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Ants are
eusocial Eusociality ( Greek 'good' and social) is the highest level of organization of sociality. It is defined by the following characteristics: cooperative brood care (including care of offspring from other individuals), overlapping generations wit ...
insect Insects (from Latin ') are Hexapoda, hexapod invertebrates of the class (biology), class Insecta. They are the largest group within the arthropod phylum. Insects have a chitinous exoskeleton, a three-part body (Insect morphology#Head, head, ...
s of the
family Family (from ) is a Social group, group of people related either by consanguinity (by recognized birth) or Affinity (law), affinity (by marriage or other relationship). It forms the basis for social order. Ideally, families offer predictabili ...
Formicidae and, along with the related
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 and
bee Bees are winged insects closely related to wasps and ants, known for their roles in pollination and, in the case of the best-known bee species, the western honey bee, for producing honey. Bees are a monophyletic lineage within the superfamil ...
s, belong to the
order Order, ORDER or Orders may refer to: * A socio-political or established or existing order, e.g. World order, Ancien Regime, Pax Britannica * Categorization, the process in which ideas and objects are recognized, differentiated, and understood ...
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 ...
. Ants evolved from vespoid wasp ancestors in the
Cretaceous The Cretaceous ( ) is a geological period that lasted from about 143.1 to 66 mya (unit), million years ago (Mya). It is the third and final period of the Mesozoic Era (geology), Era, as well as the longest. At around 77.1 million years, it is the ...
period. More than 13,800 of an estimated total of 22,000
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), ...
have been classified. They are easily identified by their geniculate (elbowed) antennae and the distinctive node-like structure that forms their slender waists. Ants form
colonies A colony is a territory subject to a form of foreign rule, which rules the territory and its indigenous peoples separated from the foreign rulers, the colonizer, and their '' metropole'' (or "mother country"). This separated rule was often or ...
that range in size from a few dozen individuals often living in small
natural Nature is an inherent character or constitution, particularly of the ecosphere or the universe as a whole. In this general sense nature refers to the laws, elements and phenomena of the physical world, including life. Although humans are part ...
cavities to highly organised colonies that may occupy large territories with sizeable nest that consist of millions of individuals or into the hundreds of millions in super colonies. Typical colonies consist of various castes of sterile, wingless females, most of which are workers (ergates), as well as soldiers (dinergates) and other specialised groups. Nearly all ant colonies also have some fertile males called "drones" and one or more fertile females called "
queens Queens is the largest by area of the Boroughs of New York City, five boroughs of New York City, coextensive with Queens County, in the U.S. state of New York (state), New York. Located near the western end of Long Island, it is bordered by the ...
" (
gyne The gyne (, from Greek γυνή, "woman") is the primary reproductive female caste of social insects (especially ants, wasps, and bees of order Hymenoptera, as well as termites). Gynes are those destined to become queens, whereas female workers ...
s). The colonies are described as
superorganism A superorganism, or supraorganism, is a group of synergetically interacting organisms of the same species. A community of synergetically interacting organisms of different species is called a '' holobiont''. Concept The term superorganism is ...
s because the ants appear to operate as a unified entity, collectively working together to support the colony. Ants have colonised almost every landmass on
Earth Earth is the third planet from the Sun and the only astronomical object known to Planetary habitability, harbor life. This is enabled by Earth being an ocean world, the only one in the Solar System sustaining liquid surface water. Almost all ...
