Crematogaster brasiliensis

Crematogaster brasiliensis
Scientific classification
Kingdom:	Animalia
Phylum:	Arthropoda
Class:	Insecta
Order:	Hymenoptera
Family:	Formicidae
Subfamily:	Myrmicinae
Tribe:	Crematogastrini
Genus:	Crematogaster
Species:	C. brasiliensis
Binomial name
Crematogaster brasiliensis Mayr, 1878
Subspecies
Crematogaster brasiliensis arawak Weber, 1938
Synonyms
Crematogaster armandi Forel, 1921 Crematogaster brasiliensis cocciphila Borgmeier, 1934 Crematogaster inca Wheeler, W.M., 1925 Crematogaster limata ludio Forel, 1912

A species of lowland wet and moist forest. Crematogaster brasiliensis are ominivores and their foragers may be encountered any time of the day or night.

Identification

Longino (2003) - The most striking variation is in worker size polymorphism. In South America various degrees of size polymorphism occur, and collections from Amazonian Brazil can be strongly polymorphic. The small workers are similar to specimens from Central America, while the large workers have heads wider than long, the scapes do not surpass the vertex margin, and the mesonotum is produced dorsally, projecting above the pronotum.

Other morphological characters vary discordantly across the range. The petiole in side view is relatively long and low in Central America, becoming shorter in South America. Specimens from South America often have longer propodeal spines, the spines may be somewhat elevated, and the dorsal face of the propodeum may be longer and more differentiated from the posterior face.

Species in the limata complex (brasiliensis, Crematogaster carinata, Crematogaster limata, and Crematogaster tenuicula in Costa Rica) all have abundant erect flexuous setae on the face, moderate length to short propodeal spines that are directed posteriorly, and elongate tapering petioles. The four species can be difficult to separate. They differ primarily in the nature of the ventral processes of the petiole and postpetiole. Crematogaster brasiliensis has distinct anteroventral processes on both the petiole and postpetiole. Crematogaster limata and carinata lack postpetiolar processes. In Costa Rica, brasiliensis and tenuicula are relatively distinct because tenuicula lacks an anteroventral petiolar process and the petiole is higher and more triangular in side view. These two species converge and become more difficult to distinguish in South America. Crematogaster brasiliensis may also be confused with Crematogaster foliocrypta, but foliocrypta has appressed rather than erect tibial pilosity.

Keys including this Species

• Key to Crematogaster of Costa Rica

Distribution

Nicaragua to Amazonian Brazil, Bolivia. (Longino 2003)

Latitudinal Distribution Pattern

Latitudinal Range: 17.3854° to -64.36°.

North Temperate	North Subtropical	Tropical	South Subtropical	South Temperate

• Source: AntMaps

Distribution based on Regional Taxon Lists

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

Distribution based on AntMaps

Distribution based on AntWeb specimens

Check data from AntWeb

Countries Occupied

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

Estimated Abundance

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

Biology

Longino (2003) - This species prefers lowland wet to moist forest habitats, and may occur in mature forest or second growth vegetation. In Costa Rica and other northern Neotropical sites that I have visited, two other limata group species, limata and carinata, are usually much more abundant. For example, at La Selva Biological Station in Costa Rica, carinata, limata, and brasiliensis occurred in 24, 11, and 4 fogging events, respectively, out of a total of 52 (Longino et al. 2002).

Foragers may be encountered during the day or at night. They are generalized omnivores, recruiting to protein or carbohydrate baits and visiting extrafloral nectaries.

The species is a generalized cavity nester, most often nesting in dead branches or dead vine stems that are lodged in low vegetation. It may also nest occasionally in live stems and in myrmecophytes such as Cordia nodosa and Triplaris, but its nesting in these cavities appears to be opportunistic rather than any specialized preference. Workers may build small amounts of thin, brittle carton that cover or restrict nest entrances. Colonies may be polydomous, and aggregations of workers and brood may be found in ephemeral cavities such as beneath aroid leaves appressed to trunks and beneath bark flaps. Most often nests contain only workers and brood, but I have twice found nests with workers, brood, and a single physogastric queen, suggesting monogyny.

