A common species that is found in a variety of habitats. Neoponera verenae nests in rotten wood and logs in twigs and under stones. (Mackay and Mackay 2010, Traniello and Hölldobler 1984, Wild 2002)
- 1 Identification
- 2 Distribution
- 3 Biology
- 4 Castes
- 5 Nomenclature
- 6 References
- 7 References based on Global Ant Biodiversity Informatics
From Mackay and Mackay (2010): The lack of erect hairs on the dorsum of the mesosoma, petiole and first tergum of the worker and the female of N. verenae would separate this species from all of the others of the genus Neoponera , except for Neoponera apicalis, Neoponera obscuricornis, Neoponera bucki and Neoponera magnifica. Neoponera verenae can be separated from the last two species by the large eyes, which occupy more than 1/3 of the side of the head. The workers and females of N. verenae can be separated from N. apicalis as the apex of the funiculus is usually dark, but may be reddish brown, but is never yellow as it is in N. apicalis. The posterior lateral edges of the petiole of the worker and female of N. verenae are sharp and may even form a carina, whereas in N. apicalis this area is mostly rounded.
This species has been consistently confused with N. obscuricornis. Wild (2005) recently resolved the confusion between the species. The two species can be easily separated as the posterior lateral margins of the petiole of N. verenae are sharp, those of N. obscuricornis are broadly rounded. Additionally N. verenae lacks the dense pubescence on the hypopygium, which is present on the hypopygidium of N. obscuricornis. Neoponera verenae is much more common than N. obscuricornis.
The male of N. obscuricornis is completely dark brown or black, including the entire antenna, which would separate it from that of N. apicalis, in which the antenna is completely pale brown. The side of the petiole lacks the rugae that are found in the male of N. apicalis. Thus it is unlikely that the male would be confused with the other two closely related species. It may be difficult to separate it from the unknown male of the rarely collected Neoponera cooki from Colombia south to Brazil.
Keys including this Species
Mexico through central South America.
Distribution based on Regional Taxon Lists
Distribution based on AntMaps
Distribution based on AntWeb specimens
Check data from AntWeb
Neoponera verenae is a common species that is found in tropical rain forests, subtropical forests, riparian habitats, riparian areas in the llanos [grasslands], gallery forests and wet forests to open urban environments and cacao plantations, from 0 to 925 m elevation. Wild (2005) notes that southern populations seem to be more commonly collected in open habitats while northern populations are more likely be found in forests. (Mackay and Mackay 2010)
Almost all the information published about N. verenae appears in the literature under the name Neoponera obscuricornis (see this species page for more about previous species determination problems). This common species exhibits great flexibility in habitat. 14 specimen records are from rainforest or other types of wet forest, seven are from forest edge habitats, five from open natural habitats such as campo cerrado or savannah, one from pasture, one from tropical scrub forest, and one from a cacao plantation. Interestingly, southern populations seem to be more commonly collected in open habitats, while northern populations are more likely to be found in forest. This species displays similar nesting habits to Neoponera apicalis and Neoponera obscuricornis. Three nest records from specimen collection data were from rotting wood, and one from a grass clump in a pasture. Traniello and Hölldobler’s (1984) study colony was collected nesting in a log in Panama, and Wild (2003) reports a nest in rotting wood in Paraguay.
Neoponera verenae is a predaceous and scavenging species. Foragers will also carry droplets of liquid held between their mandibles, a common trait in poneromorph ants (Hölldobler 1986). Longino (2004) has observed N. verenae attacking live lepidopteran larvae in Costa Rica, and captive colonies have taken crickets, cockroaches, termites, and other insect parts (Traniello & Hölldobler 1984, Oliveira & Hölldobler 1991, Gobin et al 2003). Foragers use visual cues (Duelli & Duelli-Klein 1976), and there is no recruitment to food sources (Traniello & Hölldobler 1984). Colonies are small, reportedly with fewer than 100 workers. Gobin et al (2003) collected 27 colonies from La Selva in Costa Rica with a median number of 39 workers per colony. The study colony of Fresneau (1984) contained 57 workers, and that of Traniello and Hölldobler (1984) grew to about 80–90 workers. N. verenae appears to be polygynous. Traniello and Hölldobler’s study colony had “several” fertile queens, and Fresneau (1984) found developed ovaries in five of seven dealate queens. Oliveira & Hölldobler (1991) described the agonistic interactions between workers and unmated queens in a queenless laboratory colony. These dominance interactions have a measurable energetic cost to the colony (Gobin et al 2003).
