Paraponera clavata
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''Paraponera clavata'', commonly known as the bullet ant, is a
species A species () is often defined as the largest group of organisms in which any two individuals of the appropriate sexes or mating types can produce fertile offspring, typically by sexual reproduction. It is the basic unit of Taxonomy (biology), ...
of
ant Ants are Eusociality, eusocial insects of the Family (biology), family Formicidae and, along with the related wasps and bees, belong to the Taxonomy (biology), order Hymenoptera. Ants evolved from Vespoidea, vespoid wasp ancestors in the Cre ...
known for its extremely painful sting. It inhabits humid lowland
rainforest Rainforests are forests characterized by a closed and continuous tree Canopy (biology), canopy, moisture-dependent vegetation, the presence of epiphytes and lianas and the absence of wildfire. Rainforests can be generally classified as tropi ...
s in Central and
South America South America is a continent entirely in the Western Hemisphere and mostly in the Southern Hemisphere, with a considerably smaller portion in the Northern Hemisphere. It can also be described as the southern Subregion#Americas, subregion o ...
.


Etymology

The
specific epithet In Taxonomy (biology), taxonomy, binomial nomenclature ("two-term naming system"), also called binary nomenclature, is a formal system of naming species of living things by giving each a name composed of two parts, both of which use Latin gramm ...
of the ant, ', means "club-shaped". The generic name, ''Paraponera'', translates to "near-'' Ponera''". Because of its fearsome reputation, the ant has several Native American, Spanish, and Portuguese local names in different geographical areas; perhaps the best-known of these is the Venezuelan nickname ' (the "24 ant" or "24-hour ant"), referring to the full day of pain that follows being stung; it can also refer to the time it takes to kill a human. In Brazil, the Portuguese names given by locals include ', ', or ' (big black ant); Native American–derived names include ''tocandira'', and ''tocanquibira'', from the Tupi–Guarani ''tuca-ndy'', which translates to "the one wounding deeply". Other names by which it is called include ''chacha'', ''cumanagata'', ''munuri'', ''siámña'', and ''yolosa''. In Costa Rica, ''P. clavata'' is known as ''bala'', meaning "bullet". ''P. clavata'' also has several common names; it is most commonly known as the bullet ant because of the extreme pain it delivers following a sting, similar to that of getting shot. Other names are the "lesser giant hunting ant" and "conga ant".