One of these nests was in an elongate scar on the trunk of a small understory tree (Erythrochiton lindenii, Rutaceae). The scar was covered with thin, brittle, mud-like carton. I found one physogastric queen among the workers and brood, and a thorough search did not reveal any other reproductives.

Another nest was in a live stem of a large-stemmed shrub in the Asteraceae. The trailside herb had stems that had been extensively hollowed out by a stem-boring lepidopteran larva, and one of the stems had been cut by machete, leaving a 13mm diameter, exposed, necrotic end. A brasiliensis nest occupied the cavity in this cut stem, and the workers had constructed a carton baffle over the cut end. The large lumen was almost completely closed by carton, except for a small entrance hole just large enough for workers.

Colonies may be subject to parasitism by Microdon (Syrphidae). One nest I collected at La Selva Biological Station contained three large Microdon larvae in addition to workers and brood.

Castes

Worker

Images from AntWeb

Syntype of Crematogaster limata ludio. Worker. Specimen code casent0908432. Photographer Z. Lieberman, uploaded by California Academy of Sciences.	Owned by MHNG, Geneva, Switzerland.

Syntype of Crematogaster brasiliensis. Worker. Specimen code casent0902157. Photographer Ryan Perry, uploaded by California Academy of Sciences.	Owned by NHMUK, London, UK.

Syntype of Crematogaster armandi. Worker. Specimen code casent0902155. Photographer Ryan Perry, uploaded by California Academy of Sciences.	Owned by NHMUK, London, UK.

Syntype of Crematogaster armandi. Worker. Specimen code casent0908433. Photographer Z. Lieberman, uploaded by California Academy of Sciences.	Owned by MHNG, Geneva, Switzerland.

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Cotype inca. Worker. Specimen code lacm-ent-145093. Photographer C. Richart, uploaded by University of Utah.	Owned by LACM.

Specimen code jtlc000002077. .

Queen

Images from AntWeb

Queen (alate/dealate). Specimen code jtlc000001270. Photographer C. Richart, uploaded by University of Utah.	Owned by JTLC.

