| Neoponera villosa|
A common species that is found in a variety of habitats.
|At a Glance||• Polygynous|
- 1 Identification
- 2 Distribution
- 3 Biology
- 4 Castes
- 5 Nomenclature
- 6 References
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.
Keys including this Species
- Key to Neoponera foetida species complex: Males
- Key to Neoponera foetida species complex: Queens
- Key to Neoponera foetida species complex: Workers
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.
Check distribution from AntMaps.
Distribution based on specimens
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)
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).
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.
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).
The following information is derived from Barry Bolton's New General Catalogue, a catalogue of the world's ants.
- 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.
Unless otherwise noted the text for the remainder of this section is reported from the publication that includes the original description.
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.
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.
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.
Central America; México; Brasil, Villa Nova; Brasil, Rio Constantia. Worker seen, was compared to type by Brown, MCZC (Mackay and Mackay 2010)
Holotype worker in The Natural History Museum. Labelled “Rio. Constancia.”
A specimen in The Natural History Museum previously indicated as type is labelled “Ega. 58/6” (= Brazil) and therefore is not type-material. A specimen in Oxford University Museum of Natural History has “pedunculata” on a label but no other data. All specimens represent a single species.
Holotype male in The Natural History Museum. Labelled “Brazil. Villa Nova.” There is another male in Oxford University Museum of Natural History, mounted as the holotype but without the correct data labels.
Unless otherwise noted the text for the remainder of this section is reported from the publication that includes the original description.
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)
- Bolton, B. 1995b. A new general catalogue of the ants of the world. Cambridge, Mass.: Harvard University Press, 504 pp. (page 311, Combination in Pachycondyla; revived combination)
- Caetano, F. H. 1988. Anatomia, histologia e histoquímica do sistema digestivo e excretor de operárias de formigas (Hymenoptera, Formicidae). Naturalia 13: 129-174.
- Camargo Mathias, M. I. and F. H. Caetano. 1995a. Trophic eggs in workers of Neoponera villosa ants (Hymenoptera: Ponerinae). Journal of Advanced Zoology 16:62-66.
- Camargo Mathias, M. I. and F. Caetano. 1995b. Corpora allata in females of Neoponera villosa ants (Hymenoptera: Formicidae) - relations with ovarian development. Sociobiology 26:283-289.
- Caperucci, D. and M. Camargo-Mathias. 2006. Lipids in oocytes of ants Neoponera villosa (Hymenoptera: Formicidae). Sociobiology 47:531-541.
- DaRocha, W. D., S. P. Ribeiro, F. S. Neves, G. W. Fernandes, M. Leponce, and J. H. C. Delabie. 2015. How does bromeliad distribution structure the arboreal ant assemblage (Hymenoptera: Formicidae) on a single tree in a Brazilian Atlantic forest agroecosystem? Myrmecological News. 21:83-92.
- Dejean, A. 1990 Influence de l’environnement pre-imaginal et précoce dans le choix du site de nidification de Pachycondyla (= Neoponera) villosa (Fabr) (Formicidae, Ponerinae). Behavioral Processes 21:107-125.
- Dejean, A. and Corbara, B. 1990b. Predatory behavior of a Neotropical arboricolous ant - Pachycondyla villosa (Formicidae, Ponerinae). Sociobiology 17:271-286.
- Dejean, A. and B. Corbara. 1998. Study of different foraging paths of the predatory Neotropical ponerine ant Pachycondyla (= Neoponera) villosa (Hymenoptera: Formicidae). Sociobiology 32:409-426.
- Dejean, A. and I. Olmsted. 1997. Ecological studies on Aechmea bracteata (Swartz) (Bromeliaceae) Journal of Natural History 31:1313-1334.
- Dejean, A., B. Corbara and J. Oliva-Rivera. 1990 Mise en evidence d'une forme d'apprentissage dans le comportement de capture des proies chez Pachycondyla (= Neoponera) villosa (Formicidae, Ponerinae). Behaviour 115:175-187.
- Dejean, A., I. Olmsted and J. Camal. 1992. Interaction between Atta cephalotes and arboreal ants in the Biosphere Reserve Sian Ka'an (Quintana Roo, Mexico): efficient protection of the trees (Hymenoptera: Formicidae). Sociobiology 20:57-76.
- Dejean, A., I. Olmsted and R. Snelling. 1995. Tree-epiphyte-ant relationships in the low inundated forest of Sian Ka'an Biosphere Reserve, Quintana Roo, Mexico. Biotropica 27:57-70.
