Pachycondyla harpax

A common species with a wide range, it can be found in North, Central, and South America.

Identification
Mackay and Mackay (2010): Pachycondyla harpax is the most common species in the New World and is also widely distributed. The worker can be separated from most other species in the genus as it lacks the malar carina and the eye is small but is located less than one maximum diameter from the anterior edge of the head (side view). The pronotal carina is poorly developed but forms a shiny raised line (usually), the metanotal suture is absent on the dorsum of the mesosoma and the petiole is rectangular shaped with a distinct dorsal face and the posterior lateral margin forms a sharp carina, which is barely evident as it passes to the anterior edge of the petiole. The anterior face of the postpetiole forms a right angle with the dorsal face, which may be slightly concave.

The sharply angulate anterior face of the postpetiole of P. harpax could cause confusion with Neoponera procidua, but they can be easily separated as the metanotal suture of P. procidua breaks the sculpture and the petiole of P. procidua has a vertical anterior face and broadly rounded posterior face, which meet near the anterior edge of the apex of the petiole in a sharp angle. The basically rectangular-shaped petiole of P. harpax could cause confusion with Pachycondyla crassinoda, Pachycondyla striata,  and Pachycondyla impressa. Pachycondyla harpax can be easily separated from all three of these species by its much smaller size (the workers of the other three species are usually at least 10 mm in total length). Pachycondyla harpax can be separated from smaller specimens of P. impressa, as the pronotal carina is present, at least as a shiny raised line, not nearly rounded as in P. impressa. It lacks the clypeal carina found in Pachycondyla lenis of Brasil and although the pronotal carina is poorly developed, it is still obvious (nearly always) at least as a shiny raised line against the dull sculpture, not completely absent as in P. lenis.

John Longino refers to specimens which lack the pronotal carina as JTL-003. All specimens from the southern Pacific lowlands (Osa Peninsula) lack the carina, everywhere else in Costa Rica they have it (Longino, pers. comm.). They are not sympatric with the “normal” P. harpax and may represent a separate species. Longino also has a collection of this phenotype from the state of Barinas, Venezuela. We will recognize them as only a minor geographic variant in this revision.

Distribution
Southern United States through South America, Trinidad, Grenada, Jamaica. (Mackay and Mackay 2010)

Distribution based on Regional Taxon Lists
Nearctic Region: United States. Neotropical Region: Bolivia, Brazil, Colombia, Costa Rica, Ecuador, French Guiana, Greater Antilles, Grenada, Guadeloupe, Guatemala, Guyana, Honduras, Mexico, Nicaragua, Panama, Paraguay, Peru, Suriname, Trinidad and Tobago, Venezuela.

Habitat
These ants are found in a wide variety of habitats, ranging from dry forest, late dry season gap forest, urban environments, parks, grassy areas, coffee, cacao and banana plantations, cypress swamps, to oak forests, oak riparian forests, rocky wet quebradas [streams], arid scrub, palm thorn forests, tropical deciduous forests, tropical evergreen forests, second growth tropical forests, transitional bamboo/cloud forest, old growth dry tropical forest, steep rocky forest slopes, ridge forest, wet mountain forests, riparian rain forest, montane evergreen forest, lowland forests, cloud forest and riparian tropical rain forests. This species has been collected in caves near the entrances (Reddell and Cokendolpher, 2001). They occur from sea level to 2000 meters. (Mackay and Mackay 2010)

Biology
Mackay and Mackay (2010): Nests are found in the soil often under stones (Buckley, 1866) or pieces of wood, or in rotten logs, in organic matter on top of rotten logs and under cow dung (Pergande, 1895). Logs may be very large (1+ meter diameter, 2+ meters long); stones may be very heavy (200+ kg). Soils vary from clays to sandy gravel. A nest gyne was found at a soil depth of 50 cms. One nest was found in the abandoned nest of the native fire ant Solenopsis geminata. They can nest in a wide variety of soils ranging from clay to sand. Workers are commonly collected in Winkler extractions and in pitfall traps. Brood was found in nests in July (Guatemala), August (USA) and November (Colombia). Sexuals were found in nests in February (Bolivia), March to June (Texas, Mexico, Ecuador), April (México) and November (Colombia). Alate females have been collected in February (Perú) and April (Texas, Costa Rica). Dealate females have been collected in May (Mexico), June (Mexico, Costa Rica, Colombia), July to September (Mexico, Colombia), August (USA, Ecuador), October (USA, Venezuela, Bolivia) and December (México, Colombia). Loose males have been collected in January (México), February (Ecuador), April (USA), July (México, El Salvador), between January and July (Ecuador, canopy fogging), August (México), October (Panamá) and November (México) and December (México, Costa Rica). The nests have multiple ergatomorphic females (Wheeler, 1900).

Workers are predaceous (Maes, 1989) and forage primarily on the ground. They can be collected in surface traps baited with mealworms or tuna and pitfall traps. A few workers have been collected in baits hanging in the vegetation, especially trees. This species preys on worker termites (Gnathamitermes tubiformans) in northern México (Garcia-Pérez et al., 1977). Workers are alert and fast and normally not very aggressive when the nest is disturbed. Wheeler (1900) describes interesting worker behavior: they fold their antennae and “peep out” of holes and crevices in the soil “like a rat”. Workers produce foam from the tip of the gaster (Maschwitz et al., 1981; Overal, 1987), similar to workers of Pachycondyla striata.