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Nomenclature

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

• brasiliensis. Crematogaster brasiliensis Mayr, 1878: 875 (w.) BRAZIL (no state data).
  • Type-material: lectotype worker (by designation of Longino, 2003a: 37), 1 paralectotype worker.
  • Type-locality: lectotype Brazil: Pará, Prainha (1°48’S, 53°29’W), on Tococa guianensis, #5 (J. Trail); paralectotype Brazil: Amazonas, Barcelos (0°58’S, 62°57’W), on Clidemia tococoidea, #27 (J. Trail)
  • [Notes (i): original published data by Mayr is merely Brazil: “Nord-Brasilien in der Umgegend des Amazonenstromes” (J. Trail); (ii) full type-data is provided by Longino, 2003a: 37, based on Benson & Setz, 1985 (reference not seen).]
  • Type-depository: NHMW.
  • Forel, 1912f: 217 (q.).
  • Combination in C. (Orthocrema): Emery, 1922e: 136.
  • Status as species: Dalla Torre, 1893: 80; Forel, 1895b: 131; Forel, 1899c: 83; Forel, 1912f: 217; Mann, 1916: 443; Wheeler, W.M. 1916d: 324; Emery, 1922e: 136; Wheeler, W.M. 1922c: 8; Wheeler, W.M. 1923a: 3; Wheeler, W.M. 1925a: 26; Borgmeier, 1927c: 92; Wheeler, W.M. 1942: 193; Kempf, 1961b: 503; Kempf, 1970b: 332; Kempf, 1972a: 84; Brandão, 1991: 339; Bolton, 1995b: 149; Longino, 2003a: 37 (redescription); Branstetter & Sáenz, 2012: 258; Bezděčková, et al. 2015: 116; Pedraza & Fernández, 2019: 895.
  • Senior synonym of armandi: Longino, 2003a: 37.
  • Senior synonym of cocciphila: Longino, 2003a: 37.
  • Senior synonym of inca: Longino, 2003a: 37.
  • Senior synonym of ludio: Longino, 2003a: 37.
  • Distribution: Bolivia, Brazil, Colombia, Costa Rica, Guatemala, Guyana, Nicaragua, Peru, Suriname, Trinidad.
  • Current subspecies: nominal plus arawak.
• armandi. Crematogaster armandi Forel, 1921a: 208 (w.q.) BRAZIL (Mato Grosso).
  • Type-material: syntype workers (number not stated), 1 syntype queen.
  • Type-locality: Brazil: Mato Grosso (S. Moore).
  • Type-depository: MHNG.
  • Combination in C. (Orthocrema): Kempf, 1972a: 84.
  • Status as species: Wheeler, W.M. 1942: 193; Kempf, 1972a: 84; Bolton, 1995b: 147.
  • Junior synonym of brasiliensis: Longino, 2003a: 37.
• cocciphila. Crematogaster (Orthocrema) brasiliensis var. cocciphila Borgmeier, 1934: 102 (w.q.) SURINAME.
  • Type-material: syntype workers (number not stated, “numerous”), 1 syntype queen.
  • Type-locality: Suriname (“Dutch Guiana”): Paramaribo (G.H. Bünzli).
  • Type-depositories: IBVR, MZSP.
  • Subspecies of brasiliensis: Kempf, 1972a: 84; Bolton, 1995b: 150.
  • Junior synonym of brasiliensis: Longino, 2003a: 37.
• inca. Crematogaster (Eucrema) inca Wheeler, W.M. 1925a: 27 (w.) PERU.
  • Type-material: 20 syntype workers.
  • Type-locality: Peru: Chaquimayo (N. Holmgren).
  • Type-depositories: MCZC, NHRS.
  • Status as species: Kempf, 1972a: 82; Bolton, 1995b: 154.
  • Junior synonym of brasiliensis: Longino, 2003a: 37.
• ludio. Crematogaster limata r. ludio Forel, 1912f: 217 (w.) BRAZIL (Ceará).
  • Type-material: syntype workers (number not stated).
  • Type-locality: Brazil: Ceará (F. Diaz da Rocha).
  • Type-depository: MHNG.
  • Wheeler, W.M. 1921f: 151 (q.).
  • Combination in C. (Orthocrema): Emery, 1922e: 136.
  • Subspecies of limata: Wheeler, W.M. 1916c: 12; Wheeler, W.M. 1918b: 24; Wheeler, W.M. 1921f: 151.
  • Subspecies of brasiliensis: Forel, 1913l: 233 (in text); Emery, 1922e: 136; Borgmeier, 1927c: 92; Wheeler, W.M. 1942: 194; Kempf, 1972a: 84; Bolton, 1995b: 156.
  • Junior synonym of brasiliensis: Longino, 2003a: 37.

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

Description

Worker

Longino (2003) - HL 0.646, 0.601, 0.897; HW 0.713, 0.642, 1.026; HC 0.665, 0.594, 0.995; SL 0.657, 0.622, 0.798; EL 0.165, 0.163, 0.207; A11L 0.284; A11W 0.118; A10L 0.149; A10W 0.086; A09L 0.074; A09W 0.057; A08L 0.067; A08W 0.044; WL 0.740, 0.723, 1.092; SPL 0.151, 0.135, 0.198; PTH 0.140, 0.125, 0.227; PTL 0.248, 0.264, 0.354; PTW 0.170, 0.154, 0.259; PPL 0.186, 0.133, 0.215; PPW 0.168, 0.145, 0.248; CI 110, 107, 114; OI 26, 27, 23; SI 102, 103, 89; PTHI 56, 47, 64; PTWI 69, 58, 73; PPI 90, 109, 115; SPI 20, 19, 18; ACI 1.61.