Taxonomy

''Paraponera clavata'' was first described by Danish
zoologist Zoology ( , ) is the scientific study of animals. Its studies include the structure, embryology, classification, habits, and distribution of all animals, both living and extinct, and how they interact with their ecosystems. Zoology is one ...
Johan Christian Fabricius Johann Christian Fabricius (7 January 1745 – 3 March 1808) was a Danish zoologist, specialising in "Insecta", which at that time included all arthropods: insects, arachnids, crustaceans and others. He was a student of Carl Linnaeus, and is co ...
in 1775, who named it ''Formica clavata'' in his ''Systema entomologiae''. Fabricius listed the type locality as India, incorrectly, as these ants are only found in Central and South America. In 1804, ''P. clavata'' was transferred to the genus '' Ponera'' by the French zoologist
Pierre André Latreille Pierre André Latreille (; 29 November 1762 – 6 February 1833) was a French zoology, zoologist, specialising in arthropods. Having trained as a Roman Catholic priest before the French Revolution, Latreille was imprisoned, and only regained hi ...
. The genus '' Paraponera'' was established by the British entomologist Frederick Smith in 1858, and ''P. clavata'' was designated as the
type species In International_Code_of_Zoological_Nomenclature, zoological nomenclature, a type species (''species typica'') is the species name with which the name of a genus or subgenus is considered to be permanently taxonomically associated, i.e., the spe ...
by
monotypy In biology, a monotypic taxon is a taxonomic group (taxon) that contains only one immediately subordinate taxon. A monotypic species is one that does not include subspecies or smaller, infraspecific taxa. In the case of genera, the term "unispe ...
(the creation of a taxonomic group that includes only a single
taxon In biology, a taxon (back-formation from ''taxonomy''; : taxa) is a group of one or more populations of an organism or organisms seen by taxonomists to form a unit. Although neither is required, a taxon is usually known by a particular name and ...
). In his book, Smith synonymised multiple taxa under ''Paraponera clavata'', including ''Formica armata'', ''Formica spininoda'', ''Ponera tarsalis'', and ''Ponera clavata''. Later publications would also synonymise more taxa, including ''Formica aculeata'' and ''Formica clavata''. The genus was placed in a monotypic
tribe The term tribe is used in many different contexts to refer to a category of human social group. The predominant worldwide use of the term in English is in the discipline of anthropology. The definition is contested, in part due to conflict ...
, the Paraponerini, in 1901 by the Italian entomologist
Carlo Emery Carlo Emery (25 October 1848, Naples – 11 May 1925) was an Italian entomologist. He is remembered for Emery's rule, which states that insect social parasitism (biology), social parasites are often closely related to their hosts. Early in hi ...
, who stressed the importance of certain morphological characteristics of ''Paraponera''; Emery had also placed the tribe close to the Ectatommini. This classification was accepted by the entomological community until 1958, when the American entomologist William Brown Jr. synonymised Paraponerini and transferred ''Paraponera'' to Ectatommini. It was treated as a valid tribe in 1994, but in 2003, the English myrmecologist elevated the rank of the tribe to subfamily level as Paraponerinae, part of the Poneromorph subfamilies. Under the present classification, the bullet ant is a member of the genus ''Paraponera'' in the tribe Paraponerini, subfamily Paraponerinae. It is a member of the family Formicidae, belonging to the order
Hymenoptera Hymenoptera is a large order of insects, comprising the sawflies, wasps, bees, and ants. Over 150,000 living species of Hymenoptera have been described, in addition to over 2,000 extinct ones. Many of the species are parasitic. Females typi ...
. It was once the sole member of its own genus and tribe, until the extinct '' Paraponera dieteri'' was described in 1994 by the entomologist Cesare Baroni Urbani. The ant, described from
Dominican amber Dominican amber is amber from the Dominican Republic derived from resin of the extinct tree '' Hymenaea protera''. Dominican amber differentiates itself from Baltic amber by being nearly always transparent, and it has a higher number of fossil in ...
, existed during the
Early Miocene The Early Miocene (also known as Lower Miocene) is a sub-epoch of the Miocene epoch (geology), Epoch made up of two faunal stage, stages: the Aquitanian age, Aquitanian and Burdigalian stages. The sub-epoch lasted from 23.03 ± 0.05 annum, Ma to ...
15 to 45 million years ago. ''P. dieteri'' can be distinguished from ''P. clavata'' by its much narrower head, length, pronotum width, petiole width, and other features. The good preservation of the fossil allowed comprehensive comparisons between the two species; the body sculpture of ''P. dieteri'' suggests that the genus as a whole exhibits a slow evolutionary rate. ''P. clavata'' is the only living species in its subfamily. Although ''P. dieteri'' was the first extinct relative of ''P. clavata'' to be described, another ''Paraponera'' fossil had been examined earlier in the 1980s. The fossil, which was from the
Miocene The Miocene ( ) is the first epoch (geology), geological epoch of the Neogene Period and extends from about (Ma). The Miocene was named by Scottish geologist Charles Lyell; the name comes from the Greek words (', "less") and (', "new") and mea ...
, was found embedded in
Dominican amber Dominican amber is amber from the Dominican Republic derived from resin of the extinct tree '' Hymenaea protera''. Dominican amber differentiates itself from Baltic amber by being nearly always transparent, and it has a higher number of fossil in ...
from
Hispaniola Hispaniola (, also ) is an island between Geography of Cuba, Cuba and Geography of Puerto Rico, Puerto Rico in the Greater Antilles of the Caribbean. Hispaniola is the most populous island in the West Indies, and the second-largest by List of C ...
; at the time of discovery, the ant was the largest fossil of its kind. It showed similar characteristics to ''P. clavata'', although it was considerably smaller. The fossil also has biogeographic importance. As ''P. clavata'' is not found in the
Greater Antilles The Greater Antilles is a grouping of the larger islands in the Caribbean Sea, including Cuba, Hispaniola, Puerto Rico, and Jamaica, together with Navassa Island and the Cayman Islands. Seven island states share the region of the Greater Antille ...
, but rather in Central and South America, this suggests that moister tropical forests covered the island during the
Tertiary period The Tertiary ( ) is an obsolete Period (geology), geologic period spanning 66 million to 2.6 or 1.8 million years ago. The period began with the extinction of the non-bird, avian dinosaurs in the Cretaceous–Paleogene extinction event, at t ...
. This is further supported by the fact that ''P. clavata'' is a forest ant that forages on the ground and up into bushes and trees.


