Atta colombica

Atta colombica
Scientific classification
Kingdom:	Animalia
Phylum:	Arthropoda
Class:	Insecta
Order:	Hymenoptera
Family:	Formicidae
Subfamily:	Myrmicinae
Tribe:	Attini
Genus:	Atta
Species:	A. colombica
Binomial name
Atta colombica Guérin-Méneville, 1844 Specimen Labels
Synonyms
Atta cephalotes erecta Santschi, 1929 Atta columbica tonsipes Santschi, 1929 Atta lebasii Guérin-Méneville, 1844

Photo Gallery

• 

  Atta colombica with a visitor. Barro Colorado Island, Panama. Photo by Ajay Narendra.
• 

  Atta colombica major and minor workers. Barro Colorado Island, Panama. Photo by Ajay Narendra.

Distribution

Latitudinal Distribution Pattern

Latitudinal Range: 14.624795° to -12.043°.

North Temperate	North Subtropical	Tropical	South Subtropical	South Temperate

• Source: AntMaps

Distribution based on Regional Taxon Lists

Neotropical Region: Colombia (type locality), Costa Rica, Guatemala, Nicaragua, Panama, Peru.

Distribution based on AntMaps

Distribution based on AntWeb specimens

Check data from AntWeb

Countries Occupied

Number of countries occupied by this species based on AntWiki Regional Taxon Lists. In general, fewer countries occupied indicates a narrower range, while more countries indicates a more widespread species.

Estimated Abundance

Relative abundance based on number of AntMaps records per species (this species within the purple bar). Fewer records (to the left) indicates a less abundant/encountered species while more records (to the right) indicates more abundant/encountered species.

Biology

Explore Fungus Growing  For additional details see Fungus growing ants. A handful of ant species (approx. 275 out of the known 15,000 species) have developed the ability to cultivate fungus within their nests. In most species the fungus is used as the sole food source for the larvae and is an important resource for the adults as well. Additionally, in a limited number of cases, the fungus is used to construct part of the nest structure but is not as a food source. These fungus-feeding species are limited to North and South America, extending from the pine barrens of New Jersey, United States, in the north (Trachymyrmex septentrionalis) to the cold deserts in Argentina in the south (several species of Acromyrmex). Species that use fungi in nest construction are known from Europe and Africa (a few species in the genera Crematogaster, Lasius). The details of fungal cultivation are rich and complex. First, a wide variety of materials are used as substrate for fungus cultivating. The so-called lower genera include species that prefer dead vegetation, seeds, flowers, fruits, insect corpses, and feces, which are collected in the vicinity of their nests. The higher genera include non leaf-cutting species that collect mostly fallen leaflets, fruit, and flowers, as well as the leafcutters that collect fresh leaves from shrubs and trees. Second, while the majority of fungi that are farmed by fungus-feeding ants belong to the family Lepiotaceae, mostly the genera Leucoagaricus and Leucocoprinus, other fungi are also involved. Some species utilise fungi in the family Tricholomataceae while a few others cultivate yeast. The fungi used by the higher genera no longer produce spores. Their fungi produce nutritious and swollen hyphal tips (gongylidia) that grow in bundles called staphylae, to specifically feed the ants. Finally, colony size varies tremendously among these ants. Lower taxa mostly live in inconspicuous nests with 100–1000 individuals and relatively small fungus gardens. Higher taxa, in contrast, live in colonies made of 5–10 million ants that live and work within hundreds of interconnected fungus-bearing chambers in huge subterranean nests. Some colonies are so large, they can be seen from satellite photos, measuring up to 600 m3. Based on these habits, and taking phylogenetic information into consideration, these ants can be divided into six biologically distinct agricultural systems (with a list of genera involved in each category): Nest Construction A limited number of species that use fungi in the construction of their nests. some Crematogaster some Lasius Lower Agriculture Practiced by species in the majority of fungus-feeding genera, including those thought to retain more primitive features, which cultivate a wide range of fungal species in the tribe Leucocoprineae. some Apterostigma Cyatta (1 species) some Cyphomyrmex Kalathomyrmex (1 species) Mycocepurus (6 species) Myrmicocrypta (31 species) Mycetarotes (4 species) Mycetosoritis (2 species) Mycetophylax (21 species) Paramycetophylax (1 species) Xerolitor (1 species) Coral Fungus Agriculture Practiced by species in the Apterostigma pilosum species-group, which cultivate fungi within the Pterulaceae. some Apterostigma Yeast Agriculture Practiced by species within the Cyphomyrmex rimosus species-group, which cultivate a distinct clade of leucocoprineaceous fungi derived from the lower attine fungi. some Cyphomyrmex Generalized Higher Agriculture Practiced by species in several genera of non-leaf-cutting "higher attine" ants, which cultivate a distinct clade of leucocoprineaceous fungi separately derived from the lower attine fungi. Mycetomoellerius (31 species) Paratrachymyrmex (9 species) Sericomyrmex (11 species) Trachymyrmex (9 species) Leaf-Cutter Agriculture A subdivision of higher attine agriculture practiced by species within several ecologically dominant genera, which cultivate a single highly derived species of higher attine fungus. Acromyrmex (56 species) Amoimyrmex (3 species) Atta (20 species) Note that the farming habits of Mycetagroicus (4 species) are unknown. Also, while species of Pseudoatta (2 species) are closely related to the fungus-feeding genus Acromyrmex, they are social parasites, living in the nests of their hosts and are not actively involved in fungus growing. ‎