These ants are found in the same areas as Neoponera apicalis and one nest was in the refuse of an Atta mexicana nest. Baena (1993) reported that P. harpax nests together with ants of the genus Leptogenys. Hung and Vinson (1975) describe the male reproductive system.

This species was studied in Texas by Wheeler (1900b), who found colonies under stones or logs on moderately dry hillsides, often in the shade of trees or bushes. This species "has the peculiar habit of folding its antennae and of peeping out of holes and crevices like a rat." Both winged and ergatoid queens occur. In two nests there were several dealate queens and winged queens that lost their wings "at a slight touch." This suggests that at least some winged queens have no dispersal flights. When colonies are combined, there is a period of non-lethal combat that lasts for several hours to several days, followed by the merging of the colonies. Wheeler (1900b) also includes an illustration of the larva.

Mitchell and Pierce (1912) provide notes on P. harpax in Victoria County, in southeastern Texas. They found colonies only in moist sites, such as river bottoms and lake shores. Nests were under logs, in decayed logs, or in the decomposed phloem between the outer bark and the wood. Colonies were small, with a maximum of about 150 workers. Workers were found foraging under cow dung.

Foraging
Workers employee tandem running to recruit to and forage from larger food items. Gruter et al (2018) - Studying wild colonies in Brazil, we for the first time describe tandem running in the ponerine ant Pachycondyla harpax. We asked if foragers perform tandem runs to carbohydrate- (honey) and protein-rich (cheese) food items. Furthermore, we tested whether the speed and success rate of tandem runs depend on the foraging distance. Foragers performed tandem runs to both carbohydrate food sources and protein-rich food items that exceed a certain size. The probability to perform a tandem run and the travelling speed increase with increasing foraging distances, which could help colonies monopolize more distant food sources in a competitive environment. Guiding a recruit to a food source is costly for leaders as ants are similar to 66% faster when travelling alone. If tandem runs break up (similar to 23% of all tandem runs), followers do not usually discover the food source on their own but return to the nest. Our results show that tandem running to food sources is common in P. harpax, but that foragers modify their behaviour according to the type of food and its distance from the nest. Competition with other ants was intense and we discuss how tandem running in P. harpax might help colonies to build-up a critical number of ants at large food items that can then defend the food source against competitors.

Brazil
DaRocha et al. (2015) studied the diversity of ants found in bromeliads of a single large tree of Erythrina, a common cocoa shade tree, at an agricultural research center in Ilhéus, Brazil. Forty-seven species of ants were found in 36 of 52 the bromeliads examined. Bromeliads with suspended soil and those that were larger had higher ant diversity. Pachycondyla harpax was found in 2 different bromeliads and was associated with the suspended soil and litter of the plants.

Associations with other Organisms
Foragers disperse the seeds of Calathea ovandensis (Marantaceae) approximately 75 cm from where they are collected (Horvitz and Schemske, 1986b).

This species is parasitized by the phorid fly (see Wheeler and Wheeler, 1952 for a survey of the literature) as well as

Nomenclature

 *  harpax. Formica harpax Fabricius, 1804: 401 (w.) SOUTH AMERICA. Wheeler, W.M. 1900b: 4, 17 (q.m.l.); Wheeler, G.C. & Wheeler, J. 1952c: 618 (l.). Combination in Pachycondyla: Roger, 1863b: 18. Senior synonym of concinna, dibullana, irina, montezumia (and its junior synonyms amplinoda, orizabana): Brown, 1950e: 247. See also: Mackay & Mackay, 2010: 374.
 * montezumia. Pachycondyla montezumia Smith, F. 1858b: 108 (q.m.) MEXICO. Forel, 1899c: 12 (w.); Wheeler, G.C. & Wheeler, J. 1952c: 618 (l.). Junior synonym of harpax: Roger, 1862c: 288; Roger, 1863b: 18; Dalla Torre, 1893: 34. Revived from synonym as subspecies of harpax: Forel, 1899c: 12; Emery, 1911d: 75. Senior synonym of amplinoda, orizabana: Emery, 1911d: 75. Junior synonym of harpax: Brown, 1950e: 247.
 * amplinoda. Ponera amplinoda Buckley, 1866: 171 (w.) U.S.A. Junior synonym of harpax: Emery, 1895c: 266; of villosa: Forel, 1899c: 14; of montezumia: Emery, 1911d: 75.
 * dibullana. Pachycondyla harpax var. dibullana Forel, 1901f: 347 (w.) COLOMBIA. Forel, 1912c: 39 (q.). Junior synonym of harpax: Brown, 1950e: 247.
 * concinna. Pachycondyla harpax var. concinna Wheeler, W.M. 1925a: 5 (w.q.) BRAZIL. Junior synonym of harpax: Brown, 1950e: 247.
 * irina. Pachycondyla harpax var. irina Wheeler, W.M. 1925a: 5 (w.) GUATEMALA. Junior synonym of harpax: Brown, 1950e: 247.
 * orizabana. Pachycondyla orizabana Norton, 1868c: 8 (w.) MEXICO. [P. orizabana Norton, 1868a: 64. Nomen nudum.] Junior synonym of harpax: Emery, 1892b: 167; of montezumia: Emery, 1911d: 75.

