Carebara urichi

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Carebara urichi
Scientific classification
Kingdom: Animalia
Phylum: Arthropoda
Class: Insecta
Order: Hymenoptera
Family: Formicidae
Subfamily: Myrmicinae
Tribe: Crematogastrini
Genus: Carebara
Species: C. urichi
Binomial name
Carebara urichi
(Wheeler, W.M., 1922)

Carebara urichi casent0178633 profile 1.jpg

Carebara urichi casent0178633 dorsal 1.jpg

Specimen labels


Individuals and nest series have been collected from the leaf-litter using Winkler sifting.


A member of the Neotropical Carebara concinna species complex and Afrotropical Carebara concinna species complex.

Fischer et al. (2014) - Antennae with eleven segments. Major worker: Head almost as long as wide to slightly longer, nearly subquadrate in full-face view, posterolateral corners rounded. Head with longitudinal striations, rugulae on the posterior side, propodeal spines absent or inconspicuous, petiolar ventral process large, digitiform and anteriorly directed, gaster covered by very abundant erect hairs. Minor worker: Head subrectangular, sides convex, posterior margin almost straight, posterolateral corners angulate, frons medially smooth and shiny, face with longitudinal rugulae and reticulations, propodeal spines short, acute-triangular and upwardly directed, metatibia with long, suberect hairs along outer edge, petiole anteroventrally with small, anteriorly-pointing tooth, gaster with relatively abundant, long, suberect or subdecumbent hairs.

Carebara urichi can be confused with Carebara brevipilosa, but is easily separated by the sculpture on the dorsal promesonotum, which is irregularly longitudinally rugose to rugoreticulate with few irregular longitudinal rugae in C. urichi minor workers and weakly to superficially reticulate or with few very short rugulae in C. brevipilosa. Carebara urichi and C. brevipilosa are the only two species in the C. polita group recorded for the Neotropical Region.

Keys including this Species


This species is widely distributed in the neotropics, from México to Argentina.

Distribution based on Regional Taxon Lists

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

Distribution based on AntMaps


Distribution based on AntWeb specimens

Check data from AntWeb


Mainly found in rainforest and cloud forest at elevations ranging from 20–1470 m.


Wheeler (1937) reported: "This minute ant was previously known only from the type specimens collected in nests of the Guacharos (Steatornis), in a cave in Trinidad. Very recently Prof. A. S. Pearse has sent me two specimens which he found on bat guano in a cave in Yucatan. Its occurrence in leaf-mold in Cuba is not surprising since other minute cave-insects are known to live occasionally in similar situations, but we should expect it to be more abundant in some of the many bat-infested caves of the island."

Mann (1926) noted a colony of this ant, reported under the synonymized name Erebomyrma nevermanni, was collected in a rotten log (Reventazon, Santa Clara, Costa Rica) "in company with Pseudoponera stigma."

Life History Traits

  • Mean colony size: 180 (Wilson, 1986; Beckers et al., 1989)
  • Foraging behaviour: trunk trail (Wilson, 1986; Beckers et al., 1989)


Borgmeier (1949) described and illustrated a case of “dinergatogyny” from a Costa Rican specimen of O. morai.




