Pachycondyla impressa

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Pachycondyla impressa
Scientific classification
Kingdom: Animalia
Phylum: Arthropoda
Class: Insecta
Order: Hymenoptera
Family: Formicidae
Subfamily: Ponerinae
Tribe: Ponerini
Genus: Pachycondyla
Species: P. impressa
Binomial name
Pachycondyla impressa
(Roger, 1861)

Pachycondyla impressa casent0178689 profile 1.jpg

Pachycondyla impressa casent0178689 dorsal 1.jpg

Specimen labels

Synonyms

A fairly common species within its range and preferred habitats.

Identification

From Mackay and Mackay (2010): The large size of P. impressa would separate it from most of the other species of Pachycondyla. Pachycondyla impressa could be easily separated from other large species such as Neoponera commutata and Neoponera laevigata by the dull surfaces (P. commutata and P. laevigata are mostly shiny), from Pachycondyla crassinoda by the lack of the two sharp lateral processes on the pygidium (which are present in P. crassinoda) and from Neoponera villosa and Pachycondyla striata by the relatively rounded pronotal shoulders (formed into distinct sharp carinae in N. villosa and P. striata). The lack of a metanotal suture in the smaller workers of P. impressa could cause confusion with the smaller Pachycondyla harpax (total length of worker less than 10 mm). It can be separated as the carina on the pronotal shoulder of P. harpax is relatively sharp, but not greatly raised from the surface (absent in P. impressa). Additionally the middle of the pygidium is concave in P. impressa, not rounded as in P. harpax. Some small isolated specimens are essentially impossible to separate into one species or the other. Pachycondyla impressa is superficially similar to the rare Colombian Pachycondyla fuscoatra; it can be easily separated by the depressed pygidium and the striae on the side of the pygidium, both of which are absent in P. fuscoatra. It is also slightly smaller and the posterior lateral edges of the petiole are rounded, not sharply marginate as in P. fuscoatra.

Pachycondyla impressa workers are nearly identical to those of Pachycondyla lattkei. It differs in that the dorsum of the gaster is dull and sculptured, not smooth and glossy as in P. lattkei. There are exceptions, especially in specimens from Ecuador, which have a smooth gaster, but are otherwise identical to P. impressa. The pygidium is strongly depressed in P. impressa, but rounded and not depressed in P. lattkei. The lobes on the metasternal process of P. impressa are often relatively close (usually less than 0.3 mm between the tips) and a pair of internal teeth is nearly always present and if the teeth are poorly developed, at least a swollen region is present, both of which are absent in P. lattkei.

The workers and females of Pachycondyla purpurascens from Costa Rica and Pachycondyla inca from Perú and Bolivia are nearly identical to P. impressa, except the clypeuses are slightly longer and less emarginate along the medial border. The male of P. impressa is are also very similar to those of P. inca and P. striata. The relatively small ocelli, the large size and coarse rugae along the posterior lateral edge of the petiole would separate the males of P. impressa from most of the others. The concave anterior margin of clypeus will separate P. impressa from the very similar P. lattkei. The single examined male of P. impressa was not associated with workers, but a process of elimination and the similarity of this species with the closely related P. lattkei, are strong evidence that this male is a member of P. impressa. An unidentified male near P. impressa was collected in Quincemil (Cuzco), Peru (MCZC). It differs from the male of P. impressa in having rugae on the side of the petiole (similar to P. lattkei) and in having a wide petiole (width 1.4 mm). The anterior border of the clypeus is concave as in P. lattkei. Pachycondyla harpax variety irina is a smaller P. impressa, but the sides of the pronotum are nearly rounded and the pygidium is concave, indicating that it is actually P. impressa. Pachycondyla fuscoatra montana differs from the typical P. impressa in having a slightly more developed pronotal carina. Kempf (1961) synonymized P. fuscoatra var. andicola (which was not seen) and P. fuscoatra variety cearensis, which is actually a synonym of P. inca.

Pachycondyla fuscoatra transversa has transverse striae on the dorsum of the pygidium, which are usually lacking in P. impressa. There is considerable variation in this character and we follow Kempf (1961) in considering this to be a synonym of P. impressa.