. The only places lacking
indigenous Indigenous may refer to: *Indigenous peoples *Indigenous (ecology) In biogeography, a native species is indigenous to a given region or ecosystem if its presence in that region is the result of only local natural evolution (though often populari ...
ants are
Antarctica Antarctica () is Earth's southernmost and least-populated continent. Situated almost entirely south of the Antarctic Circle and surrounded by the Southern Ocean (also known as the Antarctic Ocean), it contains the geographic South Pole. ...
and a few remote or inhospitable islands. Ants thrive in moist tropical ecosystems and may exceed the combined
biomass Biomass is a term used in several contexts: in the context of ecology it means living organisms, and in the context of bioenergy it means matter from recently living (but now dead) organisms. In the latter context, there are variations in how ...
of wild birds and mammals. Their success in so many environments has been attributed to their social organisation and their ability to modify habitats, tap resources, and defend themselves. Their long
co-evolution In biology, coevolution occurs when two or more species reciprocally affect each other's evolution through the process of natural selection. The term sometimes is used for two traits in the same species affecting each other's evolution, as well a ...
with other species has led to
mimetic Mimesis (; , ''mīmēsis'') is a term used in literary criticism and philosophy that carries a wide range of meanings, including ''imitatio'', imitation, Similarity (philosophy), similarity, receptivity, representation (arts), representation, m ...
,
commensal Commensalism is a long-term biological interaction (symbiosis) in which members of one species gain benefits while those of the other species neither benefit nor are harmed. This is in contrast with mutualism, in which both organisms benefit f ...
,
parasitic Parasitism is a close relationship between species, where one organism, the parasite, lives (at least some of the time) on or inside another organism, the host, causing it some harm, and is adapted structurally to this way of life. The ent ...
, and mutualistic relationships. Ant societies have
division of labour The division of labour is the separation of the tasks in any economic system or organisation so that participants may specialise ( specialisation). Individuals, organisations, and nations are endowed with or acquire specialised capabilities, a ...
, communication between individuals, and an ability to solve complex problems. These parallels with
human societies A society () is a group of individuals involved in persistent social interaction or a large social group sharing the same spatial or social territory, typically subject to the same political authority and dominant cultural expectations. Soc ...
have long been an inspiration and subject of study. Many human cultures make use of ants in cuisine, medication, and rites. Some species are valued in their role as
biological pest control Biological control or biocontrol is a method of controlling pests, whether pest animals such as insects and mites, weeds, or pathogens affecting animals or plants by using other organisms. It relies on predation, parasitism, herbivory, or ot ...
agents. Their ability to exploit resources may bring ants into conflict with humans, however, as they can damage crops and invade buildings. Some species, such as the
red imported fire ant ''Solenopsis invicta'', the fire ant, or red imported fire ant (RIFA), is a species of ant native to South America. A member of the genus ''Fire ant, Solenopsis'' in the subfamily Myrmicinae, it was Species description, described by Swiss ento ...
(''Solenopsis invicta'') of South America, are regarded as
invasive species An invasive species is an introduced species that harms its new environment. Invasive species adversely affect habitats and bioregions, causing ecological, environmental, and/or economic damage. The term can also be used for native spec ...
in other parts of the world, establishing themselves in areas where they have been introduced accidentally.


Etymology

The word ''ant'' and the archaic word ''emmet'' are derived from ', ' of
Middle English Middle English (abbreviated to ME) is a form of the English language that was spoken after the Norman Conquest of 1066, until the late 15th century. The English language underwent distinct variations and developments following the Old English pe ...
, which come from ' of
Old English Old English ( or , or ), or Anglo-Saxon, is the earliest recorded form of the English language, spoken in England and southern and eastern Scotland in the Early Middle Ages. It developed from the languages brought to Great Britain by Anglo-S ...
; these are all related to
Low Saxon Low Saxon (), also known as West Low German () are a group of Low German dialects spoken in parts of the Netherlands, northwestern Germany and southern Denmark (in North Schleswig by parts of the German-speaking minority). It is one of two di ...
', ' and varieties (
Old Saxon Old Saxon (), also known as Old Low German (), was a Germanic language and the earliest recorded form of Low German (spoken nowadays in Northern Germany, the northeastern Netherlands, southern Denmark, the Americas and parts of Eastern Eur ...
') and to
German German(s) may refer to: * Germany, the country of the Germans and German things **Germania (Roman era) * Germans, citizens of Germany, people of German ancestry, or native speakers of the German language ** For citizenship in Germany, see also Ge ...
