SI: 98.37.

Head: mandibles with 15 teeth on masticatory border; ocelli well developed; malar carina reaching eye; clypeus concave medially.

Mesosoma: pronotal carina sharp and well developed; scutum large and thickened; scutellum reduced; scutello-metanotal groove well developed; deep metanotal-propodeal grooves; propodeal carina poorly developed in dorsal view, with posteropropodeum nearly concave between carina in dorsal view.

Petiole: anterior face vertical, lateral face with posterolateral carina evident in lateral view; dorsal face broadly convex and rounded; sternopetiolar process forming anterior projection separated by carina.

Gaster: sternopostpetiolar process without carina.

Sculpture, pilosity and color: integument of head black and brown, finely punctate; scape covered by golden pubescence; clypeus with long and short erect golden hairs; mandibles opaque, with few punctures; masticatory border with short golden hairs and fine striation on dorsal surface; integument of mesosoma black and brown, abundant erect golden and silver hairs and pubescence distributed on all surfaces, more evident on dorsum; legs brown, covered with sparse silver and golden pubescence; anterior, medial and posterior basitarsi and tarsi brown, dense golden pubescence on ventral surface; medial and posterior tarsi with row of golden setae on ventral surface; tarsal claws not armed with tooth; arolium brown; integument of petiole brown below, black above, with erect golden hairs abundant on all surface, most evident on dorsum; integument of gaster brown weakly punctate, erect golden hairs present on dorsum and appressed gold and silver pubescence present on all surfaces; tergites covered by golden pubescence and long golden hairs; hypopygidium and pygidium punctate. Appressed golden and silver pubescence distributed on most surfaces, especially on mesosoma, petiole and gaster. Erect golden hairs abundant on most surfaces, including the dorsal and ventral surfaces of the head, the antennal scapes, the posterior border of the head, the dorsum of the mesosoma, dorsum of the petiole, all surfaces of the gaster. The hairs on the legs either erect or suberect.

From Mackay and Mackay (2010): The female is larger (about 17 mm total length) than the worker, usually black with reddish brown femora and appearing somewhat golden due to the abundant appressed pubescence. The anterior border of the clypeus is slightly concave medially and the malar carina is well developed. The distance between the insertion of the mandible and the anterior border of the clypeus is greater than the maximum diameter of the eye (side view). The ocelli are well developed and the scape extends slightly past the posterior lateral corner. The pronotal carina is very well developed, sharp and overhangs the side of pronotum. The metanotal suture is well developed and forms a well-defined metanotum. The petiole is shaped as in the worker. The wing is similar to that of other members of the genus.

The pilosity and sculpture are similar to that of the worker.

Male
From Mackay and Mackay (2010) - The male is a large (total length 14 mm) predominantly black specimen. The anterior border of the clypeus is slightly concave. The ocelli are moderately large (maximum diameter of the median ocellus is 0.19 mm, located 0.19 mm from the lateral ocellus, whose diameter is 0.21 mm). The pronotal shoulder is swollen, the propodeal spiracle is slit-shaped. The petiole is thick when viewed in profile with a nearly straight anterior face and a broadly rounded posterior face, which form a blunt rounded apex. The subpetiolar process consists of an angulate lobe, gradually diminishing in size posteriorly. The stridulatory file is present on the second pretergite, the arolia are well developed.

Erect hairs are moderately abundant on the head, mesosoma, petiole and gaster; appressed golden pubescence is dense on most surfaces.

Most surfaces are finely coriaceous and moderately shining.

The males come in two color forms: completely black and black with yellow legs and a yellow gaster. It is possible that two species are involved.

Fernandes et al. (2014) - (n = 65): HW: 1.92-2.01; HL: 1.50-1.55; PW: 1.40-1.48; WL: 4.00-4.12; NLd: 0.65-0.70; NWd: 0.70-0.75; NLl: 0.50-0.54; NHl: 0.85-0.93; NIl: 58.06-58.82.

