Paraponera clavata

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Paraponera clavata
Scientific classification
Kingdom: Animalia
Phylum: Arthropoda
Class: Insecta
Order: Hymenoptera
Family: Formicidae
Subfamily: Paraponerinae
Genus: Paraponera
Species: P. clavata
Binomial name
Paraponera clavata
(Fabricius, 1775)

Paraponera clavata casent0006789 profile 1.jpg

Paraponera clavata casent0006789 dorsal 1.jpg

Specimen Label

Synonyms

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.

At a Glance • Tandem running  

Photo Gallery

  • Paraponera clavata worker, Pacaya Samiria National Reserve, Peru. Photo by Michael Angelo.
  • Foraging worker. Photo by Piotr Naskrecki.
  • Trophallaxis between workers. Photo by Piotr Naskrecki.
  • Paraponera clavata worker, Pacaya Samiria National Reserve, Peru. Photo by Michael Angelo.
  • Paraponera clavata worker. Image by Alex Wild.
  • Paraponera clavata queen. Photo by Alex Wild.
  • Paraponera clavata infected by the fungus Ophiocordycceps ponerinarum. In this association, the fungus leads the ant to climb up on a tree, to die attached to the trunk. Photo by João P. M. Araújo.
  • Paraponera clavata attached by a Ophiocordyceps fungus species. Photo by Piotr Naskrecki.
  • Paraponera clavata queen, Ophiocordyceps sp. fungus. Orellana Province, Ecuador, Catta.
  • A crab spider (top left) has killed a bullet ant queen and at least 2 species of small flies (Milichiidae & Phoridae) have arrived to feed from the carcass and/or lay their eggs. Amazonian Peru. Photo by Alex Wild.

Identification

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

Distribution

Latitudinal Distribution Pattern

Latitudinal Range: 50.674633° to -29.755°.

       
North
Temperate 		North
Subtropical 		Tropical South
Subtropical 		South
Temperate

Distribution based on Regional Taxon Lists

Neotropical Region: Argentina, 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.

Distribution based on AntMaps

AntMapLegend.png

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.
pChart

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.

Young & Hermann (1980) found that tandem running occurs in this species.

Association with Other Organisms

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Diptera

  • This species is a host for the phorid fly Apocephalus paraponerae (a parasite) (phorid.net) (attacked).
  • This species is a host for the phorid fly Apocephalus paraponerae (a parasite) (Brown et al., 2015) (attracted to injured).
  • This species is a host for the phorid fly Apocephalus tanyurus (a parasite) (phorid.net) (attacked).
  • This species is a host for the phorid fly Apocephalus tanyurus (a parasite) (Brown et al., 2015) (injured).
  • This species is a host for the phorid fly Beckerina sp. (a parasite) (Brown et al., 2015) (injured).
  • This species is a host for the phorid fly Beckerina sp. (a parasitoid) (Quevillon, 2018) (encounter mode primary; direct transmission; transmission outside nest).
  • This species is a host for the phorid fly Megaselia aurea (a parasite) (Brown et al., 2015) (injured).
  • This species is a host for the phorid fly Megaselia sp. (a parasite) (Brown et al., 2015) (injured).
  • This species is a host for the phorid fly Megaselia sp. (a parasitoid) (Quevillon, 2018) (encounter mode primary; direct transmission; transmission outside nest).

Nematode

  • This species is a host for the nematode Mermithidae (unspecified "Mermix") (a parasite) in Neotropics (Wheeler, 1928; Laciny, 2021).

Fungi

  • This species is a host for the fungus Beauveria bassiana (a parasitoid) (Quevillon, 2018) (encounter mode primary; direct transmission; transmission within nest).
  • This species is a host for the fungus Ophiocordyceps kniphofioides (a parasitoid) (Quevillon, 2018) (encounter mode primary; direct transmission; transmission outside nest).
  • This species is a host for the fungus Cordyceps doiana (a pathogen) in Colombia (Kobayasi, 1981; Araujo et al., 2018; Sanjuan et al., 2015; Shrestha et al., 2017).
  • This species is a host for the fungus Ophiocordyceps australis (a pathogen) (Araujo et al., 2018; Sanjuan et al., 2015; Shrestha et al., 2017).
  • This species is a host for the fungus Ophiocordyceps ponerinarum (a pathogen) (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).

Laboratory colony of P. clavata, showing dimorphism between founding queen (bottom right) and its first workers. From Villa Carmen, Peru. Photo by Christian Peeters.

Flight Period

X X X X X X X X X X X X
Jan Feb Mar Apr May Jun Jul Aug Sep Oct Nov Dec

Source: Kannowski, 1991.

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Castes

  • Paraponera clavata worker from Jatun Sacha reserve, Ecuador. Photo by Alex Wild.

Specimens from Villa Carmen, near Pilcopata (Peru), imaged by Mônica Antunes Ulysséa

Queen

(abdomen partly removed for dissection of ovaries)

Paraponera Queen L Mônica Antunes Ulysséa.jpg.jpg
Paraponera Queen D Mônica Antunes Ulysséa.jpg
Paraponera Queen F Mônica Antunes Ulysséa.jpg

Worker

Paraponera Worker L Mônica Antunes Ulysséa.jpg
Paraponera Worker D Mônica Antunes Ulysséa.jpg
Paraponera Worker F Mônica Antunes Ulysséa.jpg.jpg
Paraponera-clav-Gyne.jpgParaponera clavata worker head, antennal grooves.jpgParaponera clavata worker thorax.jpgParaponera clavata worker petiole.jpgParaponera clavata worker front leg.jpg
.

Worker

Images from AntWeb

Paraponera clavata casent0006789 head 1.jpgParaponera clavata casent0006789 profile 1.jpgParaponera clavata casent0006789 dorsal 1.jpgParaponera clavata casent0006789 label 1.jpg
Worker. Specimen code casent0006789. Photographer April Nobile, uploaded by California Academy of Sciences. Owned by CAS, San Francisco, CA, USA.
Paraponera clavata casent0106092 head 1.jpgParaponera clavata casent0106092 profile 1.jpgParaponera clavata casent0106092 dorsal 1.jpgParaponera clavata casent0106092 label 1.jpg
Worker. Specimen code casent0106092. Photographer Michael Branstetter, uploaded by California Academy of Sciences. Owned by UCDC, Davis, CA, USA.
Paraponera clavata casent0003165 head 1.jpgParaponera clavata casent0003165 profile 1.jpgParaponera clavata casent0003165 dorsal 1.jpgParaponera clavata casent0003165 label 1.jpg
Worker. Specimen code casent0003165. Photographer April Nobile, uploaded by California Academy of Sciences. Owned by CAS, San Francisco, CA, USA.

ParaponeraEconomo-header (arilab.unit.oist.jp).png  X-ray micro-CT scan 3D model of Paraponera clavata (worker) prepared by the Economo lab at OIST.

Paraponera clavata. This a large, ponerine ant occurs in Central and South American rainforests. Due to its large and painful stinger it is also known as “bullet ant” or “bala” (located at OIST: CASENT0745697). See on Sketchfab. See list of 3D images.

Nomenclature

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

  • clavata. 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.

Description

Karyotype

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  • n = 33, 2n = 66, karyotype = 66A? (Brazil) (Costa et al., 2005; Mariano et al., 2015).
  • n = 27, 2n = 54, karyotype = 12M + 42A (Brazil) (Costa et al., 2005; Mariano et al., 2015).

References

References based on Global Ant Biodiversity Informatics

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