Distribution

Central America through central South America, Trinidad. (Mackay and Mackay 2010)

Distribution based on Regional Taxon Lists

Neotropical Region: Bolivia, Brazil, Colombia (type locality), Costa Rica, Ecuador, French Guiana, Guatemala, Guyana, Honduras, Mexico, Panama, Peru, Suriname, Trinidad and Tobago, Venezuela.


Distribution based on AntMaps

AntMapLegend.png

Distribution based on AntWeb specimens

Check data from AntWeb

Habitat

This common species is found in a variety of habitats, ranging from lowland rain forest, at a pasture edge, dry tropical forest, the edge of second growth rain forest, cloud forest, thick wet cloud forest, low montane rain forest, upper montane cloud forest, tropical evergreen forest, a forest slope, montane hardwood forest, cloud forest, a roadside in a sunny spot, a cacao plantation, montane tropical forest, to a guano cave; at elevations ranging from 10 - 2350 meters. (Mackay and Mackay 2010)

Biology

From Mackay and Mackay (2010): Pachycondyla impressa nests in and under rotten logs and under stones. Brood was present in nests in July (Colombia) and August (Costa Rica). Females were collected in nests in January (Panamá) and April (Brasil). Dealate females have been collected in February and March (Costa Rica), April (Panamá), May (Panamá, Perú), June (Panamá), July (Perú) and August (Venezuela). Kempf (1961) mentions isolated females from Brasil were collected in March, May and October.

Individuals have been collected in intercept traps, pitfall traps and foraging on the ground. Workers were extracted from cacao leaf litter and other similar materials. Foragers are attracted to peanut butter baits (Kaspari and Weiser, 2000). One series was collected in the stomach of a Bufo toad. Workers are alert and fast. Sexuals are attracted to lights. They are parasitized by the head decapitating phorid fly Apocephalus (Brown and Feener, 1991).

Sanjuan et al. (2015) - This species is parasitized by the fungal pathogen Ophiocordyceps evansii.

De la Mora et al. (2016) found this species nesting in logs found in coffee farms and forests in the Soconusco region of Chiapas, Mexico. A small percentage of pupae were found to be parasitized by eucharitid wasps.

Fungi

This species is a host for the fungus Ophiocordyceps evansii (a pathogen) (Sanjuan et al., 2015; Shrestha et al., 2017).

Castes

Nomenclature

The following information is derived from Barry Bolton's Online Catalogue of the Ants of the World.