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

  • urichi. Spelaeomyrmex urichi Wheeler, W.M. 1922c: 9, fig. 1 (w.) TRINIDAD.
    • Fernández, 2004a: 206 (s.q.m.).
    • Combination in Erebomyrma: Wilson, 1962a: 63; Wilson, 1986b: 61;
    • combination in Oligomyrmex: Ettershank, 1966: 124; Bolton, 1995b: 300;
    • combination in Carebara: Fernández, 2004a: 205.
    • Status as species: Wheeler, W.M. 1937b: 446; Wheeler, W.M. 1938: 252; Wilson, 1962a: 62; Ettershank, 1966: 124; Kempf, 1972a: 173; Brandão, 1991: 343; Bolton, 1995b: 300; Fernández, 2004a: 205 (redescription); Branstetter & Sáenz, 2012: 257; Fischer, et al. 2014: 99 (redescription); Bezděčková, et al. 2015: 115; Fernández & Serna, 2019: 822.
    • Senior synonym of eidmanni: Fernández, 2004a: 205; Fischer, et al. 2014: 99.
    • Senior synonym of morai: Fernández, 2004a: 205; Fischer, et al. 2014: 99.
    • Senior synonym of nevermanni: Fernández, 2004a: 205; Fischer, et al. 2014: 99.
  • eidmanni. Erebomyrma eidmanni Menozzi, in Eidmann, 1936a: 47 (s.w.q.) BRAZIL (Rio de Janeiro).
    • Combination in Erebomyrma: Wilson, 1962a: 63; Wilson, 1986b: 61;
    • combination in Oligomyrmex: Ettershank, 1966: 123; Bolton, 1995b: 299.
    • Status as species: Wilson, 1962a: 63; Ettershank, 1966: 123; Kempf, 1972a: 172; Wilson, 1986b: 61; Brandão, 1991: 343; Bolton, 1995b: 299.
    • Junior synonym of urichi: Fernández, 2004a: 205; Fischer, et al. 2014: 99.
  • morai. Erebomyrma morai Menozzi, 1931b: 271, fig. 7 (w.) COSTA RICA.
    • Borgmeier, 1949: 207 (ergatoid q.).
    • Combination in Erebomyrma: Wilson, 1962a: 63; Brandão, 1991: 343;
    • combination in Oligomyrmex: Ettershank, 1966: 124; Bolton, 1995b: 299.
    • Status as species: Borgmeier, 1949: 207; Wilson, 1962a: 63; Ettershank, 1966: 124; Kempf, 1972a: 172; Brandão, 1991: 343; Bolton, 1995b: 299.
    • Junior synonym of urichi: Fernández, 2004a: 205; Fischer, et al. 2014: 99.
  • nevermanni. Erebomyrma nevermanni Mann, 1926: 103 (w.) COSTA RICA.
    • Combination in Erebomyrma: Wilson, 1986b: 61;
    • combination in Oligomyrmex: Ettershank, 1966: 124; Bolton, 1995b: 299.
    • Status as species: Ettershank, 1966: 124; Kempf, 1972a: 172; Brandão, 1991: 343; Bolton, 1995b: 299.
    • Junior synonym of urichi: Fernández, 2004a: 205; Fischer, et al. 2014: 99.

Unless otherwise noted the text for the remainder of this section is reported from the publication that includes the original description.



Fischer et al. (2014) - Major (n=4): HW 0.97–0.99 (0.98), HL 1.01–1.06 (1.04), SL 0.41–0.44 (0.43), MDL 0.53–0.54 (0.53), EL 0.08, WL 0.97–1.02 (0.99), PNH 0.53–0.57 (0.54), PNW 0.56–0.61 (0.58), MNH 0.64–0.68 (0.66), PDH 0.42–0.46 (0.45), PTL 0.42–0.46 (0.44), PPL 0.25–0.26 (0.26), PTH 0.35–0.38 (0.37), PPH 0.26–0.30 (0.28), PTW 0.31–0.35 (0.33), PPW 0.38–0.43 (0.41), PSL 0.11–0.14 (0.13), MFL 0.58–0.61 (0.60), MTL 0.44–0.46 (0.45), CI 93–95 (94), SI 43–44 (44), MDI 54–55 (55), EI 8–9 (8), FI 60–62 (61), PSLI 12–14 (13), LPpI 88–96 (91), DPpI 152–167 (161), PpWI 124–132 (127), PpLI 57–60 (58), PpHI 74–80 (77).

Head slightly longer than wide (CI 93–95), in full-face view nearly subquadrate. Posterior margin of head with deep, V-shaped emargination, posterolateral corners rounded, sides subparallel and almost straight. Mandibles with five teeth. Anterior margin of clypeus nearly straight, sides convex. Frontal carinae absent to inconspicuous. Antenna with eleven segments, scapes short and reaching to about midlength of head (SI 43–44). Eyes present, multi-faceted, and comparatively large (EI 8–9).

In profile, promesonotum queen-like with moderately high pronotum, comparatively long scutum, and short scutellum, the latter slightly to strongly raised above convex to weakly convex dorsal outline of mesosoma. Promesonotal suture on dorsum present as broad impression, metanotum small and obliquely raised posteriorly. Propodeum higher than long in profile, with dorsal face oblique, declining posteriorly, propodeal spines and posterolateral lamella absent to reduced to slightly raised, posterior corners bluntly angulate, posterior declivity subvertical with well-developed lobe. Propodeal spiracle roundly convex and situated closer to dorsum of propodeum than to its center, not close to posterior declivity.

Petiole with relatively short peduncle, anteroventrally concave, with very conspicuous, short to relatively long, finger-like anterior process, posteroventrally convex, petiole node high, anteriorly oblique to weakly concave, posteriorly vertical and weakly concave, dorsum straight with a rounded angle anteriorly and a sharp right angle posteriorly. Postpetiole in profile squarish with rounded corners and lower than petiole (PpHI 74-80), with shallow, anteriorly angulate ventral process, anterior face almost vertical, dorsum almost straight and posterior face short and oblique. In dorsal view, petiole node much wider than long and anteroposteriorly flattened, anteriorly slightly convex, posterioly transverse and sharply marginate, postpetiole with wide, lamellate processes extending lateroventrally (PpWI 124-132), the node itself narrower than petiole and slightly wider than long, with convex sides, and anterior and posterior margins almost transverse.