Color red brown; workers monomorphic in size.

Mandibles smooth and shining; clypeus varying from feebly striate, with 5 or more longitudinal carinulae, to nearly smooth and shining; head about as long as wide, subquadrate, with broadly convex sides and flat to weakly emarginate posterior border; antenna with terminal two segments enlarged to form a club, third segment from end somewhat enlarged, blurring distinction between two and three-segmented club; scapes with abundant long erect setae; when scapes laid back from antennal insertions, they surpass margin of vertex; face largely smooth and shining, with variable extent of striated region between antennal insertion and eye, and whorled above antennal insertion; face covered with abundant long flexuous white setae, no appressed pubescence; in face view abundant setae project from lateral and posterior margins.

Promesonotum in profile somewhat flattened dorsally, short anterior face of pronotum rises to dorsal face, dorsal faces of pronotum and mesonotum subequal in length, horizontal, forming single flat surface or meeting at a slightly produced angle, dorsal and posterior faces of mesonotum meeting at distinct angle, posterior face dropping to propodeal suture; propodeal suture deep in dorsal view but obscured in profile due to lateral carinulae that bridge the suture; lateral carinulae often with minute triangular tooth at propodeal suture; propodeal spines medium length, projecting posteriorly; in lateral view propodeum appears to have distinct dorsal and posterior faces because profile is horizontal from propodeal suture onto propodeal spines, but medially there is a single declivity from propodeal suture to petiolar insertion; pronotal dorsum with variably developed longitudinal carinulae, strongest laterally, becoming weaker medially, interspaces smooth and shining, in some cases almost entirely smooth and shining with no carinulae; anterodorsal face of mesonotum with weak, subparallel lateral carinae, these continue onto posterodorsal face as stronger carinae that converge posteriorly, interspace concave, smooth and shining; propodeal declivity smooth and shining; side of pronotum smooth and shining; katepisternum weakly to distinctly punctate or punctatorugose; side of propodeum very faintly sculptured; mesosomal dorsum with abundant long flexuous white setae, setae on pronotal humeri longest; femora and tibiae with abundant long erect setae.

Petiole in side view elongate, trapezoidal, weakly punctate to nearly smooth; anteroventral margin with a short right angle to somewhat rounded tooth; posterior ring-like aperture that receives postpetiole large, dorsal margin at nearly same level as posterolateral lobes of dorsal face when petiole viewed in profile with ventral margin horizontal (in contrast to tenuicula, in which posterior aperture is smaller and posterolateral lobes of dorsal face are distinctly higher than dorsal margin of aperture); dorsal face of petiole smooth and shining, elongate, widest posteriorly, regularly tapering anteriorly, with long flexuous setae along posterior border; postpetiole with distinct, subacute, short anteroventral tooth, globular in dorsal view, with abundant erect setae; fourth abdominal tergite smooth and shining, with abundant long flexuous erect white setae, no appressed pubescence.

Queen

Longino (2003) - A normal queen (dorsal face of propodeum drops steeply from postscutellum and much of propodeum appears ventral to scutellum and postscutellum) with general shape, sculpture, and pilosity characters of the worker; size characters as in Figures.

Type Material

Longino (2003) - Syntype workers: Brazil, Pará, Prainha, 1°48'S 53°29'W, on Tococa guianensis Aubl. (Trail #5); Brazil, Amazonas, Barcelos, 0°58'S 62°57'W, on Clidemia tococoidea (DC.) Gleason (Trail #27) [type data from Benson and Setz 1985] Naturhistorisches Museum Wien, Vienna (examined, Trail #5 worker here designated LECTOTYPE).

