Cephalotes pusillus

C. pusillus is one of the commonest and most widespread species of the genus, though its distribution is narrower than that of other common species like Cephalotes minutus and Cephalotes atratus. (de Andrade and Baroni Urbani 1999)

Identification
A member of the pusillus clade characterised in the worker and soldier by the less regular body foveae and by the narrow, semi-transparent gastral lamellae. (de Andrade and Baroni Urbani 1999)

Distribution based on Regional Taxon Lists
Neotropical Region: Argentina, Bolivia, Brazil, Colombia, Ecuador, French Guiana, Guyana, Paraguay, Peru, Suriname, Trinidad and Tobago, Venezuela.

Biology
DaRocha et al. (2015) studied the diversity of ants found in bromeliads of a single large tree of Erythrina, a common cocoa shade tree, at an agricultural research center in Ilhéus, Brazil. Forty-seven species of ants were found in 36 of 52 the bromeliads examined. Bromeliads with suspended soil and those that were larger had higher ant diversity. Cephalotes pusillus was found in 5 different bromeliads but was associated with twigs and bark cavities, rather than suspended soil or litter, of the plants.

Koch et al. (2018) sampled this species in Caryocar barsiliense trees, in southeastern Brazil cerrado, as part of a study examining species interactions in ant-plants.

Spider Mimic and Predator
Aphantochilus rogersi is an an-mimicking spider that preys exclusively on cephalotine ants. The spiders oviposit in close proximity to nests of the model ant Cephalotes pusillus, and emergent spiderlings tend to remain in the vicinity of natal egg sacs. Females of A. rogersi actively defend their egg sacs against approaching workers of C. pusillus, but the latter may sometimes destroy the eggs. Feeding specialization on these ants is confirmed by more than 300 observations of young and adult A. rogersi carrying ant corpses in the field. Although A. rogersi possesses several behavioural traits which may reduce the risk of being injured by ants during subjugation, field and laboratory observations showed that social defence by C. pusillus may cause mutilation to the spiders. Tests in captivity revealed an ontogenetic change in the prey-capture techniques employed by A. rogersi. Early-instar spiderlings can apparently only seize the ant’s petiole tightly if they are able to approach the ant from the front. As the ant is paralysed, the spiderling positions itself vertically in relation to the substratum. Larger spiders, on the other hand, attack ants most frequently from behind, and seem better equipped to seize the ant’s petiole firmly with their larger chelicerae. Owing to their greater strength, late-instar spiders are able to lift the struggling ant aloft. The selection of a suitable oviposition site, the mother’s ability to defend herself and the eggs from nearby ants, and the capacity to capture and subdue ants safely from emergence to maturity, are regarded as crucial traits inherent in the mimetic and feeding specialization by A. rogersi. (Castanho & Oliveira, 1997)