Dijkstra and Boomsma (2006) investigated the viability of worker produced eggs in Atta cephalotes, Atta sexdens and Atta colombica. Most Atta workers have rudimentary, non-functional ovaries in a queenright colony but a few, typically tending the queen, can produce trophic eggs (Dijkstra et al., 2005). These eggs are feed to the queen. It was not known if any worker eggs can produce males. Most worker eggs of A. colombica did not contain yolk but a few did and yielded morphologically correct but very small males. They hypothesize that worker reproduction in orphaned Atta field colonies is almost never successful because the last workers die before their sons can be raised to adulthood, but the importance of worker-laid trophic eggs for queen feeding has precluded the evolutionary loss of worker ovaries.

Foraging

Bochynek et al. (2017) studied foraging rhythms in a colony of A. colombica in Panama: Foraging on the control (leaf) trail was diurnal and occurred in discrete periods approximately 10 h long. Activity began at dawn, rose to a peak in early afternoon and declined toward evening, with no foraging at night, a typical pattern for Atta leaf foraging (Lewis et al., 1974a). Leaf foraging on the berry/leaf trail followed a similar pattern. The peak harvesting rates, about 100–150 laden ant returns per min., were similar to the mean rate of 108 laden returns per min. observed by Bruce and Burd (2012) for colonies of A. colombica and Atta cephalotes in Panama and Costa Rica. Thus, leaf foraging was typical of colonies at other times and places.

Fruit foraging was fundamentally different. Harvesting was continuous with no periods of inactivity. There was, nonetheless, a diel rhythm. Throughout most of a 24 h cycle, fruit carriers returned at approximately the same rate as leaf harvesters at peak activity, but activity slowed to about 50 returns per min. in early morning, at 07.00 hours, before returning to a plateau of about 100–150 returns per min. for the remainder of the cycle.

Association with Other Organisms

 Explore: Show all Associate data or Search these data. See also a list of all data tables or learn how data is managed.

• This species is a host for the phorid fly Apocephalus colombicus (a parasite) (phorid.net) (attacked).
• This species is a host for the phorid fly Apocephalus attophilus (a parasitoid) (Quevillon, 2018) (encounter mode primary; direct transmission; transmission outside nest).
• This species is a host for the phorid fly Eibesfeldtphora pala (a parasitoid) (Quevillon, 2018) (encounter mode primary; direct transmission; transmission outside nest).
• This species is a host for the fungus Ophiocordyceps subramanianii (a parasitoid) (Quevillon, 2018) (encounter mode primary; direct transmission; transmission outside nest).

Castes

Images from AntWeb

Syntype of Atta cephalotes erecta. Worker. Specimen code casent0912490. Photographer Z. Lieberman, uploaded by California Academy of Sciences.	Owned by NHMB, Basel, Switzerland.