References

• Albuquerque, E., Prado, L., Andrade-Silva, J., Siqueira, E., Sampaio, K., Alves, D., Brandão, C., Andrade, P., Feitosa, R., Koch, E., Delabie, J., Fernandes, I., Baccaro, F., Souza, J., Almeida, R., Silva, R. 2021. Ants of the State of Pará, Brazil: a historical and comprehensive dataset of a key biodiversity hotspot in the Amazon Basin. Zootaxa 5001, 1–83 (doi:10.11646/zootaxa.5001.1.1).
• Cantone S. 2017. Winged Ants, The Male, Dichotomous key to genera of winged male ants in the World, Behavioral ecology of mating flight (self-published).
• Emery, C. 1922c. Hymenoptera. Fam. Formicidae. Subfam. Myrmicinae. [part]. Genera Insectorum 174B: 95-206 (page 136, Combination in C. (Orthocrema))
• Forel, A. 1912g. Formicides néotropiques. Part III. 3me sous-famille Myrmicinae (suite). Genres Cremastogaster et Pheidole. Mém. Soc. Entomol. Belg. 19: 211-237 (page 217, queen described)
• Franco, W., Ladino, N., Delabie, J.H.C., Dejean, A., Orivel, J., Fichaux, M., Groc, S., Leponce, M., Feitosa, R.M. 2019. First checklist of the ants (Hymenoptera: Formicidae) of French Guiana. Zootaxa 4674, 509–543 (doi:10.11646/zootaxa.4674.5.2).
• Longino, J.T. 2003a. The Crematogaster of Costa Rica. Zootaxa 151: 1-150.
• Mayr, G. 1878 [1877]. Formiciden gesammelt in Brasilien von Professor Trail. Verh. K-K. Zool.-Bot. Ges. Wien 27: 867-878 (page 875, worker described)
• Przybyszewski, K.R., Silva, R.J., Vicente, R.E., Garcia Freitas, J.V., Pereira, M.J.B., Izzo, T.J., Tonon, D.S. 2020. Can baited pitfall traps for sampling dung beetles replace conventional traps for sampling ants? Sociobiology 67, 376-387 (doi:10.13102/sociobiology.v67i3.5201).
• Tibcherani, M., Aranda, R., Mello, R.L. 2020. Time to go home: The temporal threshold in the regeneration of the ant community in the Brazilian savanna. Applied Soil Ecology 150, 103451 (doi:10.1016/j.apsoil.2019.103451).
• Ulysséa, M.A., Brandão, C.R.F. 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, 217–224 (doi:10.1590/s0085-56262013005000002).

