Paraponera clavata

Paraponera clavata is a very large (> 2 cm long) and potentially aggressive ant of the New World tropics. Known within its range as the "bala" ant they pack a powerful sting. Workers forage arboreally at all heights in the canopy; nests are subterranean at the bases of trees, or occasionally in humus accumulations in the canopy; workers forage on live prey and extrafloral nectar; males are frequent at blacklights.

Identification
Its large size, conspicuous antennal scrobes, and the uniquely shaped petiole make this monotypic genus immediately recognizable.

Distribution based on Regional Taxon Lists
Neotropical Region: Bolivia, Brazil, Colombia, Costa Rica, Ecuador, French Guiana, Guyana, Honduras, Nicaragua, Panama, Paraguay, Peru, Suriname.

It is found in Ecuador, Brazil, Bolivia, Colombia, Peru, Suriname, Paraguay, French Guiana, Guyana, Honduras, Nicaragua, Panama and Costa Rica.

Biology
Wetterer (1994) - P. clavata usually nest at the base of trees, but some have arboreal nests (Breed & Harrison 1989). Colonies can grow to have more than 2000 workers (Breed & Harrison 1988). Workers usually ascend their nest tree to forage in the foliage of the canopy and understory. Foragers most commonly return to the nest carrying drops of liquid in their mandibles, but they also bring back plant parts and captured invertebrate prey (Young & Hermann 1980, Bennet & Breed 1985). Foragers collect plant extrafloral nectaries (Young 1977). Janzen and Carroll (1983) observed P. clavata workers guarding and collecting nectar from extrafloral nectaries of Pentaclethra macroloba and other plants.

Morrison (2018) found cane toads were a major predator of P. clavata. Toads were observed sitting adjacent to nests and eating up to an ant a minute. The predation was deemed to have caused the demise of 5 of 12 observed nests in the Panama study area.

Fungi
This taxon is a host for the fungi , and (Araujo et al., 2018; Sanjuan et al., 2015; Shrestha et al., 2017).

Sting
Alex Wild provides the following notes on the sting of this ant:

I was stung by a bullet ant last week in Costa Rica. On purpose. How did it feel? Bearable. Given this species’ fearsome reputation, I was expecting worse. It certainly hurt, though.

It wasn’t just the initial sear from the sting’s penetration, imparting all the sharpness one would anticipate from a relatively large hymenopteran, but the way the pain sank beneath the skin.

The bullet ant has a reputation for feeling like a firearm wound. Having never been shot, I can’t make much of the comparison. I imagine an actual shooting would be far more traumatic, but all the same I understand where the name comes from. A Paraponera sting feels more profound than the average insect sting. Like tissue or bone damage, it is a deep throbbing ache that crescendos over several hours. Unlike a honey bee sting, whose sharpness gives way quickly to a dull itch, the bullet ant’s sting is the gift that keeps on giving. Less a gunshot, I suppose, than the lasting pain following a solid crowbar to the arm. Although bearable, mine still ached when I went to bed 8 hours later. All pain was gone in the morning.

We tend not to make much of where on the body we’re stung, but stings are like real estate. Location, location, location. The forearm is a relatively mild substrate, a safe place to experiment with stings. I was once zinged on the tip of the nose by a common honey bee. Holy bejeezus. I’ll take twelve bullet ants to the arm before I wish to relive that one.

Independent colony foundation
Founding queens need to hunt before the first generation of workers become adult (i.e. non-claustral foundation). Trophic eggs were observed to be laid by first workers and given directly to medium and mature larvae (Peeters 2017).



Queen
(abdomen partly removed for dissection of ovaries)