Neoponera verenae has been the subject of considerable research on gland structure. Abdominal glands in the male were described by Hölldobler and Engel-Siegel (1982), the pygidial gland by Traniello and Hölldobler (1984), the metapleural gland was briefly investigated by Hölldobler and Engel-Siegel (1985), and the ultrastructure of the labial gland was reported by Lommelen et al (2002, 2003).
Tandem-running, a stereotyped behavior where an ant recruits a single nestmate at a time, was investigated in Neoponera verenae by Traniello and Hölldobler (1984). N. verenae was found to employ tandem-running during nest relocation, mediated by a pheromone originating in the pygidial gland of the lead ant and spread to the hind-legs by a self-grooming behavior.
There is one record in Museum of Comparative Zoology of N. verenae in the gut contents of Bufo coniferus Cope in Nicaragua.
Lange et al. (2015) - This species was found in semideciduous mesophytic forest pitfall-trap samples. The study examined the ant community within a patch (11.2 hectares) of forest in an urban park (Araguari, Brazil).
Association with Other Organisms
- Oplitis apicalis
- Petalomium verenae - Da Silva et al. (2017) - Phoretic females found attached to the hairs between the first (mainly) and second pairs of coxae of N. verenae workers from the municipalities of Itamaraju, Barro Preto, São José da Vitória and Camacan, south of the state of Bahia, Northeastern Brazil, were collected from the ants in 2011.
- This species is a host for the strepsipteran Myrmecolax incautus (a parasite) (Cook, 2019).
The following information is derived from Barry Bolton's Online Catalogue of the Ants of the World.
- verenae. Neoponera apicalis var. verenae Forel, 1922: 90 (w.) PANAMA.
- Type-material: lectotype worker (by designation of Mackay & Mackay, 2010: 561), 1 paralectotype worker.
- [Note: Wild, 2005: 10, records 3 syntype workers in MHNG.]
- Type-locality: lectotype Panama (no further data) (Christophersen); paralectotype with same data.
- Type-depository: MHNG.
- Mackay & Mackay, 2010: 562 (q.m.).
- Combination in Neoponera: Schmidt, C.A. & Shattuck, 2014: 152.
- Junior synonym of apicalis: Brown, 1957e: 230; Kempf, 1972a: 161; Bolton, 1995b: 311.
- Status as species: Wild, 2005: 10 (redescription); Wild, 2007b: 40; Mackay, Mackay, et al. 2008: 199; Mackay & Mackay, 2010: 561 (redescription); Branstetter & Sáenz, 2012: 263; Bezděčková, et al. 2015: 124; Feitosa, 2015c: 99; Fernández & Guerrero, 2019: 534.
- Distribution: Bolivia, Brazil, Colombia, Costa Rica, Ecuador, El Salvador, French Guiana, Guatemala, Guyana, Honduras, Mexico, Nicaragua, Panama, Paraguay, Peru, Venezuela.
Panamá. Lectotype worker, 1 paralectotype worker designated, Musee d'Histoire Naturelle Genève (Mackay and Mackay 2010) Unless otherwise noted the text for the remainder of this section is reported from the publication that includes the original description.
