Neoponera commutata

This is a relatively common ground surface dweller, found mostly in a variety of habitats from the Amazon–Orinoco watershed, where the majority of the records are known, and have been examined in this study. Individuals of all castes have been collected throughout the year, however, research is still pending in regards to colony life cycle and production of alate forms (but see Schmidt & Overal 2009). Nothing is known in regards to its flight phenology. Several techniques have been used to collect this species including (from the most to the least effective) ground surface pitfall traps, hand collections, Malaise (mostly males), litter sifting and Winkler, and very unusually with canopy fogging. Neoponera commutata nests underground (Wheeler 1936; Mill 1984; Schmidt & Overal 2009; A. Troya pers. obs.) with colonies varying in population size from approximately 400–1000 individuals (Mill 1984; Schmidt & Overal 2009). Adrian Troya observed various individuals of N. commutata coming in and out from an (apparently) empty, ca. 1m diameter, dome–shaped, termite nest. The structure was attached to a live tree trunk in a lowland Amazonian forest in Ecuador, and seemed to host an ant colony. After ca. 15 minutes of observation no prey of any kind was seen in the mandibles of the workers. Although no guarding behavior was recognized at what would be their nest entrance located laterally on the dome, the hypothesis of reutilization of termite nests by N. commutata needs further confirmation. (Troya & Lattke, 2022)

Identification
From Mackay and Mackay (2010): The worker and female of N. commutata are easily recognized by their large size and mostly shining integument. Neoponera commutata can be separated from the three other shiny black species (Neoponera carbonaria, Neoponera marginata and Neoponera laevigata) by their larger size. Occasionally workers of N. laevigata are nearly as large, but differ in the shape of the petiole. The apex of the petiole of N. commutata is highest near the posterior edge; the dorsum of the petiole of N. laevigata is at approximately the same height over the entire dorsal face. The mandibles of N. commutata are dull and covered with striae; they are smooth and glossy in N. laevigata.

Distribution
Central America through central South America (Mackay and Mackay 2010)

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

Habitat
Specimens have been collected in tropical rain forests along jungle trails (Hermann, 1968), in open savannah and on the shady forest floor (Wheeler, 1936) and tropical semi-humid forest (Mill, 1983). One series was collected at 250 m, another at 580m. (Mackay and Mackay 2010)

Biology
From Mackay and Mackay (2010): Mill (1984) found a density of 10 colonies per hectare in Brasil. A dealate female was collected in December (Ecuador).

This is a group raiding species in which stridulation causes an escape behavior among the other workers (Hermann, 1968). Nests have about 400 workers and they are group mass recruitment foragers, which prey exclusively on three species termites in the leaf-cutting genus Syntermes spp. (Wheeler, 1936), especially S. molestus (Mill, 1982a, 1982b, 1984). The other two species of termites are S. solidus and S. calvus (Mill, 1984). Wheeler (1936) reported they feed on S. chaquemayensis, S. territus, S. snyderi, S. grandis and S. brasiliensis. Foragers employ two different modes of searching for termite prey (Mill, 1982a). The first involves a forager encountering a foraging column of termites. She returns to the nest laying a pheromone trail. She leads other workers back to attack the termites. They stockpile the dead and wounded termites and each ant often returns to the nest with two or more termites (Mill, 1984). The second method involves a group of foragers (20 - 117 workers - Mill, 1984), which work together, or may separate into smaller groups of individuals (Mill, 1984). When a worker encounters termites she stridulates to attract the other ants and they attack as a group. Stridulation by a worker may result in the other foragers scattering into the surrounding litter (Mill, 1984). Termites instantly become immobile when stung (Mill, 1984).

Troya & Lattke, 2022: Mill (1982a, 1984), as part of his research on termites, registered the group recruitment, predatory behavior of N. commutata on several termite groups, including the genus Syntermes Holmgren, some of the species of which are considered exclusive prey of these ants, for example S. calvus Emerson, S. molestus (Burmeister), S. spinosus (Latreille) [= S. solidus Emerson] (Wheeler 1936; Hermann 1968?; Mill 1984), all of them broadly distributed in Amazonia, except for S. calvus which has only been found in Brazil, French Guiana, and Guyana (Constantino 2020). Most of this data was recorded in a few sites, such as Kartabo and Kaieteur in Guyana, and Vista Alegre, state of Amazonas and Ilha de Maracá state of Roraima, in Brazil. John Lattke observed nocturnal hunting of Syntermes in forests of Cerro Los Pijiguaos, southern Venezuela. A loose column of 6–8 workers making their way through the litter was seen, and a little over an hour later, presumably the same column was seen returning, each worker carrying 2–3 termites in their mandibles. Based on this pattern of specific predatory behavior Wheeler (1936) proposed that the distribution of N. commutata should follow that of their prey. Thus far, this hypothesis has not been challenged, nor tested. N. commutata is an efficient predator, according to Mill (1984) estimations, only three colonies can consume as much as three times the worker and soldier populations of the above–mentioned termite species per hectare, annually. Mill (1982b) also informed on the nest migration strategy of N. commutata in an Amazonian rain forest.