Worker
Minor

Images from AntWeb
Major

Nomenclature

 * . Cryptocerus pusillus Klug, 1824: 202 (w.) BRAZIL (Pará, Rio de Janeiro), GUYANA.
 * Type-material: syntype workers (number not stated).
 * Type-localities: Brazil: Pará (“ex Museo de Hoffmannsegg”), Brazil: Rio de Janeiro (“sent by Dr de Olfers”), Guyana: Demerara (“from Hemmerich”).
 * Type-depository: MNHU (perhaps also in MRSN).
 * Emery, 1890b: 75 (s.q.); Emery, 1896h: 636 (m.); Wheeler, G.C. & Wheeler, J. 1954b: 156 (l.).
 * Combination in Cryptocerus (Paracryptocerus): Emery, 1915i: 192;
 * combination in Paracryptocerus: Kempf, 1951: 176;
 * combination in Paracryptocerus (Paracryptocerus): Kempf, 1958a: 14;
 * combination in Zacryptocerus: Brandão, 1991: 388;
 * combination in Cephalotes: Baroni Urbani, 1998: 316; De Andrade & Baroni Urbani, 1999: 172.
 * Status as species: Guérin-Méneville, 1844a: 426; Smith, F. 1853: 220; Smith, F. 1858b: 190; Smith, F. 1862d: 410; Mayr, 1863: 406; Roger, 1863b: 38; Dalla Torre, 1893: 144; Forel, 1895b: 134; Emery, 1896h: 636; Forel, 1905b: 158; Emery, 1906c: 170; Forel, 1906d: 234; Forel, 1907e: 3; Forel, 1908c: 354; Forel, 1908e: 69; Forel, 1909a: 257; Forel, 1912e: 201; Bruch, 1914: 218; Wheeler, W.M. 1916c: 12; Wheeler, W.M. 1916d: 326; Mann, 1916: 450; Crawley, 1916b: 371; Santschi, 1916e: 383; Luederwaldt, 1918: 41; Wheeler, W.M. 1918b: 26; Santschi, 1921h: 127; Wheeler, W.M. 1922c: 11; Wheeler, W.M. 1923a: 4; Emery, 1924d: 307; Wheeler, W.M. 1925a: 36; Borgmeier, 1927c: 116; Menozzi, 1935b: 197; Eidmann, 1936b: 83; Wheeler, W.M. 1942: 209; Kempf, 1951: 176 (redescription); Kempf, 1958a: 14; Kempf, 1961b: 514; Kempf, 1972a: 179; Brandão, 1991: 388; Bolton, 1995b: 427; De Andrade & Baroni Urbani, 1999: 172 (redescription); Wild, 2007b: 32; Bezděčková, et al. 2015: 116; Sandoval-Gómez & Sánchez-Restrepo, 2019: 913.
 * Senior synonym of brevispinosa: Kempf, 1951: 176; Kempf, 1958a: 14; Kempf, 1972a: 180; Brandão, 1991: 388; Bolton, 1995b: 427; De Andrade & Baroni Urbani, 1999: 172.
 * Senior synonym of caustica: Emery, 1924d: 307; Borgmeier, 1927c: 116; Kempf, 1951: 176; Kempf, 1958a: 14; Kempf, 1972a: 180; Brandão, 1991: 388; Bolton, 1995b: 427; De Andrade & Baroni Urbani, 1999: 172.
 * Senior synonym of elongatus: Roger, 1862c: 297; Mayr, 1863: 406; Roger, 1863b: 38; Dalla Torre, 1893: 144; Forel, 1895b: 134; Emery, 1924d: 307; Borgmeier, 1927c: 116; Kempf, 1951: 176; Kempf, 1972a: 180; Brandão, 1991: 388; Bolton, 1995b: 427; De Andrade & Baroni Urbani, 1999: 172.
 * Senior synonym of obtusus: Forel, 1911e: 262; Emery, 1924d: 307; Borgmeier, 1927c: 116; Kempf, 1951: 176; Kempf, 1972a: 180; Brandão, 1991: 388; Bolton, 1995b: 427; De Andrade & Baroni Urbani, 1999: 172.
 * Material of the unavailable name rufescens referred here by Kempf, 1951: 177; Kempf, 1958a: 14; Kempf, 1972a: 180; Brandão, 1991: 388.
 * Distribution: Argentina, Bolivia, Brazil, Colombia, Ecuador, Guyana, Paraguay, Peru, Trinidad, Venezuela.
 * brevispinosa. Cryptocerus pusillus var. brevispinosa Santschi, 1921h: 127, fig. 2 (w.) ARGENTINA (Misiones, Córdoba), PARAGUAY, BRAZIL (Rio de Janeiro).
 * Type-material: syntype workers (number not stated).
 * Type-localities: Argentina: Misiones, vic. San Ignacio, Villa Lutecia (E.R. Wagner), Argentina: Cerro Cora (von Steiger), Argentina: Córdoba (C. Bruch), Paraguay: (no further data) (Fiebrig), Brazil: Rio de Janeiro, Alto da Boa Vista (E.R. Wagner).
 * Type-depository: NHMB.
 * Subspecies of pusillus: Borgmeier, 1927c: 116.
 * Junior synonym of pusillus: Kempf, 1951: 176; Kempf, 1958a: 14; Kempf, 1972a: 180; Brandão, 1991: 388; Bolton, 1995b: 424; De Andrade & Baroni Urbani, 1999: 172.
 * caustica. Formica caustica Kollar, in Pohl & Kollar, 1832: 17, fig. 12 (w.) BRAZIL (no state data).
 * Type-material: holotype (?) worker.
 * [Note: no indication of number of specimens is given.]
 * Type-locality: Brazil: (no further data).
 * Type-depository: unknown.
 * [Note: neither Pohl nor Kollar are mentioned in Horn & Kahle, 1935, 1936.]
 * Combination in Cryptocerus: Guérin-Méneville, 1844a: 426.
 * Status as species: Smith, F. 1853: 222; Smith, F. 1858b: 191; Mayr, 1862: 764; Smith, F. 1862d: 410; Mayr, 1863: 405; Roger, 1863b: 38; Emery, 1878a: ix (in list).
 * Junior synonym of minutus: Emery, 1890b: 74; Dalla Torre, 1893: 143; Forel, 1895b: 134; Forel, 1899c: 50.
 * Junior synonym of pusillus: Emery, 1924d: 307; Borgmeier, 1927c: 116; Kempf, 1951: 176; Kempf, 1958a: 14; Kempf, 1972a: 180; Brandão, 1991: 388; Bolton, 1995b: 424; De Andrade & Baroni Urbani, 1999: 172.
 * [Note: De Andrade & Baroni Urbani, 1999: 172, 194, enter caustica as a junior synonym under both pusillus and minutus. It rests here as a junior synonym of pusillus, though its true identity may never be known.]
 * elongatus. Cryptocerus elongatus Klug, 1824: 214 (q.) BRAZIL (Pará, Rio de Janeiro), GUYANA.
 * Type-material: syntype queens (number not stated).
 * Type-localities: Brazil: Pará (“sent by Sieber, ex Mus. Com. De Hoffmannsegg”), Brazil: Rio de Janeiro (“sent by Sello”), Guyana: Demerara (“from Hemmerich”).
 * Type-depository: MNHU.
 * Smith, F. 1860c: 75 (w.); Smith, F. 1862d: 409 (s.).
 * Status as species: Guérin-Méneville, 1844a: 426; Smith, F. 1853: 218; Smith, F. 1858b: 190; Smith, F. 1860c: 75; Smith, F. 1862d: 409.
 * Junior synonym of pusillus: Roger, 1862c: 297; Mayr, 1863: 406; Roger, 1863b: 38; Dalla Torre, 1893: 144; Forel, 1895b: 134; Emery, 1924d: 307; Borgmeier, 1927c: 116; Kempf, 1951: 176; Kempf, 1972a: 180; Brandão, 1991: 388; Bolton, 1995b: 425; De Andrade & Baroni Urbani, 1999: 172.
 * obtusus. Cryptocerus obtusus Smith, F. 1858b: 191 (w.) BRAZIL (Pará).
 * Type-material: holotype (?) worker.
 * [Note: no indication of number of specimens is given.]
 * Type-locality: Brazil: Santarem (H.W. Bates).
 * [Note: De Andrade & Baroni Urbani, 1999: 172, record a second syntype (in ZSBS) from Amazonas, but this locality is not mentioned in the original description.]
 * Type-depository: BMNH.
 * Status as species: Smith, F. 1862d: 410; Mayr, 1862: 764; Mayr, 1863: 406; Roger, 1863b: 38; Dalla Torre, 1893: 143; Forel, 1895b: 134.
 * Junior synonym of pusillus: Forel, 1911e: 262; Emery, 1924d: 307; Borgmeier, 1927c: 116; Kempf, 1951: 176; Kempf, 1972a: 180; Brandão, 1991: 388; Bolton, 1995b: 426; De Andrade & Baroni Urbani, 1999: 172.