Etymology
The name of this species is derived from the Latin word villosus, meaning hairy, referring to the condition of this species. (Mackay and Mackay 2010)

References based on Global Ant Biodiversity Informatics

 * Adams B. J., S. A. Schnitzer, and S. P. Yanoviak. 2019. Connectivity explains local ant community structure in a Neotropical forest canopy: a large-scale experimental approach. Ecology 100(6): e02673.
 * Alatorre-Bracamontes, C.E. and M Vasquez-Bolanos. 2010. Lista comentada de las hormigas (Hymenoptera: Formicidae) del norte de México. Dugesiana 17(1):9-36
 * Amat-G G., M. G. Andrade-C. and F. Fernández. (eds.) 1999. Insectos de Colombia. Volumen II. Bogotá: Academia Colombiana de Ciencias Exactas, Físicas y Naturales, 433 pp. 131975
 * Antoniazzi R., R. N. S. L. Garoo, W. Dattilo, S. P. Ribeiro, and F. S. Neves. 2019. Ant species richness and interactions in canopies of two distinct successional stages in a tropical dry forest. The Science of Nature 106: 20
 * Arruda F. V., M. A. Pesquero, D. G. Marcelino, G. A. Leiter, J. H. C. Delabie, and R. Fagundes. 2015. Size and condition of bamboo as structural factors behind the vertical stratification of the bamboo-nesting ant community. Insectes Sociaux DOI 10.1007/s00040-015-0440-4
 * Baena, M.L. 1993. Hormigas cazadoras del genero Pachcondyla (Hymenoptera: Ponerinae) de la Isla Gorgona y la Planicie Pacifica Colombiana. Bol. Mus. Ent. Univ. Valle 1(1):13-21
 * Bezdeckova K., P. Bedecka, and I. Machar. 2015. A checklist of the ants (Hymenoptera: Formicidae) of Peru. Zootaxa 4020 (1): 101–133.
 * 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.
 * Borgmeier T. 1934. Contribuição para o conhecimento da fauna mirmecológica dos cafezais de Paramaribo, Guiana Holandesa (Hym. Formicidae). Archivos do Instituto de Biologia Vegetal (Rio de Janeiro) 1: 93-111.
 * Byk J., K. Del-Claro. 2010. Nectar- and pollen-gathering Cephalotes ants provide no protection against herbivory: a new manipulative experiment to test ant protective capabilities. Acta Ethol. 13: 33-38.
 * Cameron P. 1891. Appendix. Hym. Formicidae. Pp. 89-95 in: Whymper, E. Travels amongst the Great Andes. London: J. Murray, xxii + 147 pp.
 * Castano-Meneses G., R. De Jesus Santos, J. R. Mala Dos Santos, J. H. C. Delabie, L. L. Lopes, and C. F. Mariano. 2019. Invertebrates associated to Ponerine ants nests in two cocoa farming systems in the southeast of the state of Bahia, Brazil. Tropical Ecology 60: 52–61.
 * Castano-Meneses, G., M. Vasquez-Bolanos, J. L. Navarrete-Heredia, G. A. Quiroz-Rocha, and I. Alcala-Martinez. 2015. Avances de Formicidae de Mexico. Universidad Nacional Autonoma de Mexico.
 * Christianini A. V., A. J. Mayhé-Nunes, and P. S. Oliveira. 2012. Exploitation of Fallen Diaspores By Ants: Are There Ant-Plant Partner Choices? Biotropica 44: 360-367.
 * Christianini A. V., and P. S. Oliveira. 2009. The relevance of ants as seed rescuers of a primarily bird-dispersed tree in the neotropical cerrado savanna. Oecologia 160: 735745.
 * Christianini A. V., and P. S. Oliveira. 2013. Edge effects decrease ant-derived benefits to seedlings in a neotropical savanna. Arthropod-Plant Interactions 7: 191-199.