Mandibles smooth and shiny, laterally weakly striate and with scattered punctures. Clypeus smooth and shiny, with several weak lateral carinae. Sculpture on frons and anterior sides of head with strong, irregular rugae, posterior parts of face coarsely rugoreticulate, interspaces weakly to superficially punctate, ventral side of head with long, parallel, longitudinal rugae. Mesosoma mostly coarsely and very irregularly rugoreticulate with punctate ground sculpture, rugoreticulate sculpture can be reduced on lateral surfaces, anepisternum sometimes with smooth and shiny area posteriorly. Petiole node and postpetiole dorsally smooth and shiny to faintly punctate, gaster shagreened or superficially punctate. Head and body usually with abundant, erect to suberect hairs of varying length and shorter decumbent to suberect pilosity, scapes and tibiae with appressed to decumbent pilosity and long suberect to erect hairs along outer edge. Color reddish brown, appendages and gaster light brown.

Minor (n=5): HW 0.36–0.44 (0.40), HL 0.41–0.51 (0.46), SL 0.29–0.34 (0.32), MDL 0.23–0.29 (0.26), EL 0.02, WL 0.45–0.59 (0.53), PNH 0.17–0.22 (0.19), PNW 0.23–0.27 (0.25), MNH 0.25–0.31 (0.28), PDH 0.17–0.22 (0.19), PTL 0.15–0.18 (0.16), PPL 0.11–0.12 (0.11), PTH 0.09–0.11 (0.10), PPH 0.08–0.09 (0.08), PTW 0.07–0.08 (0.08), PPW 0.08–0.10 (0.09), PSL 0.07–0.08 (0.07), MFL 0.30–0.39 (0.34), MTL 0.26–0.32 (0.29), CI 86–88 (87), SI 76–91 (81), MDI 62–66 (65), EI 4–6 (5), FI 80–91 (84), PSLI 17–19 (18), LPpI 139–160 (147), DPpI 69–81 (75), PpWI 109–122 (115), PpLI 67–80 (71), PpHI 71–83 (77).

Head almost as wide as long (CI 93–95), posterior margin nearly straight to weakly concave, sides convex. Mandible with four well-defined teeth, apical and preapical tooth larger than others. Anterior margin of clypeus very weakly concave, at each side with a triangular tooth. Frontal carinae moderately long, sometimes reaching posterior third of head. Antenna with eleven segments, scape short and reaching about 6/7 towards posterior margin of head (SI 76–91). Eyes consisting of one ommatidium (EI 4–6).

In profile, dorsum of promesonotum weakly convex, anteriorly sharply marginate and right-angled, posteriorly curving slightly downwards toward widely, but shallowly impressed metanotal groove. Dorsum of propodeum convex, softly declining posteriorly and slightly shorter than posterior declivity, anterodorsal corner convex, propodeal spines relatively short, acute (PSLI 14–15) and lamellate, lamella proceeding ventrally, ending in large propodeal lobes. Propodeal spiracle rounded and situated close to base of spines.

Petiole in profile with peduncle about as long as petiole node, anteroventrally concave, with small, acute, anteriorly pointing ventral process, posteroventrally weakly convex, petiole node low and dorsally rounded. Postpetiole in profile dorsally convex, ventrally almost straight, on average 1.5 times longer than high (LPpI 139–160), lower than petiole (PpHI 71–83). In dorsal view, petiole node almost as wide as long, anteriorly roundly convex, posteriorly nearly straight or weakly convex, postpetiole longer than wide (DPpI 69–81), slightly wider than petiole (PpWI 109–122), anteriorly tapering and posteriorly weakly concave. Gaster slender in dorsal view, its anterior margin straight to faintly convex.

Mandibles, clypeus and center of frons smooth and shiny, remainder of face coarsely rugoreticulate, with scattered punctures. Promesonotum dorsally with several irregular longitudinal rugae, interspaces superficially punctate to smooth, remainder of mesosoma alveolate, alveolae on basal lateropronotum sometimes strongly effaced. Gaster and dorsum of petiole and postpetiole smooth and shiny, remainder of waist segments weakly alveolate.

Head and body with long, suberect to erect hairs and shorter subdecumbent to decumbent pilosity. Scapes and tibiae with decumbent to subdecumbent pilosity, the latter with longer suberect hairs along outer edge. Color orange to dark orange with lighter colored antennae, legs and gaster.