Nomenclature

 * . Formica clavata Fabricius, 1775: 394 (w.) (no state data, “Habitat in India”).
 * Type-material: holotype (?) worker.
 * [Note: no indication of number of specimens is given.]
 * Type-locality: “Habitat in India” (error).
 * [Note: type-locality incorrect as distribution of this species is Neotropical. Mayr, 1862: 731, records material from Brazil, Suriname, and French Guiana.]
 * Type-depository: ZMUC.
 * Lepeletier, 1835: 189 (w.q.); Smith, F. 1858b: 101 (m.); Mayr, 1862: 731 (q.m.); Wheeler, G.C. & Wheeler, J. 1952a: 117 (l.).
 * Combination in Ponera: Latreille, 1809: 128; Latreille, 1818c: 570;
 * combination in Paraponera: Smith, F. 1858b: 100.
 * [Combination in Camponotus: Mayr, 1884: 30 (error).]
 * Status as species: Christ, 1791: 507; Fabricius, 1793: 360; Fabricius, 1804: 410; Lepeletier de Saint-Fargeau, 1835: 188; Smith, F. 1858b: 100; Roger, 1861a: 19; Mayr, 1862: 731; Roger, 1863b: 18; Mayr, 1863: 440; Emery, 1890b: 42; Dalla Torre, 1893: 18; Emery, 1894k: 47; Forel, 1895b: 111; Forel, 1899c: 10; Emery, 1904b: 599; Wheeler, W.M. 1905b: 120; Forel, 1907e: 1; Forel, 1908c: 340; Forel, 1909b: 58; Emery, 1911d: 27; Santschi, 1913h: 33; Mann, 1916: 402; Wheeler, W.M. 1916c: 2; Crawley, 1916b: 377; Luederwaldt, 1918: 34; Wheeler, W.M. 1923a: 2; Borgmeier, 1923: 52; Wheeler, W.M. 1925a: 2; Menozzi, 1935b: 190; Santschi, 1939f: 160; Brown, 1958g: 205; Kempf, 1970b: 324; Kempf, 1972a: 181; Fernández, 1993: 253; Bolton, 1995b: 312; Wild, 2007b: 38; Arias-Penna, 2008c: 120; Bezděčková, et al. 2015: 122; Feitosa, 2015c: 98; Delsinne, Sonet & Arias-Penna, T.M. 2019: 637.
 * Senior synonym of aculeata: Latreille, 1802c: 207; Lepeletier de Saint-Fargeau, 1835: 188; Mayr, 1863: 440; Roger, 1863b: 18; Dalla Torre, 1893: 19; Forel, 1895b: 111; Forel, 1899c: 10; Emery, 1911d: 28; Santschi, 1913h: 33; Borgmeier, 1923: 52; Kempf, 1972a: 181; Bolton, 1995b: 312.
 * Senior synonym of tarsalis: Smith, F. 1858b: 100; Mayr, 1863: 440; Roger, 1863b: 18; Dalla Torre, 1893: 19; Forel, 1895b: 111; Forel, 1899c: 10; Emery, 1911d: 28; Santschi, 1913h: 33; Borgmeier, 1923: 52; Kempf, 1972a: 181; Bolton, 1995b: 312.
 * Material of the unavailable name spininoda referred here by Latreille, 1802c: 207; Fabricius, 1804: 410; Lepeletier de Saint-Fargeau, 1835: 188; Smith, F. 1858b: 100; Mayr, 1863: 440; Roger, 1863b: 18; Dalla Torre, 1893: 19; Forel, 1895b: 111; Forel, 1899c: 10; Emery, 1911d: 28; Borgmeier, 1923: 52; Kempf, 1972a: 181; Bolton, 1995b: 312.
 * Distribution: Bolivia, Brazil, Colombia, Costa Rica, Ecuador, French Guiana, Guyana, Nicaragua, Panama, Paraguay, Peru, Suriname, Venezuela.
 * aculeata. Formica aculeata Olivier, 1792: 498 (w.) FRENCH GUIANA.
 * Type-material: holotype (?) worker.
 * [Note: no indication of number of specimens is given.]
 * Type-locality: French Guiana: Cayenne (Tugni?).
 * Type-depository: MNHN.
 * [Misspelled as armata by Mayr, 1863: 440.]
 * Junior synonym of clavata: Latreille, 1802c: 207; Lepeletier de Saint-Fargeau, 1835: 188; Mayr, 1863: 440; Roger, 1863b: 18; Dalla Torre, 1893: 19; Forel, 1895b: 111; Forel, 1899c: 10; Emery, 1911d: 28; Santschi, 1913h: 33; Borgmeier, 1923: 52; Kempf, 1972a: 181; Bolton, 1995b: 312.
 * tarsalis. Ponera tarsalis Perty, 1833: 135, pl. 27, fig. 2 (w.) BRAZIL (no state data).
 * Type-material: holotype (?) worker.
 * [Note: no indication of number of specimens is given.]
 * Type-locality: Brazil: “Habitat in sylvis secundum flumen Solimoes”.
 * Type-depository: MNHN.
 * Junior synonym of clavata: Smith, F. 1858b: 100; Mayr, 1863: 440; Roger, 1863b: 18; Dalla Torre, 1893: 19; Forel, 1895b: 111; Forel, 1899c: 10; Emery, 1911d: 28; Santschi, 1913h: 33; Borgmeier, 1923: 52; Kempf, 1972a: 181; Bolton, 1995b: 312.