- De Medeiros, M. A., Fowler, H. G. and Bueno, O.C. 1995. Ant (Hym., Formicidae) mosaic stability in Bahian cocoa plantations: implications for management. Journal of Applied Entomology 119:411-414.
- D’Ettorre, P., K. Kellner, J. Delabie and J. Heinze. 2005. Number of queens in founding associations of the ponerine ant Pachycondyla villosa. Insectes Sociaux 52:327-332.
- Emery, C. 1901b. Notes sur les sous-familles des Dorylines et Ponérines (Famille des Formicides). Ann. Soc. Entomol. Belg. 45: 32-54 (page 43, Combination in Neoponera)
- Fabricius, J. C. 1804. Systema Piezatorum secundum ordines, genera, species, adjectis synonymis, locis, observationibus, descriptionibus. Brunswick: C. Reichard, xiv + 15-439 + 30 pp. (page 409, worker described)
- Fernandes, I.O., De Oliveira, M.L. & Delabie, J.H.C. 2014. Description of two new species in the Neotropical Pachycondyla foetida complex (Hymenoptera: Formicidae: Ponerinae) and taxonomic notes on the genus. Myrmecological News 19, 133-163.
- Flores-Maldonado, K. Y.; Phillips, S. A., Jr.; Sánchez-Ramos, G. 1999. The myrmecofauna (Hymenoptera: Formicidae) along an altitudinal gradient in the Sierra Madre Oriental of northeastern Mexico. Southwest. Nat. 44: 457-461 (page 457, Record in Mexico)
- Fresneau, D. 1985. Individual foraging and path fidelity in a ponerine ant. Insectes Sociaux 32:109-116.
- Gallardo, A. 1918c. Las hormigas de la República Argentina. Subfamilia Ponerinas. An. Mus. Nac. Hist. Nat. B. Aires 30: 1-112 (page 56, see also)
- García-Pérez, J. A. 1989. Estrategia del comportamiento de apaciguamiento de Pachycondyla villosa (Hymenoptera: Ponerinae) durante un encuentro con los raids de Eciton burchelli en la naturaleza. XXIV Congreso Nacional de Entomología [México], p. 136.
- Heinze, J., B. Trunzer, P. Oliveira and B. Hölldobler. 1996. Regulation of reproduction in the Neotropical ponerine ant, Pachycondyla villosa. Journal of Insect Behavior 9:441-450.
- Hölldobler, B. 1985. Liquid food transmission and antennation signals in ponerine ants. Israel Journal of Entomology 19:89-99.
- Kellner, K., A. Trindl., J. Heinze and P. D’Ettorre. 2007. Polygyny and polyandry in small ant societies. Molecular Ecology 16:2363-2369.
- Kolmer, K. and J. Heinze. 2000b. Rank orders and division of labour among unrelated co-founding ant queens. Proceedings of the Royal Society of London B 267:1729-1734.
- Leopoldo Heitor Henrique, Karim Christina Scopinho Furquim & Maria Izabel Camargo Mathias. 2011. Histochemistry and Protein Profile of The Venom Glands of Workers of Neoponera villosa Ants (Hymenoptera: Ponerinae). Sociobiology 58: 749-768.
- Lucas, C.; Fresneau, D.; Kolmer, K.; Heinze, J.; Delabie, J. H. C.; Pho, D. B. 2002. A multidisciplinary approach to discriminating different taxa in the species complex Pachycondyla villosa (Formicidae). Biol. J. Linn. Soc. 75: 249-259 (page 249, see also, subversa as provisional species)
- Mackay, W. P., and E. E. Mackay 2010. The Systematics and Biology of the New World Ants of the Genus Pachycondyla (Hymenoptera: Formicidae). Edwin Mellon Press, Lewiston. Information from this publication is used with permission from the authors.
- Maes, J.M. 1989. Catálogo de los insectos controladores Biológicas en Nicaragua. Volumen I. Insectos depredadores (Primera parte). Revista Nicaraguense de Entomología 8:1-106.
- Majer, J. D., J. H. Delabie and M. Smith. 1994. Arboreal ant community patterns in Brazilian cocoa farms. Biotropica 26:73-83.
- Mariano, C., S. Pompolo and J. Delabie. 2000. Citogenética das espécies gêmeas e simpátricas Pachycondyla villosa e Pachycondyla sp. ‘inversa’ (Ponerinae). Naturalia 24:215-217.