Type Material


de Andrade and Baroni Urbani (1999):

Worker. Type locality: South America. Type material: Syntype workers presumably in the Museum fur Naturkunde der Humboldt-Universitat Berlin (Roger, 1862: 297), and in the Museo regionale di Scienze Naturali, Torino (Casolari & Casolari Moreno, 1980: 84) (not available for the present study).

Cryptocerus elongates Gyne. Type locality: South America. Type material: a gyne presumably in the Museum fur Naturkunde der Humboldt-Universitat Berlin (Roger, 1862: 297) (not available for the present study).

Formica caustica Worker. Type locality: Brazil. Type material not available for the present study. Synonymy by De Dalla Torre, 1893: 143. Note: synonym of minutus in Emery (1890) and Forel (1895).

Cryptocerus obtusus Soldier. Type locality: Santarem (Brazil). Type material: a syntype worker labelled "Santarem, 54 63, 1 74, C. obtusus (type) Sm.", other label (probably more recent): "pusillus, Rio", examined; a syntype worker labelled "Amazonas, H. W. Bates" in the (Forel, 1911b: 262), examined.

Cryptocerus pusillus var. brevispinosa Worker. Type localities: S. Ignacio, Villa Lutecia, Cerro Cora in the govern. of Misiones, Cordoba (Argentina), Paraguay, Alto da Boa Vista (Rio de Janeiro, Brazil). Type material: 3 syntype workers from Cordoba, Misiones and Paraguay in (examined).

Worker
Kempf (1951) - Length 4.4 mm. Median head length 1.14 mm; Weber's length of thorax 1.29 mm. Black; the following ferruginous: apex of mandibles, anterior portion of frontal carinae, apex of last funicular segment, apex of femora and basitarsi, distal half of the following tarsal segments, anterolateral border of first gastral tergite, apex of lateral projections of peduncular segments.

Head subopaque, about as long as broad; interocular distance subequal to maximum length of head (57:58). Sides of head subparallel, posterior margin between the internal corners of the short, obliquely truncate, infuscated occipital lamellae evenly curved and emarginate. Apical border of occipital lamellae straight. Cheeks strongly marginate beneath, densely covered with flat, silvery hairs, which lie in grooves on the posterior portion of the vertex. Eyes small, their maximum diameter less than 1 /3 of the median head length.

Thorax opaque. Anterior border arcuate, shoulders angulate. Pronotum somewhat expanded laterally behind into a narrow, denticulate crest, with two short teeth on each side, the posterior tooth distinctly shorter than the anterior; posterior corner of pronotum rectangular and projecting. Promesonotal suture distinct. Mesonotum on each side with a very small, subrectangular tooth. Mesoepinotal suture vestigial, obsolete mesad. Basal face of epinotum in the same plane as mesonotum, transverse, very short, with a short lateral tooth, longer than the second pronotal tooth, and a very long, subcylindrical posterior spine arising from the posterior corner, diverging about 45° degrees from the longitudinal axis. Dorsum of thorax moderately both longitudinally and transversely convex. Declivous face not excavate mesad, nor crested laterad. The entire thorax finely reticulate-punctate, with rather dense, sharply impressed, more or less elongate foveolae, each containing an appressed silvery, elongate glistening scale. Declivous face without nacrosculpture and without scales. Laterotergite of pronotum and mesopleura more or less regularly rugose and sparsely scaled.

Petiole opaque, transverse, with a very small somewhat recurved lateral denticule. Postpetiole opaque, as broad as petiole, with a broad, apically truncate, posteriorly subdentate-projection on each side, curving slightly forward. Both peduncular segments finely reticulatepunctate with sparse, glistening appressed hairs.

Gaster opaque, elliptical, emarginate anteriorly mesad. First gastral tergite with a very narrow, not distinctly set off nor crested antero-Iateral border. Tergites and sternites finely and rather sharply reticulate-punctate. Scales of gaster sparser, more slender than on head and thorax, not lying in distinctly excavated foveolae. Erect pile confined to mandibles and to the 2-4th tergites and the sternites of the gaster.

de Andrade and Baroni Urbani (1999) - Measurements (in mm) and indices: TL 3.84-5.52; HL 0.98-1.32; HW 1.24-1.76; EL 0.30-0.36; PW 0.92-1.38; PeW 0.37-0.57; PpW 0.44-0.66; HBaL 0.46-0.58; HBaW 0.10-0.15; CI 121.6-133.3; PI 127.5-134.8; PPeI 200.0-259.4; PPpI 204.2-209.1; HBaI 20.4-25.9.

