Neoponera villosa

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

Identification
From Mackay and Mackay (2010): Pachycondyla villosa vies with Pachycondyla harpax for being the most common and widely distributed member of the genus in the New World. Pachycondyla villosa is much larger than P. harpax (total length of  the worker, female and male of P. harpax < 10 mm). Workers of P. villosa can be easily confused with Pachycondyla holcotyle, Pachycondyla  theresiae,  and Pachycondyla foetida. The side of the petiole of P. 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), P. foetida is approximately the same size. Pachycondyla foetida also has horizontal striae on the posterior face of the petiole, which are lacking in P. villosa. Pachycondyla villosa could also misidentified as Pachycondyla bugabensis, as both lack striae on the sides and posterior face of the petiole. It is much larger (total length of workers of P. bugabensis is < 10 mm) and the legs are dark red (ferrugineous red in color in P. 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 Pachycondyla 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 Pachycondyla commutata, Pachycondyla laevigata  and Pachycondyla marginata. Pachycondyla fisheri is a similar size, but lacks the preocular and pronotal carinae.

It is similar to the closely related Pachycondyla curvinodis and Pachycondyla 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 P. villosa and P. inversa.

The males of three common species are very similar: P. villosa, Pachycondyla striata and P. impressa. Pachycondyla 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 P. 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 P. curvinodis and P. inversa are unknown, but it could be expected that the anterior faces of the petioles of these two species would be concave.

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
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).

Pachycondyla 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. Pachycondyla 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).

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 P. 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). Pachycondyla 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 Pachycondyla 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. 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.

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.

Queen
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.

Type Material
worker seen, was compared to type by Brown, MCZC (Mackay and Mackay 2010)

Type Locality Information
Central America; México; Brasil, Villa Nova; Brasil, Rio Constantia (Mackay and Mackay 2010)

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)

Additional References

 * 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.


 * 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.


 * [[Media: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.pdf|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.]]


 * Fresneau, D. 1985. Individual foraging and path fidelity in a ponerine ant. Insectes Sociaux 32:109-116.


 * 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.




 * 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.


 * 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.


 * 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.


 * 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.


 * 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, 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.
 * 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)