Syntype of Atta colombica tonsipes. Worker. Specimen code casent0912491. Photographer Z. Lieberman, uploaded by California Academy of Sciences.	Owned by NHMB, Basel, Switzerland.

Phylogeny

Atta	Atta mexicana Atta insularis Atta texana Atta cephalotes Atta colombica Atta robusta Atta sexdens Atta saltensis Atta vollenweideri Atta bisphaerica Atta goiana Atta laevigata Atta capiguara Atta opaciceps

Based on Barrera, C.A. et al., 2021. Note that only selected species are included.

Nomenclature

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

• colombica. Atta colombica Guérin-Méneville, 1844a: 422 (w.) COLOMBIA.
  • Type-material: holotype (?) worker.
  • [Note: no indication of number of specimens is given.]
  • Type-locality: Colombia: (no further data).
  • [Notes (i): Borgmeier, 1959b: 346, cites Colombia: Santa Marta as type-locality; (ii) according to Horn & Kahle, 1935: 99, Guérin’s Hymenoptera material was split between MNHN, MSNG, and ZSBS, but some are also present in NNML.]
  • Type-depository: MNHN.
  • [Misspelled as columbica by Roger, 1863b: 35, Mayr, 1865: 81, Forel, 1885a: 362, Emery, 1890b: 55, Dalla Torre, 1893: 152, Forel, 1899c: 33, Forel, 1912e: 179, Emery, 1913b: 259, Wheeler, W.M. 1923c: 5, Emery, 1924d: 353, Menozzi, 1927c: 268, Santschi, 1929f: 92, Weber, 1938b: 205, Borgmeier, 1939: 422, Bolton, 1995b: 75, and others.]
  • Forel, 1913l: 239 (m.); Borgmeier, 1959b: 348 (q.); Wheeler, G.C. 1949: 681 (l.).
  • Combination in Oecodoma: Smith, F. 1858b: 184; Mayr, 1863: 438;
  • combination in Atta: Roger, 1863b: 35.
  • Subspecies of lebasii: Emery, 1890b: 55.
  • Subspecies of cephalotes: Emery, 1913b: 259.
  • Status as species: Smith, F. 1858b: 184; Roger, 1863b: 35; Mayr, 1863: 437; Mayr, 1865: 81; Forel, 1885a: 362; Dalla Torre, 1893: 152; Emery, 1894k: 58; Forel, 1899c: 33; Forel, 1908b: 40; Forel, 1908c: 348; Forel, 1912e: 179; Forel, 1913l: 239; Forel, 1914e: 10; Wheeler, W.M. 1923c: 5; Emery, 1924d: 353; Wheeler, W.M. 1925a: 36; Menozzi, 1927c: 268; Weber, 1938b: 205; Borgmeier, 1939: 422 (in list); Weber, 1941b: 127; Gonçalves, 1942: 346; Borgmeier, 1950d: 259; Weber, 1958a: 11; Borgmeier, 1959b: 345 (redescription); Kempf, 1972a: 26; Cherrett & Cherrett, 1989: 53; Bolton, 1995b: 75; Branstetter & Sáenz, 2012: 257; Fernández, et al. 2015: 137 (redescription); Guénard & Economo, 2015: 227; Fernández & Serna, 2019: 842.