Soldier
Kempf (1951) - Length 7.5 mm. Median head length 1 .95 mm; Weber's length of thorax 2.12 mm. Black; the following ferruginous: the four apical tarsal segments of the fore legs and the two apical segments of the mid and hind legs. Tips of last funicular segments orange-brown.

Head subopaque, somewhat shorter than wide. Frontal carinae strongly converging anteriorly, prolonged behind above the eyes in the form of a raised carinule, fading out before reaching the occipital corner. Upper surface of head distinctly convex, vertex with a pair of faint, not tuberculate median swellings on the posterior border. Occiput truncate, immarginate above; occipital angles form a scarcely projecting blunt, strong tubercle. Cheeks not marginate beneath in front; a vestigial margination beneath the eye is present. Eyes very little convex. Upper and lower surface of head finely and shallowly reticulate, the upper surface with rather dense, small foveolae in which a minute, not shiny, scale is visible. foveolae slightly more crowded, longer on occiput. Still larger and more deeply impressed on occiput and lower surface of head, containing a short decumbent, somewhat glistening seta.

Thorax subopaque. Anterior border arcuate, shoulder subangulate. Sides of pronotum diverging until reaching the lateral end of the transverse crest, having before the crest a stout, apically rounded tooth, projecting obliquely forwars and laterad. Transverse pronotal crest broadly interrupted mesad, and obsolescent towards the sides. Lateral border of pronotum angulate at the crest, and converging posteriorly behind the crest. Promesonotum, in profile, greatly convex. Promesonotal suture more or less distinct laterad, vestigial mesad. Sides of mesonotum subparallel with a small posterior, acute, lateral tooth. Mesoepinotal suture impressed and distinct. Basal face of epinotum transversely convex, with a small, blunt tooth antero- Iaterad, and a much longer and stronger, blunt spine on the posterior corner, slightly more divergent than in worker. Entire thorax finely and sharply reticulate-punctate. Dorsum of thorax and laterotergite of pronotum covered with rather dense, elongate deeply impressed setiferous foveolae. declivous face and posterior portion of the sides of the thorax with more or less vestigial, irregular, fine rugosities.

Petiole opaque, transverse, its sides subparallel, with a minute denticule. Postpetiole opaque, slightly broader than petiole, with a tapering, plate-like lateral projection, the apex of which is rounded in front, sub dentate behind, curving slightly forward. Both segments finely reticulate-punctate, covered with vestigial foveolae and rugosities.

Gaster subopaque, elliptical, finely reticulate-punctate, not foveolate, deeply emarginate in front mesad. First gastral tergite anterolaterally sharply marginate, not forming a lamellate border.

All foveolae bear a decumbent seta, which is slightly flattened and glistening on thorax and peduncle. Erect setae on the terminal tergitcs and on the sternites of the gaster. First gastral tergite covered with minute, decumbent setae laterad, almost scaleless discad.

de Andrade and Baroni Urbani (1999) - Measurements (in mm) and indices: TL 6.44-7.76; HL 1.60-1.90; HW 2.24-2.36; EL 0.45-0.48; PW 1.80-2.04; PeW 0.62-0.78; PpW 0.71-0.89; HBaL 0.64-0.66; HBaW 0.18-0.19; CI 124.2-130.0; PI 115.5-115.7; PPeI 261.5-290.3; PPpI 229.2-253.5; HBaI 28.1-28.8.

Queen
Kempf (1951) - Length 10.5 mm. Median head length 2.13 mm; interocular width 2.27 mm; Weber's length of thorax 3. 1 7 mm. Black; tip of last funicular segment fuscous ferruginous. In general resembling minutus, from which it differs by the appreciably larger size, the black coloration of the appendages, the lack of the yellow maculae on the first gastral tergite. The head is shorter than wide, the upper face subopaque, greatly convex, with a distinct carinule laterad, below the eyes. Occiput subtruncate, immargin ate above. Pronotum with a distinct transverse crest. Sides of petiole without a spine, or tooth; subparallel. Sides of thorax more extensively foveolate.

de Andrade and Baroni Urbani (1999) - Measurements (in mm) and indices: TL 9.77-11.16; HL 1.76-1.96; HW 2.10-2.36; EL 0.45-049; PW 1.96-2.26; PeW 0.87-0.91; PpW 0.99-1.12; HBaL 0.72-0.75; HBaW 0.19; CI 119.3-127.3; PI 103.7-107.1; PPeI 222.7-248.3; PPpI 198.0-201.8; HBaI 25.3-26.4.