Troya & Lattke, 2022: The venom of N. commutata contains antimicrobial, insecticidal and haemolytic peptides, according to Aili et al. (2016). These proteins are known as ponericins and it is assumed these help prevent diseases originated from the prey organisms brought by the ants inside their colonies (Orivel et al. 2001). Aili et al. (2016) also found in the venom of N. commutata the highest number of proteins as compared to the venoms of the ants Ectatomma brunneum Smith, Myrmecia gulosa (Fabricius), Neoponera apicalis, and Odontomachus hastatus (Fabricius). Schmidt and Overal (2009) found that raiders and nest defenders are equipped with more amounts of venom and that also this is more lethal than in other worker nest mates who do not get involved in foraging activities and colony defense.

This species is associated with menarche and female initiation rites in the Tupi-Guarani language family in South America (Balée, 2000).

Association with Other Organisms
They may be parasitized by a phorid fly (see Wheeler, 1936:174). The workers from different colonies are antagonistic and individuals may be killed (Mill, 1984).



Nomenclature

 * . Ponera commutata Roger, 1860: 311 (w.) (no state data, “amerikanische”).
 * Type-material: 1 syntype worker, 1 syntype queen.
 * [Note: perhaps more syntypes originally, but Latreille, 1802: 220, pl. 7, figs. 44A, 44B, illustrates one of each caste.]
 * Type-locality: none given.
 * Type-depository: MNHN (if syntypes still in existence).
 * [Note: in 1860 Roger nominated commutata as the new species name for the Neotropical specimens of Formica tarsata Fabricius, sensu Latreille, 1802c: 200 (misidentification). Neither Latreille nor Roger cited a Neotropical type-locality for the taxon. Two years earlier Smith, F. 1858b: 94, had observed that this was a Neotropical species, not Afrotropical (see his discussion), and gave Guyana and Brazil (Pará) as localities.]
 * Wheeler, W.M. 1936d: 170 (q.); Borgmeier, 1959a: 313 (m.).
 * Combination in Termitopone (Syntermitopone): Wheeler, W.M. 1936d: 169;
 * combination in Pachycondyla: Emery, 1890a: 72; Brown, in Bolton, 1995b: 304;
 * combination in Neoponera: Emery, 1901a: 47; Schmidt, C.A. & Shattuck, 2014: 151.
 * Status as species: Mayr, 1863: 447; Mayr, 1884: 32; Emery, 1890a: 72 (in key); Emery, 1892b: 163; Emery, 1906c: 114; Emery, 1911d: 71; Mann, 1916: 410; Wheeler, W.M. 1916c: 2; Wheeler, W.M. 1918b: 24; Forel, 1921b: 133; Borgmeier, 1923: 65; Wheeler, W.M. 1923a: 2; Wheeler, W.M. 1925a: 5; Menozzi, 1935b: 190; Wheeler, W.M. 1936d: 169 (redescription); Santschi, 1939f: 160; Borgmeier, 1959a: 313; Kempf, 1961b: 494; Kempf, 1970b: 327; Kempf, 1972a: 249; Brandão, 1991: 381; Bolton, 1995b: 304; Wild, 2002: 7; Wild, 2007b: 39; Mackay, Mackay, et al. 2008: 189; Mackay & Mackay, 2010: 257 (redescription); Bezděčková, et al. 2015: 123; Feitosa, 2015c: 99; Fernández & Guerrero, 2019: 533.
 * [Note: Emery, 1892b: 163, suggests that commutata may be synonymous with Formica ovata Reich, 1793: 132, but the latter is unidentifiable to genus (Bolton, 1995b: 304).]
 * Distribution: Bolivia, Brazil, Colombia, Costa Rica, Ecuador, French Guiana, Guyana, Paraguay, Peru, Suriname, Venezuela.

Worker
From Mackay and Mackay (2010): The worker is a large (total length 17 mm) shining black ant. The mandibles have many teeth, the anterior border of the clypeus is broadly convex, the eye is large (maximum diameter 1 mm), much greater in length than the distance between the anterior edge of the eye and the anterior edge of the head (side view). The scapes extend about the first two funicular segments past the posterior lateral corner of the head. The malar carina is present, but the pronotal carina is absent. The mesosoma is depressed at the metanotal suture, which is well marked on the dorsum. The propodeal spiracle is elongated. The petiole is wide when viewed in profile with the anterior and posterior faces being almost parallel and with the dorsal face rounded and forming the highest point near the posterior edge of the petiole. The posterior lateral carina of the petiole is well developed and sharp. The stridulatory file on the second pretergite is present, the arolia are poorly developed.