From Mackay and Mackay (2010): The workers are moderate sized (total length 8 - 9 mm) black ants. The mandibles are mostly smooth and glossy; the remainder of the ant is dull, which forms a strong contrast. The malar carina is well developed and sharp. The eyes are large, occupying more than ⅓ of the length of the side of the head. The scapes extend about ⅓ length past the posterior lateral corner. The dorsum of the mesosoma is nearly straight, but is slightly depressed at the metanotal suture. The shoulder of the pronotum is swollen but does not form a distinct carina. The propodeal spiracle is elongated and slit-shaped. The petiole is thick when viewed in profile with both the anterior and posterior faces being convex and rounded and meeting at a rounded apex. The posterior lateral borders of the petiole meet at a sharp margin, which nearly forms a carina. The metasternal process is formed of two wide lobes, which are more separated near the apex than at the base. The lower inner margin has oblique striae.
Erect hairs are sparse, with a few on the mandibles, clypeus, coxae, femora, tibiae (many of the hairs on the legs are suberect), ventral surface of the gaster and posterior ⅓ of the gaster. Erect and suberect hairs are absent on the side of the head (a few may be present on the cheeks or on the malar area) the dorsum of the mesosoma, the petiole, the first and on most of the second terga.
The mandibles of the worker are finely striate and moderately shining. The remainder of the ant is either very finely striate or very finely punctate and dull.
From Mackay and Mackay (2010): The female (undescribed) is similar to the worker, lacking erect hairs on the dorsum of the mesosoma, the petiole and most of the dorsum of the gaster. The large eyes and the well-developed carina anterior to the eye would further assist in recognizing this species. The posterior lateral edges of the petiole form a sharp carina. The total length is 12 mm, the head length is 2.4 mm and the head width is 1.7 mm. The eyes are large (0.86 mm maximum diameter) located less than one diameter from the anterior edge of the head. The ocelli are small (the diameters of the medial and lateral ocelli are 0.09 mm). The scape (2.7 mm) extends about ⅓ length past the posterior lateral corner of the head. The carina on the side of the pronotum is sharp and slightly overhangs the side of the pronotum. The propodeal spiracle is slit-shaped and the stridulatory file is well developed. The sculpture is similar to that of the worker.
From Mackay and Mackay (2010): The male (undescribed) is a moderately large (total length about 12 mm) black specimen. The cavity at the base of the mandible is relatively large with the depressed area being more than ½ the length of the mandible. The head length is 1.5 - 1.6 mm; the head width 1.3 - 1.4 mm. The eyes are relatively small with a diameter of 0.8 mm (side view). The ocelli (maximum diameters of the medial and lateral ocelli 0.2 mm) are separated by about one diameter. The pronotal shoulder is swollen but does not form a margin. The Mayrian sutures are developed but do not connect in the center of the scutum; the parapsidal sutures are well developed. The propodeal spiracle is slit-shaped. The petiole is rounded dorsally and the two faces (anterior and posterior) are about equal in length and shape. The posterior lateral regions of the petiole are broadly rounded and not sharply bent as they are in the worker and the female. The stridulatory file on the second pretergite is well developed and even slightly wider than it is in the worker and female.
Most of the bodily surfaces including the sides of the head, the dorsum of the mesosoma, the dorsum of the petiole and all surfaces of the gaster are covered with short (most about 0.25 mm in length) flexuous golden or silver hairs.
The tiny mandibles are weakly shining and the remainder of the body is dull.
It would be difficult to recognize the male without workers or females.
- 2n = 58, karyotype = 14M+44A (Brazil) (Mariano et al., 2007; Delabie et al., 2008) (Different karyotype reported in Mariano 2007).
- 2n = 60 (Brazil) (Mariano et al., 2007; Delabie et al., 2008) (Different karyotype reported in Mariano 2007).
- 2n = 64, karyotype = 12M+52A (Brazil) (Mariano et al., 2007; Delabie et al., 2008).
- 2n = 62, karyotype = 14M+48A (Brazil) (Mariano et al., 2007; Delabie et al., 2008).
- 2n = 42, karyotype = 30M+12A (Brazil) (Mariano et al., 2007; Delabie et al., 2008).
