Nomamyrmex esenbeckii

Identification
Watkins (1977) - N. esenbecki. Workers: postoccipital sulcus distinct; dorsum of petiole with longitudinal rugae. Males: median border of head behind ocellar peduncle with a narrow lamella; first gastric tergite usually with longitudinal rugae; width of, blade of stipes about one-half its length; volsella blunt, gradually tapered or snout-shaped.

N. esenbecki s. str. Workers: transversely keeled posterior margin of mesonotum usually straight or slightly convex in the middle; anterior corners of postpetiole prominent and abruptly rounded causing the postpetiole to appear subquadrate with an anterior face as broad or broader than the petiole; entire body usually dark reddish brown with gaster sometimes slightly lighter. The mesonotal keel is occasionally slightly concave, but usually not in all specimens in a series. The posterior edge of the propodeum is often only slightly indented between the propodeal “teeth,” but a few specimens may be deeply indented (dorsal view). Males: setae on fifth gastric tergite broadly separated along the midline (i.e., the separation is about one half the width of the tergite near its base; distal portion of volsella distinctly snout-shaped, but the “snout” varies somewhat in detail.

N. esenbecki crassicornis. Workers: transversely keeled posterior margin of mesonotum usually straight or slightly convex in the middle; anterior corners of the postpetiole not prominent with the sides gradually curving inward to the anterior face which is slightly narrower than the petiole causing the postpetiole to appear trapezoidal; gaster usually yellowish brown, remainder of body reddish brown. A few workers from Panama have slightly concave mesonotal keels; however, these workers have postpetioles and yellowish brown gasters typical for crassicornis. Males: setae on fifth gastric tergite not distinctly separated along the midline; longitudinal rugae distinct on the second gastric tergite; apex of volsella slightly enlarged and rounded. The entire posterior half of the fifth tergite is usually covered with long thick setae; however, some specimens from Panama have the setae along the midline more confined to the posterior border and these setae are shorter than the more lateral setae. The distal portion of the volsella is slightly upturned with a gradually rounded apex in specimens from Panama, almost straight with a bluntly rounded apex in specimens from Trinidad, and somewhat more downturned in specimens from Peru.

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

Biology


Jack Longino: Nomamyrmex esenbeckii forages in columns during the day; I do not know if they forage at night. The workers are robust and their columns very conspicuous. In Costa Rica this species is uncommon relative to several species of Eciton and Labidus.

Nomamyrmex esenbeckii appears to be a specialist raider of Atta colonies (Swartz 1998 and included references). Swartz reviewed the earlier literature, in which workers reported Nomamyrmex raids on Atta colonies in Mexico and various sites in Brazil. She also observed and carefully documented a raid on an Atta cephalotes colony in Corcovado National Park, Costa Rica.

During work in Corcovado National Park in the early 1980's, I saw this species twice with prey and three times in the process of raiding ant colonies. Four of these observations involved attines. I observed Atta brood as prey in one column, and Acromyrmex brood as prey in another. I observed two attacks on mature Atta cephalotes colonies. Both these raids were observed mid-morning. In the first instance, Nomamyrmex were entering through a collapsed Atta chamber in the middle of a trail, taking away brood. I saw very little response on the Atta's part. There were two dead soldiers near the raiding column. In the second case, Atta workers were abundant on the surface of their mound, and had brought piles of brood to the surface of the largest, most active mounds. Nomamyrmex columns were carrying adult Atta soldiers and brood down into Atta nest entrances, suggesting a largely subterranean raid. I observed one raid on a Pheidole nest.

Jack Longino: I have once observed this species with prey. A column was entering and issuing from fissures in a trail, carrying abundant larvae and pupae of Atta, including pupae of fairly large workers.

Kjetil Aasen, a student working at La Selva Biological Station during the summer of 1997, made the following observation of a Nomamyrmex raid on Atta cephalotes:

On the evening on my last day at La Selva, I went to take a look at the Atta cephalotes colony between the bridge and the library and I was surprised to find a fierce battle in progress. It was dark and it was raining and thousands of Atta soldiers and Nomamyrmex esenbeckii were fighting in the area surrounding the main entrances of the Atta nest. They were all emerging from two closely spaced nest entrances and most of the Nomamyrmex were in the mandibles of some large leafcutter. In most cases, the Nomamyrmex were caught by their antenna, but they still managed to sting Atta, and they seemed to be very good at this because there were many dead or dying leafcutters on the battlefield. I did not see many dead Nomamyrmex.