 * Cokendolpher J. C., and O. F. Francke. 1990. The ants (Hymenoptera, Formicidae) of western Texas. Part II. Subfamilies Ecitoninae, Ponerinae, Pseudomyrmecinae, Dolichoderinae, and Formicinae. Special Publications, the Museum. Texas Tech University 30:1-76.
 * Correa M. M., W. D. Fernandes, and I. R. Leal. 2006. Ant Diversity (Hymenoptera: Formicidae) from Capões in Brazilian Pantanal: Relationship between Species Richness and Structural Complexity. Neotropical Entomology 35(6): 724-730.
 * Cuezzo, F. 1998. Formicidae. Chapter 42 in Morrone J.J., and S. Coscaron (dirs) Biodiversidad de artropodos argentinos: una perspectiva biotaxonomica Ediciones Sur, La Plata. Pages 452-462.
 * Dattilo W. et al. 2019. MEXICO ANTS: incidence and abundance along the Nearctic-Neotropical interface. Ecology https://doi.org/10.1002/ecy.2944
 * Davidson, D.W. 2005. Ecological stoichiometry of ants in a New World rain forest. Oecologia 142:221-231
 * Davidson, D.W., J.A. Arias and J. Mann. 2006. An experimental study of bamboo ants in western Amazonia. Insectes Sociaux 53:108-114
 * Davidson, D.W., J.A. Arias and J. Mann. 2006. An experimental study of bamboo ants in western Amazonia. Insectes Sociaux 53:108114.
 * Dejean A., B. Corbara, J. Orivel, R. R. Snelling, J. H. C. Delabie, and M. Belin-Depoux. 2000. The importance of ant gardens in the pioneer vegetal formations of French Guiana (Hymenoptera: Formicidae). Sociobiology 35(3): 425-439.
 * Dejean, A., S. Durou, I. Olmsted, R.R. Snelling and J. Orivel. 2003. Nest Site Selection by Ants in a Flooded Mexican Mangrove, with Special Reference to the Epiphytic Orchid Myrmecophila christinae. Journal of Tropical Ecology 19(3) :325-331
 * Dejean, A., S. Durou, I. Olmsted, R.R. Snelling and J. Orivel. 2003. Nest Site Selection by Ants in a Flooded Mexican Mangrove, with Special Reference to the Epiphytic Orchid Myrmecophila christinae. Journal of Tropical Ecology 19(3):325-331
 * Del Toro, I., M. Vazquez, W. Mackay, P. Rojas, and R. Zapata-Mata. "Hormigas (Hymenoptera: Formicidae) de Tabasco: explorando la diversidad de la mirmecofauna en las selvas tropicales de baja altitud." Dugesiana 16, no. 1 (2009): 1-14.
 * Del Toro, I., M. Vázquez, W.P. Mackay, P. Rojas and R. Zapata-Mata. Hormigas (Hymenoptera: Formicidae) de Tabasco: explorando la diversidad de la mirmecofauna en las selvas tropicales de baja altitud. Dugesiana 16(1):1-14.
 * Emery C. 1896. Formiciden, gesammelt in Paraguay von Dr. J. Bohls. Zoologische Jahrbücher. Abteilung für Systematik, Geographie und Biologie der Tiere 9: 625-638.
 * Emery C. 1906. Studi sulle formiche della fauna neotropica. XXVI. Bullettino della Società Entomologica Italiana 37: 107-194.
 * Emery C. 1911. Hymenoptera. Fam. Formicidae. Subfam. Ponerinae. Genera Insectorum 118: 1-125.
 * 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
 * Fernandes I. O., M. L. de Oliveira, and J. H. C. Delabie. 2014. Description of two new sepcies in the Neotropical Pachycondyla foetida complex (Hymenoptera: Formicidae: Ponerinae) and taxonomic notes on the genus. Myrmecological News 19: 133-163.
 * Fernández, F. and S. Sendoya. 2004. Lista de las hormigas neotropicales. Biota Colombiana Volume 5, Number 1.
 * Field Museum Collection, Chicago, Illinois (C. Moreau)
 * Flores-Maldonado K. Y., S. A. Phillips, and G. Sanchez-Ramos. 1999. The myrmecofauna (Hymenoptera: Formicidae) along an altitudinal gradient in the Sierra Madre Oriental of Northeastern Mexico. The Southwestern Naturalist 44(4): 457-461.