Type Material
Lectotype worker, 3 paralectotype workers, 3 paralectotype females here designated, ; 3 syntypes seen, ; two syntype workers seen,, 3 syntypes workers, , 6 worker syntypes , 5 worker cotypes, 1 female cotype (Mackay and Mackay 2010)

Pachycondyla montezumia

One syntype queen in (syntype male not found). Labelled “Mex. 56/143.” Acc. Reg.: “1856 no. 143. Mexico. Purchased of Cuming. Collected by M. Sallé on and around the volcano of Orizaba, two or three are from St Thomas in the West Indies.”

Worker
Mackay and Mackay (2010): The workers are moderately large (total length up to 10 mm) black ants with dark brown appendages. The mandibles have 9 or 10 teeth; the anterior border of the clypeus is usually convex, although it may be nearly straight or even slightly concave medially. The central region of the clypeus is swollen and may even form a longitudinal carina posteriorly. The region anterior to the eye is without a malar carina, although it may be slightly raised. The eyes are relatively large (0.34 mm maximum diameter) and are separated from the insertion of the mandibles by less than one maximum diameter (side view). The scape extends only slightly past the posterior lateral corner of the head. The pronotal shoulder is nearly always formed into a definite carina, which passes forward on the anterior lateral surface of the pronotum. The metanotal suture is absent on the dorsum of the mesosoma or is only slightly developed. The petiole is rectangular-shaped with the anterior and posterior faces being nearly parallel, the posterior surface forming definite bend with the dorsal surface. Posteriorly the petiole has sharp lateral borders which define the posterior face. The anterior face of the post-petiole may be slightly concave.

Erect hairs are abundant on most surfaces, including the mandibles, clypeus, scapes, dorsum of the head, region near the eyes, ventral surface of the head, dorsum of the mesosoma, petiole and all surfaces of the gaster, the hairs on the tibiae are mostly suberect. Appressed golden pubescence is abundant on most surfaces.

The mandibles are smooth and glossy with scattered punctures, most of the dorsum of the head is covered with dense punctures, as is the mesosoma and petiole, the punctures on the dorsum of the gaster are more scattered and much of the gaster is smooth and glossy.

Queen
Mackay and Mackay (2010): The female is slightly larger (total length over 10 mm) and is similar to the worker except the mesosoma is more massive and some structures, especially the side of the pronotum, mesopleuron and the side of the propodeum, are striate. The metanotal suture is well developed, forming a definite metanotum. The wing has the typical venation of members of the genus.

Erect hairs are abundant on essentially all of the surfaces, including the mandibles, dorsal and ventral surfaces of the head, sides of the head, clypeus, scapes (often longer than the diameter of the scape), mesosoma, petiole and gaster. Fine golden appressed pubescence is present on all surfaces except the mandibles and is especially obvious on the head and gaster.

The mandibles are finely sculptured and mostly smooth and glossy, the head and mesosoma are densely punctured and weakly shining, the gaster is finely punctured and moderately smooth and glossy.

Male
Mackay and Mackay (2010): The male is also a moderate sized specimen (total length about 9 mm) black with brown legs. The eye is large, covering more than half of the side of the head. The pronotal carina, which is well developed in both the worker and female, is lacking. The metanotal suture is well developed, as is the metanotum. The petiole is somewhat rectangular shaped but is definitely narrowed toward the apex, with a rounded node (not developed into a definite dorsal surface). The posterior face of the petiole is not defined by a carina. There is considerable variability in the shape of the subpetiolar process.

Erect hairs are abundant and scattered on most surfaces; the hairs on the tibiae are mostly suberect.

Most surfaces are coriaceous and at least weakly shining, although most of the surface is hidden by appressed golden pubescence.

Etymology
The name of this species may be based on the Latin word harpe, meaning sword and possibly referring to the sting. (Mackay and Mackay 2010)