  • impressa. Ponera (Pachycondyla) impressa Roger, 1861a: 6 (w.) COLOMBIA.
    • Type-material: syntype workers (number not stated, “several”).
    • Type-locality: Colombia (no further data)
    • Type-depository: MNHU.
    • Mackay & Mackay, 2010: 395 (q.m.).
    • Combination in Pachycondyla (Pachycondyla): Emery, 1901a: 45;
    • combination in Pachycondyla: Roger, 1863b: 18; Mayr, 1863: 439; Schmidt, C.A. & Shattuck, 2014: 155.
    • Status as species: Roger, 1863b: 18; Mayr, 1863: 439; Emery, 1890a: 58, 71 (in key); Dalla Torre, 1893: 34; Forel, 1905b: 156; Emery, 1911d: 75; Wheeler, W.M. 1916d: 323; Wheeler, W.M. 1922c: 3; Kempf, 1961b: 493; Kempf, 1961c: 195; Kempf, 1964e: 52 (in key); Kempf, 1972a: 174; Bolton, 1995b: 306; Mackay, Mackay, et al. 2008: 194; Mackay & Mackay, 2010: 393 (redescription); Branstetter & Sáenz, 2012: 263; Bezděčková, et al. 2015: 124; Feitosa, 2015c: 99; Fernández & Guerrero, 2019: 541.
    • Senior synonym of andicola: Kempf, 1961c: 195; Kempf, 1972a: 174; Bolton, 1995b: 306; Mackay & Mackay, 2010: 393.
    • Senior synonym of irina: Mackay & Mackay, 2010: 393.
    • Senior synonym of montana: Kempf, 1961c: 195; Kempf, 1972a: 174; Bolton, 1995b: 306; Mackay & Mackay, 2010: 393.
    • Senior synonym of transversa: Kempf, 1961c: 195; Kempf, 1972a: 174; Bolton, 1995b: 306; Mackay & Mackay, 2010: 393.
    • Distribution: Bolivia, Brazil, Colombia, Costa Rica, Ecuador, French Guiana, Guatemala, Guyana, Honduras, Mexico, Nicaragua, Panama, Peru, Trinidad.
  • andicola. Pachycondyla fuscoatra var. andicola Santschi, 1913h: 34 (w.) ECUADOR.
    • Type-material: holotype worker.
    • Type-locality: Ecuador: Santo Domingo de los Colorados (no collector’s name).
    • Type-depository: NHMB.
    • Junior synonym of impressa: Kempf, 1961c: 195; Kempf, 1972a: 174; Bolton, 1995b: 302; Mackay & Mackay, 2010: 393.
  • irina. Pachycondyla harpax var. irina Wheeler, W.M. 1925a: 5 (w.) GUATEMALA, COSTA RICA, COLOMBIA, GUYANA, PERU.
    • Type-material: lectotype worker (by designation of Mackay & Mackay, 2010: 393), 1 paralectotype worker.
    • Type-locality: lectotype Guatemala: Quirigua (W.M. Wheeler); paralectotype with same data; other original syntypes: Guatemala: Escuintla (W.M. Wheeler), Costa Rica: Cartago (W.M. Wheeler), Colombia: Santa Marta (E.M. Gaige), Guyana: Camaria, nr Kartabo (W.M. Wheeler), Peru: El Campamiento, Perene (J.C. Bradley), Peru: Rio Charape (C.H.T. Townsend), and Peru: Llinquipata (N. Holmgren).
    • Type-depository: USNM (lectotype, paralectotype).
    • [Note: Mackay & Mackay, 2010: 374, 393, mention other original syntypes in AMNH, LACM, MCZC, USNM; probably also present in NHRS.]
    • Junior synonym of harpax: Brown, 1950e: 247; Kempf, 1972a: 174; Smith, D.R. 1979: 1340; Bolton, 1995b: 306.
    • Junior synonym of impressa: Mackay & Mackay, 2010: 393.
    • [Note: irina is entered under the synonymy of harpax: Mackay & Mackay, 2010: 374, and also under the synonymy of impressa: Mackay & Mackay, 2010: 393, because of mixed series containing both species. As the lectotype of irina is listed under impressa, this name has seniority.]
  • montana. Pachycondyla fuscoatra var. montana Forel, 1912c: 39 (w.) COLOMBIA.
    • Type-material: lectotype worker (by designation of Mackay & Mackay, 2010: 393), 1 paralectotype worker.
    • Type-locality: lectotype Colombia: Sierra Nevada de Santa Marta, San Antonio (A. Forel); paralectotype with same data.
    • [Note: Mackay & Mackay, 2010: 393, report specimens in MHNG labelled as syntypes from two localities in Costa Rica, and one in Colombia: Rio Frio. These were not recorded by Forel in the original description and are not type-material.]
    • Type-depository: MHNG.
    • Junior synonym of impressa: Kempf, 1961c: 195; Kempf, 1972a: 174; Bolton, 1995b: 307; Mackay & Mackay, 2010: 393.
  • transversa. Pachycondyla fuscoatra r. transversa Emery, 1890a: 58 (in text) (w.q.) COSTA RICA.
    • Type-material: lectotype worker (by designation of Mackay & Mackay, 2010: 393), 2 paralectotype workers.
    • [Note: in the orginal description Emery also mentions a queen.]
    • Type-locality: Costa Rica: (no further data in original description).
    • [Note:Type- locality: lectotype Costa Rica: Alajuela, Juan Viñas (data from Emery, 1890b: 42, repeated in Mackay & Mackay, 2010: 393); paralectotype with same data.]
    • Type-depository: MSNG.
    • [Also described as new by Emery, 1890b: 42; Emery, 1894k: 48.]
    • Combination in Pachycondyla (Pachycondyla): Emery, 1901a: 45.
    • Forel, 1908b: 38 (q.).
    • Status as species: Dalla Torre, 1893: 35.
    • Subspecies of fuscoatra: Emery, in Dalla Torre, 1893: 34 (footnote); Forel, 1908b: 38; Forel, 1908b: 38; Emery, 1911d: 74; Forel, 1914e: 10.
    • Junior synonym of impressa: Kempf, 1961c: 195; Kempf, 1972a: 174; Bolton, 1995b: 311; Mackay & Mackay, 2010: 393.