' (
Old High German Old High German (OHG; ) is the earliest stage of the German language, conventionally identified as the period from around 500/750 to 1050. Rather than representing a single supra-regional form of German, Old High German encompasses the numerous ...
'). All of these words come from West Germanic ''*'', and the original meaning of the word was "the biter" (from
Proto-Germanic Proto-Germanic (abbreviated PGmc; also called Common Germanic) is the linguistic reconstruction, reconstructed proto-language of the Germanic languages, Germanic branch of the Indo-European languages. Proto-Germanic eventually developed from ...
', "off, away" + ' "cut"). The family name ''Formicidae'' is derived from the
Latin Latin ( or ) is a classical language belonging to the Italic languages, Italic branch of the Indo-European languages. Latin was originally spoken by the Latins (Italic tribe), Latins in Latium (now known as Lazio), the lower Tiber area aroun ...
' ("ant") from which the words in other
Romance languages The Romance languages, also known as the Latin or Neo-Latin languages, are the languages that are Language family, directly descended from Vulgar Latin. They are the only extant subgroup of the Italic languages, Italic branch of the Indo-E ...
, such as the Portuguese ', Italian ', Spanish ', Romanian ', and French ' are derived. The study of ants is called ''
myrmecology Myrmecology (; from Greek: μύρμηξ, ''myrmex'', "ant" and λόγος, ''logos'', "study") is a branch of entomology focusing on the study of ants. Ants continue to be a model of choice for the study of questions on the evolution of social ...
'', from
Ancient Greek Ancient Greek (, ; ) includes the forms of the Greek language used in ancient Greece and the classical antiquity, ancient world from around 1500 BC to 300 BC. It is often roughly divided into the following periods: Mycenaean Greek (), Greek ...
μύρμηξ ''mýrmēx'' ("ant"). It has been hypothesised that a
Proto-Indo-European Proto-Indo-European (PIE) is the reconstructed common ancestor of the Indo-European language family. No direct record of Proto-Indo-European exists; its proposed features have been derived by linguistic reconstruction from documented Indo-Euro ...
word *morwi- was the root for
Sanskrit Sanskrit (; stem form ; nominal singular , ,) is a classical language belonging to the Indo-Aryan languages, Indo-Aryan branch of the Indo-European languages. It arose in northwest South Asia after its predecessor languages had Trans-cultural ...
''vamrah'', Greek μύρμηξ ''mýrmēx'', Latin ',
Old Church Slavonic Old Church Slavonic or Old Slavonic ( ) is the first Slavic languages, Slavic literary language and the oldest extant written Slavonic language attested in literary sources. It belongs to the South Slavic languages, South Slavic subgroup of the ...
''mraviji'',
Old Irish Old Irish, also called Old Gaelic (, Ogham, Ogham script: ᚌᚑᚔᚇᚓᚂᚉ; ; ; or ), is the oldest form of the Goidelic languages, Goidelic/Gaelic language for which there are extensive written texts. It was used from 600 to 900. The ...
''moirb'',
Old Norse Old Norse, also referred to as Old Nordic or Old Scandinavian, was a stage of development of North Germanic languages, North Germanic dialects before their final divergence into separate Nordic languages. Old Norse was spoken by inhabitants ...
''maurr'', Dutch ''mier'', Swedish ''myra'', Danish ''myre'',
Middle Dutch Middle Dutch is a collective name for a number of closely related West Germanic dialects whose ancestor was Old Dutch. It was spoken and written between 1150 and 1500. Until the advent of Modern Dutch after 1500 or , there was no overarching sta ...
''miere'', and
Crimean Gothic Crimean Gothic was a Germanic languages, Germanic, probably East Germanic languages, East Germanic, language spoken by the Crimean Goths in some isolated locations in Crimea until the late 18th century. Crimea was inhabited by the Goths in Late ...
'' miera''.


Taxonomy and evolution

The family Formicidae belongs 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 ...