References based on Global Ant Biodiversity Informatics

 * 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.
 * Andrade T., G. D. V. Marques, K. Del-Claro. 2007. Diversity of ground dwelling ants in Cerrado: an analysis of temporal variations and distinctive physiognomies of vegetation (Hymenoptera: Formicidae). Sociobiology 50(1): 121-134.
 * Antoniazzi R., R. N. S. L. Garoo, W. Dattilo, S. P. Ribeiro, and F. S. Neves. 2019. Ant species richness and interactions in canopies of two distinct successional stages in a tropical dry forest. The Science of Nature 106: 20
 * Araujo L. M., and G. W. Fernandes. 2003. Altitudinal patterns in a tropical ant assemblage and variation in species richness between habitats. Lundiana 4(2): 103-109.
 * Arruda F. V., M. A. Pesquero, D. G. Marcelino, G. A. Leiter, J. H. C. Delabie, and R. Fagundes. 2015. Size and condition of bamboo as structural factors behind the vertical stratification of the bamboo-nesting ant community. Insectes Sociaux DOI 10.1007/s00040-015-0440-4
 * 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., R. Silvestre, and A. Reis-Menezes> Influencia das interacoes comportamentais entre especies de formigas em levantamentos faunisticos em communidades de Cerrado> pp 371-404> In martins R. P., T. M. Lewinsohn, and M. S. Barbeitos (eds). Ecologia e comportamento de insetos. Serie Oecologia Brasiliensis, vol. VIII. PPGE-URFJ, Rio de Janeiro, Brasil.
 * Brandao, C.R.F. 1991. Adendos ao catalogo abreviado das formigas da regiao neotropical (Hymenoptera: Formicidae). Rev. Bras. Entomol. 35: 319-412.
 * Bruch C. 1914. Catálogo sistemático de los formícidos argentinos. Revista del Museo de La Plata 19: 211-234.
 * Byk J., K. Del-Claro. 2010. Nectar- and pollen-gathering Cephalotes ants provide no protection against herbivory: a new manipulative experiment to test ant protective capabilities. Acta Ethol. 13: 33-38.
 * Bächtold A., K. Del-Claro, L. A. Kaminski, A. V. L. Freitas, and P. S. Oliveira. 2012. Natural history of an ant-plant-butterfly interaction in a Neotropical savanna. Journal of Natural History 46: 943-954.
 * Caldart V. M., S. Iop, J. A. Lutinski, and F. R. Mello Garcia. 2012. Ants diversity (Hymenoptera, Formicidae) of the urban perimeter of Chapecó county, Santa Catarina, Brazil. Revista Brasileira de Zoociências 14 (1, 2, 3): 81-94.
 * Carvalho K. S., M. A. F. Carneiro, I. S. Nascimento, A. K. Saha, and E. M. Bruna. Lower ant diversity on earth mounds in a semi-arid ecosystem: natural variation or a sign of degradation? Sociobiology 63(4): 1022-1030.
 * Castanho L. M., and P. S. Oliveira. 1997. Biology and behaviour of the neotropical ant-mimicking spider Aphantochilus rogersi (Araneae: Aphantochilidae): nesting, maternal care and ontogeny of ant-hunting techniques. Journal of Zoology 242: 643-650.
 * Christianini A. V., A. J. Mayhé-Nunes, and P. S. Oliveira. 2012. Exploitation of Fallen Diaspores By Ants: Are There Ant-Plant Partner Choices? Biotropica 44: 360-367.
 * Coelho M. S., G. W. Fernandes, J. C. Santos, and J. H. C. Delabie. 2009. Ants (Hymenoptera: Formicidae) as bioindicators of land restoration in a Brazilian Atlantic forest fragment. Sociobiology 54(1): 51-63.
 * Costa-Milanez C. B., F. F. Ribeiro, P. T. A. Castro, J. D. Majer, S. P. Ribeiro. 2015. Effct of fire on ant assemblages in Brazilian Cerrado in areas containing Vereda wetlands. Sociobiology 62(4): 494-505.
 * Costa-Milanez C. B., G. Lourenco-Silva, P. T. A. Castro, J. D. Majer, and S. P. Ribeiro. 2014. Are ant assemblages of Brazilian veredas characterised by location or habitat type? Braz. J. Biol. 74(1): 89-99.
 * Crawley W. C. 1916. Ants from British Guiana. Ann. Mag. Nat. Hist. 8(17): 366-378.
 * 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.
 * Del-Claro K., and P. S. Oliveira. 1996. Honeydew flicking by treehoppers provides cues to potential tending ants. Animal Behaviour 51: 1071-1075.
 * Emery C. 1906. Studi sulle formiche della fauna neotropica. XXVI. Bullettino della Società Entomologica Italiana 37: 107-194.
 * Escalante Gutiérrez J. A. 1993. Especies de hormigas conocidas del Perú (Hymenoptera: Formicidae). Revista Peruana de Entomología 34:1-13.
 * Escalante J. A. 1975. Hormigas de la Provincia de la Convencionm Cusco. Revista Peruana de Entomologia 18:125-126.
 * Favretto M. A., E. Bortolon dos Santos, and C. J. Geuster. 2013. Entomofauna from West of Santa Catarina State, South of Brazil. EntomoBrasilis 6 (1): 42-63.
 * Fernandes G. W., W. J. Boecklen, R. P. Martins, and A. G. Castro. 1988. Ants associated with a coleopterous leaf-bud gall on Xylopa aromatica (Annonaceae). Proc. Entomol. Soc. Wash. 91(1): 81-87.