Castes
Primarily wingless queen described by Borgmeier 1958

Nomenclature

 * . Labidus esenbeckii Westwood, 1842: 75, pl. 20, fig. 4 (m.) BRAZIL (Goiás).
 * Type-material: holotype male.
 * Type-locality: Brazil: Rio Vendinha, 10.ix.1828, in the evening (W. Burchell).
 * Type-depository: OXUM.
 * Combination in Eciton (Labidus): Emery, 1890b: 39; Santschi, 1920d: 364; Creighton, 1950a: 64;
 * combination in E. (Holopone): Santschi, 1925b: 11;
 * combination in E. (Nomamyrmex): Borgmeier, 1936: 56;
 * combination in Nomamyrmex: Borgmeier, 1953: 14.
 * Status as species: Smith, F. 1859b: 7; Roger, 1863b: 41; Mayr, 1863: 424; Emery, 1890b: 39; Dalla Torre, 1893: 2; Emery, 1894k: 46; Forel, 1895b: 119; Forel, 1899c: 25; Emery, 1900a: 177 (in key); Forel, 1901h: 47; Wheeler, W.M. 1908e: 409 (redescription); Emery, 1910b: 23; Wheeler, W.M. 1910g: 562; Forel, 1911c: 288; Mann, 1916: 422; Santschi, 1920d: 364; Borgmeier, 1923: 43; Borgmeier, 1936: 56; Creighton, 1950a: 64; Smith, M.R. 1951a: 779; Borgmeier, 1953: 14; Borgmeier, 1955: 139 (redescription); Borgmeier, 1958: 201; Kempf, 1970b: 322; Kempf, 1972a: 164; Kempf & Lenko, 1976: 46; Watkins, 1976: 7 (in key); Watkins, 1977a: 204; Watkins, 1982: 215 (in key); Bolton, 1995b: 292; Palacio, 1999: 161 (in key); Wild, 2007b: 26, 43; Branstetter & Sáenz, 2012: 254; Guénard & Economo, 2015: 226; Shattuck & Cover, 2016: 19; Palacio, 2019: 626.
 * Senior synonym of afflictus: Borgmeier, 1955: 139; Kempf, 1972a: 164; Bolton, 1995b: 292.
 * Senior synonym of crassicornis: Wild, 2007b: 43.
 * Senior synonym of crudele: Borgmeier, 1955: 139; Kempf, 1972a: 164; Bolton, 1995b: 292.
 * Senior synonym of fimbriatum: Borgmeier, 1953: 14; Borgmeier, 1955: 139; Kempf, 1972a: 164; Bolton, 1995b: 292.
 * Senior synonym of hirsutipodex: Wild, 2007b: 43 (by implication).
 * Senior synonym of indomabile: Borgmeier, 1955: 139; Kempf, 1972a: 164; Bolton, 1995b: 292.
 * Senior synonym of interrupta: Borgmeier, 1953: 14; Borgmeier, 1955: 139; Kempf, 1972a: 164; Bolton, 1995b: 292.
 * Senior synonym of intrepidum: Wild, 2007b: 43 (by implication).
 * Senior synonym of mordax: Shattuck & Cover, 2016: 19.
 * Senior synonym of terrificum: Borgmeier, 1953: 14; Borgmeier, 1955: 139; Kempf, 1972a: 164; Bolton, 1995b: 292.
 * Senior synonym of wilsoni: Smith, M.R., 1951a: 779; Shattuck & Cover, 2016: 19.
 * Distribution: Argentina, Belize, Bolivia, Brazil, Colombia, Costa Rica, French Guiana, Guatemala, Guyana, Honduras, Mexico, Panama, Paraguay, Peru, Suriname, Trinidad, U.S.A., Venezuela.
 * afflictus. Acanthostichus afflictus Gallardo, 1919a: 239, fig. 3 (m.) ARGENTINA (Formosa).
 * Type-material: 2 syntype males.
 * Type-locality: Argentina: Formosa, Rio Pilcomayo, vii.1907, no. 7653 (no collector’s name).
 * Type-depository: MACN.
 * Combination in Eciton: Emery, 1920a: 60.