References based on Global Ant Biodiversity Informatics

 * Achury R., and A.V. Suarez. 2017. Richness and composition of ground-dwelling ants in tropical rainforest and surrounding landscapes in the Colombian Inter-Andean valley. Neotropical Entomology https://doi.org/10.1007/s13744-017-0565-4
 * Ahuatzin D. A., E. J. Corro, A. Aguirre Jaimes, J. E. Valenzuela Gonzalez, R. Machado Feitosa, M. Cezar Ribeiro, J. Carlos Lopez Acosta, R. Coates, W. Dattilo. 2019. Forest cover drives leaf litter ant diversity in primary rainforest remnants within human-modified tropical landscapes. Biodiversity and Conservation 28(5): 1091-1107.
 * 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
 * Alonso L. E., J. Persaud, and A. Williams. 2016. Biodiversity assessment survey of the south Rupununi Savannah, Guyana. BAT Survey Report No.1, 306 pages.
 * 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.
 * Campos R. B. F., J. H. Schoereder, and C. F. Sperber. 2007. Small-scale patch dynamics after disturbance in litter ant communities. Basic and Applied Ecology 8: 3643.
 * Cancino, E.R., D.R. Kasparan, J.M.A. Coronado Blanco, S.N. Myartseva, V.A. Trjapitzin, S.G. Hernandez Aguilar and J. Garcia Jimenez. 2010. Himenópteros de la Reserva El Cielo, Tamaulipas, México. Dugesiana 17(1):53-71
 * 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.
 * Clemes Cardoso D., and J. H. Schoereder. 2014. Biotic and abiotic factors shaping ant (Hymenoptera: Formicidae) assemblages in Brazilian coastal sand dunes: the case of restinga in Santa Catarina. Florida Entomologist 97(4): 1443-1450.
 * Clemes Cardoso D., and M. Passos Cristiano. 2010. Myrmecofauna of the Southern Catarinense Restinga sandy coastal plain: new records of species occurrence for the state of Santa Catarina and Brazil. Sociobiology 55(1b): 229-239.
 * 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.
 * Cokendolpher J.C., Reddell J.R., Taylor S.J, Krejca J.K., Suarez A.V. and Pekins C.E. 2009. Further ants (Hymenoptera: Formicidae) from caves of Texas [Hormigas (Hymenoptera: Formicdae) adicionales de cuevas de Texas]. Texas Memorial Museum Speleological Monographs, 7. Studies on the cave and endogean fauna of North America, V. Pp. 151-168
 * Corassa J. N., I. C. Magistrali, J. C. Moreno, E. B. Cantarelli, and A. Corassa. Effect of formicid granulated baits on non-target ants biodiversity in eucalyptus plantations litter. Comunicata Scientiae 4(1): 35-42.
 * Dattilo W. et al. 2019. MEXICO ANTS: incidence and abundance along the Nearctic-Neotropical interface. Ecology https://doi.org/10.1002/ecy.2944
 * De La Mora, A., and S. M. Philpott. 2010. Wood-nesting ants and their parasites in forests and coffee agroecosystems. Environmental Entomology 39: 1473-1481.
 * De la Mora, A., C. J. Murnen, and S. M. Philpott. 2013. Local and landscape drivers of ant-communities in Neotropical coffee landscapes. Biodiversity and Conservation 22: 871-888.
 * De la Mora, A., G. Perez-Lachaud, J. P.Lachaud, and S. M. Philpott. 2015. Local and landscape drivers of ant parasitism in a coffee landscape. Environmental Entomology 44: 939-950.
 * 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.
 * Delabie J. H. C., R. Céréghino, S. Groc, A. Dejean, M. Gibernau, B. Corbara, and A. Dejean. 2009. Ants as biological indicators of Wayana Amerindian land use in French Guiana. Comptes Rendus Biologies 332(7): 673-684.
 * Dias N. D. S., R. Zanetti, M. S. Santos, M. F. Gomes, V. Peñaflor, S. M. F. Broglio, and J. H. C. Delabie. 2012. The impact of coffee and pasture agriculture on predatory and omnivorous leaf-litter ants. Journal of Insect Science 13:29. Available online: http://www.insectscience.org/13.29
 * Dias N. S., R. Zanetti, M. S. Santos, J. Louzada, and J. H. C. Delabie. 2008. Interaction between forest fragments and adjacent coffee and pasture agroecosystems: responses of the ant communities (Hymenoptera, Formicidae). Iheringia, Sér. Zool., Porto Alegre, 98(1): 136-142.
 * Donoso D. A. 2014. Assembly mechanisms shaping tropical litter ant communities. Ecography 37 doi: 10.1111/j.1600-0587.2013.00253.x
 * Emery C. 1890. Voyage de M. E. Simon au Venezuela (Décembre 1887 - Avril 1888). Formicides. Annales de la Société Entomologique de France (6)10: 55-76.
 * Emery C. 1911. Hymenoptera. Fam. Formicidae. Subfam. Ponerinae. Genera Insectorum 118: 1-125.
 * Favretto M. A., E. Bortolon dos Santos, and C. J. Geuster. 2013. Entomofauna from West of Santa Catarina State, South of Brazil. EntomoBrasilis 6 (1): 42-63.