- 2n = 58 (Brazil) (Mariano et al., 2007; Delabie et al., 2008).
This species was named in honor of an unspecified woman named Verena. (Mackay and Mackay 2010)
- Mackay, W. P., and E. E. Mackay 2010. The Systematics and Biology of the New World Ants of the Genus Pachycondyla (Hymenoptera: Formicidae). Edwin Mellon Press, Lewiston. Information from this publication is used with permission from the authors.
- Araujo, E.S., Koch, E.B.A., Delabie, J.H.C., Zeppelini, D., DaRocha, W.D., Castaño-Meneses, G., Mariano, C.S.F. 2019. Diversity of commensals within nests of ants of the genus Neoponera (Hymenoptera: Formicidae: Ponerinae) in Bahia, Brazil. Annales de la Société entomologique de France (N.S.), 1–9. (doi:10.1080/00379271.2019.1629837).
- Baena, M. L. 1993. Hormigas cazadoras del género Pachycondyla (Hymenoptera: Ponerinae) de la Isla Gorgona y la planicie Pacifica Colombiana. Boletin. Del Museo. Entomológica de la Universidad del Valle 1:13-21.
- Brown, B. and D. Feener. 1998. Parasitic phorid flies (Diptera: Phoridae) associated with army ants (Hymenoptera: Formicidae: Ecitoninae, Dorylinae) and their conservation biology. Biotropica 30:482-487.
- Brown, W. L., Jr. 1957e. Biological investigations in the Selva Lacandona, Chiapas. 4. Ants from Laguna Ocotal (Hymenoptera: Formicidae). Bull. Mus. Comp. Zool. 116: 228-237 (page 230, Junior synonym of apicalis)
- Calazans, E.G., da Costa, R.V., Cristiano,M.P., Cardoso, D.C. 2020. Daily dynamics of an ant community in a mountaintop ecosystem. Environmental Entomology (doi:10.1093/ee/nvaa011).
- Cook, J.L. 2019. Annotated catalog of the order Strepsiptera of the world. Transactions of the American Entomological Society 144: 121-267 (doi:10.3157/061.145.0202).
- Da Silva, R. A., A. A. Khaustov, J. M. S. Lopes, J. H. C. Delabie, and A. R. Oliveira. 2017. A new species of Petalomium from Brazil with a redescription of Petalomium gottrauxi Mahunka 1977 (Acari: Heterostigmatina: Neopygmephoridae). Systematic and Applied Acarology. 22:1800-1812. doi:10.11158/saa.22.11.2
- Delabie JHC, Mariano CSF, Mendes LF, Pompolo SG, Fresneau D (2008). Problemas apontados por estudos morfológicos, ecológicos e citogenéticos no gênero Pachycondyla na região Neotropical: o caso do complexo apicalis. In: Vilela EF, Santos IA, Schoereder JH, Neto JL, Serrão JE, Campos LAO, editors. Insetos sociais: da biologia à aplicação. Viçosa: Editora UVF. pp. 197-222.
- dos Santos Lopes, J. M., A. R. Oliveira, J. H. C. Delabie, and H. Klompen. 2015. A new species of myrmecophile mite of the genus Oplitis (Acari: Mesostigmata: Oplitidae) from Brazil. International Journal of Acarology. 41:676-680. doi:10.1080/01647954.2015.1096960
- Duelli, P. and R. Duelli-Klein. 1976. Freilandversuche zum Heimfindevermögen südamerikanischer Ameisen (Formicidae: Ponerinae, Dolichoderinae, Formicinae). Studia Entomologica 19:409-419.
- Düssmann, O., C. Peeters and B. Hölldobler. 1996. Morphology and reproductive behaviour of inter-castes in the ponerine ant Pachycondyla obscuricornis. Insectes Sociaux 43:421-425. (= N. verenae)
- Forel, A. 1922b. Glanures myrmécologiques en 1922. Rev. Suisse Zool. 30: 87-102 (page 90, worker described)
- Fresneau, D. 1984. Dévelopment ovarien et status sociale chez une fourmi primitive Neoponera obscuricornis Emery (Hym. Formicidae, Ponerinae). Insectes Sociaux 31:387-402.