 * 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 T. T., R. R. Silva, D. Rodrigues de Souza-Campana, O. Guilherme Morais da Silva, and M. Santina de Castro Morini. 2019. Winged ants (Hymenoptera: Formicidae) presence in twigs on the leaf litter of Atlantic Forest. Biota Neotropica 19(3): http://dx.doi.org/10.1590/1676-0611-bn-2018-0694
 * Fernández, F. and S. Sendoya. 2004. Lista de las hormigas neotropicales. Biota Colombiana Volume 5, Number 1.
 * Fleck M. D., E. Bisognin Cantarelli, and F. Granzotto. 2015. Register of new species of ants (Hymenoptera: Formicidae) in Rio Grande do Sul state. Ciencia Florestal, Santa Maria 25(2): 491-499.
 * Forel A. 1905. Miscellanea myrmécologiques II (1905). Ann. Soc. Entomol. Belg. 49: 155-185.
 * Forel A. 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. 1909. Ameisen aus Guatemala usw., Paraguay und Argentinien (Hym.). Deutsche Entomologische Zeitschrift 1909: 239-269.
 * 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.
 * Gallego-Ropero M.C., R.M. Feitosa & J.R. Pujol-Luz, 2013. Formigas (Hymenoptera, Formicidae) Associadas a Ninhos de Cornitermes cumulans Kollar (Isoptera, Termitidae) no Cerrado do Planalto Central do Brasil. EntomoBrasilis, 6(1): 97-101.
 * 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.
 * Iop S., V. M. Caldart, J. A. Lutinski, and F. R. Mello Garcia. 2009. Formigas urbanas da cidade de Xanxerê, Santa Catarina, Brasil. Biotemas 22(2): 55-64.
 * Jaffe, K., et al. 2007. Comparing the ant fauna in a tropical and a temperat forest canopy. Ecotropicos 20(2):74-81
 * Kamura, C.M., M.S.C. Morini, C.J. Figueiredo, O.C. Bueno, and A.E.C. Campos-Farinha. 2007. Comunidades de formigas (Hymenoptera: Formicidae) em um ecossistema urbano próximo à Mata Atlântica. Brazilian Journal of Biology 67(4): 635-641
 * Kempf W. W. 1951. A taxonomic study on the ant tribe Cephalotini (Hymenoptera: Formicidae). Revista de Entomologia (Rio de Janeiro) 22:1-244
 * 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. 1972. Catalago abreviado das formigas da regiao Neotropical (Hym. Formicidae) Studia Entomologica 15(1-4).
 * Kusnezov N. 1953. La fauna mirmecológica de Bolivia. Folia Universitaria. Cochabamba 6: 211-229.
 * Luederwaldt H. 1918. Notas myrmecologicas. Rev. Mus. Paul. 10: 29-64.
 * Lutinski J. A., B. C. Lopes, and A. B. B.de Morais. 2013. Diversidade de formigas urbanas (Hymenoptera: Formicidae) de dez cidades do sul do Brasil. Biota Neotrop. 13(3): 332-342.
 * Lutinski J. A., F. R. Mello Garcia, C. J. Lutinska, and S. Iop. 2008. Ants diversity in Floresta Nacional de Chapecó in Santa Catarina State, Brazil. Ciência Rural, Santa Maria 38(7): 1810-1816.
 * Lutinski J. A., and F. R. Mello Garcia. 2005. Análise faunística de Formicidae (Hymenoptera: Apocrita) em ecossistema degradado no município de Chapecó, Santa Catarina. Biotemas 18(2): 73 - 86.
 * Maciel L., J. Iantas, F. C. Gruchowski-W, and D. R. Holdefer. 2011. INVENTORY OF FAUNA OF ANTS (HYMENOPTERA: FORMICIDAE) IN ECOLOGICAL SUCCESSION FLORISTIC ENVIRONMENT IN THE CITY OF UNION OF VICTORIA, PARANA, BRAZIL. Biodiversidade Pampeana Pucrs, Uruguiana 9(1): 38-43.
 * Mann W. M. 1916. The Stanford Expedition to Brazil, 1911, John C. Branner, Director. The ants of Brazil. Bulletin of the Museum of Comparative Zoology 60: 399-490
 * Marinho C. G. S., R. Zanetti, J. H. C. Delabie, M. N. Schlindwein, and L. de S. Ramos. 2002. Ant (Hymenoptera: Formicidae) Diversity in Eucalyptus (Myrtaceae) Plantations and Cerrado Litter in Minas Gerais, Brazil. Neotropical Entomology 31(2): 187-195.
 * Marques G. D. V., and K. Del-Claro. 2006. The Ant Fauna in a Cerrado area: The Influence of Vegetation Structure and Seasonality (Hymenoptera: Formicidae). Sociobiology 47(1): 1-18.
 * Menozzi C. 1926. Neue Ameisen aus Brasilien. Zoologischer Anzeiger. 69: 68-72.
 * Menozzi C. 1935. Spedizione del Prof. Nello Beccari nella Guiana Britannica. Hymenoptera-Formicidae. Redia. 21: 189-203.
 * Mentone T. O., E. A. Diniz, C. B. Munhae, O. C. Bueno, and M. S. C. Morini. 2011. Composition of ant fauna (Hymenoptera: Formicidae) at litter in areas of semi-deciduous forest and Eucalyptus spp., in Southeastern Brazil. Biota Neotrop. 11(2): http://www.biotaneotropica.org.br/v11n2/en/abstract?inventory+bn00511022011.
 * 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.
 * Neves F. S., R. F. Braga, M. M. do Espirito-Santo, J. H. C. Delabie, G. Wilson Fernandes, and G. A. Sanchez-Azofeifa. 2010. Diversity of Arboreal Ants In a Brazilian Tropical Dry Forest: Effects Of Seasonality and Successional Stage. Sociobiology 56(1): 1-18.