 * Flores-Maldonado, K. Y., S. A. Phillips-Jr, and G. Sanchez-Ramos. 1999. The myrmecofauna (Hymenoptera: Formicidae) along an altitudinal gradient in the Sierra Madre Oriental of Northeastern Mexico. The Southwestern Naturalist 44: 457-461.
 * Forel A. 1907. Formicides du Musée National Hongrois. Ann. Hist.-Nat. Mus. Natl. Hung. 5: 1-42.
 * 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.
 * Forel A. 1912. Formicides néotropiques. Part I. Annales de la Société Entomologique de Belgique. 56: 28-49.
 * Forel A. 1921. Quelques fourmis des environs de Quito (Ecuador) récoltées par Mlle Eléonore Naumann. Bulletin de la Société Vaudoise des Sciences Naturelles 54: 131-135.
 * Franco W., N. Ladino, J. H. C. Delabie, A. Dejean, J. Orivel, M. Fichaux, S. Groc, M. Leponce, and R. M. Feitosa. 2019. First checklist of the ants (Hymenoptera: Formicidae) of French Guiana. Zootaxa 4674(5): 509-543.
 * Gomez V. E. S., and G. Z. González. 2007. Catalogo de Las Hormigas Presentes en El Museo de Historia Natural de la Universidad del Cauca. Popayán : 1-58.
 * Groc S., J. H. C. Delabie, F. Fernandez, F. Petitclerc, B. Corbara, M. Leponce, R. Cereghino, and A. Dejean. 2017. Litter-dwelling ants as bioindicators to gauge the sustainability of small arboreal monocultures embedded in the Amazonian rainforest. Ecological Indicators 82: 43-49.
 * Henaut, Y., B. Corbara, L. Pelozuelo, F. Azemar, R. Cereghino, B. Herault, and A. Dejean, 2014. A tank bromeliad favors spider presence in a neotropical inundated forest. Plos One 9: e114592.
 * INBio Collection (via Gbif)
 * 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.
 * Jaffe, K., et al. 2007. Comparing the ant fauna in a tropical and a temperat forest canopy. Ecotropicos 20(2):74-81
 * Kempf W. W., and K. Lenko. 1976. Levantamento da formicifauna no litoral norte e ilhas adjacentes do Estado de São Paulo, Brasil. I. Subfamilias Dorylinae, Ponerinae e Pseudomyrmecinae (Hym., Formicidae). Studia Entomologica 19: 45-66.
 * Kempf, W.W. 1972. Catalago abreviado das formigas da regiao Neotropical (Hym. Formicidae) Studia Entomologica 15(1-4).
 * Kusnezov N. 1952. El género Camponotus en la Argentina (Hymenoptera, Formicidae). Acta Zoologica Lilloana 12: 183-252.
 * Kusnezov N. 1953. La fauna mirmecológica de Bolivia. Folia Universitaria. Cochabamba 6: 211-229.
 * Kusnezov N. 1978. Hormigas argentinas: clave para su identificación. Miscelánea. Instituto Miguel Lillo 61:1-147 + 28 pl.
 * LeBrun E. G., R. M. Plowes, and L. E. Gilbert. 2015. Imported fire ants near the edge of their range: disturbance and moisture determine prevalence and impact of an invasive social insect. Journal of Animal Ecology,81: 884–895.
 * Leal I. R. and B. Cortes Lopes. 1992. Estrutura das comunidades de formigas (Hymenoptera: Formicidae) de solo e vegetacao no morro da lagoa de conceicao, Ilha de Santa Catarina, SC. Capa 5(1): 107-122.
 * Leponce M., J. H. C. Delabie, J. Orivel, J. Jacquemin, M. Calvo Martin, and A. Dejean. 2019. Tree-dwelling ant survey (Hymenoptera, Formicidae) in Mitaraka, French Guiana, in Touroult J. (ed.), “Our Planet Reviewed” 2015 large-scale biotic survey in Mitaraka, French Guiana. Zoosystema 41 (10): 163-179.