 * Junior synonym of esenbeckii: Borgmeier, 1955: 139; Kempf, 1972a: 164; Bolton, 1995b: 292.
 * crassicornis. Eciton crassicornis Smith, F. 1855c: 163, pl. 13, figs. 1, 2 (w.) BRAZIL (Amazonas).
 * Type-material: syntype workers (number not stated).
 * Type-locality: Brazil: Villa Nova (H.W. Bates).
 * Type-depository: BMNH.
 * Borgmeier, 1958: 202 (q.).
 * Combination in E. (Labidus): Emery, 1910b: 23;
 * combination in E. (Holopone): Santschi, 1925b: 12;
 * combination in E. (Nomamyrmex): Borgmeier, 1936: 55;
 * combination in Nomamyrmex: Borgmeier, 1953: 16.
 * Status as species: Smith, F. 1858b: 151; Mayr, 1863: 409; Roger, 1863b: 36; Mayr, 1865: 77 (in key); Norton, 1868b: 45; Mayr, 1886b: 115 (in key); Emery, 1890a: 55; Emery, 1890b: 38; Dalla Torre, 1893: 2; Emery, 1894c: 179; Emery, 1894k: 45; Forel, 1895b: 141; Emery, 1896h: 625; Forel, 1899c: 24; Emery, 1900a: 185; Forel, 1901c: 124; Forel, 1906d: 249; Forel, 1908b: 40; Forel, 1908c: 346; Forel, 1908e: 64; Forel, 1909a: 254; Emery, 1910b: 23; Mann, 1916: 422; Wheeler, W.M. 1916d: 324; Luederwaldt, 1918: 37; Santschi, 1920d: 367; Wheeler, W.M. 1922c: 1; Borgmeier, 1923: 42; Wheeler, W.M. 1925a: 2; Santschi, 1929a: 414; Borgmeier, 1936: 55; Borgmeier, 1948b: 460; Creighton, 1950a: 63; Smith, M.R. 1951a: 779; Borgmeier, 1953: 16.
 * Subspecies of esenbeckii: Borgmeier, 1955: 149 (redescription); Borgmeier, 1958: 202; Kempf, 1972a: 164; Watkins, 1977a: 205; Bolton, 1995b: 292.
 * Junior synonym of esenbeckii: Wild, 2007b: 43.
 * Material of the unavailable name furiosum referred here by Borgmeier, 1955: 139; Kempf, 1972a: 164.
 * crudele. Eciton (Holopone) crassicorne subsp. crudele Santschi, 1929a: 414, figs. c, g (w.) PARAGUAY, BOLIVIA.
 * Type-material: holotype worker.
 * Type-locality: Paraguay: (no further data) (Fiebrig).
 * [Note: Santschi also includes non-type material from Bolivia: Caranditi (Lizer & Deletang).]
 * Type-depository: NHMB.
 * Subspecies of crassicorne: Santschi, 1929f: 85.
 * Junior synonym of esenbeckii: Borgmeier, 1955: 139; Kempf, 1972a: 164; Bolton, 1995b: 292.
 * fimbriatum. Eciton (Labidus) fimbriatum Santschi, 1920d: 363, figs. 1, 2 (m.) FRENCH GUIANA.
 * Type-material: 8 paratype males.
 * Type-locality: French Guiana: St Jean du Maroni (Le Moult).
 * Type-depositories: MNHN, NHMB.
 * Status as species: Eidmann, 1936a: 29.
 * Junior synonym of esenbeckii: Borgmeier, 1953: 14; Borgmeier, 1955: 139; Kempf, 1972a: 164; Bolton, 1995b: 292.
 * hirsutipodex. Eciton (Labidus) esenbecki st. hirsutipodex Santschi, 1920d: 364, fig. 2a (m.) MEXICO (Yucatan).
 * Type-material: holotype male.
 * Type-locality: Mexico: Merida (no collector’s name).
 * Type-depository: NHMB.
 * Junior synonym of crassicornis: Borgmeier, 1955: 139; Kempf, 1972a: 164; Bolton, 1995b: 292.
 * Junior synonym of esenbeckii: Wild, 2007b: 43 (by implication).
 * indomabile. Eciton (Holopone) crassicorne var. indomabile Santschi, 1930e: 82 (w.) BRAZIL (Pernambuco).