 * Nunes F. A., G. B. Martins Segundo, Y. B. Vasconcelos, R. Azevedo, and Y. Quinet. 2011. Ground-foraging ants (Hymenoptera: Formicidae) and rainfall effect on pitfall trapping in a deciduous thorn  woodland (Caatinga), Northeastern Brazil. Rev. Biol. Trop. 59 (4): 1637-1650.
 * Oliveira Mentone T. de, E. A. Diniz, C. de Bortoli Munhae, O. Correa Bueno and M. S. de Castro Morini. 2012. Composition of ant fauna (Hymenoptera: Formicidae) at litter in areas of semi-deciduous forest and Eucalyptus spp., in Southeastern Brazil. Biota Neotrop 11(2): 237-246.
 * Oliveira P. S., and I. Sazima,. 1984. The adaptive bases of ant-mimicry in a neotropical aphantochild spider (Araneae: Aphantochilidae). Biological Journal of the Linnean Society. 22: 145-155.
 * Pacheco R., H. L. Vasconcelos, S. Groc, G. P. Camacho, and T. L. M. Frizzo. 2013. The importance of remnants of natural vegetation for maintaining ant diversity in Brazilian agricultural landscapes. Biodivers. Conserv. DOI 10.1007/s10531-013-0463-y
 * Pacheco R., and H. L. Vasconcelos. 2012. Subterranean Pitfall Traps: Is ItWorth Including Them in Your Ant Sampling Protocol? Psyche doi:10.1155/2012/870794
 * Passos, L. and P.S. Oliveira. 2003. Interactions between ants, fruits and seeds in a restinga forest in south-eastern Brazil. Journal of Tropical Ecology 19(3):261-270.
 * Pereira J. C., J. H. C. Delabie, L. R. S. Zanette, and Y. Quinet. 2014. Studies on an Enigmatic Blepharidatta Wheeler Population (Hymenoptera: Formicidae) from the Brazilian Caatinga. Sociobiology 61(1): 52-59.
 * Pignalberi C. T. 1961. Contribución al conocimiento de los formícidos de la provincia de Santa Fé. Pp. 165-173 in: Comisión Investigación Científica; Consejo Nacional de Investigaciones Científicas y Técnicas (Argentina) 1961. Actas y trabajos del primer Congreso Sudamericano de Zoología (La Plata, 12-24 octubre 1959). Tomo III. Buenos Aires: Librart, 276 pp.
 * 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.
 * Powell, S. 2008. Ecological specialization and the evolution of a specialized caste in Cephalotes ant. Functional Ecology 22:902-911
 * 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
 * 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.
 * Ramos L. de S., C. G. S. Marinho, R. Zanetti, and J. H. C. Delabie. 2003. Impacto de iscas formicidas granuladas sobre a mirmecofauna não-alvo em eucaliptais segundo duas formas de aplicacação / Impact of formicid granulated baits on non-target ants in eucalyptus plantations according to two forms of application. Neotropical Entomology 32(2): 231-237.
 * Ramos L. de S., R. Zanetti, C. G. S. Marinho, J. H. C. Delabie, M. N. Schlindwein, and R. P. Almado. 2004. Impact of mechanical and chemical weedings of Eucalyptus grandis undergrowth on an ant community (Hymenoptera: Formicidae). Rev. Árvore 28(1): 139-146.
 * Reis P. C. J., W. D. Darocha, L. Falcao, T. J. Guerra, and F. S. Neves. 2013. Ant Fauna on Cecropia pachystachya Trécul (Urticaceae) Trees in an Atlantic Forest Area, Southeastern Brazil. Sociobiology 60(3): 222-228.
 * Ribas C. R., J. H. Schoereder, M. Pic, and S. M. Soares. 2003. Tree heterogeneity, resource availability, and larger scale processes regulating arboreal ant species richness. Austral Ecology 28(3): 305-314.
 * Ribas C. R., R. R. C. Solar, R. B. F. Campos, F. A. Schmidt, C. L. Valentim, and J. H. Schoereder. 2012. Can ants be used as indicators of environmental impacts caused by arsenic? Insect Conserv 16: 413421.
 * Ribas C. R., and J. H. Shoereder. 2007. Ant communities, environmental characteristics and their implications for conservation in the Brazilian Pantanal. Biodivers. Conserv. 16: 1511-1520.
 * Ribeiro L. F., R. R. C. Solar, T. G. Sobrinho, D. C. Muscardi, J. H. Schoereder, and A. N. Andersen. 2019. Different trophic groups of arboreal ants show differential responses to resource supplementation in a neotropical savanna. Oecologia 190(2): 433-443.
 * Rosa da Silva R. 1999. Formigas (Hymenoptera: Formicidae) do oeste de Santa Catarina: historico das coletas e lista atualizada das especies do Estado de Santa Catarina. Biotemas 12(2): 75-100.
 * Santos J. C., and Del-Claro K. 2001. Interactions between ants, herbivores and extrafloral nectaries in Tocoyena formosa (Cham & Schlechtd.) K. Schum. (Rubiaceae) in cerrado vegetation. Rev. Bras. Zoociencias Juiz de Fora 3(1): 77-92.
 * Santos Lopes J. F., N. Martins dos Reis Hallack, T. Archanjo de Sales, M. Silva Brugger, L. F. Ribeiro, I. N. Hastenreiter, and R. da Silva Camargo. 2012. Comparison of the Ant Assemblages in Three Phytophysionomies: Rocky Field, Secondary Forest, and Riparian ForestA Case Study in the State Park of Ibitipoca, Brazil. Psyche doi:10.1155/2012/928371