  • Senior synonym of erecta: Borgmeier, 1959b: 346; Kempf, 1972a: 27; Bolton, 1995b: 75; Fernández, et al. 2015: 137.
  • Senior synonym of lebasii: Dalla Torre, 1893: 152; Forel, 1899c: 33, Emery, 1924d: 353; Kempf, 1972a: 27; Bolton, 1995b: 75; Fernández, et al. 2015: 137.
  • Senior synonym of tonsipes: Borgmeier, 1959b: 346; Kempf, 1972a: 27; Bolton, 1995b: 75; Fernández, et al. 2015: 137.
  • Distribution: Colombia, Costa Rica, Guatemala, Panama, Peru.
• erecta. Atta cephalotes var. erecta Santschi, 1929f: 92 (diagnosis in key) (w.) COSTA RICA.
  • Type-material: holotype (?) worker.
  • [Note: no indication of number of specimens is given.]
  • Type-locality: Costa Rica: Santa Clara, Columbiana, 2.i.1924 (G.C. Wheeler).
  • Type-depository: NHMB.
  • Subspecies of cephalotes: Borgmeier, 1939: 422 (in list); Gonçalves, 1942: 345; Borgmeier, 1950d: 243.
  • Junior synonym of colombica: Borgmeier, 1959b: 346; Kempf, 1972a: 27; Bolton, 1995b: 76; Fernández, et al. 2015: 137.
• lebasii. Atta lebasii Guérin-Méneville, 1844a: 422 (w.) COLOMBIA.
  • Type-material: holotype (?) worker.
  • [Note: no indication of number of specimens is given.]
  • Type-locality: Colombia: (no further data) (Lebas).
  • Type-depository: unknown (perhaps MNHN).
  • [Note: according to Horn & Kahle, 1935: 99, Guérin’s Hymenoptera material was split between MNHN, MSNG, and ZSBS, but some are also present in NNML.]
  • Emery, 1890b: 54 (q.m.).
  • Combination in Oecodoma: Smith, F. 1858b: 184;
  • combination in Atta: Roger, 1863b: 35.
  • Status as species: Smith, F. 1858b: 184; Roger, 1863b: 35; Mayr, 1863: 438; Mayr, 1865: 81; Emery, 1890b: 54.
  • Junior synonym of colombica: Dalla Torre, 1893: 152; Forel, 1899c: 33, Emery, 1924d: 353; Kempf, 1972a: 27; Bolton, 1995b: 76; Fernández, et al. 2015: 137.
• tonsipes. Atta columbica var. tonsipes Santschi, 1929f: 92 (diagnosis in key) (w.) PANAMA.
  • Type-material: syntype workers (number not stated).
  • Type-locality: Panama: Bellavista (W.M. Wheeler).
  • Type-depository: NHMB (perhaps also MCZC).
  • Subspecies of colombica: Borgmeier, 1939: 422 (in list); Weber, 1941b: 127; Gonçalves, 1942: 346; Borgmeier, 1950d: 243; Weber, 1958a: 12.
  • Junior synonym of colombica: Borgmeier, 1959b: 346; Kempf, 1972a: 27; Bolton, 1995b: 77; Fernández, et al. 2015: 137.