Erect and suberect hairs are abundant on most surfaces; appressed golden pubescence is sparse, except near the insertions of the antennae, on the anterior edge of the pronotum and on the middle and posterior coxae.

Most surfaces are shiny black, striae cover much of the dorsum of the head, diverging posteriorly and converging on the ventral surface of the head. The side of the propodeum has obvious oblique striae.

Queen
From Mackay and Mackay (2010): The female is a large (total length 19 mm) shining black ant. The head and the side of the mesosoma are covered with striae. The eyes are large, covering about one third of the side of the head and located about one half of their maximum diameter from the anterior margin of the head. The malar carina is well developed, but the pronotal carina is absent. The propodeal spiracle is slit-shaped. The petiole is similar to that of the worker with the anterior face being nearly straight and meeting the broadly rounded posterior face at the apex and with the posterior lateral margins forming sharp carinae.

The pilosity and sculpture are similar to those of the worker.

Male
Males are not known for this species.

Type Material
South America

Etymology
Apparently the name of this species is derived from the Greek words kommos meaning embellishment and the intensifier tatos, meaning they are really shiny. (Mackay and Mackay 2010)

References based on Global Ant Biodiversity Informatics

 * Aldana delq Torre R. C., and P. Chacon de Ulloa. 1999. Megadiversidad de hormigas (Hymenoptera: Formicidae) de la cuenca media del rio Calima. Revista Colombiana de Entomologia 25(1-2): 37-47.
 * 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.
 * Borgmeier T. 1923. Catalogo systematico e synonymico das formigas do Brasil. 1 parte. Subfam. Dorylinae, Cerapachyinae, Ponerinae, Dolichoderinae. Archivos do Museu Nacional (Rio de Janeiro) 24: 33-103.
 * Borgmeier T. 1959. Myrmecologische Studien. II. Anais da Academia Brasileira de Ciencias 31: 309-319.
 * Brandao, C.R.F. 1991. Adendos ao catalogo abreviado das formigas da regiao neotropical (Hymenoptera: Formicidae). Rev. Bras. Entomol. 35: 319-412.
 * Emery C. 1906. Studi sulle formiche della fauna neotropica. XXVI. Bullettino della Società Entomologica Italiana 37: 107-194.
 * Emery C. 1911. Hymenoptera. Fam. Formicidae. Subfam. Ponerinae. Genera Insectorum 118: 1-125.
 * Escalante Gutiérrez J. A. 1993. Especies de hormigas conocidas del Perú (Hymenoptera: Formicidae). Revista Peruana de Entomología 34:1-13.
 * 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., E. E. Palacio, W. P. Mackay, and E. S. MacKay. 1996. Introducción al estudio de las hormigas (Hymenoptera: Formicidae) de Colombia. Pp. 349-412 in: Andrade M. G., G. Amat García, and F. Fernández. (eds.) 1996. Insectos de Colombia. Estudios escogidos. Bogotá: Academia Colombiana de Ciencias Exactas, Físicas y Naturales, 541 pp
 * Fernández F., and T. M. Arias-Penna. 2008. Las hormigas cazadoras en la región Neotropical. Pp. 3-39 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.
 * Fernández, F. and S. Sendoya. 2004. Lista de las hormigas neotropicales. Biota Colombiana Volume 5, Number 1.
 * Forel A. 1921. Quelques fourmis des environs de Quito (Ecuador) récoltées par Mlle Eléonore Naumann. Bulletin de la Société Vaudoise des Sciences Naturelles 54: 131-135.
 * 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.
 * Gomez V. E. S., and G. Z. González. 2007. Catalogo de Las Hormigas Presentes en El Museo de Historia Natural de la Universidad del Cauca. Popayán : 1-58.
 * 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).
 * 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.
 * 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.
 * Menozzi C. 1935. Spedizione del Prof. Nello Beccari nella Guiana Britannica. Hymenoptera-Formicidae. Redia. 21: 189-203.
 * 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.
 * 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.
 * 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.
 * Souza J. L. P., C. A. R. Moura, A. Y. Harada, and E. Franklin. 2007. Diversity of species of the genera Crematogaster, Gnamptogenys and Pachycondyla, (Hymenoptera: Formicidae) and complementarity of sampling methods during the dry season in an ecological station in the Brazilian state of Pará. Acta Amazonica 37(4): 649-656.
 * 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
 * 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., 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
 * 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. 1936. Ecological relations of ponerine and other ants to termites. Proceedings of the American Academy of Arts and Sciences. 71: 159-243.
 * Wild A. L. 2002. The genus Pachycondyla (Hymenoptera: Formicidae) in Paraguay. Boletín del Museo Nacional de Historia Natural del Paraguay 14: 1-18
 * 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.