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.
 * Alonso L. E., and J. A. Helms. 2013. A Rapid Assessment of the Ants of the Grensgebergte and Kasikasima Regions of Southeastern Suriname. A Rapid Biological Assessment of the Upper Palumeu River Watershed (Grensgebergte and Kasikasima) of Southeastern Suriname: 109-118.
 * Arias-Penna T. M. 2008. Subfamilia Paraponerinae. Pp. 119-122 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.
 * Astruc C., J. F. Julien, C. Errard, and A. Lenoir. 2004. Phylogeny of ants based on morphology and DNA sequence data. Molecular Phylogenetics and Evolution 31: 880-893.
 * 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.
 * 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.
 * Boudinot B. E. 2015. Contributions to the knowledge of Formicidae (Hymenoptera, Aculeata): a new diagnosis of the family, the first global male-based key to subfamilies, and a treatment of early branching lineages. European Journal of Taxonomy 120: 1-62.
 * Breed, M.D. 2002. Allometry in the giant tropical ant, Paraponera clavata. Insectes Sociaux 49:125-128
 * Bustos H., J. 1994. Contribucion al conocimiento de al fauna de hormigas (Hymenoptera: Formicidae) del occidente del Departamento de Narino (Colombia). Bol. Mus. Ent. Univ. Valle 2(1,2):19-30
 * COSTA, M.S.; MARIANO, C.S.F.; FERNANDES, A. M.; POMPOLO, S. G.; DELABIE, J. H. C. 2005. Variação cariotípica em populações distantes de Paraponeraclavata(Fabricius) (Formicidae, Paraponerinae). In: XVII SIMPÓSIO DE MIRMECOLOGIA, Campo Grande. Resumos XVII Simpósio de Mirmecologia Biodiversidade e Conservação. Campo Grande: 2005, p. 1-3.
 * Crawley W. C. 1916. Ants from British Guiana. Ann. Mag. Nat. Hist. 8(17): 366-378.
 * Dejean A., A. Compin, J. H. C. Delabie, F. Azemar, B. Corbara, and M. Leponce. 2019. Biotic and abiotic determinants of the formation of ant mosaics in primary Neotropical rainforests. Ecological Entomology https://doi-org.eproxy.lib.hku.hk/10.1111/een.12735
 * Emery C. 1896. Formiche raccolte dal dott. E. Festa nei pressi del golfo di Darien. Bollettino dei Musei di Zoologia ed Anatomia Comparata della Reale Università di Torino 11(229): 1-4.
 * 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.
 * Escalante J. A. 1975. Hormigas de la Provincia de la Convencionm Cusco. Revista Peruana de Entomologia 18:125-126.
 * Fernández F. 1993. Hormigas de Colombia III: los géneros Acanthoponera Mayr, Heteroponera Mayr y Paraponera Fr. Smith (Formicidae: Ponerinae: Ectatommini). Caldasia 17: 249-258.
 * 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.
 * Field Museum Collection, Chicago, Illinois (C. Moreau)
 * 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.
 * 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.
 * INBio Collection (via Gbif)
 * Jaffe K., P. Horchler, M. Verghaagh, C. Gomez, R. Sievert, R. Jaffe, and W. Morawetz. 2007. Comparing the ant fauna in a tropical and a temperate forest canopy. Ecotropicos 20(2): 74-81.
 * Jaffe, K., et al. 2007. Comparing the ant fauna in a tropical and a temperat forest canopy. Ecotropicos 20(2):74-81
 * Kempf W. W. 1978. A preliminary zoogeographical analysis of a regional ant fauna in Latin America. 114. Studia Entomologica 20: 43-62.
 * Kempf, W.W. 1972. Catalago abreviado das formigas da regiao Neotropical (Hym. Formicidae) Studia Entomologica 15(1-4).
 * Kugler C. 1991. Stings of ants of the tribe Ectatommini (Formicidae: Ponerinae). Insecta Mundi 5: 153-166.
 * Kusnezov N. 1953. La fauna mirmecológica de Bolivia. Folia Universitaria. Cochabamba 6: 211-229.