References based on Global Ant Biodiversity Informatics

• Adams B. J., S. A. Schnitzer, and S. P. Yanoviak. 2016. Trees as islands: canopy ant species richness increases with the size of liana-free trees in a Neotropical forest. Ecography doi: 10.1111/ecog.02608
• Adams B. J., S. A. Schnitzer, and S. P. Yanoviak. 2019. Connectivity explains local ant community structure in a Neotropical forest canopy: a large-scale experimental approach. Ecology 100(6): e02673.
• 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.
• Bezdeckova K., P. Bedecka, and I. Machar. 2015. A checklist of the ants (Hymenoptera: Formicidae) of Peru. Zootaxa 4020 (1): 101–133.
• Chacon de Ulloa P., S. Valdes-Rodriguez, A. Hurtado-Giraldo, and M. Cleopatra Pimienta. 2014. Arboreal ants of Gorgona National Park (Pacific of Colombia). Rev. Biol. Trop. 62(1): 277-287.
• Dattilo W. et al. 2019. MEXICO ANTS: incidence and abundance along the Nearctic-Neotropical interface. Ecology https://doi.org/10.1002/ecy.2944
• Davidson, D.W. 2005. Ecological stoichiometry of ants in a New World rain forest. Oecologia 142:221-231
• Davidson, D.W., J.A. Arias and J. Mann. 2006. An experimental study of bamboo ants in western Amazonia. Insectes Sociaux 53:108-114
• Felizardo S. P. S., and A. Y. Harada. 2007. The genus Crematogaster Lund, 1831 (Formicidae: Myrmicinae: Crematogastrini) at ant collection from Emílio Goeli Paraense Museum (MPEG). Biológico, São Paulo 69(2): 425-427.
• 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.
• Fernández, F. and S. Sendoya. 2004. Lista de las hormigas neotropicales. Biota Colombiana Volume 5, Number 1.
• Forel A. 1912. Formicides néotropiques. Part III. 3me sous-famille Myrmicinae (suite). Genres Cremastogaster et Pheidole. Mémoires de la Société Entomologique de Belgique. 19: 211-237.
• 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.
• Hahn, D.A. and D.E. Wheeler. 2002. Seasonal Foraging Activity and Bait Preferences of Ants on Barro Colorado Island, Panama. Biotropica 34(3):348-356
• INBio Collection (via Gbif)
• Kempf W. W. 1961. A survey of the ants of the soil fauna in Surinam (Hymenoptera: Formicidae). Studia Entomologica 4: 481-524.
• Kempf W. W. 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).
• Leponce M., J. H. C. Delabie, J. Orivel, J. Jacquemin, M. Calvo Martin, and A. Dejean. 2019. Tree-dwelling ant survey (Hymenoptera, Formicidae) in Mitaraka, French Guiana, in Touroult J. (ed.), “Our Planet Reviewed” 2015 large-scale biotic survey in Mitaraka, French Guiana. Zoosystema 41 (10): 163-179.
• 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. et al. ADMAC project. Accessed on March 24th 2017 at https://sites.google.com/site/admacsite/
• Longino, J.T. 2003. The Crematogaster (Hymenoptera, Formicidae, Myrmicinae) of Costa Rica. Zootaxa 151:1-150
• Longino, J.T. 2010. Personal Communication. Longino Collection Database
• Lopes M. C., G. P. A. Lamarre, C. Baraloto, P. V. A. Fine, A. Vincentini, and F. B. Baccaro. 2019. The Amazonas-trap: a new method for sampling plant-inhabiting arthropod communities in tropical forest understory. Entomologia Experimentalis et Applicata https://doi.org/10.1111/eea.12797
• Miranda P. N., F. B. Baccaro, E. F. Morato, M. A. Oliveira. J. H. C. Delabie. 2017. Limited effects of low-intensity forest management on ant assemblages in southwestern Amazonian forests. Biodivers. Conserv. DOI 10.1007/s10531-017-1368-y
• 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.
• 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.
• Salazar F., and D. A. Donoso. 2013. New ant (Hymenoptera: Formicidae) records for Ecuador deposited at the Carl Rettenmeyer ant collection in the QCAZ Museum. Boletín Tecnico 11, Serie Zoológica 8-9: 151 – 177.
• Santos P. P., A. Vasconcelos, B. Jahyny, and J. H. C. Delabie. 2010. Ant fauna (Hymenoptera, Formicidae) associated to arboreal nests of Nasutitermes spp. (Isoptera, Termitidae) in a cacao plantation in southeastern Bahia, Brazil. Revista Brasileira de Entomologia 54(3): 450–-454.
• 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
• Vásquez-Bolaños M. 2011. Lista de especies de hormigas (Hymenoptera: Formicidae) para México. Dugesiana 18: 95-133
• 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. 1921. The Tachigalia ants. Zoologica (New York) 3: 137-168.
• Wheeler W. M. 1922. The ants of Trinidad. American Museum Novitates 45: 1-16.
• 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
• da Silva de Oliveira A. B., and F. A. Schmidt. 2019. Ant assemblages of Brazil nut trees Bertholletia excelsa in forest and pasture habitats in the Southwestern Brazilian Amazon. Biodiversity and Conservation 28(2): 329-344.