- Mayr, G. 1862. Myrmecologische Studien. Verh. K-K. Zool.-Bot. Ges. Wien 12: 649-776 (page 720, Combination in Pachycondyla)
- McIver, J. and G. Stonedahl. 1993. Myrmecomorphy: Morphological and behavioral mimicry of ants. Annual Review of Entomology 38:351-377.
- Paul, J. and F. Roces. 2003. Fluid intake rates in ants correlate with their feeding habits. Journal of Insect Physiology 49:347-357.
- Pérez-Bautista, M., J.P. Lachaud and D. Fresneau. 1985. La división del trabajo en la hormiga primitiva Neoponera villosa (Hymenoptera: Formicidae). Folia Entomológica Mexicana 65:119-130.
- Petralia, R. S.; Vinson, S. B. 1980 . Comparative anatomy of the ventral region of ant larvae, and its relation to feeding behavior. Psyche (Camb.) 86: 375-394 (page 378, larva described)
- Quiroz-Robledo, L. and J. Valenzuela-González. 1995. A comparison of ground ant communities in a tropical rainforest and adjacent grassland in Los Tuxtlas, Veracruz, Mexico. Southwestern Entomologist 20:203-213.
- Reddell, J. and J. Cokendolpher. 2001 Ants (Hymenoptera: Formicidae) from the caves of Belize, Mexico and California and Texas (U.S.A) Texas Memorial Museum, Speleological Monographs 5:129-154.
- Reiskind, J. 1977. Ant-mimicry in Panamian clubionid and salticid spiders (Araneae: Clubionidae and Salticidae). Biotropica 9:1-8.
- Roger, J. 1861a. Die Ponera-artigen Ameisen (Schluss). Berl. Entomol. Z. 5: 1-54 (page 2, worker, queen described)
- Roger, J. 1861a. Die Ponera-artigen Ameisen (Schluss). Berl. Entomol. Z. 5: 1-54 (page 1, senior synonym of bicolor, pedunculata and pilosa)
- Roger, J. 1862c. Synonymische Bemerkungen. 1. Ueber Formiciden. Berl. Entomol. Z. 6: 283-297 (page 288, senior synonym of bicolor, pedunculata and pilosa)
- Roth, D.S., I. Perfecto and B. Rathcke. 1994 The effects of management systems on ground-foraging ant diversity in Costa Rica. Ecological Applications 4:423-436.
- Schmidt, C.A. & Shattuck, S.O. 2014. The higher classification of the ant subfamily Ponerinae (Hymenoptera: Formicidae), with a review of ponerine ecology and behavior. Zootaxa. 3817, 1–242 (doi:10.11646/zootaxa.3817.1.1)
- Trunzer, B., J. Heinze and B. Hölldobler. 1998. Cooperative colony founding and experimental primary polygyny in the ponerine ant Pachycondyla villosa. Insectes Sociaux 45:267-276.
- Trunzer, B., J. Heinze and B. Hölldobler. 1999. Social status and reproductive success in queenless ant colonies. Behaviour 136:1093-1105.
- Wheeler, G. C.; Wheeler, J. 1952c. The ant larvae of the subfamily Ponerinae - Part II. Am. Midl. Nat. 48: 604-672 (page 615, larva described)
- Wheeler, W. M. 1908h. The ants of Texas, New Mexico and Arizona. (Part I.). Bull. Am. Mus. Nat. Hist. 24: 399-485 (page 404, see also)
- Wheeler, W. 1942. Studies of Neotropical ant-plants and their ants. Bulletin of the Museum of Comparative Zoology 90:1-262 + 57 plates.
- Whittaker, P. L. 1984. The insect fauna of mistletoe (Phoradendron tomentosum, Loranthaceae) in southern Texas. Southwestern Naturalist 29:435-444.
- Wild, A. 2002. The genus Pachycondyla (Hymenoptera: Formicidae) in Paraguay. Boletín del Museo Nacional de Historia Natural de Paraguay 14:1-18.
- Wild, A. 2005. Taxonomic revision of the Pachycondyla apicalis species complex (Hymenoptera: Formicidae). Zootaxa 834:1-25.
- Zara, F., A. Cabrera, K. Jaffé and F. Caetano. 2002. Chemical analysis of different larval instars and tissues of Pachycondyla (= Neo-ponera) villosa (Hymenoptera: Formicidae, Ponerinae). Sociobiology 39:89-101.
- Zara, F., F. Caetano, A. Cabrera and K. Jaffé. 2003. Ultrastructure of last larval instar fat body cells of Pachycondyla (= Neoponera) villosa (Formicidae: Ponerinae): cytochemical and chemical analysis. Animal Biology 53:1-16.