Description

Worker ants are long and resemble stout, reddish-black wingless
wasp A wasp is any insect of the narrow-waisted suborder Apocrita of the order Hymenoptera which is neither a bee nor an ant; this excludes the broad-waisted sawflies (Symphyta), which look somewhat like wasps, but are in a separate suborder ...
s. ''Paraponera'' is
predator Predation is a biological interaction in which one organism, the predator, kills and eats another organism, its prey. It is one of a family of common List of feeding behaviours, feeding behaviours that includes parasitism and micropredation ...
y, and like all primitive poneromorphs, does not display polymorphism in the worker caste; the
queen ant A queen ant (also known as a gyne) is an adult, reproducing female ant in an ant colony; she is usually the mother of all the other ants in that colony. Some female ants, such as the '' Cataglyphis'', do not need to mate to produce offspring, rep ...
is not much larger than the workers. They are not aggressive ants but are vicious when defending the nest, they produce a stridulating sound and sting with ferocity.


Distribution

''Paraponera'' is distributed throughout Central and
South America South America is a continent entirely in the Western Hemisphere and mostly in the Southern Hemisphere, with a considerably smaller portion in the Northern Hemisphere. It can also be described as the southern Subregion#Americas, subregion o ...
, commonly found in the wet
Neotropical realm The Neotropical realm is one of the eight biogeographic realms constituting Earth's land surface. Physically, it includes the tropics, tropical Ecoregion#Terrestrial, terrestrial ecoregions of the Americas and the entire South American temperat ...
. These ants are found in
Honduras Honduras, officially the Republic of Honduras, is a country in Central America. It is bordered to the west by Guatemala, to the southwest by El Salvador, to the southeast by Nicaragua, to the south by the Pacific Ocean at the Gulf of Fonseca, ...
,
El Salvador El Salvador, officially the Republic of El Salvador, is a country in Central America. It is bordered on the northeast by Honduras, on the northwest by Guatemala, and on the south by the Pacific Ocean. El Salvador's capital and largest city is S ...
,
Nicaragua Nicaragua, officially the Republic of Nicaragua, is the geographically largest Sovereign state, country in Central America, comprising . With a population of 7,142,529 as of 2024, it is the third-most populous country in Central America aft ...
,
Costa Rica Costa Rica, officially the Republic of Costa Rica, is a country in Central America. It borders Nicaragua to the north, the Caribbean Sea to the northeast, Panama to the southeast, and the Pacific Ocean to the southwest, as well as Maritime bo ...
and
Panama Panama, officially the Republic of Panama, is a country in Latin America at the southern end of Central America, bordering South America. It is bordered by Costa Rica to the west, Colombia to the southeast, the Caribbean Sea to the north, and ...
from the north, and in
Venezuela Venezuela, officially the Bolivarian Republic of Venezuela, is a country on the northern coast of South America, consisting of a continental landmass and many Federal Dependencies of Venezuela, islands and islets in the Caribbean Sea. It com ...
,
Colombia Colombia, officially the Republic of Colombia, is a country primarily located in South America with Insular region of Colombia, insular regions in North America. The Colombian mainland is bordered by the Caribbean Sea to the north, Venezuel ...
,
Ecuador Ecuador, officially the Republic of Ecuador, is a country in northwestern South America, bordered by Colombia on the north, Peru on the east and south, and the Pacific Ocean on the west. It also includes the Galápagos Province which contain ...
,
Peru Peru, officially the Republic of Peru, is a country in western South America. It is bordered in the north by Ecuador and Colombia, in the east by Brazil, in the southeast by Bolivia, in the south by Chile, and in the south and west by the Pac ...
,
Bolivia Bolivia, officially the Plurinational State of Bolivia, is a landlocked country located in central South America. The country features diverse geography, including vast Amazonian plains, tropical lowlands, mountains, the Gran Chaco Province, w ...
, and
Brazil Brazil, officially the Federative Republic of Brazil, is the largest country in South America. It is the world's List of countries and dependencies by area, fifth-largest country by area and the List of countries and dependencies by population ...
from the south. Colonies are found in lowland areas, at elevations ranging from sea level to . However, specimens have been collected at elevations of in La Amistad International Park.