 * Levings S. C. 1983. Seasonal, annual, and among-site variation in the ground ant community of a deciduous tropical forest: some causes of patchy species distributions. Ecological Monographs 53(4): 435-455.
 * Longino J. T. 2013. Ants of Nicargua. Consulted on 18 Jan 2013. https://sites.google.com/site/longinollama/reports/ants-of-nicaragua
 * Longino J. T. L., and M. G. Branstetter. 2018. The truncated bell: an enigmatic but pervasive elevational diversity pattern in Middle American ants. Ecography 41: 1-12.
 * Longino J. et al. ADMAC project. Accessed on March 24th 2017 at https://sites.google.com/site/admacsite/
 * Lozano-Zambrano F. H., E. Jimenez, T. M. Arias-Penna, A. M. Arcila, J. Rodriguez, and D. P. Ramirez. 2008. Biogeografía de las hormigas cazadoras de Colombia. Pp. 349-406. in: Jiménez, E.; Fernández, F.; Arias, T.M.; Lozano-Zambrano, F. H. (eds.) 2008. Sistemática, biogeografía y conservación de las hormigas cazadoras de Colombia. Bogotá: Instituto de Investigación de Recursos Biológicos Alexander von Humboldt, xiv + 609 pp.
 * 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.
 * MacKay, W.P. and E.E. MacKay. 2010. The systematics and biology of the New World ants of the genus Pachycondyla (Hymenoptera: Formicidae). Edwin Mellen Press Lewiston, NY
 * Maes, J.-M. and W.P. MacKay. 1993. Catalogo de las hormigas (Hymenoptera: Formicidae) de Nicaragua. Revista Nicaraguense de Entomologia 23.
 * Majer, J.D., J.H.C. Delabie and M.R.B. Smith. 1994. Arboreal ant community patterns in Brazilian cocoa farms. Biotropica 26(1): 73-83
 * Mann W. M. 1916. The Stanford Expedition to Brazil, 1911, John C. Branner, Director. The ants of Brazil. Bulletin of the Museum of Comparative Zoology 60: 399-490
 * Maravalhas J., J. H. C. Delabie, R. G. Macedo, and H. C. Morais. 2012. Tree-dwelling ants: contrasting two Brazilian Cerrado plant species without extrafloral nectaries. Psyche doi:10.1155/2012/172739
 * 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.
 * Mirmecofauna de la reserva ecologica de San Felipe Bacalar
 * Neves F. S., K. S. Queiroz-Dantas, W. D. da Rocha, and J. H. C. Delabie. 2013. Ants of Three Adjacent Habitats of a Transition Region Between the Cerrado and Caatinga Biomes: The Effects of Heterogeneity and Variation in Canopy Cover. Neotrop Entomol 42: 258268.
 * Neves F. S., R. F. Braga, M. M. do Espirito-Santo, J. H. C. Delabie, G. Wilson Fernandes, and G. A. Sanchez-Azofeifa. 2010. Diversity of Arboreal Ants In a Brazilian Tropical Dry Forest: Effects Of Seasonality and Successional Stage. Sociobiology 56(1): 1-18.
 * O'Keefe S. T., J. L. Cook, T. Dudek, D. F. Wunneburger, M. D. Guzman, R. N. Coulson, and S. B. Vinson. 2000. The Distribution of Texas Ants. The Southwestern Entomologist 22: 1-92.
 * Perez-Lachaud, G., M. A. Jervis, M. Reemer, and J. P. Lachaud. 2014. An unusual, but not unexpected, evolutionary step taken by syrphid flies: the first record of true primary parasitoidism of ants by Microdontinae. Biological Journal of the Linnean Society 111: 462-472.
 * Pergande, T. 1895. Mexican Formicidae. Proceedings of the California Academy of Sciences Ser. 2 :850-896