- Gobin, B., J. Heinze, M. Straetz and F. Roces. 2003. The energetic cost of reproductive conflicts in the ant Pachycondyla obscuricornis. Journal of Insect Physiology 49:747-752.
- Hölldobler, B. 1985. Liquid food transmission and antennation signals in ponerine ants. Israel Journal of Entomology 19:89-99.
- Hölldobler, B. and H. Engel-Siegel. 1982. Tergal and sternal glands in male ants. Psyche 89:113-132.
- Hölldobler, B. and H. Engel-Siegel. 1984. On the metapleural gland of the ants. Psyche 91:201-224.
- Lange, D., A. A. Vilela, G. Erdogmus, A. B. Barbosa, S. C. Costa, and V. Stefani. 2015. Temporal dynamic of foraging of epigeic ants in an urban forest fragment. Bioscience Journal. 31:1501-1511.
- Lommelen, E., E. Schoeters and J. Billen. 2002. Ultrastructure of the labial gland in the ant Pachycondyla obscuricornis (Hymenoptera, Formicinae). Netherlands Journal of Zoology 52:61-68.
- Lommelen, E., E. Schoeters and J. Billen. 2003. Development of the labial gland of the ponerine ant Pachycondyla obscuricornis (Hymenoptera, Formicidae) during the pupil stage. Arthropods Structure and Development 32:209-217.
- Longino, J. T. website www. evergreen. edu/ants/genera/ Pachycondyla/SPECIES.
- Oliveira, P. S. and B. Hölldobler. 1991. Agnostic interactions and reproductive dominance in Pachycondyla obscuricornis (Hymenoptera: Formicidae). Psyche 98:215-225.
- Reiskind, J. 1977. Ant-mimicry in Panamian clubionid and salticid spiders (Araneae: Clubionidae and Salticidae). Biotropica 9:1-8.
- Schmidt, C.A. & Shattuck, S.O. 2014. The higher classification of the ant subfamily Ponerinae (Hymenoptera: Formicidae), with a review of ponerine ecology and behavior. Zootaxa 3817, 1–242 (doi:10.11646/zootaxa.3817.1.1).
- Traniello, J. F. 1981. Enemy deterrence in the recruitment strategy of a termite: soldier-organized foraging in Nasutitermes costalis. Proceedings of the National Academy of Science 78:1976-1979.
- Traniello, J. F. and Hölldobler, B. 1984. Chemical communication during tandem running in Pachycondyla obscuricornis (Hymenoptera: Formicidae). Journal of Chemical Ecology 10:783-794
- Wild, A. 2002. The genus Pachycondyla (Hymenoptera: Formicidae) in Paraguay. Boletín del Museo Nacional de Historia Natural de Paraguay 14:1-18.
- Wild, A. L. 2005. Taxonomic revision of the Pachycondyla apicalis species complex (Hymenoptera: Formicidae). Zootaxa 834: 1-25 (page 10, raised to species)
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 https://doi.org/10.1007/s13744-017-0565-4
- 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.
- Araujo Castilho G., F. Barbosa Noll, E. R. da Silva, and E. F. dos Santos. 2011. Diversidade de Formicidae (Hymenoptera) em um fragmento de Floresta Estacional Semidecídua no Noroeste do estado de São Paulo, Brasil. R. bras. Bioci., Porto Alegre 9(2): 224-230.
- Biagiotti G., J. A. Alves Pereira, C. Rodrigues Ribas, V. Korasaki, R. Zanetti, A. C. Medeiros de Queiroz. 2013. Richness and species composition of ants in the recovery process of a Gully erosion. Cerne, Lavras 19(4): 661-668.