 * Fernandes I., and J. de Souza. 2018. Dataset of long-term monitoring of ground-dwelling ants (Hymenoptera: Formicidae) in the influence areas of a hydroelectric power plant on the Madeira River in the Amazon Basin. Biodiversity Data Journal 6: e24375.
 * Fernandes, P.R. XXXX. Los hormigas del suelo en Mexico: Diversidad, distribucion e importancia (Hymenoptera: Formicidae).
 * Fernández F. 2008. Subfamilia Ponerinae s.str. Pp. 123-218 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.
 * Fernández F., E. E. Palacio, W. P. MacKay, and E. S. MacKay. 1996. Introducción al estudio de las hormigas (Hymenoptera: Formicidae) de Colombia. Pp. 349-412 in: Andrade M. G., G. Amat García, and F. Fernández. (eds.) 1996. Insectos de Colombia. Estudios escogidos. Bogotá: Academia Colombiana de Ciencias Exactas, Físicas y Naturales, 541 pp
 * Fernández F., and T. M. Arias-Penna. 2008. Las hormigas cazadoras en la región Neotropical. Pp. 3-39 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.
 * Fernández, F. and S. Sendoya. 2004. Lista de las hormigas neotropicales. Biota Colombiana Volume 5, Number 1.
 * Fichaux M., B. Bechade, J. Donald, A. Weyna, J. H. C. Delabie, J. Murienne, C. Baraloto, and J. Orivel. 2019. Habitats shape taxonomic and functional composition of Neotropical ant assemblages. Oecologia 189(2): 501-513.
 * Fleck M. D., E. Bisognin Cantarelli, and F. Granzotto. 2015. Register of new species of ants (Hymenoptera: Formicidae) in Rio Grande do Sul state. Ciencia Florestal, Santa Maria 25(2): 491-499.
 * Forel A. 1905. Miscellanea myrmécologiques II (1905). Ann. Soc. Entomol. Belg. 49: 155-185.
 * Forel A. 1911. Ameisen des Herrn Prof. v. Ihering aus Brasilien (Sao Paulo usw.) nebst einigen anderen aus Südamerika und Afrika (Hym.). Deutsche Entomologische Zeitschrift 1911: 285-312.
 * 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.
 * Galkowski C. 2016. New data on the ants from the Guadeloupe (Hymenoptera, Formicidae). Bull. Soc. Linn. Bordeaux 151, 44(1): 25-36.
 * Gomes E. C. F., G. T. Ribeiro, T. M. S. Souza, and L. Sousa-Souto. 2014. Ant assemblages (Hymenoptera: Formicidae) in three different stages of forest regeneration in a fragment of Atlantic Forest in Sergipe, Brazil. Sociobiology 61(3): 250-257.
 * Goodnight, C. J., and M. L. Goodnight. 1956. Some observations in a tropical rain forest in Chiapas, Mexico. Ecology 37: 139-150.
 * Gove, A. D., J. D. Majer, and V. Rico-Gray. 2009. Ant assemblages in isolated trees are more sensitive to species loss and replacement than their woodland counterparts. Basic and Applied Ecology 10: 187-195.
 * 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.
 * Groc S., J. H. C. Delabie, F. Fernandez, M. Leponce, J. Orivel, R. Silvestre, Heraldo L. Vasconcelos, and A. Dejean. 2013. Leaf-litter ant communities (Hymenoptera: Formicidae) in a pristine Guianese rainforest: stable functional structure versus high species turnover. Myrmecological News 19: 43-51.
 * Groc S., J. Orivel, A. Dejean, J. Martin, M. Etienne, B. Corbara, and J. H. C. Delabie. 2009. Baseline study of the leaf-litter ant fauna in a French Guianese forest. Insect Conservation and Diversity 2: 183-193.
 * Horvitz, C.C. and A.J. Beattie. 1980. Ant Dispersal of Calathea (Marantaceae) Seeds by Carnivorous Ponerines (Formicidae) in a Tropical Rain Forest. American Journal of Botany 67(3):321-326.
 * Horvitz, C.C. and D.W. Schemske. 1986. Seed Dispersal of a Neotropical Myrmecochore: Variation in Removal Rates and Dispersal Distance. Biotropica 18(4):319-323.
 * 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.
 * Iop S., V. M. Caldart, J. A. Lutinski, and F. R. Mello Garcia. 2009. Formigas urbanas da cidade de Xanxerê, Santa Catarina, Brasil. Biotemas 22(2): 55-64.
 * Kaspari M., D. Donoso, J. A. Lucas, T. Zumbusch, and A. D. Kay. 2012. Using nutritional ecology to predict community structure: a field test in Neotropical ants. Ecosphere 3(11): art.93.
 * Kempf W. W. 1961. A survey of the ants of the soil fauna in Surinam (Hymenoptera: Formicidae). Studia Entomologica 4: 481-524.
 * Kempf W. W. 1961. As formigas do gênero Pachycondyla Fr. Smith no Brasil (Hymenoptera: Formicidae). Revista Brasileira de Entomologia 10: 189-204.
 * Kempf, W.W. 1972. Catalago abreviado das formigas da regiao Neotropical (Hym. Formicidae) Studia Entomologica 15(1-4).
 * Kusnezov N. 1953. La fauna mirmecológica de Bolivia. Folia Universitaria. Cochabamba 6: 211-229.
 * Kusnezov N. 1978. Hormigas argentinas: clave para su identificación. Miscelánea. Instituto Miguel Lillo 61:1-147 + 28 pl.
 * Lachaud J. P., P. Cerdan, and G. Perez-Lachaud. 2012. Poneromorph ants associated with parasitoid wasps of the genus Kapala Cameron (Hymenoptera: Eucharitidae) in French Guiana. Psyche doi:10.1155/2012/393486.