, which also includes
sawflies Sawflies are wasp-like insects that are in the suborder Symphyta within the order Hymenoptera, alongside ants, bees, and wasps. The common name comes from the saw-like appearance of the ovipositor, which the females use to cut into the plant ...
,
bee Bees are winged insects closely related to wasps and ants, known for their roles in pollination and, in the case of the best-known bee species, the western honey bee, for producing honey. Bees are a monophyletic lineage within the superfamil ...
s, and
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. Ants evolved from a lineage within the stinging wasps, and a 2013 study suggests that they are a sister group of the
Apoidea The superfamily Apoidea is a major group (of over 30 000 species) within the Hymenoptera, which includes two traditionally recognized lineages, the "sphecoid" wasps, and the bees. Molecular phylogeny demonstrates that the bees arose from ...
. However, since Apoidea is a superfamily, ants must be upgraded to the same rank. A more detailed basic taxonomy was proposed in 2020. Three species of the extinct mid-
Cretaceous The Cretaceous ( ) is a geological period that lasted from about 143.1 to 66 mya (unit), million years ago (Mya). It is the third and final period of the Mesozoic Era (geology), Era, as well as the longest. At around 77.1 million years, it is the ...
genera '' Camelomecia'' and '' Camelosphecia'' were placed outside of the Formicidae, in a separate
clade In biology, a clade (), also known as a Monophyly, monophyletic group or natural group, is a group of organisms that is composed of a common ancestor and all of its descendants. Clades are the fundamental unit of cladistics, a modern approach t ...
within the general superfamily
Formicoidea Ants are eusocial insects of the family Formicidae and, along with the related wasps and bees, belong to the order Hymenoptera. Ants evolved from vespoid wasp ancestors in the Cretaceous period. More than 13,800 of an estimated total of 22 ...
, which, together with Apoidea, forms the higher-ranking group Formicapoidina. Fernández et al. (2021) suggest that the common ancestors of ants and apoids within the Formicapoidina probably existed as early as in the end of the
Jurassic The Jurassic ( ) is a Geological period, geologic period and System (stratigraphy), stratigraphic system that spanned from the end of the Triassic Period million years ago (Mya) to the beginning of the Cretaceous Period, approximately 143.1 Mya. ...
period, before divergence in the Cretaceous. In 1966,
E. O. Wilson Edward Osborne Wilson (June 10, 1929 – December 26, 2021) was an American biologist, naturalist, ecologist, and entomologist known for developing the field of sociobiology. Born in Alabama, Wilson found an early interest in nature and frequ ...
and his colleagues identified the
fossil A fossil (from Classical Latin , ) is any preserved remains, impression, or trace of any once-living thing from a past geological age. Examples include bones, shells, exoskeletons, stone imprints of animals or microbes, objects preserve ...
remains of an ant (''
Sphecomyrma ''Sphecomyrma'' is an Extinction, extinct genus of ants which existed in the Cretaceous approximately 79 to 92 million years ago. The first specimens were collected in 1966, found embedded in amber which had been exposed in the cliffs of Cliffwo ...
'') that lived in the Cretaceous period. The specimen, trapped in amber
dating Dating is a stage of Romance (love), romantic relationships in which individuals engage in activity together, often with the intention of evaluating each other's suitability as a partner in a future intimate relationship. It falls into the cate ...
back to around 92 million years ago, has features found in some wasps, but not found in modern ants. The oldest fossils of ants date to the mid-Cretaceous, around 113-100 million years ago, which belong to extinct
stem-group In phylogenetics, the crown group or crown assemblage is a collection of species composed of the living representatives of the collection, the most recent common ancestor of the collection, and all descendants of the most recent common ancestor. ...
s such as the
Haidomyrmecinae Haidomyrmecinae, occasionally called hell ants, are an extinct subfamily of ants (Formicidae) known from Cretaceous fossils primarily found in amber from North America, South America, Europe, and Asia, spanning the late Aptian to Campanian, aroun ...