Fernández (2004) - (n=2): HW 1.17-1.18; HL 1.15-1.25; SL 0.51-0.56; EL 0.24-0.25; WL 1.69-1.75; GL 1.93; TL 6.02-6.06; CI 95-100; SI 44-47.

General habitus as soldier with several differences: Posterior border of head slightly concave. Mandible with 5 tooth, with a space between subapical and remainder. Eyes big, bulging. Three ocellae present. Mesosoma modified as in myrmicine queens. Metanotum sharply defined. Propodeum armed with triangular teeth. Propodeal spiracle big, round, backward. Subpetiolar process well-defined. Sculpture and general pilosity as soldier.


Fernández (2004) - HW 0.78; HL 0.63; SL 0.14; EL 0.34; WL 1.33; GL 1.17; TL 3.81; CI 124; SI 18.

Head subsphaerical. Eyes large. Mandibles narrow, not touching when closed, with two teeth, the apical twice as big as basal. Clypeus medially bulging. Ocellae turreted, the frontal round and prominent. Mesosternum bulging. Propodeum unarmed. Petiole larger that postpetiole, their peduncle running evenly in the dorsal face of node. Petiole ventrally without subpetiolar process, and with several irregular longitudinal carinae. Postpetiole low, broadly attached to the gaster. Genitalia strongly protruding, with aedeagus and parameres long and narrow. Body smooth and shining. Posterior half of head with transverse striation, anterior half more or less longitudinal striate. Most of promesonotum striatopunctate. Mesosoma punctate. Body dark brown, appendanges lighter. Wings dark.

Type Material

Fischer et al. (2014) - Spelaeomyrmex urichi W.M. Wheeler, 1922b: 9. Syntypes (11 minor workers) TRINIDAD: “Guacharo Cave” (F.M. Urich) [not examined].