Habitat

Colonies consist of several hundred individuals and are usually situated at the bases of trees. Workers
forage Forage is a plant material (mainly plant leaves and stems) eaten by grazing livestock. Historically, the term ''forage'' has meant only plants eaten by the animals directly as pasture, crop residue, or immature cereal crops, but it is also used m ...
arboreally for small
arthropods Arthropods ( ) are invertebrates in the phylum Arthropoda. They possess an arthropod exoskeleton, exoskeleton with a cuticle made of chitin, often Mineralization (biology), mineralised with calcium carbonate, a body with differentiated (Metam ...
and
nectar Nectar is a viscous, sugar-rich liquid produced by Plant, plants in glands called nectaries, either within the flowers with which it attracts pollination, pollinating animals, or by extrafloral nectaries, which provide a nutrient source to an ...
in the area directly above the nest, often as far as the upper
canopy Canopy may refer to: Plants * Canopy (biology), aboveground portion of plant community or crop (including forests) * Canopy (grape), aboveground portion of grapes Religion and ceremonies * Baldachin or canopy of state, typically placed over an a ...
; little foraging occurs on the forest floor.
Nectar Nectar is a viscous, sugar-rich liquid produced by Plant, plants in glands called nectaries, either within the flowers with which it attracts pollination, pollinating animals, or by extrafloral nectaries, which provide a nutrient source to an ...
, carried between the mandibles, is the most common food taken back to the nest by foragers. Two studies in Costa Rica and on Barro Colorado Island (BCI) found about four bullet ant nests per
hectare The hectare (; SI symbol: ha) is a non-SI metric unit of area equal to a square with 100-metre sides (1 hm2), that is, square metres (), and is primarily used in the measurement of land. There are 100 hectares in one square kilometre. ...
of forest. On BCI, the nests were found under 70 species of trees, six species of shrubs, two species of
liana A liana is a long-Plant stem, stemmed Woody plant, woody vine that is rooted in the soil at ground level and uses trees, as well as other means of vertical support, to climb up to the Canopy (biology), canopy in search of direct sunlight. T ...
s, and one species of palm. Nests were most common beneath the canopies of ''
Faramea ''Faramea'' is a genus of plants in the family Rubiaceae Rubiaceae () is a family (biology), family of flowering plants, commonly known as the coffee, madder, or bedstraw family. It consists of terrestrial trees, shrubs, lianas, or herbs tha ...
occidentalis'' and ''
Trichilia ''Trichilia'' is a flowering plant genus in the family (biology), family Meliaceae. These plants are particularly diverse in sub-Saharan Africa and tropical South America. Several species are used in folk medicine and shamanism – e.g. ''Tr ...
tuberculata'', but these trees are also the most abundant in the forest. Nests were present under '' Alseis blackiana'', '' Tabernaemontana arborea'', '' Virola sebifera'', '' Guarea guidonia'', and '' Oenocarpus mapora'' more frequently than would be expected from abundance of these tree species. The large number of nest plants suggests little active selection of nest sites by bullet ants. Small shrubs, however, are underused, probably because they do not provide access to the forest canopy. The study on BCI concluded that bullet ants may select trees with
buttresses A buttress is an architectural structure built against or projecting from a wall which serves to support or reinforce the wall. Buttresses are fairly common on more ancient (typically Gothic) buildings, as a means of providing support to act a ...
and
extrafloral nectaries Nectar is a viscous, sugar-rich liquid produced by plants in glands called nectaries, either within the flowers with which it attracts pollinating animals, or by extrafloral nectaries, which provide a nutrient source to animal mutualists, ...
.