,
Sphecomyrminae Sphecomyrminae is an extinct subfamily of ants in family Formicidae known from a series of Cretaceous fossils found in North America, Europe, and Asia. Sphecomyrminae contains eight genera, divided into two tribes Sphecomyrmini and Zigrasimeciin ...
and
Zigrasimeciinae Zigrasimeciinae is a subfamily of ants, known from the Cretaceous period, originally named as the tribe Zigrasimeciini within the subfamily Sphecomyrminae by Borysenko, 2017, it was elevated to full subfamily in 2020. It contains three describe ...
, with modern ant subfamilies appearing towards the end of the Cretaceous around 80–70 million years ago. Ants diversified extensively during the Angiosperm Terrestrial Revolution and assumed ecological dominance around 60 million years ago. Some groups, such as the
Leptanillinae Leptanillinae is a subfamily of primitive 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 ...
and
Martialinae ''Martialis heureka'' is a species of ant discovered in 2000 from the Amazon rainforest near Manaus, Brazil. It was described as a new species and placed as the sole member of a new subfamily, Martialinae. The generic name means "from Mars" ...
, are suggested to have diversified from early primitive ants that were likely to have been predators underneath the surface of the soil. During the Cretaceous period, a few species of primitive ants ranged widely on the
Laurasia Laurasia () was the more northern of two large landmasses that formed part of the Pangaea supercontinent from around ( Mya), the other being Gondwana. It separated from Gondwana (beginning in the late Triassic period) during the breakup of Pa ...
n supercontinent (the
Northern Hemisphere The Northern Hemisphere is the half of Earth that is north of the equator. For other planets in the Solar System, north is defined by humans as being in the same celestial sphere, celestial hemisphere relative to the invariable plane of the Solar ...
). Their representation in the fossil record is poor, in comparison to the populations of other insects, representing only about 1% of fossil evidence of insects in the era. Ants became dominant after
adaptive radiation In evolutionary biology, adaptive radiation is a process in which organisms diversify rapidly from an ancestral species into a multitude of new forms, particularly when a change in the environment makes new resources available, alters biotic int ...
at the beginning of the
Paleogene period The Paleogene Period ( ; also spelled Palaeogene or Palæogene) is a geologic period and system that spans 43 million years from the end of the Cretaceous Period Ma (million years ago) to the beginning of the Neogene Period Ma. It is the fir ...
. By the
Oligocene The Oligocene ( ) is a geologic epoch (geology), epoch of the Paleogene Geologic time scale, Period that extends from about 33.9 million to 23 million years before the present ( to ). As with other older geologic periods, the rock beds that defin ...
and
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 ...
, ants had come to represent 20–40% of all insects found in major fossil deposits. Of the species that lived in the
Eocene The Eocene ( ) is a geological epoch (geology), epoch that lasted from about 56 to 33.9 million years ago (Ma). It is the second epoch of the Paleogene Period (geology), Period in the modern Cenozoic Era (geology), Era. The name ''Eocene'' comes ...
epoch, around one in 10 genera survive to the present. Genera surviving today comprise 56% of the genera in
Baltic amber Baltic amber or succinite is amber from the Baltic region, home of its largest known deposits. It was produced sometime during the Eocene epoch, but exactly when is controversial. It has been estimated that this forested region provided the re ...
fossils (early Oligocene), and 92% of the genera 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 ...
fossils (apparently early Miocene).Hölldobler & Wilson (1990), pp. 23–24
Termite Termites are a group of detritivore, detritophagous Eusociality, eusocial cockroaches which consume a variety of Detritus, decaying plant material, generally in the form of wood, Plant litter, leaf litter, and Humus, soil humus. They are dist ...
s live in colonies and are sometimes called "white ants", but termites are only distantly related to ants. They are the sub-order
Isoptera Termites are a group of detritophagous eusocial cockroaches which consume a variety of decaying plant material, generally in the form of wood, leaf litter, and soil humus. They are distinguished by their moniliform antennae and the soft-bo ...