 * Lachaud J. P., P. Cerdan, and G. Perez-Lachaud. 2012. Poneromorph ants associated with parasitoid wasps of the genus Kapala Cameron (Hymenoptera: Eucharitidae) in French Guiana. Psyche doi:10.1155/2012/393486.
 * Longino J. et al. ADMAC project. Accessed on March 24th 2017 at https://sites.google.com/site/admacsite/
 * 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
 * Lozano-Zambrano F. H., E. Jimenez, T. M. Arias-Penna, A. M. Arcila, J. Rodriguez, and D. P. Ramirez. 2008. Biogeografía de las hormigas cazadoras de Colombia. Pp. 349-406. 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.
 * Luna Bello G. O. 2005. Composicion y estructura de la comunidad de hormigas a lo largo de un gradiente de intensificacion agricola en zonas de bosque tropical humedo en la region autonoma Atlantico Sur, Nicaragua. PhD Universidad Centroamericana, Facultad de Ciencia y Tecnologia del Ambiente carrera de Ecologi y Desarrollo. 127 pages
 * Maes, J.-M. and W.P. MacKay. 1993. Catalogo de las hormigas (Hymenoptera: Formicidae) de Nicaragua. Revista Nicaraguense de Entomologia 23.
 * 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
 * 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.
 * Mertl A. L., J. F. A. Traniello, K. Ryder Wilkie, and R. Constantino. 2012. Associations of two ecologically significant social insect taxa in the litter of an amazonian rainforest: is there a relationship between ant and termite species richness? Psyche doi:10.1155/2012/312054
 * 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.
 * Radoszkowsky O. 1884. Fourmis de Cayenne Française. Trudy Russkago Entomologicheskago Obshchestva 18: 30-39.
 * Robson Simon Ant Collection, 05-Sept-2014
 * Ryder Wilkie K.T., A. L. Mertl, and J. F. A. Traniello. 2010. Species Diversity and Distribution Patterns of the Ants of Amazonian Ecuador. PLoS ONE 5(10): e13146.doi:10.1371/journal.pone.0013146
 * Sandoval V. E., and G. Zambrano. 2007. Catálogo de las hormigas presentes en el Museo de Historia Natural de la Universidad del Cauca. Taller Editorial de la Universidad del Cauca, Popayán. 60 pp.
 * Silva T. S. R., and R. M. Feitosa. 2019. Using controlled vocabularies in anatomical terminology: A case study with Strumigenys (Hymenoptera: Formicidae). Arthropod Structure and Development 52: 1-26.
 * 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.
 * Smith M. A., W. Hallwachs, D. H. Janzen. 2014. Diversity and phylogenetic community structure of ants along a Costa Rican elevational gradient. Ecography 37(8): 720-731.
 * Tobin E. J. 1997. Competition and coexistence of ants in a small patch of Rainforest Canopy in Peruvian Amazonia. J. New York Entomol. Soc. 105(1-2): 105-122
 * Ulloa Chacon P., M. L. Baena, J. Bustos, R. C. Aldana, J. A. Aldana, and M. A. Gamboa. 1996. Fauna de hormigas del departamento del Valle del Cauca (Colombia). Pp. 413-451. In Andrade-C M. G., G. Amat Garcia, and F. Fernandez. Insectoss de Colombia, estudios escogidos.
 * 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.
 * 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.
 * Wild, A. L. "A catalogue of the ants of Paraguay (Hymenoptera: Formicidae)." Zootaxa 1622 (2007): 1-55.
 * Wilson, E.O. 1987. The Arboreal Ant Fauna of Peruvian Amazon Forests: A First Assessment. Biotropica 19(3):245-251.
 * 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 Souza J. L. P., F. B. Baccarob, V. L. Landeirob, E. Franklinc, and W. E. Magnussonc. 2012. Trade-offs between complementarity and redundancy in the use of different sampling techniques for ground-dwelling ant assemblages. Applied Soil Ecology 56: 63 73.