 * Pires de Prado L., R. M. Feitosa, S. Pinzon Triana, J. A. Munoz Gutierrez, G. X. Rousseau, R. Alves Silva, G. M. Siqueira, C. L. Caldas dos Santos, F. Veras Silva, T. Sanches Ranzani da Silva, A. Casadei-Ferreira, R. Rosa da Silva, and J. Andrade-Silva. 2019. An overview of the ant fauna (Hymenoptera: Formicidae) of the state of Maranhao, Brazil. Pap. Avulsos Zool. 59: e20195938.
 * Quiroz Robledo L. N., and J. E. Valenzuela Gonzalez. 1993. Contribucion al conocimiento de la mirmecofauna del estado de Hidalgo, Mexico (Hymenoptera: Formicidae). En: Villavicencio-Nieto (ed) Flora y Fauna del Estado de Hidalgo. Universidad Autónoma de Hidalgo. p. 340-393. ISBN 968-63 40-36-X
 * Quiroz-Robledo, L.N. and J. Valenzuela-Gonzalez. 2007. Distribution of poneromorph ants (Hymenoptera: Formicidae) in the Mexican state of Morelos. Florida Entomologist 90(4):609-615
 * 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.
 * Reynoso-Campos J. J., J. A. Rodriguez-Garza, and M. Vasquez-Bolanos. 2015. Hormigas (Hymenoptera: Formicidae) de la Isla Cozumel, Quintana Roo, Mexico (pp. 27-39). En: Castaño Meneses G., M. Vásquez-Bolaños, J. L. Navarrete-Heredia, G. A. Quiroz-Rocha e I. Alcalá-Martínez (Coords.). Avances de Formicidae de  México.  UNAM,  Universiad  de  Guadalajara, Guadalajara, Jalisco.
 * 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.
 * Ribeiro L. F., R. R. C. Solar, T. G. Sobrinho, D. C. Muscardi, J. H. Schoereder, and A. N. Andersen. 2019. Different trophic groups of arboreal ants show differential responses to resource supplementation in a neotropical savanna. Oecologia 190(2): 433-443.
 * 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.
 * Rosa da Silva R., and B. Cortes Lopes. 1997. Ants (Hymenoptera: Formicidae) from Atlantic rainforest at Santa Catarina Island, Brazil: two years of sampling. Rev. Biol. Trop. 45(4): 1641-1648.
 * Rosumek, F.B., M.A. Ulyssea, B.C. Lopes, J. Steiner. 2008. Formigas de solo e de bromélias em uma área de Mata Atlântica, Ilha de Santa Catarina, sul do Brasil: Levantamento de espécies e novos registros. Revista Biotemas 21(4):81-89.
 * Salazar F., and D. A. Donoso. 2013. New ant (Hymenoptera: Formicidae) records for Ecuador deposited at the Carl Rettenmeyer ant collection in the QCAZ Museum. Boletín Tecnico 11, Serie Zoológica 8-9: 151  177.
 * Sanchez-Pena S. R. 2010. Some fungus-growing ants (Hymenoptera: Formicidae) from northeastern Mexico. Florida Entomologist 93(4): 501-504.
 * Santos J. C., and Del-Claro K. 2001. Interactions between ants, herbivores and extrafloral nectaries in Tocoyena formosa (Cham & Schlechtd.) K. Schum. (Rubiaceae) in cerrado vegetation. Rev. Bras. Zoociencias Juiz de Fora 3(1): 77-92.
 * Santos P. P., A. Vasconcelos, B. Jahyny, and J. H. C. Delabie. 2010. Ant fauna (Hymenoptera, Formicidae) associated to arboreal nests of Nasutitermes spp. (Isoptera, Termitidae) in a cacao plantation in southeastern Bahia, Brazil. Revista Brasileira de Entomologia 54(3): 450-454.
 * Santos-Junior L. C., J. M. Saraiva, R. Silvestre, and W. F. Antonialli-Junior. 2014. Evaluation of Insects that Exploit Temporary Protein Resources Emphasizing the Action of Ants (Hymenoptera, Formicidae) in a Neotropical Semi-deciduous Forest. Sociobiology 61(1): 43-51
 * Santschi F. 1921. Ponerinae, Dorylinae et quelques autres formicides néotropiques. Bulletin de la Société Vaudoise des Sciences Naturelles 54: 81-103.