, and together with
cockroach Cockroaches (or roaches) are insects belonging to the Order (biology), order Blattodea (Blattaria). About 30 cockroach species out of 4,600 are associated with human habitats. Some species are well-known Pest (organism), pests. Modern cockro ...
es, they form the order
Blattodea Blattodea is an order (biology), order of insects that contains cockroaches and termites. Formerly, termites were considered a separate order, Isoptera, but genetics, genetic and molecular evidence suggests they evolved from within the cockroach ...
. Blattodeans are related to
mantid Mantidae is one of the largest families in the order of praying mantises, based on the type species ''Mantis religiosa''; most genera are tropical or subtropical. Historically, this was the only family in the order, and many references still ...
s,
crickets Crickets are orthopteran insects which are related to bush crickets and more distantly, to grasshoppers. In older literature, such as Imms,Imms AD, rev. Richards OW & Davies RG (1970) ''A General Textbook of Entomology'' 9th Ed. Methuen 886 ...
, and other winged insects that do not undergo
complete metamorphosis Holometabolism, also called complete metamorphosis, is a form of insect development which includes four life stages: egg, larva, pupa, and imago (or adult). Holometabolism is a synapomorphic trait of all insects in the clade Holometabola. Immature ...
. Like ants, termites are
eusocial Eusociality ( Greek 'good' and social) is the highest level of organization of sociality. It is defined by the following characteristics: cooperative brood care (including care of offspring from other individuals), overlapping generations wit ...
, with sterile workers, but they differ greatly in the genetics of reproduction. The similarity of their social structure to that of ants is attributed to
convergent evolution Convergent evolution is the independent evolution of similar features in species of different periods or epochs in time. Convergent evolution creates analogous structures that have similar form or function but were not present in the last comm ...
. Velvet ants look like large ants, but are wingless female
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.


Distribution and diversity

Ants have a
cosmopolitan distribution In biogeography, a cosmopolitan distribution is the range of a taxon that extends across most or all of the surface of the Earth, in appropriate habitats; most cosmopolitan species are known to be highly adaptable to a range of climatic and en ...
. They are found on all continents except
Antarctica Antarctica () is Earth's southernmost and least-populated continent. Situated almost entirely south of the Antarctic Circle and surrounded by the Southern Ocean (also known as the Antarctic Ocean), it contains the geographic South Pole. ...
, and only a few large islands, such as
Greenland Greenland is an autonomous territory in the Danish Realm, Kingdom of Denmark. It is by far the largest geographically of three constituent parts of the kingdom; the other two are metropolitan Denmark and the Faroe Islands. Citizens of Greenlan ...
,
Iceland Iceland is a Nordic countries, Nordic island country between the Atlantic Ocean, North Atlantic and Arctic Oceans, on the Mid-Atlantic Ridge between North America and Europe. It is culturally and politically linked with Europe and is the regi ...
, parts of
Polynesia Polynesia ( , ) is a subregion of Oceania, made up of more than 1,000 islands scattered over the central and southern Pacific Ocean. The indigenous people who inhabit the islands of Polynesia are called Polynesians. They have many things in ...
and the
Hawaiian Islands The Hawaiian Islands () are an archipelago of eight major volcanic islands, several atolls, and numerous smaller islets in the Pacific Ocean, North Pacific Ocean, extending some from the Hawaii (island), island of Hawaii in the south to nort ...
lack native ant species. Ants occupy a wide range of
ecological niche In ecology, a niche is the match of a species to a specific environmental condition. Three variants of ecological niche are described by It describes how an organism or population responds to the distribution of Resource (biology), resources an ...
s and exploit many different food resources as direct or indirect herbivores, predators and scavengers. Most ant species are omnivorous generalists, but a few are specialist feeders. There is considerable variation in ant abundance across habitats, peaking in the moist tropics to nearly six times that found in less suitable habitats. Their ecological dominance has been examined primarily using estimates of their
biomass Biomass is a term used in several contexts: in the context of ecology it means living organisms, and in the context of bioenergy it means matter from recently living (but now dead) organisms. In the latter context, there are variations in how ...