Type Material

Colombia; Costa Rica, Alajuela, Juan Viñas; Dalla Torre; Colombia, Sierra Nevada de Santa Marta, San Antonio; Ecuador, Santo Domingo de los Colorados; Guatemala, Quirigua. Lectotype worker and 2 paralectotype workers here designated, Museo Civico di Storia Naturale, Genoa; lectotype worker, 1 paralectotype worker labeled, others labeled as syntypes from Suerre and El Aragua, Costa Rica and Río Frio, Colombia, Musee d'Histoire Naturelle Genève; lectotype worker and 1 paralectotype worker seen, National Museum of Natural History, 3 syntypes workers, Museum of Comparative Zoology, 6 worker syntypes American Museum of Natural History (Mackay and Mackay 2010) Unless otherwise noted the text for the remainder of this section is reported from the publication that includes the original description.

Description

Worker

From Mackay and Mackay (2010): The workers are usually large (total length about 11 mm) black ants with reddish brown appendages. Most surfaces are dull and punctate; the mandibles are relatively smooth and glossy. The eyes are relatively small (maximum diameter 0.5 mm) and separated from the anterior edge of the head (side view) by less than one diameter. The malar carina is not developed anterior to the eye. The pronotal shoulder is swollen but usually does not form a carina, the dorsum of the mesosoma is broadly convex and the metanotal suture is poorly marked or absent on the dorsal surface of the mesosoma. The propodeal spiracle is slit-shaped. The petiole is large and thick when viewed in profile, rectangular-shaped with the apex sometimes being slightly broader than the region near the base of the peduncle. The anterior face of the postpetiole is slightly concave and forms a relatively sharp angle with the dorsal surface. The metasternal process consists of two triangular lobes with small teeth on the interior surfaces. Erect hairs are moderately abundant, including on the malar area, the shaft of the scape, the dorsal and ventral surfaces of the head, the dorsum of the mesosoma, the legs (many hairs are suberect), the dorsum of the petiole and all surfaces of the gaster.

The mandibles are finely sculptured and moderately to strongly shining, the head and mesosoma are dull and punctate with some of the punctures in rows that appear as poorly defined striae, especially on the dorsum of the mesosoma and the mesopleuron. The petiole is roughly punctate; the gaster has scattered punctures and is moderately smooth and glossy.

Queen

From Mackay and Mackay (2010): The female is similar to the worker, but is somewhat larger (total length 16 mm). The remainder of the ant is nearly identical to that of the worker, except ocelli are present, the mesosoma is swollen and the metanotum is well developed.

The pilosity and sculpture are similar to those of the workers.

Male

From Mackay and Mackay (2010): (not associated with workers or females.) The male (undescribed) is a large (total length 15 mm) black specimen. The mandibles are tiny with well-developed depressions near the bases. The medial anterior margin of the clypeus is strongly concave, the surface of the clypeus is convex, but it does not overhang the anterior half when the head is seen from side. The head length is 1.80 mm, the head width is 1.84 mm. The eye is moderate in size (maximum diameter 0.83 mm) located less than one diameter from the lateral ocellus (oblique side view). The medial ocellus (diameter 0.15 mm) is located more than one diameter (distance of 0.24 mm) from the lateral ocellus (0.16 mm).

The pronotum is barely swollen at the shoulder. The Mayrian furrows are well developed as are the parapsidal sutures. The scutellum is swollen and rounded when viewed from side. The spiracle is slit-shaped. The petiole is wide when viewed in profile with sloping (not parallel) anterior and posterior faces, which narrow dorsally and form a broadly rounded dorsal face. The subpetiolar process is poorly developed and consists of a rounded lobe anteriorly with the remainder of the process gradually diminishing in width posteriorly. The anterior face of the postpetiole is broadly rounded; the postpetiolar process is not developed. The pygidium is not depressed or concave as in the worker and female. Erect hairs are mostly short (0.1 - 0.35 mm) and are present on the clypeus, dorsal surface of the head (ventral surface cannot be seen), dorsum of the mesosoma, dorsum of the petiole, ventral surface of the petiole and all surfaces of the gaster. The hairs on the legs are mostly appressed, especially on the tibiae, a few hairs on the coxae are erect and suberect. Appressed golden pubescence is scattered on all surfaces.