 * Lapola D. M., and H. G. Fowler. 2008. Questioning the implementation of habitat corridors: a case study in interior São Paulo using ants as bioindicators. Braz. J. Biol., 68(1): 11-20.
 * LeBrun E. G., 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.
 * 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/
 * Lopes D. T., J. Lopes, I. Cardoso do Nascimento, and J. H. Delabie. 2010. Epigeic ants diversity (Hymenoptera, Formicidae) in three environments in Mata dos Godoy State Park, Londrina, State of Paraná, Brazil. Iheringia, Sér. Zool., Porto Alegre, 100(1): 84-90.
 * 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.
 * 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
 * Marinho C. G. S., R. Zanetti, J. H. C. Delabie, M. N. Schlindwein, and L. de S. Ramos. 2002. Ant (Hymenoptera: Formicidae) Diversity in Eucalyptus (Myrtaceae) Plantations and Cerrado Litter in Minas Gerais, Brazil. Neotropical Entomology 31(2): 187-195.
 * Medina U. C. A., F. Fernandez, and M. G. Andrade-C. 2010. Insectos: escarabajos coprofagos, hormigas y mariposas. Capitulo 6. Pp 197-215. En: Lasso, C. A., J. S. Usma, F. Trujillo y A. Rial (eds.). 2010. Biodiversidad de la cuenca del Orinoco: bases científicas para la identificación de áreas prioritarias para la conservación y uso sostenible de la biodiversidad. Instituto de Investigación de Recursos Biológicos Alexander von Humboldt, WWF Colombia, Fundación Omacha, Fundación La Salle e Instituto de Estudios de la Orinoquia (Universidad Nacional de Colombia). Bogotá, D. C., Colombia.
 * Mentone T. O., E. A. Diniz, C. B. Munhae, O. C. Bueno, and M. S. C. Morini. 2011. Composition of ant fauna (Hymenoptera: Formicidae) at litter in areas of semi-deciduous forest and Eucalyptus spp., in Southeastern Brazil. Biota Neotrop. 11(2): http://www.biotaneotropica.org.br/v11n2/en/abstract?inventory+bn00511022011.
 * Mirmecofauna de la reserva ecologica de San Felipe Bacalar
 * Morrison, L.W. 2002. Long-Term Impacts of an Arthropod-Community Invasion by the Imported Fire Ant, Solenopsis invicta. Ecology 83(8):2337-2345
 * Navarro, E.V. Vergara, H. Echavarria Sanchez, F.J. Serna Cardona. 2007. Hormigas (Hymenoptera: Formicidae) asociadas al arboretum de la Universidad Nacional de Colombia, sede Medellin. Boletín Sociedad Entomológica Aragonesa 40:497-505.
 * 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.
 * Oliveira Mentone T. de, E. A. Diniz, C. de Bortoli Munhae, O. Correa Bueno and M. S. de Castro Morini. 2012. Composition of ant fauna (Hymenoptera: Formicidae) at litter in areas of semi-deciduous forest and Eucalyptus spp., in Southeastern Brazil. Biota Neotrop 11(2): 237-246.
 * Orsolon-Souza G., C. E. L. Esberard, A. J. Mayhe-Nunes, A. B. Vargas, S. Veiga-Ferreira, and E. Folly-Ramos. 2011. Comparison between Winklers extractor and pitfall traps to estimate leaf litter ants richness (Formicidae) at a rainforest site in southest Brazil. Braz. J. Biol. 71(4): 873-880.
 * Pacheco R., H. L. Vasconcelos, S. Groc, G. P. Camacho, and T. L. M. Frizzo. 2013. The importance of remnants of natural vegetation for maintaining ant diversity in Brazilian agricultural landscapes. Biodivers. Conserv. DOI 10.1007/s10531-013-0463-y
 * Pacheco R., and H. L. Vasconcelos. 2007. Invertebrate conservation in urban areas: ants in the Brazilian Cerrado. Landscape and Urban Planning 81: 193199.
 * 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., and J. Valenzuela-Gonzalez. 1995. A comparison of ground ant communities in a tropical rainforest and adjacent grassland in Los Tuxtlas, Veracruz, Mexico. Southwestern Entomologist 20(2): 203-213.
 * Ramos L. de S., C. G. S. Marinho, R. Zanetti, and J. H. C. Delabie. 2003. Impacto de iscas formicidas granuladas sobre a mirmecofauna não-alvo em eucaliptais segundo duas formas de aplicacação / Impact of formicid granulated baits on non-target ants in eucalyptus plantations according to two forms of application. Neotropical Entomology 32(2): 231-237.
 * Ramos L. de S., R. Zanetti, C. G. S. Marinho, J. H. C. Delabie, M. N. Schlindwein, and R. P. Almado. 2004. Impact of mechanical and chemical weedings of Eucalyptus grandis undergrowth on an ant community (Hymenoptera: Formicidae). Rev. Árvore 28(1): 139-146.
 * Ramón G., A. Barragán, and D. A. Donoso. 2013. Can clay banks increase the local ant species richness of a montane forest? In press: Métodos en Ecología y Sistemática 8: 37  53.
 * Reddell J. R., and J. C. Cokendolpher. 2001. Ants (Hymenoptera: Formicidae) from caves of Belize, Mexico, and California and Texas (U.S.A.) Texas. Texas Memorial Museum Speleological Monographs 5: 129-154.