- 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., R. De Jesus Santos, J. R. Mala Dos Santos, J. H. C. Delabie, L. L. Lopes, and C. F. Mariano. 2019. Invertebrates associated to Ponerine ants nests in two cocoa farming systems in the southeast of the state of Bahia, Brazil. Tropical Ecology 60: 52–61.
- Dattilo W. et al. 2019. MEXICO ANTS: incidence and abundance along the Nearctic-Neotropical interface. Ecology https://doi.org/10.1002/ecy.2944
- De Souza Holanda P. M. 2016. Efeitos da variacao do lencol: freatico em assembleias de formigas (Hymenoptera: Formicidae) que vivem na serapilheira na reserva Adolpho Ducke, Manaus, Am. Master's Thesis Universidade Federal do Amazonas, 48 pages.
- Delabie J. H. C., R. Céréghino, S. Groc, A. Dejean, M. Gibernau, B. Corbara, and A. Dejean. 2009. Ants as biological indicators of Wayana Amerindian land use in French Guiana. Comptes Rendus Biologies 332(7): 673-684.
- 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. 2008. Subfamilia Ponerinae s.str. Pp. 123-218 in: Jiménez, E.; Fernández, F.; Arias, T.M.; Lozano-Zambrano F. H. (eds.) 2008. Sistemática, biogeografía y conservación de las hormigas cazadoras de Colombia. Bogotá: Instituto de Investigación de Recursos Biológicos Alexander von Humboldt, xiv + 609 pp.
- Forel A. 1922. Glanures myrmécologiques en 1922. Revue Suisse de Zoologie. 30: 87-102.
- 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.
- Groc S., J. H. C. Delabie, F. Fernandez, F. Petitclerc, B. Corbara, M. Leponce, R. Cereghino, and A. Dejean. 2017. Litter-dwelling ants as bioindicators to gauge the sustainability of small arboreal monocultures embedded in the Amazonian rainforest. Ecological Indicators 82: 43-49.
- Gutierrez-Martínez P. R., and D. Arauz Naranjo. 2016. Hormigas (Hymenoptera: Formicidae) en el bosque seco del Parque Nacional Diria, Costa Rica. Cuadernos de Investigación UNED 8(2): 185-188.
- INBio Collection (via Gbif)
- 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/
- Mackay, W.P. and E.E. MacKay. 2010. The systematics and biology of the New World ants of the genus Pachycondyla (Hymenoptera: Formicidae). Edwin Mellen Press Lewiston, NY
- 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.
- Ottonetti L., L. Tucci, F. Frizzi, G. Chelazzi, and G. Santini. 2010. Changes in ground-foraging ant assemblages along a disturbance gradient in a tropical agricultural landscape. Ethology Ecology & Evolution 22: 7386.
- 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
- 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.
- 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.
- Santos-Junior L. C., J. M. Saraiva, R. Silvestre, and W. F. Antonialli-Junior. 2014. Evaluation of Insects that Exploit Temporary Protein Resources Emphasizing the Action of Ants (Hymenoptera, Formicidae) in a Neotropical Semi-deciduous Forest. Sociobiology 61(1): 43-51
- 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.
- Valdes-Rodriguez S., P. Chacon de Ulloa, and I. Armbrecht. 2014. Soil ant species in Gorgona Island, Colombian Pacific. Rev. Biol. Trop. 62 (1): 265-276.
- Wild A. L. 2005. Taxonomic revision of the Pachycondyla apicalis species complex (Hymenoptera: Formicidae). Zootaxa 834: 1-25.
- Wild, A. L. "A catalogue of the ants of Paraguay (Hymenoptera: Formicidae)." Zootaxa 1622 (2007): 1-55.
- 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.
- de Almeida Soares S., Y. R. Suarez, W. D. Fernandes, P. M. Soares Tenorio, J. H. C. Delabie, and W. F. Antonialli-Junior. 2013. Temporal variation in the composition of ant assemblages (Hymenoptera, Formicidae) on trees in the Pantanal floodplain, Mato Grosso do Sul, Brazil. Rev. Bras. entomol. 57: 84-90