The head is very finely punctate and weakly shining, the mesosoma is similar, but some of the surfaces, especially the mesopleuron, are moderately shiny. The side of the propodeum and the side of the petiole have poorly developed rugae with well-developed rugae near the posterior lateral edges and on the apex. The gaster is finely punctate and moderately shining.

Karyotype

  • 2n = 94, karyotype = 8M+86A (Brazil) (Mariano et al., 2012).

Etymology

The name of this species is from the Latin word impressio, meaning a pressing into, referring to the impressed pygidium. (Mackay and Mackay 2010)

References

  • Brown, B. V. and D. Feener. 1991. Behavior and host location cues of Apocephalus paraponerae (Diptera, Phoridae), a parasitoid of the giant tropical ant, Paraponera clavata (Hymenoptera: Formicidae). Biotropica 23:182-187.
  • 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. doi:10.1093/ee/nvv071
  • Kaspari, M. and M. Weiser. 2000. Ant activity along moisture gradients in a Neotropical forest. Biotropica 32:703-711.
  • Kempf, W. W. 1961e. As formigas do gênero Pachycondyla Fr. Smith no Brasil (Hymenoptera: Formicidae). Rev. Bras. Entomol. 10: 189-204 (page 195, Senior synonym of andicola, cearensis, inca, montana, purpurascens and transversa)
  • Roger, J. 1861a. Die Ponera-artigen Ameisen (Schluss). Berl. Entomol. Z. 5: 1-54 (page 6, worker described)
  • Roger, J. 1863b. Verzeichniss der Formiciden-Gattungen und Arten. Berl. Entomol. Z. 7(B Beilage: 1-65 (page 18, Combination in Pachycondyla)
  • Sanjuan, T. I., A. E. Franco-Molano, R. M. Kepler, J. W. Spatafora, J. Tabima, A. M. Vasco-Palacios, and S. Restrepo. 2015. Five new species of entomopathogenic fungi from the Amazon and evolution of neotropical Ophiocordyceps. Fungal Biology. 119:901-916. doi:10.1016/j.funbio.2015.06.010
  • Shrestha B, Tanaka E, Hyun MW, Han JG, Kim CS, Jo JW, Han SK, Oh J, Sung JM, Sung GH. 2017. Mycosphere Essay 19. Cordyceps species parasitizing hymenopteran and hemipteran insects. Mycosphere 8(9): 1424–1442 (DOI 10.5943/mycosphere/8/9/8).