Description

Karyotype

• See additional details at the Ant Chromosome Database.

 Explore: Show all Karyotype data or Search these data. See also a list of all data tables or learn how data is managed.

• n = 11, 2n = 22, karyotype = 12M+6SM+4A (Panama) (Murakami et al., 1998; Barros et al., 2014) (Atta columbica is a writing mistake).

References

• Guérin-Méneville, F. E. 1844a. Iconographie du règne animal de G. Cuvier, ou représentation d'après nature de l'une des espèces les plus remarquables, et souvent non encore figurées, de chaque genre d'animaux. Insectes. Paris: J. B. Baillière, 576 pp.
• Armitage, S., Boomsma, J., Bear, B. 2010. Diploid male production in a leaf-cutting ant. Ecological Entomology 35, 175–182 (doi:10.1111/j.1365-2311.2009.01167.x).
• Baer, B. 2011. The copulation biology of ants (Hymenoptera: Formicidae). Myrmecological News 14: 55-68.
• Barrera, C.A., Sosa-Calvo, J., Schultz, T.R., Rabeling, C., Bacci, M., Jr 2021. Phylogenomic reconstruction reveals new insights into the evolution and biogeography of Atta leaf-cutting ants (Hymenoptera: Formicidae). Systematic Entomology 47: 13-35 (doi:10.1111/syen.12513).
• Barros, L.A.C., Teixeira, G.A., de Aguiar, J.A.C., Mariano, C.D.S.F., Delabie, J.H.C., Pompolo, S.D.G. 2014. Banding Patterns of Three Leafcutter Ant Species of the Genus Atta (Formicidae: Myrmicinae) and Chromosomal Inferences. Florida Entomologist 97(4): 1694-1701.
• Bochynek, T., J. L. Tanner, B. Meyer, and M. Burd. 2017. Parallel foraging cycles for different resources in leaf-cutting ants: a clue to the mechanisms of rhythmic activity. Ecological Entomology. 42:849-852. doi:10.1111/een.12437
• Bustamante, S., Amarillo-Suárez, A. 2019. Foraging plasticity of Atta cephalotes (Hymenoptera: Formicidae) in response to temperature differences between forest and pasture. Revista de Biología Tropical 67(4), 963-974.
• Cardoso, D. C., Cristiano, M. P. 2021. Karyotype diversity, mode, and tempo of the chromosomal evolution of Attina (Formicidae: Myrmicinae: Attini): Is there an upper limit to chromosome number? Insects 1212, 1084 (doi:10.3390/insects12121084).
• Dahan, R.A., Grove, N.K., Bollazzi, M., Gerstner, B.P., Rabeling, C. 2021. Decoupled evolution of mating biology and social structure in Acromyrmex leaf-cutting ants. Behavioral Ecology and Sociobiology 76, 7 (doi:10.1007/s00265-021-03113-1).
• de Bekker, C., Will, I., Das, B., Adams, R.M.M. 2018. The ants (Hymenoptera: Formicidae) and their parasites: effects of parasitic manipulations and host responses on ant behavioral ecology. Myrmecological News 28: 1-24 (doi:10.25849/myrmecol.news_028:001).
• Dijkstra, M. B. and J. J. Boomsma. 2006. Are workers of Atta leafcutter ants capable of reproduction? Insectes Sociaux. 53(2):136-140. doi:10.1007/s00040-005-0848-3
• Dijkstra, M. B., D. R. Nash, and J. J. Boomsma. 2005. Self-restraint and sterility in workers of Acromyrmex and Atta leafcutter ants. Insectes Sociaux. 52(1):67-76. doi:10.1007/s00040-004-0775-8
• Farias, A.P., Camargo, R.da S., Caldato, N., Forti, L.C. 2020. Nest architecture development of grass-cutting ants, Atta capiguara (Hymenoptera: Formicidae). Revista Brasileira de Entomologia 64, e20200034 (doi:10.1590/1806-9665-rbent-2020-0034).
• Fleischmann, P.N., Grob, R., Rössler, W. 2020. Magnetoreception in Hymenoptera: importance for navigation. Animal Cognition (doi:10.1007/s10071-020-01431-x).