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
  • 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.
  • Alonso L. E. 2010. A preliminary survey of the ants of the Kwamalasamutu region, SW Suriname. In: O’Shea, B.J., L.E. Alonso, & T.H. Larsen, (eds.). 2011. A Rapid Biological Assessment of the Kwamalasamutu region, Southwestern Suriname. RAP Bulletin of Biological Assessment 63. Conservation International, Arlington, VA.
  • 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.
  • 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.
  • 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
  • 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.
  • Dattilo W. et al. 2019. MEXICO ANTS: incidence and abundance along the Nearctic-Neotropical interface. Ecology
  • 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.
  • Donoso D. A. 2014. Assembly mechanisms shaping tropical litter ant communities. Ecography 37 doi: 10.1111/j.1600-0587.2013.00253.x
  • Ettershank G. 1966. A generic revision of the world Myrmicinae related to Solenopsis and Pheidologeton (Hymenoptera: Formicidae). Aust. J. Zool. 14: 73-171.
  • 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. 2004. The American species of the myrmicine ant genus Carebara Westwood (Hymenoptera: Formicidae). Caldasia 26: 191-238.
  • Fernández F., and E. E. Palacio. 1995. Hormigas de Colombia IV: nuevos registros de géneros y especies. Caldasia 17: 587-596.
  • 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.
  • Fischer G., F. Azorsa, F. Fernandez, and B. L. Fisher. 2014. The ant genus Carebara Westwood (Hymenoptera, Formicidae): synonymisation of Pheidologeton Mayr under Carebara, establishment and revision of the C. polita species group. Zookeys doi: 10.3897/zookeys.@@.7922
  • Fontanla Rizo J.L. 1997. Lista preliminar de las hormigas de Cuba. Cocuyo 6: 18-21.
  • Fontenla J. L., and J. Alfonso-Simonetti. 2018. Classification of Cuban ants (Hymenoptera: Formicidae) into functional groups. Poeyana Revista Cubana de Zoologia 506: 21-30.
  • Fontenla Rizo J. L. 1997. Lista preliminar de las hormigas de Cuba (Hymenoptera: Formicidae). Cocuyo 6: 18-21.
  • 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-Martinez M. A., V. Vanoye-Eligio, O. R. Leyva-Ovalle, P. Zetina-Cordoba, M. J. Aguilar-Mendez, and M. Rosas-Mejia. 2019. Diversity of ants (Hymenoptera: Formicidae) in a sub-montane and sub-tropical cityscape of Northeastern Mexico. Sociobiology 66(3): 440-447.
  • 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.
  • Jacquemin J., T. Drouet, T. Delsinne, Y. Roisin, and M. Leponce. 2012. Soil properties only weakly affect subterranean ant distribution at small spatial scales. Applied Soil Ecology 62: 163-169.
  • 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. Labidus coecus as a cave ant. Studia Entomologica 4: 551-552.
  • 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).
  • Kugler C. 1986. Stings of ants of the tribe Pheidologetini (Myrmicinae). Insecta Mundi 1: 221-230.
  • Kusnezov N. 1963. Zoogeografia de las hormigas en sudamerica. Acta Zoologica Lilloana 19: 25-186
  • 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. 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. T., and R. K. Colwell. 2011. Density compensation, species composition, and richness of ants on a neotropical elevational gradient. Ecosphere 2(3): 16pp.
  • Longino J. et al. ADMAC project. Accessed on March 24th 2017 at
  • Longino, J.T. 2010. Personal Communication. Longino Collection Database
  • Maes, J.-M. and W.P. MacKay. 1993. Catalogo de las hormigas (Hymenoptera: Formicidae) de Nicaragua. Revista Nicaraguense de Entomologia 23.
  • Mann W. M. 1926. Some new neotropical ants. Psyche (Cambridge) 33: 97-107.
  • 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.
  • Menozzi C. 1931. Contribuzione alla conoscenza del microgenton di Costa Rica. III. Hymenoptera - Formicidae. Bollettino del Laboratorio di Zoologia Generale e Agraria della Reale Scuola Superiore d'Agricoltura. Portici. 25: 259-274.
  • Mertl A. L., J. F. A. Traniello, K. Ryder Wilkie, and R. Constantino. 2012. Associations of two ecologically significant social insect taxa in the litter of an amazonian rainforest: is there a relationship between ant and termite species richness? Psyche doi:10.1155/2012/312054
  • Mirmecofauna de la reserva ecologica de San Felipe Bacalar
  • Neves F. S., K. S. Queiroz-Dantas, W. D. da Rocha, and J. H. C. Delabie. 2013. Ants of Three Adjacent Habitats of a Transition Region Between the Cerrado and Caatinga Biomes: The Effects of Heterogeneity and Variation in Canopy Cover. Neotrop Entomol 42: 258–268.
  • Pacheco R., and H. L. Vasconcelos. 2007. Invertebrate conservation in urban areas: ants in the Brazilian Cerrado. Landscape and Urban Planning 81: 193–199.
  • Pacheco R., and H. L. Vasconcelos. 2012. Subterranean Pitfall Traps: Is ItWorth Including Them in Your Ant Sampling Protocol? Psyche doi:10.1155/2012/870794
  • 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.
  • Ramos L. S., R. Z. B. Filho, J. H. C. Delabie, S. Lacau, M. F. S. dos Santos, I. C. do Nascimento, and C. G. S. Marinho. 2003. Ant communities (Hymenoptera: Formicidae) of the leaf-litter in cerrado “stricto sensu” areas in Minas Gerais, Brazil. Lundiana 4(2): 95-102.
  • 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.
  • 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.
  • Ryder Wilkie K.T., A. L. Mertl, and J. F. A. Traniello. 2010. Species Diversity and Distribution Patterns of the Ants of Amazonian Ecuador. PLoS ONE 5(10): e13146.doi:10.1371/journal.pone.0013146
  • Schmidt F. A., and R. R. C. Solar. Is it important to collect hypogaeic ants? How to collect them? Biológico, São Paulo 69(2): 267-270.
  • Schmidt, F.A. and R.R.C Solar. 2010. Hypogaeic pitfall traps: methodological advances and remarks to improve the sampling of a hidden ant fauna. Insectes Sociaux 57:261-266.
  • 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
  • Siqueira de Castro F., A. B. Gontijo, P. de Tarso Amorim Castro, and S. Pontes Ribeiro. 2012. Annual and Seasonal Changes in the Structure of Litter-Dwelling Ant Assemblages (Hymenoptera: Formicidae) in Atlantic Semideciduous Forests. Psyche doi:10.1155/2012/959715
  • Siqueira de Castro F., A. B. Gontijo, W. Duarte da Rocha, and S. Pontes Ribeiro. 2011. As comunidades de formigas de serapilheira nas florestas semidecíduas do Parque Estadual do Rio Doce, Minas Gerais. MG.BIOTA, Belo Horizonte 3(5): 5-24.
  • 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.
  • 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.
  • 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 W. M. 1937. Ants mostly from the mountains of Cuba. Bulletin of the Museum of Comparative Zoology. 81: 439-465.
  • Wilson E. O. 1986. Caste and division of labor in Erebomyrma, a genus of dimorphic ants (Hymenoptera: Formicidae: Myrmicinae). Insectes Sociaux 33: 59-69.