 * Type-material: 19 syntype workers.
 * Type-locality: Brazil: Pernambuco, Olinda (Reichensperger).
 * Type-depository: NHMB.
 * Subspecies of crassicornis: Santschi, 1931c: 270.
 * Junior synonym of esenbeckii: Borgmeier, 1955: 139; Kempf, 1972a: 164; Bolton, 1995b: 292.
 * interrupta. Eciton fimbriatum var. interrupta Santschi, 1923c: 249 (m.) BRAZIL (Minas Gerais).
 * Type-material: holotype male.
 * Type-locality: Brazil: Minas Gerais, Piracicabo (Luja).
 * Type-depository: NHMB.
 * Junior synonym of esenbeckii: Borgmeier, 1953: 14; Borgmeier, 1955: 139; Kempf, 1972a: 164; Bolton, 1995b: 292.
 * intrepidum. Eciton (Holopone) crassicorne var. intrepidum Santschi, 1929a: 414, figs. b, f (w.) VENEZUELA.
 * Type-material: syntype workers (number not stated).
 * Type-locality: Venezuela: between Carupano and Cariaco, 1896 (no collector’s name).
 * Type-depository: unknown.
 * [Note: Baroni Urbani, 1977e: 68-71, does not record the presence of type-material of this taxon in NHMB.]
 * Junior synonym of crassicornis: Borgmeier, 1955: 139; Kempf, 1972a: 164; Bolton, 1995b: 292.
 * Junior synonym of esenbeckii: Wild, 2007b: 43 (by implication).
 * mordax. Eciton (Holopone) crassicorne st. mordax Santschi, 1929a: 415, figs. d, h (w.) MEXICO (Morelos).
 * Type-material: syntype workers (number not stated).
 * Type-locality: Mexico: Cuernavaca (A. Forel).
 * Type-depository: NHMB.
 * Watkins, 1977a: 210 (m.).
 * Combination in Nomamyrmex: Borgmeier, 1953: 13.
 * Subspecies of crassicorne: Menozzi, 1935b: 189.
 * Junior synonym of wilsoni: Borgmeier, 1955: 139; Smith, M.R. 1958c: 108; Kempf, 1972a: 165; Smith, D.R. 1979: 1329.
 * Subspecies of esenbeckii: Watkins, 1977a: 205; Brandão, 1991: 361; Bolton, 1995b: 292.
 * Junior synonym of esenbeckii: Shattuck & Cover, 2016: 19.
 * terrificum. Eciton (Holopone) fimbriatum var. terrificum Santschi, 1930e: 82 (m.) BRAZIL (no state data).
 * Type-material: holotype male.
 * Type-locality: Brazil: (no further data), 23.ix.1929 (Reichensperger).
 * Type-depository: NHMB.
 * Junior synonym of esenbeckii: Borgmeier, 1953: 14; Borgmeier, 1955: 139; Kempf, 1972a: 164; Bolton, 1995b: 292.
 * wilsoni. Eciton (Labidus) esenbecki st. wilsoni Santschi, 1920d: 366 (m.) U.S.A. (Texas).
 * Type-material: holotype male.
 * Type-locality: U.S.A.: Texas, Las Borragas, nr Brownsville, vi. (C. Schaeffer).
 * Type-depository: BMNY.
 * [Note: holotype may now be in AMNH, or if retained by Wheeler, in MCZC.]
 * Combination in E. (Nomamyrmex): Smith, M.R. 1951a: 779;
 * combination in Nomamyrmex: Borgmeier, 1953: 13.
 * Subspecies of esenbeckii: Borgmeier, 1955: 147; Smith, M.R. 1958c: 108; Kempf, 1972a: 165; Watkins, 1977a: 205; Smith, D.R. 1979: 1329; Watkins, 1985: 481; Bolton, 1995b: 292.
 * Junior synonym of esenbeckii: Smith, M.R. 1951a: 779; Shattuck & Cover, 2016: 19.