• Horna-Lowell, E., Neumann, K.M., O’Fallon, S., Rubio, A., Pinter-Wollman, N. 2021. Personality of ant colonies (Hymenoptera: Formicidae) – underlying mechanisms and ecological consequences. Myrmecological News 31: 47-59 (doi:10.25849/MYRMECOL.NEWS_031:047).
• Hull-Sanders, H. M., and Howard, J. J. 2003. Impact of Atta colombica colonies on understory vegetation and light availability in a Neotropical forest. Biotropica 35(3): 441–445
• Jacobs, S. 2020. Population genetic and behavioral aspects of male mating monopolies in Cardiocondyla venustula (Ph.D. thesis).
• Lau, M.K., Ellison, A.M., Nguyen, A., Penick, C., DeMarco, B., Gotelli, N.J., Sanders, N.J., Dunn, R.R., Helms Cahan, S. 2019. Draft Aphaenogaster genomes expand our view of ant genome size variation across climate gradients. PeerJ 7, e6447 (doi:10.7717/PEERJ.6447).
• Li, H., Young, S.E., Poulsen, M., Currie, C.R. 2020. Symbiont-mediated digestion of plant biomass in fungus-farming insects. Annual Review of Entomology 66:16.1–16.20 (doi:10.1146/annurev-ento-040920-061140).
• Matte, A., Billen, J. 2012. Flight muscle histolysis in Lasius niger queens. Asian myrmecology 13, e013003 (doi:10.20362/am.013003).
• Mueller, U.G., Ishak, H.D., Bruschi, S.M., Smith, C.C., Herman, J.J., Solomon, S.E., Mikheyev, A.S., Rabeling, C., Scott, J.J., Cooper, M., Rodrigues, A., Ortiz, A., Brandão, C.R.F., Lattke, J.E., Pagnocca, F.C., Rehner, S.A., Schultz, T.R., Vasconcelos, H.L., Adams, R.M.M., Bollazzi, M., Clark, R.M., Himler, A.G., LaPolla, J.S., Leal, I.R., Johnson, R.A., Roces, F., Sosa-Calvo, J., Wirth, R., Bacci, M. 2017. Biogeography of mutualistic fungi cultivated by leafcutter ants. Molecular Ecology 26, 6921–6937 (doi:10.1111/mec.14431).
• Mueller, U.G., Schultz, T.R., Currie, C.R., Adams, R.M.M., Malloch, D. 2001. The origin of the attine ant-fungus mutualism. The Quarterly Review of Biology 76, 169-197.
• Narendra, A., Ramirez-Esquivel, F., Ribi, W.A. 2016. Compound eye and ocellar structure for walking and flying modes of locomotion in the Australian ant, Camponotus consobrinus. Scientific Reports 6, 22331 (doi:10.1038/srep22331).
• Nehring, V., Boomsma, J.J., d'Ettorre, P. 2012. Wingless virgin queens assume helper roles in Acromyrmex leaf-cutting ants. Current Biology 22, R671–R673 (doi:10.1016/j.cub.2012.06.038).
• Pokorny, T., Sieber, L.-M., Hofferberth, J.E., Bernadou, A., Ruther, J. 2020. Age-dependent release of and response to alarm pheromone in a ponerine ant. The Journal of Experimental Biology 223, jeb218040. (doi:10.1242/JEB.218040).
• Ramalho, M.de O., Martins, C., Morini, M.S.C., Bueno, O.C. 2020. What can the bacterial community of Atta sexdens (Linnaeus, 1758) tell us about the habitats in which this ant species evolves? Insects 11, 332. (doi:10.3390/INSECTS11060332).
• Ronque, M.U.V., Lyra, M.L., Migliorini, G.H., Bacci, M., Oliveira, P.S. 2020. Symbiotic bacterial communities in rainforest fungus-farming ants: evidence for species and colony specificity. Scientific Reports 10, 10172 (doi:10.1038/S41598-020-66772-6).
• Toledo-Hernández, E., Hernández-Flores, J., Sotelo-Leyva, C., Alvear-García, A., Peña-Chora, G. 2021. A new enemy of Apis mellifera (Hymenoptera: Apidae): First report of Nomamyrmex esenbeckii (Hymenoptera: Formicidae) attacking honey bee colonies. Journal of Apicultural Research, 1–2 (doi:10.1080/00218839.2021.1987740).