Enemies

The ant is a predator of '' Greta oto'', the glasswing butterfly. This butterfly attempts to combat ''P. clavata'' by producing chemical extracts during the
larva A larva (; : larvae ) is a distinct juvenile form many animals undergo before metamorphosis into their next life stage. Animals with indirect development such as insects, some arachnids, amphibians, or cnidarians typically have a larval phase ...
l stage that are unpalatable to these ants.


Parasites

The small (1.5- to 2.0-mm-long) phorid fly '' Apocephalus paraponerae'' is a
parasite Parasitism is a Symbiosis, close relationship between species, where one organism, the parasite, lives (at least some of the time) on or inside another organism, the Host (biology), host, causing it some harm, and is Adaptation, adapted str ...
of injured workers of ''P. clavata'', of which the supply is constant because frequent aggressive encounters occur between neighbouring ant colonies, resulting in maimed workers. The flies are able to parasitise healthy ants if the ants are artificially restrained, but healthy ants are agile and able to repel them. Both male and female flies are attracted by the scent of injured ants; the females lay eggs, as well as feeding, and the males feed and possibly mate with the females. The flies are attracted to a crushed ant within two to three minutes, and 10 or more flies may be attracted to each ant. Each ant can harbour 20 fly
larvae A larva (; : larvae ) is a distinct juvenile form many animals undergo before metamorphosis into their next life stage. Animals with indirect developmental biology, development such as insects, some arachnids, amphibians, or cnidarians typical ...
. Carl Rettenmeyer observed ''P. clavata'' actively trying to attack ''A. paraponerae'' when they approached the entrance to their nest.


Relationship with humans


Sting

The bullet ant's sting currently ranks the highest of all insect stings on Justin O. Schmidt's informal sting pain index, at 4.0+. According to Schmidt, the pain is like " lking over flaming charcoal with a three-inch nail embedded in your heel". Some victims compared the pain to that of being shot, hence the name of the insect. It is described as causing "waves of burning, throbbing, all-consuming pain that continues unabated for up to 24 hours".
Lymphadenopathy Lymphadenopathy or adenopathy is a disease of the lymph nodes, in which they are abnormal in size or consistency. Lymphadenopathy of an inflammatory type (the most common type) is lymphadenitis, producing swollen or enlarged lymph nodes. In c ...
,
edema Edema (American English), also spelled oedema (British English), and also known as fluid retention, swelling, dropsy and hydropsy, is the build-up of fluid in the body's tissue (biology), tissue. Most commonly, the legs or arms are affected. S ...
,
tachycardia Tachycardia, also called tachyarrhythmia, is a heart rate that exceeds the normal resting rate. In general, a resting heart rate over 100 beats per minute is accepted as tachycardia in adults. Heart rates above the resting rate may be normal ...
, and fresh blood appearing in human victim feces are common symptoms from even a single sting. Poneratoxin, a paralyzing
neurotoxic Neurotoxicity is a form of toxicity in which a biological, chemical, or physical agent produces an adverse effect on the structure or function of the central and/or peripheral nervous system. It occurs when exposure to a substance – specifical ...
peptide Peptides are short chains of amino acids linked by peptide bonds. A polypeptide is a longer, continuous, unbranched peptide chain. Polypeptides that have a molecular mass of 10,000 Da or more are called proteins. Chains of fewer than twenty am ...
isolated from the
venom Venom or zootoxin is a type of toxin produced by an animal that is actively delivered through a wound by means of a bite, sting, or similar action. The toxin is delivered through a specially evolved ''venom apparatus'', such as fangs or a sti ...
, affects voltage-dependent sodium ion channels and blocks the
synaptic transmission Neurotransmission (Latin: ''transmissio'' "passage, crossing" from ''transmittere'' "send, let through") is the process by which signaling molecules called neurotransmitters are released by the axon terminal of a neuron (the presynaptic neuron) ...
in the
central nervous system The central nervous system (CNS) is the part of the nervous system consisting primarily of the brain, spinal cord and retina. The CNS is so named because the brain integrates the received information and coordinates and influences the activity o ...
. It is being investigated for possible medical applications.