References based on Global Ant Biodiversity Informatics

 * Baroni Urbani C. 1977. Katalog der Typen von Formicidae (Hymenoptera) der Sammlung des Naturhistorischen Museums Basel (2. Teil). Mitt. Entomol. Ges. Basel (n.s.) 27: 61-102.
 * Basset Y., L. Cizek, P. Cuenoud, R. K. Didham, F. Guilhaumon, O. Missa, V. Novotny, F. Odegaards, T. Roslin, J. Schmidl et al. 2012. Arthropod diversity in a tropical forest. Science 338(6113): 1481-1484.
 * 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.
 * Borgmeier T. 1953. Vorarbeiten zu einer Revision der neotropischen Wanderameisen. Studia Entomologica 2: 1-51.
 * Borgmeier T. 1955. Die Wanderameisen der neotropischen Region. Studia Entomologica 3: 1-720.
 * Borgmeier T. 1957. Die Maxillar- und Labialtaster der neotropischen Dorylinen (Hym. Formicidae). Revista Brasileira de Biologia 17: 387-394.
 * Brandao, C.R.F. 1991. Adendos ao catalogo abreviado das formigas da regiao neotropical (Hymenoptera: Formicidae). Rev. Bras. Entomol. 35: 319-412.
 * Branstetter M. G. and L. Sáenz. 2012. Las hormigas (Hymenoptera: Formicidae) de Guatemala. Pp. 221-268 in: Cano E. B. and J. C. Schuster. (eds.) 2012. Biodiversidad de Guatemala. Volumen 2. Guatemala: Universidad del Valle de Guatemala, iv + 328 pp
 * Brown W. L. Jr. 1960. The release of alarm and attack behavior in some New World army ants. Psyche (Cambridge) 66: 25-27.
 * 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.
 * Cuezzo, F. 1998. Formicidae. Chapter 42 in Morrone J.J., and S. Coscaron (dirs) Biodiversidad de artropodos argentinos: una perspectiva biotaxonomica Ediciones Sur, La Plata. Pages 452-462.
 * Dattilo W. et al. 2019. MEXICO ANTS: incidence and abundance along the Nearctic-Neotropical interface. Ecology https://doi.org/10.1002/ecy.2944
 * Davidson, D.W. 2005. Ecological stoichiometry of ants in a New World rain forest. Oecologia 142:221-231
 * 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. 1896. Formiciden, gesammelt in Paraguay von Dr. J. Bohls. Zoologische Jahrbücher. Abteilung für Systematik, Geographie und Biologie der Tiere 9: 625-638.
 * Emery C. 1910. Hymenoptera. Fam. Formicidae. Subfam. Dorylinae. Genera Insectorum 102: 1-34.
 * Fernandes, P.R. XXXX. Los hormigas del suelo en Mexico: Diversidad, distribucion e importancia (Hymenoptera: Formicidae).
 * 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. 1901. Formiciden des Naturhistorischen Museums zu Hamburg. Neue Calyptomyrmex-, Dacryon-, Podomyrma- und Echinopla-Arten. Mitt. Naturhist. Mus. Hambg. 18: 43-82.
 * Forel A. 1906. Fourmis néotropiques nouvelles ou peu connues. Annales de la Société Entomologique de Belgique 50: 225-249.
 * Forel A. 1908. Ameisen aus Sao Paulo (Brasilien), Paraguay etc. gesammelt von Prof. Herm. v. Ihering, Dr. Lutz, Dr. Fiebrig, etc. Verhandlungen der Kaiserlich-Königlichen Zoologisch-Botanischen Gesellschaft in Wien 58: 340-418.
 * Forel A. 1908. Catálogo systemático da collecção de formigas do Ceará. Boletim do Museu Rocha 1(1): 62-69.
 * 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.
 * 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.
 * Hernandez, F. Varela and G. Castano-Meneses. 2010. Checklist, Biological Notes and Distribution of Ants (Hymenoptera: Formicidae) from Barranca de Metztitlán Biosphere Reserve, Hidalgo, Mexico. Sociobiology 56(2):397-434
 * Hua Li-zhong. 2006. List of Chinese insects Vol. IV. Pages 262-273. Sun Yat-sen university Press, Guangzhou. 539 pages.
 * INBio Collection (via Gbif)
 * Kempf W. W., and K. Lenko. 1976. Levantamento da formicifauna no litoral norte e ilhas adjacentes do Estado de São Paulo, Brasil. I. Subfamilias Dorylinae, Ponerinae e Pseudomyrmecinae (Hym., Formicidae). Studia Entomologica 19: 45-66.
 * Kempf, W.W. 1972. Catalago abreviado das formigas da regiao Neotropical (Hym. Formicidae) Studia Entomologica 15(1-4).
 * Lattke J. E., M. Kaspari, S. ODonnell, and S. Powell. 2007. Las hormigas ecitoninas de Venezuela (Hymenoptera: Formicidae: Ecitoninae): elenco preliminar. Entomotropica Vol. 22(3): 153-170.
 * 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.