References based on Global Ant Biodiversity Informatics

• Anderson, C. and J.L.V. Jadin. 2001. The adaptive benefit of leaf transfer in Atta colombica. Insectes Sociaux 48:404-405
• Baer, B., S. P. A. den Boer, D. J. C. Kronauer, D. R. Nash and J. J. Boomsma. 2009. Fungus gardens of the leafcutter ant Atta colombica function as egg nurseries for the snake Leptodeira annulata. Insectes Sociaux 56(3):289-291
• Borgmeier T. 1939. Nova contribuição para o conhecimento das formigas neotropicas (Hym. Formicidae). Revista de Entomologia (Rio de Janeiro) 10: 403-428.
• Borgmeier T. 1950. Atta-Studien (Hym. Formicidae). Memórias do Instituto Oswaldo Cruz. Rio de Janeiro 48: 265-292.
• Borgmeier T. 1950. Estudos sôbre Atta (Hym. Formicidae). Memórias do Instituto Oswaldo Cruz. Rio de Janeiro 48: 239-263.
• 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
• Davidson D. W., S. C. Cook, R. R. Snelling and T. H. Chua. 2003. Explaining the Abundance of Ants in Lowland Tropical Rainforest Canopies. Science 300: 969-972.
• Davidson, D.W. 2005. Ecological stoichiometry of ants in a New World rain forest. Oecologia 142:221-231
• Emery C. 1890. Studii sulle formiche della fauna neotropica. Bull. Soc. Entomol. Ital. 22: 38-8
• Emery C. 1894. Estudios sobre las hormigas de Costa Rica. Anales del Museo Nacional de Costa Rica 1888-1889: 45-64.
• 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 S. Sendoya. 2004. Lista de las hormigas neotropicales. Biota Colombiana Volume 5, Number 1.
• Forel A. 1908. Fourmis de Costa-Rica récoltées par M. Paul Biolley. Bulletin de la Société Vaudoise des Sciences Naturelles 44: 35-72.
• Forel A. 1912. Formicides néotropiques. Part II. 3me sous-famille Myrmicinae Lep. (Attini, Dacetii, Cryptocerini). Mémoires de la Société Entomologique de Belgique. 19: 179-209.
• Gonçalves C. R. 1942. Contribuiça~o para o conhecimento do gênero Atta Fabr., das formigas saúvas. Bol. Soc. Bras. Agron. 5: 333-358.
• Helmkampf, M., J. Gadau and H. Feldhaar. 2008. Population- and sociogenetic structure of the leaf-cutter ant Atta colombica (Formicidae, Myrmicinae). Insectes Sociaux 55(4):434-442.
• INBio Collection (via Gbif)
• Johnson R. Personnal Database. Accessed on February 5th 2014 at http://www.asu.edu/clas/sirgtools/resources.htm
• Kaspari, M. and M.D. Weiser. Ant Activity along Moisture Gradients in a Neotropical Forest Ant Activity along Moisture Gradients in a Neotropical Forest. Biotropica, Vol. 32, No. 4a (Dec., 2000), pp. 703-711
• Kempf, W.W. 1972. Catalago abreviado das formigas da regiao Neotropical (Hym. Formicidae) Studia Entomologica 15(1-4).
• Kooij P. W., B. M. Dentinger, D. A. Donoso, J. Z. Shik, and E. Gaya. 2018. Cryptic diversity in Colombian edible leaf-cutting ants (Hymenoptera: Formicidae). Insects 9: 191.
• Kost, C., E. Gama de Oliveira, T.A. Knoch, R. Wirth. 2005. Spatio-Temporal Permanence and Plasticity of Foraging Trails in Young and Mature LeafCutting Ant Colonies (Atta spp.). Journal of Tropical Ecology 21(6):677-688
• Kusnezov N. 1953. La fauna mirmecológica de Bolivia. Folia Universitaria. Cochabamba 6: 211-229.
• Longino J. et al. ADMAC project. Accessed on March 24th 2017 at https://sites.google.com/site/admacsite/
• Menozzi C. 1927. Formiche raccolte dal Sig. H. Schmidt nei dintorni di San José di Costa Rica. Entomologische Mitteilungen. Berlin-Dahlem. 16: 266-277.
• Murakami T., A. Fujiwara,and M. C. Yoshida. 1999. Cytogenetics of ten ant species of the tribe Attini (Hymenoptera, Formicidae) in Barro Colorado Island, Panama. Chromosome Science 2(3): 135-139.
• Powell, S. and E. Clark. 2004. Combat between large derived societies: A subterranean army ant established as a predator of mature leaf-cutting ant colonies. Insectes Sociaux 51(4):342-351.
• Sturup, M., S.P.A. den Boer, D.R. Nash, J.J. Boomsma and B. Baer. 2011. Variation in male body size and reproductive allocation in the leafcutter ant Atta colombica: estimating variance components and possible trade-offs. Insectes Sociaux 58:47-55
• 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.
• Weber N. A. 1941. The biology of the fungus-growing ants. Part VII. The Barro Colorado Island, Canal Zone, species. Rev. Entomol. (Rio J.) 12: 93-130.
• Weber N. A. 1958. Nomenclatural notes on Proatta and Atta (Hym.: Formicidae). Entomological News 69: 7-13.
• Weber N. A. 1968. The Panamanian Atta species (Hymenoptera: Formicidae). Proceedings of the Entomological Society of Washington 70: 348-350.
• Weber N. A. 1969. Ecological relations of three Atta species in Panama. Ecology 50: 141-147.