 * Resende J. J., G. M. de M. Santos, I. C. do Nascimento, J. H. C. Delabie, and E. M. da Silva. 2011. Communities of ants (Hymenoptera  Formicidae) in different Atlantic rain forest phytophysionomies. Sociobiology 58(3): 779-799.
 * Riera-Valera M. A., A. J. Pérez-Sánchez, and J. Perozo. 2009. Ants (Hymenoptera: Formicidae) and termites (Termitidae: Isoptera), Moron River basin, Carabobo, Venezuela: Preliminary data. Checklist, Campinas 5(4): 855-859.
 * Rodrigues de Souza D., E. Stingel, L. C. de Almeida, M. A. Lazarini, C. de Bortoli Munhae, A. J. Mayhe-Nunes, O. Correa Bueno, and M. Santina de C. Morini. 2010. Ant Diversity in a Sugarcane Culture without the Use of Straw Burning in Southeast, São Paulo, Brazil. American Journal of Agricultural and Biological Sciences 5 (2): 183-188.
 * Rodriguez E. R., and J. E. Lattke. 2012. Diversidad de hormigas en un gradiente altitudinal de la cordillera de la Costa, Venezuela. Boletín de la Sociedad Entomológica Aragonesa (S.E.A.) 50: 295?304.
 * 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.
 * Salinas P. J. 2010. Catalogue of the ants of the Táchira State, Venezuela, with notes on their biodiversity, biogeography and ecology (Hymenoptera: Formicidae: Amblyioponinae, Ponerinae, Proceratiinae, Myrmicinae, Ecitoninae, Formicinae, Pseudomyrmecinae, Dolichoderinae). Boletín de la SEA 47: 315-328.
 * Santos M. S., J. N. C. Louzada, N. Dias, R. Zanetti, J. H. C. Delabie, and I. C. Nascimento. 2006. Litter ants richness (Hymenoptera, Formicidae) in remnants of a semi-deciduous forest in the Atlantic rain forest, Alto do Rio Grande region, Minas Gerais, Brazil. Iheringia, Sér. Zool., Porto Alegre, 96(1): 95-101.
 * Silva F. H. O., J. H. C. Delabie, G. B. dos Santos, E. Meurer, and M. I. Marques. 2013. Mini-Winkler Extractor and Pitfall Trap as Complementary Methods to Sample Formicidae. Neotrop Entomol 42: 351358.
 * Silva R. R., R. S. Machado Feitosa, and F. Eberhardt. 2007. Reduced ant diversity along a habitat regeneration gradient in the southern Brazilian Atlantic Forest. Forest Ecology and Management 240: 61-69.
 * Silva R.R., and C. R. F. Brandao. 2014. Ecosystem-Wide Morphological Structure of Leaf-Litter Ant Communities along a Tropical Latitudinal Gradient. PLoSONE 9(3): e93049. doi:10.1371/journal.pone.0093049
 * 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.
 * Silvestre R., M. F. Demetrio, and J. H. C. Delabie. 2012. Community Structure of Leaf-Litter Ants in a Neotropical Dry Forest: A Biogeographic Approach to Explain Betadiversity. Psyche doi:10.1155/2012/306925
 * Smith M. R. 1936. A list of the ants of Texas. Journal of the New York Entomological Society 44: 155-170.
 * Sobrinho T. G., and J. H. Schoereder. 2007. Edge and shape effects on ant (Hymenoptera: Formicidae) species richness and composition in forest fragments. Biodivers Conserv 16: 14591470.
 * Sobrinho T., J. H. Schoereder, C. F. Sperber, and M. S. Madureira. 2003. Does fragmentation alter species composition in ant communities (Hymenoptera: Formicidae)? Sociobiology 42(2): 329-342.
 * Sosa-Calvo J. 2007. Ants of the leaf litter of two plateaus in Eastern Suriname. In Alonso, L.E. and J.H. Mol (eds.). 2007. A rapid biological assessment of the Lely and Nassau plateaus, Suriname (with additional information on the Brownsberg Plateau). RAP Bulletin of Biological Assessment 43. Conservation International, Arlington, VA, USA.
 * Suguituru S. S., D. R. de Souza, C. de Bortoli Munhae, R. Pacheco, and M. S. de Castro Morini. 2011. Diversidade e riqueza de formigas (Hymenoptera: Formicidae) em remanescentes de Mata Atlântica na Bacia Hidrográfica do Alto Tietê, SP. Biota Neotrop. 13(2): 141-152.
 * Suguituru S. S., M. Santina de Castro Morini, R. M. Feitosa, and R. Rosa da Silva. 2015. Formigas do Alto Tiete. Canal 6 Editora 458 pages
 * Suguituru S. S., R. Rosa Silva, D. R. de Souza, C. de Bortoli Munhae, and M. Santina de Castro Morini. Ant community richness and composition across a gradient from Eucalyptus plantations to secondary Atlantic Forest. Biota Neotrop. 11(1): 369-376.
 * Theunis, L., M. Gilbert, Y. Roisin and M. Leponce. 2005. Spatial structure of litter-dwelling ant distribution in a subtropical dry forest. Insectes Sociaux. 52:366377.
 * Ulloa Chacon P., M. L. Baena, J. Bustos, R. C. Aldana, J. A. Aldana, and M. A. Gamboa. 1996. Fauna de hormigas del departamento del Valle del Cauca (Colombia). Pp. 413-451. In Andrade-C M. G., G. Amat Garcia, and F. Fernandez. Insectoss de Colombia, estudios escogidos.