 * Li Z.h. 2006. List of Chinese Insects. Volume 4. Sun Yat-sen University Press
 * 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/
 * 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.
 * Luederwaldt H. 1918. Notas myrmecologicas. Rev. Mus. Paul. 10: 29-64.
 * Maes, J.-M. and W.P. MacKay. 1993. Catalogo de las hormigas (Hymenoptera: Formicidae) de Nicaragua. Revista Nicaraguense de Entomologia 23.
 * Mann W. M. 1925. New beetle guests of army ants. Journal of the Washington Academy of Sciences 15: 73-77.
 * 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.
 * Menozzi C. 1935. Spedizione del Prof. Nello Beccari nella Guiana Britannica. Hymenoptera-Formicidae. Redia. 21: 189-203.
 * Mirmecofauna de la reserva ecologica de San Felipe Bacalar
 * O'Donnell S., J. Lattke, S. Powell, and M. Kaspari. 2007. Army ants in four forests: geographic variation in raid rates and species composition. Journal of Animal Ecology 76: 580-589.
 * 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.
 * 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.N. and J. Valenzuela-Gonzalez. 2006. Las hormigas Ecitoninae (Hymenoptera: Formicidae) de Morelos, México. Revista Biologia Tropical 54(2):531-552
 * 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.
 * Santschi F. 1923. Solenopsis et autres fourmis néotropicales. Revue Suisse de Zoologie 30: 245-273.
 * Santschi F. 1929. Sur quelques nouvelles fourmis du Brésil (Hym. Form.). Deutsche Entomologische Zeitschrift 1928: 414-416.
 * Santschi F. 1931. Fourmis de Cuba et de Panama. Revista de Entomologia (Rio de Janeiro). 1: 265-282.
 * Solomon, S.E. and A.S. Mikheyev. 2005. The ant (Hymenoptera: Formicidae) fauna of Cocos Island, Costa Rica. Florida Entomologist 88(4):415-423
 * 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
 * Torres, I. L., M. A. G. Martinez, H. O. Rivera, M. E. G. Tovar, H. E. L. Espinosa, and J. M. Gonzalez. 2018. Comparacion de dos muestreos de hormigas del suelo en la barranca de Metlac, Fortin de las Flores, Veracruz, Mexico. Revista Cientifica UDO Agricola 10: 173-178.
 * Ulyssea M. A., and C. R. F. Brandao. 2013. Ant species (Hymenoptera, Formicidae) from the seasonally dry tropical forest of northeastern Brazil: a compilation from field surveys in Bahia and literature records. Revista Brasileira de Entomologia 57(2): 217224.
 * Ulysséa M. A., C. R. F. Brandão. 2013. Ant species (Hymenoptera, Formicidae) from the seasonally dry tropical forest of northeastern Brazil: a compilation from field surveys in Bahia and literature records. Revista Brasileira de Entomologia 57(2): 217-224.
 * Varela-Hernandez, F., M. Rocha-Ortega, R. W. Jones, and W. P. Mackay. 2016. Insectos: Hormigas (Formicidae) del estado de Queretaro, Mexico. Pages 397-404 in W. Jones., and V. Serrano-Cardenas, editors. Historia Natural de Queretaro. Universidad Autonoma de Queretaro, Mexico.
 * 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.
 * Vasquez-Bolanos M. 2011. Checklist of the ants (Hymenoptera: Formicidae) from Mexico. Dugesiana 18(1): 95-133.
 * Vittar, F. 2008. Hormigas (Hymenoptera: Formicidae) de la Mesopotamia Argentina. INSUGEO Miscelania 17(2):447-466
 * Vásquez-Bolaños M. 2011. Lista de especies de hormigas (Hymenoptera: Formicidae) para México. Dugesiana 18: 95-133
 * Watkins II, J.F. 1982.The army ants of Mexico (Hymenoptera: Formicidae: Ecitoninae). Journal of the Kansas Entomological Society 55(2): 197-247.
 * Watkins J. F., II 1976. The identification and distribution of New World army ants (Dorylinae: Formicidae). Waco, Texas: Baylor University Press, 102 pp
 * Watkins J. F., II 1977. The species and subspecies of Nomamyrmex (Dorylinae: Formicidae). Journal of the Kansas Entomological Society 50: 203-214.
 * Wheeler W. M. 1922. The ants of Trinidad. American Museum Novitates 45: 1-16.
 * Wheeler, G.C. and J. Wheeler. 1985. A checklist of Texas ants. Prairie Naturalist 17:49-64.
 * 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. "A catalogue of the ants of Paraguay (Hymenoptera: Formicidae)." Zootaxa 1622 (2007): 1-55.
 * do Nascimento, I.C. 2006. Fenologia dos Voos de Acasalamento em Formigas Tropicais