: myrmecologist
E. O. Wilson Edward Osborne Wilson (June 10, 1929 – December 26, 2021) was an American biologist, naturalist, ecologist, and entomologist known for developing the field of sociobiology. Born in Alabama, Wilson found an early interest in nature and frequ ...
had estimated in 2009 that at any one time the total number of ants was between one and ten
quadrillion Depending on context (e.g. language, culture, region), some large numbers have names that allow for describing large quantities in a textual form; not mathematical. For very large values, the text is generally shorter than a decimal numeric repres ...
(
short scale The long and short scales are two powers of ten number naming systems that are consistent with each other for smaller numbers, but are contradictory for larger numbers. Other numbering systems, particularly in East Asia and South Asia, hav ...
) (i.e., between 1015 and 1016) and using this estimate he had suggested that the total
biomass Biomass is a term used in several contexts: in the context of ecology it means living organisms, and in the context of bioenergy it means matter from recently living (but now dead) organisms. In the latter context, there are variations in how ...
of all the ants in the world was approximately equal to the total biomass of the entire
human Humans (''Homo sapiens'') or modern humans are the most common and widespread species of primate, and the last surviving species of the genus ''Homo''. They are Hominidae, great apes characterized by their Prehistory of nakedness and clothing ...
race. More careful estimates made in 2022 which take into account regional variations puts the global ant contribution at 12 megatons of dry carbon, which is about 20% of the total human contribution, but greater than that of the wild birds and mammals combined. This study also puts a conservative estimate of the ants at about 20 × 1015 (20 quadrillion). Ants range in size from ,Hölldobler & Wilson (1990), p. 589 the largest species being the fossil '' Titanomyrma giganteum'', the queen of which was long with a wingspan of . Ants vary in colour; most ants are yellow to red or brown to black, but a few species are green and some tropical species have a metallic lustre. More than 13,800 species are currently knownAntWeb
/ref> (with upper estimates of the potential existence of about 22,000; see the article List of ant genera), with the greatest diversity in the tropics. Taxonomic studies continue to resolve the classification and systematics of ants. Online databases of ant species, including AntWeb and the Hymenoptera Name Server, help to keep track of the known and newly described species. The relative ease with which ants may be sampled and studied in
ecosystem An ecosystem (or ecological system) is a system formed by Organism, organisms in interaction with their Biophysical environment, environment. The Biotic material, biotic and abiotic components are linked together through nutrient cycles and en ...
s has made them useful as
indicator species A bioindicator is any species (an indicator species) or group of species whose function, population, or status can reveal the qualitative status of the environment. The most common indicator species are animals. For example, copepods and other sma ...
in
biodiversity Biodiversity is the variability of life, life on Earth. It can be measured on various levels. There is for example genetic variability, species diversity, ecosystem diversity and Phylogenetics, phylogenetic diversity. Diversity is not distribut ...
studies.


Morphology

Ants are distinct in their
morphology Morphology, from the Greek and meaning "study of shape", may refer to: Disciplines *Morphology (archaeology), study of the shapes or forms of artifacts *Morphology (astronomy), study of the shape of astronomical objects such as nebulae, galaxies, ...
from other insects in having geniculate (elbowed) antennae,
metapleural gland Metapleural glands (also called metasternal or metathoracic glands) are secretory glands that were considered unique to ants and basal in the evolutionary history of ants. They are responsible for the production of an antibiotic fluid that then col ...
s, and a strong constriction of their second
abdominal The abdomen (colloquially called the gut, belly, tummy, midriff, tucky, or stomach) is the front part of the torso between the thorax (chest) and pelvis in humans and in other vertebrates. The area occupied by the abdomen is called the abdominal ...
segment into a node-like petiole. The body is divided into three distinct sections (formally known as '' tagmata''): the head,
mesosoma The mesosoma is the middle part of the body, or tagma, of arthropods whose body is composed of three parts, the other two being the prosoma and the metasoma. It bears the legs, and, in the case of winged insects, the wings. Wasps, bees and a ...