 * Santos Pereira M. P., M. Souza Schutte, J. Marcal Queiroz, and A. Barbosa Vargas. 2012. Riqueza, composição e atividade diurna da mirmecofauna sobre Triplaris americana L. (Polygonaceae). Revista Brasileira de Zoociências 14 (1, 2, 3): 135-142.
 * Santschi F. 1916. Formicides sudaméricains nouveaux ou peu connus. Physis (Buenos Aires). 2: 365-399.
 * Santschi F. 1921. Quelques nouveaux Cryptocerus de l'Argentine et pays voisins. Anales de la Sociedad Cientifica Argentina 92: 124-128.
 * Schoereder J. H., T. G. Sobrinho, M. S. Madureira, C. R. Ribas, and P. S. Oliveira. 2010. The arboreal ant community visiting extrafloral nectaries in the Neotropical cerrado savanna. Terrestrial Arthropod Reviews 3: 3-27.
 * Sendoya S. F., A. V. L. Freitas, and P. S. Oliveira. 2009. Egg-laying butterflies distinguish predaceous ants by sight. American Naturalist 174: 134-140.
 * Silva F. H. O., J. H. C. Delabie, G. B. dos Santos, E. Meurer, and M. I. Marques. 2013. Mini-Winkler Extractor and Pitfall Trap as Complementary Methods to Sample Formicidae. Neotrop Entomol 42: 351358.
 * Silvestre R., C. R. F. Brandão, and R. R. Silva da 2003. Grupos funcionales de hormigas: el caso de los gremios del cerrado. Pp. 113-148 in: Fernández, F. (ed.) 2003. Introducción a las hormigas de la región Neotropical. Bogotá: Instituto de Investigación de Recursos Biológicos Alexander von Humboldt, xxvi + 424 pp.
 * 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.
 * Soares S. A., D. Lange, and W. F. Antoniali Junior. 2006. Communities of Epigaeic ants (Hymenoptera: Formicidae) in an area of reforestation and in native forest. Sociobiology 49(3): 251-263.
 * Soares S. A., W. F. Antoniali Junior, and S. E. Lima-Junior. 2010. Diversidade de formigas epigéicas (Hymenoptera, Formicidae) em dois ambientes no Centro-Oeste do Brasil. Revista Brasileira de Entomologia 54(1): 7681.
 * 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.
 * 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
 * Tavares A. A., P. C. Bispo, and A. C. Zanzini. 2008. Effect of Collect Time on Communities of Epigaeic Ants (Hymenoptera: Formicidae) in Areas of Eucalyptus cloeziana and Cerrado. Neotropical Entomology 37(2): 126-130.
 * 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.
 * Vargas A. B., A. J. Mayhé-Nunes, J. M. Queroz, G. O. Souza, and E. F. Ramos. 2007. Effects of Environmental Factors on the Ant Fauna of Restinga Community in Rio de Janeiro, Brazil. Neotropical Entomology 36(1): 028-037
 * Vasconcelos H. L., J. M. S. Vilhena, W. E. Magnusson, and A. L. K. M. Albernaz. 2006. Long-term effects of forest fragmentation on Amazonian ant communities. Journal of Biogeography 33: 1348-1356.
 * Vasconcelos, H.L. and J.M.S. Vilhena. 2006. Species turnover and vertical partitioning of ant assemblages in the Brazilian Amazon: A comparison of forests and savannas. Biotropica 38(1):100-106.
 * Vasconcelos, H.L., J.M.S. Vilhena, W.E. Magnusson and A.L.K.M. Albernaz. 2006. Long-term effects of forest fragmentation on Amazonian ant communities. Journal of Biogeography 33:1348-1356
 * Vittar, F. 2008. Hormigas (Hymenoptera: Formicidae) de la Mesopotamia Argentina. INSUGEO Miscelania 17(2):447-466
 * Vittar, F., and F. Cuezzo. "Hormigas (Hymenoptera: Formicidae) de la provincia de Santa Fe, Argentina." Revista de la Sociedad Entomológica Argentina (versión On-line ISSN 1851-7471) 67, no. 1-2 (2008).
 * Wheeler W. M. 1916. Ants collected in British Guiana by the expedition of the American Museum of Natural History during 1911. Bulletin of the American Museum of Natural History 35: 1-14.
 * Wheeler W. M. 1918. Ants collected in British Guiana by Mr. C. William Beebe. Journal of the New York Entomological Society 26: 23-28.
 * Wheeler W. M. 1922. The ants of Trinidad. American Museum Novitates 45: 1-16.
 * Wheeler W. M. 1925. Neotropical ants in the collections of the Royal Museum of Stockholm. Arkiv för Zoologi 17A(8): 1-55.
 * Wheeler W. M. 1942. Studies of Neotropical ant-plants and their ants. Bulletin of the Museum of Comparative Zoology 90: 1-262.
 * 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.
 * da Silva, R.R., C.R.F. Brandao, and R. Silvestre. 2004. Similarity Between Cerrado Localities in Central and Southeastern Brazil Based on the Dry Season Bait Visitors Ant Fauna. Studies on Neotropical Fauna and Environment 39(3):191-199.
 * de Andrade, M.L. & C. Baroni Urbani. 1999. Diversity and Adaptation in the ant genus Cephalotes, past and present. Stuttgarter Beitrage zur Naturkunde Serie B 271. 893 pages, Stuttgart
 * de Souza D. R., S. G. dos Santos, C. de B. Munhae, and M. S. de C. Morini. 2012. Diversity of Epigeal Ants (Hymenoptera: Formicidae) in Urban Areas of Alto Tietê. Sociobiology 59(3): 703-117.