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
  • Aldana delq Torre R. C., and P. Chacon de Ulloa. 1999. Megadiversidad de hormigas (Hymenoptera: Formicidae) de la cuenca media del rio Calima. Revista Colombiana de Entomologia 25(1-2): 37-47.
  • Amat-G G., M. G. Andrade-C. and F. Fernández. (eds.) 1999. Insectos de Colombia. Volumen II. Bogotá: Academia Colombiana de Ciencias Exactas, Físicas y Naturales, 433 pp. 131975
  • Armbrecht I., and H. Armbrecht. 1997. Observaciones sobre la variacion espacial y temporal de hormigas en un bosque del Choco Colombiano (Arusi). Bol. Mus. Ent. Univ. Valle 5(2): 15-33.
  • 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
  • Bustos H., J. 1994. Contribucion al conocimiento de al fauna de hormigas (Hymenoptera: Formicidae) del occidente del Departamento de Narino (Colombia). Bol. Mus. Ent. Univ. Valle 2(1,2):19-30
  • Chacon de Ulloa P., S. Valdes-Rodriguez, A. Hurtado-Giraldo, and M. Cleopatra Pimienta. 2014. Arboreal ants of Gorgona National Park (Pacific of Colombia). Rev. Biol. Trop. 62(1): 277-287.
  • 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 biodiversity of four groups of ants in coffee landscapes. Biodiv. Conserv. 22: 871-888.
  • 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.
  • Donoso D. A. 2014. Assembly mechanisms shaping tropical litter ant communities. Ecography 37 doi: 10.1111/j.1600-0587.2013.00253.x
  • Emery C. 1896. Formiche raccolte dal dott. E. Festa nei pressi del golfo di Darien. Bollettino dei Musei di Zoologia ed Anatomia Comparata della Reale Università di Torino 11(229): 1-4.
  • Emery C. 1911. Hymenoptera. Fam. Formicidae. Subfam. Ponerinae. Genera Insectorum 118: 1-125.
  • 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., 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.
  • Field Museum Collection, Chicago, Illinois (C. Moreau)
  • Forel A. 1905. Miscellanea myrmécologiques II (1905). Ann. Soc. Entomol. Belg. 49: 155-185.
  • 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.
  • Garcia Cardenas R., G. A. Zabala, and J. E. Botero. 2008. Hormigas cazadoras (Formicidae: grupos Poneroide y Ectatomminoide) en paisajes cafeteros de Colombia. Pp. 461-478. In Sistemática, biogeografía y conservación de las hormigas cazadoras de Colombia. Bogotá: Instituto de Investigación de Recursos Biológicos Alexander von Humboldt, xiv + 609 pp.
  • Gonzales-Valvidia N. A., G. Gonzales-Escolastico, E. Barba, S. Hernandez-Daumas, and S. Ochoa-Gaona. 2013. Mirmecofauna associated with agroforestry systems in the Mesoamerican Biological Corridor in Tabasco, Mexico. Revista Mexicana de Biodiversidad 84: 306-317.
  • INBio Collection (via Gbif)
  • 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. 1978. A preliminary zoogeographical analysis of a regional ant fauna in Latin America. 114. Studia Entomologica 20: 43-62.
  • Kempf, W.W. 1972. Catalago abreviado das formigas da regiao Neotropical (Hym. Formicidae) Studia Entomologica 15(1-4).
  • 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., J. Coddington, and R. K. Colwell. 2002. The ant fauna of a tropical rain forest: estimating species richness three different ways. Ecology 83: 689-702.
  • Longino J. et al. ADMAC project. Accessed on March 24th 2017 at https://sites.google.com/site/admacsite/
  • 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
  • Philpott, S.M., P. Bichier, R. Rice, and R. Greenberg. 2007. Field testing ecological and economic benefits of coffee certification programs. Conservation Biology 21: 975-985.
  • 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.
  • Pérez-Sánchez A. J., J. E. Lattke, and M. A. Riera-Valera. 2014. The Myrmecofauna (Hymenoptera: Formicidae) of the Macanao Semi-arid Peninsula in Venezuela: An Altitudinal Variation Glance. J Biodivers Biopros Dev 1: 116. doi:10.4172/ijbbd.1000116
  • 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.
  • Scott-Santos, C.P., F.A. Esteves, C.R.F. Brandao. 2008. Catalogue of "Poneromorph" ant type specimens (Hymenoptera, Formicidae) deposited in the Museu de Zoologia da Universidade de Sao Paulo, Brazil. Papeis Avulsos de Zoologia 48(11):75-88.
  • Souza J. L. P., C. A. R. Moura, A. Y. Harada, and E. Franklin. 2007. Diversity of species of the genera Crematogaster, Gnamptogenys and Pachycondyla, (Hymenoptera: Formicidae) and complementarity of sampling methods during the dry season in an ecological station in the Brazilian state of Pará. Acta Amazonica 37(4): 649-656.
  • 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.
  • Vieira de Oliveira J. A., D. Martins da Silva, and F. A. Santana. 2014. Ant species diversity in ciliary forest and gallery forest areas in central Brazil. Advances in Entomology 2(1): 24-32.
  • Vásquez-Bolaños M. 2011. Lista de especies de hormigas (Hymenoptera: Formicidae) para México. Dugesiana 18: 95-133
  • Wheeler W. M. 1922. The ants of Trinidad. American Museum Novitates 45: 1-16.
  • 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
  • Yanoviak S. P., and M. Kaspari. 2000. Community structure and the habitat templet: ants in the tropical forest canopy and litter. Oikos 89: 259-266.