 * 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.
 * Valdes-Rodriguez S., P. Chacon de Ulloa, and I. Armbrecht. 2014. Soil ant species in Gorgona Island, Colombian Pacific. Rev. Biol. Trop. 62 (1): 265-276.
 * Varela-Hernandez, F., M. Rocha-Ortega, W. P. MacKay, and R. W. Jones. 2016. Lista preliminar de las hormigas (Hymenoptera: Formicidae) del estado de Queretaro, Mexico. Pages 429-435 in . W. Jones., and V. Serrano-Cardenas, editors. Historia Natural de Queretaro. Universidad Autonoma de Queretaro, Mexico.
 * 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
 * Vasconcelos, H.L. and J.M.S. Vilhena. 2006. Species turnover and vertical partitioning of ant assemblages in the Brazilian Amazon: A comparison of forests and savannas. Biotropica 38(1):100-106.
 * Vasconcelos, H.L., J.M.S. Vilhena, W.E. Magnusson and A.L.K.M. Albernaz. 2006. Long-term effects of forest fragmentation on Amazonian ant communities. Journal of Biogeography 33:1348-1356
 * Vasquez-Bolanos M. 2011. Checklist of the ants (Hymenoptera: Formicidae) from Mexico. Dugesiana 18(1): 95-133.
 * Vásquez-Bolaños M. 2011. Lista de especies de hormigas (Hymenoptera: Formicidae) para México. Dugesiana 18: 95-133
 * Weber N. A. 1938. New ants from stomachs of Bufo marinus L. and Typhlops reticulatus (L). Ann. Entomol. Soc. Am. 31: 207-210.
 * Weber N. A. 1938. The food of the giant toad, Bufo marinus (L.), in Trinidad and British Guiana with special reference to the ants. Annals of the Entomological Society of America 31: 499-503.
 * Wetterer J. K. 2016. Geographic range of Pachycondyla harpax (Fabricius) (Hymenoptera: Formicidae). Sociobiology 63(1): 623-627.
 * Wheeler W. M. 1900. A study of some Texan Ponerinae. Biological Bulletin (Woods Hole). 2: 1-31.
 * Wheeler W. M. 1901. The compound and mixed nests of American ants. Part II. The known cases of social symbiosis among American ants. American Naturalist. 35: 513-539.
 * Wheeler W. M. 1907. A collection of ants from British Honduras. Bulletin of the American Museum of Natural History 23: 271-277.
 * 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 W. M. 1909. Ants collected by Prof. F. Silvestri in Mexico. Bollettino del Laboratorio di Zoologia Generale e Agraria della Reale Scuola Superiore d'Agricoltura. Portici 3: 228-238.
 * Wheeler W. M. 1922. The ants of Trinidad. American Museum Novitates 45: 1-16.
 * Wheeler W. M. 1925. Neotropical ants in the collections of the Royal Museum of Stockholm. Arkiv för Zoologi 17A(8): 1-55.
 * Wheeler, G.C. and J. Wheeler. 1985. A checklist of Texas ants. Prairie Naturalist 17:49-64.
 * Wheeler, William Morton. 1911. Additions to the Ant-Fauna of Jamaica. Bulletin American Museum of Natural History. 30:21-29.
 * Wheeler, William Morton. 1916. Ants Collected in Trinidad by Professor Roland Thaxter, Mr. F. W. Urich, and Others. Bulletin of the Museum of Comparitive Zoology at Harvard University. 40(8):322-330
 * Wild A. L. 2002. The genus Pachycondyla (Hymenoptera: Formicidae) in Paraguay. Boletín del Museo Nacional de Historia Natural del Paraguay 14: 1-18
 * Yanoviak S. P., S, M. Berghoff, K. E. Linsenmair, and G. Zotz. 2011. Effects of an Epiphytic Orchid on Arboreal Ant Community Structure in Panama. Biotropica 43(6): 731-737.
 * da Silva R. R., and R. Silvestre. 2000. Diversidade de formigas (Hymenoptera: Formicidae) em Seara, oeste de Santa Catarina. Biotemas 13(2): 85-105.
 * da Silva de Oliveira A. B., and F. A. Schmidt. 2019. Ant assemblages of Brazil nut trees Bertholletia excelsa in forest and pasture habitats in the Southwestern Brazilian Amazon. Biodiversity and Conservation 28(2): 329-344.
 * da Silva, R.R. and R. Silvestre. 2004. Riqueza da fauna de formigas (Hymenoptera: Formicidae) que habita as camadas superficiais do solo em Seara, Santa Catarina. Papéis Avulsos de Zoologia (São Paulo) 44(1): 1-11
 * 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 Souza D. R., S. G. dos Santos, C. de B. Munhae, and M. S. de C. Morini. 2012. Diversity of Epigeal Ants (Hymenoptera: Formicidae) in Urban Areas of Alto Tietê. Sociobiology 59(3): 703-117.
 * de Souza J. L. P., F. B. Baccarob, V. L. Landeirob, E. Franklinc, and W. E. Magnussonc. 2012. Trade-offs between complementarity and redundancy in the use of different sampling techniques for ground-dwelling ant assemblages. Applied Soil Ecology 56: 63 73.
 * do Nascimento, I.C. 2006. Fenologia dos Voos de Acasalamento em Formigas Tropicais
 * dos Santos Bastos A. H., and A. Y. Harada. 2011. Leaf-litter amount as a factor in the structure of a ponerine ants community (Hymenoptera, Formicidae, Ponerinae) in an eastern Amazonian rainforest, Brazil