Initiation rites

The Sateré-Mawé people of
Brazil Brazil, officially the Federative Republic of Brazil, is the largest country in South America. It is the world's List of countries and dependencies by area, fifth-largest country by area and the List of countries and dependencies by population ...
use intentional bullet ant stings as part of their initiation rites to become warriors or leaders. The ants are first rendered unconscious by submersion in a natural sedative, and then 80 of them are woven into gloves (which resemble large oven mitts) made of vines or leaves,
stinger A stinger (or sting) is a sharp organ found in various animals (typically insects and other arthropods) capable of injecting venom, usually by piercing the epidermis of another animal. An insect sting is complicated by its introduction of ve ...
s facing inward. When the ants regain consciousness, an initiator repeatedly blows smoke at the ants, with the objective of making them agitated and aggressive. Once this is done, the initiate has the gloves put on his hands and keeps them on for 5 to 10 minutes or longer. Afterward, the boy's hand and part of his arm are temporarily paralyzed because of the ant venom, and he may shake uncontrollably for days. The only "protection" provided is a coating of charcoal on the hands, supposedly to confuse the ants and inhibit their stinging. To fully complete the initiation, a boy or man must go through the ordeal 20 times over the course of several months or even years. Video of initiation rite by ''
National Geographic ''National Geographic'' (formerly ''The National Geographic Magazine'', sometimes branded as ''Nat Geo'') is an American monthly magazine published by National Geographic Partners. The magazine was founded in 1888 as a scholarly journal, nine ...
'', showing preparation of glove and its use.


See also

* '' Myrmecia'', often called bull ants or bulldog ants, also renowned for their powerful sting * '' Diacamma rugosum'', also known as the Asian bullet ant * Fire ants * List of ant genera (alphabetical)


References


Further reading

* * * Lattke, JE (2003). Subfamilia Ponerinae, ''Introducción a las Hormigas de la Région Neotropical''. Von Humboldt Institute, Bogotá, Colombia. * *


External links


Brief article about Paraponera clavata

Short article on the bullet ant and poneratoxin


article with images * ttps://www.youtube.com/watch?v=ZGIZ-zUvotM YouTube video of initiation ritual
Paraponera clavata
at AntWeb
Data for specimens
collected in
Gracias a Dios Department Gracias a Dios (; "Thanks to God" or "Thank God") is one of the 18 departments (''departamentos'') into which Honduras is divided. The departmental capital is Puerto Lempira; until 1975 it was Brus Laguna. Etymology The department is named in ...
, Honduras, the northernmost extent of its range. * {{Taxonbar, from=Q73901 Paraponerinae Hymenoptera of South America Insects of Central America Fauna of Paraguay Insects described in 1775 Taxa named by Johan Christian Fabricius Fauna of Suriname