 * 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.
 * Silvestre R., C. R. F. Brandão, and R. R. Silva da 2003. Grupos funcionales de hormigas: el caso de los gremios del cerrado. Pp. 113-148 in: Fernández, F. (ed.) 2003. Introducción a las hormigas de la región Neotropical. Bogotá: Instituto de Investigación de Recursos Biológicos Alexander von Humboldt, xxvi + 424 pp.
 * Smith M. A., W. Hallwachs, D. H. Janzen. 2014. Diversity and phylogenetic community structure of ants along a Costa Rican elevational gradient. Ecography 37(8): 720-731.
 * Trunzer, B., J. Heinze and B. Holldobler. 1998. Cooperative colony founding and experimental primary polygyny in the ponerine ant Pachycondyla villosa. Insectes Sociaux 45:267-276
 * Ulyssea M. A., and C. R. F. Brandao. 2013. Ant species (Hymenoptera, Formicidae) from the seasonally dry tropical forest of northeastern Brazil: a compilation from field surveys in Bahia and literature records. Revista Brasileira de Entomologia 57(2): 217224.
 * 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.
 * Ulysséa M. A., C. R. F. Brandão. 2013. Ant species (Hymenoptera, Formicidae) from the seasonally dry tropical forest of northeastern Brazil: a compilation from field surveys in Bahia and literature records. Revista Brasileira de Entomologia 57(2): 217-224.
 * Valenzuela-Gonzalez, J., A. Lopez Mendez, and A. Garcia Ballinas. 1994. Ciclo de actividad y provisionamiento de Pachycondyla villosa (Hymenoptera, Formicidae) en agroecosistemas cacaoteros del Soconusco, Chiapas, Mexico. Folia Entomologica 91: 9-21.
 * Vargas A. B., A. J. Mayhé-Nunes, J. M. Queroz, G. O. Souza, and E. F. Ramos. 2007. Effects of Environmental Factors on the Ant Fauna of Restinga Community in Rio de Janeiro, Brazil. Neotropical Entomology 36(1): 028-037
 * Vásquez-Bolaños M. 2011. Lista de especies de hormigas (Hymenoptera: Formicidae) para México. Dugesiana 18: 95-133
 * Wheeler W. M. 1905. The ants of the Bahamas, with a list of the known West Indian species. Bulletin of the American Museum of Natural History 21: 79-135.
 * Wheeler W. M. 1908. The ants of Porto Rico and the Virgin Islands. Bulletin of the American Museum of Natural History 24: 117-158.
 * Wheeler W. M. 1908. The ants of Texas, New Mexico and Arizona. (Part I.). Bulletin of the American Museum of Natural History 24: 399-485.
 * Wheeler, G.C. and J. Wheeler. 1985. A checklist of Texas ants. Prairie Naturalist 17:49-64.
 * Wild A. L. 2002. The genus Pachycondyla (Hymenoptera: Formicidae) in Paraguay. Boletín del Museo Nacional de Historia Natural del Paraguay 14: 1-18
 * Wild, A. L. "A catalogue of the ants of Paraguay (Hymenoptera: Formicidae)." Zootaxa 1622 (2007): 1-55.
 * da Silva, R.R., C.R.F. Brandao, and R. Silvestre. 2004. Similarity Between Cerrado Localities in Central and Southeastern Brazil Based on the Dry Season Bait Visitors Ant Fauna. Studies on Neotropical Fauna and Environment 39(3):191-199.
 * 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 Miranda M., V. Barbosa de Andrade, G. D. Vieira Marques, and V. S. Sul Moreira. 2006. Ant fauna (Formicidae, Hymenoptera) in an urban fragment area of semideciduous mesophitic forest. Zoociencias 8(1): 49-54.
 * do Nascimento, I.C. 2006. Fenologia dos Voos de Acasalamento em Formigas Tropicais