, and
metasoma The metasoma is the posterior part of the body, or tagma (biology), tagma, of arthropods whose body is composed of three parts, the other two being the prosoma and the mesosoma. In insects, it contains most of the digestive tract, respiratory sy ...
. The petiole forms a narrow waist between their mesosoma (
thorax The thorax (: thoraces or thoraxes) or chest is a part of the anatomy of mammals and other tetrapod animals located between the neck and the abdomen. In insects, crustaceans, and the extinct trilobites, the thorax is one of the three main di ...
plus the first abdominal segment, which is fused to it) and gaster (abdomen less the abdominal segments in the petiole). The petiole may be formed by one or two nodes (the second alone, or the second and third abdominal segments). Tergosternal fusion, when the tergite and sternite of a segment fuse together, can occur partly or fully on the second, third and fourth abdominal segment and is used in identification. Fourth abdominal tergosternal fusion was formerly used as character that defined the poneromorph subfamilies, Ponerinae and relatives within their clade, but this is no longer considered a
synapomorphic In phylogenetics, an apomorphy (or derived trait) is a novel character or character state that has evolved from its ancestral form (or plesiomorphy). A synapomorphy is an apomorphy shared by two or more taxa and is therefore hypothesized to hav ...
character. Like other arthropods, ants have an
exoskeleton An exoskeleton () . is a skeleton that is on the exterior of an animal in the form of hardened integument, which both supports the body's shape and protects the internal organs, in contrast to an internal endoskeleton (e.g. human skeleton, that ...
, an external covering that provides a protective casing around the body and a point of attachment for muscles, in contrast to the internal skeletons of humans and other
vertebrate Vertebrates () are animals with a vertebral column (backbone or spine), and a cranium, or skull. The vertebral column surrounds and protects the spinal cord, while the cranium protects the brain. The vertebrates make up the subphylum Vertebra ...
s. Insects do not have
lung The lungs are the primary Organ (biology), organs of the respiratory system in many animals, including humans. In mammals and most other tetrapods, two lungs are located near the Vertebral column, backbone on either side of the heart. Their ...
s;
oxygen Oxygen is a chemical element; it has chemical symbol, symbol O and atomic number 8. It is a member of the chalcogen group (periodic table), group in the periodic table, a highly reactivity (chemistry), reactive nonmetal (chemistry), non ...
and other gases, such as
carbon dioxide Carbon dioxide is a chemical compound with the chemical formula . It is made up of molecules that each have one carbon atom covalent bond, covalently double bonded to two oxygen atoms. It is found in a gas state at room temperature and at norma ...
, pass through their exoskeleton via tiny valves called spiracles. Insects also lack closed
blood vessel Blood vessels are the tubular structures of a circulatory system that transport blood throughout many Animal, animals’ bodies. Blood vessels transport blood cells, nutrients, and oxygen to most of the Tissue (biology), tissues of a Body (bi ...
s; instead, they have a long, thin, perforated tube along the top of the body (called the "dorsal aorta") that functions like a heart, and pumps
haemolymph Hemolymph, or haemolymph, is a fluid, similar to the blood in invertebrates, that circulates in the inside of the arthropod's body, remaining in direct contact with the animal's tissues. It is composed of a fluid plasma in which hemolymph ce ...
toward the head, thus driving the circulation of the internal fluids. The
nervous system In biology, the nervous system is the complex system, highly complex part of an animal that coordinates its behavior, actions and sense, sensory information by transmitting action potential, signals to and from different parts of its body. Th ...
consists of a
ventral nerve cord The ventral nerve cord is a major structure of the invertebrate central nervous system. It is the functional equivalent of the vertebrate spinal cord. The ventral nerve cord coordinates neural signaling from the brain to the body and vice ve ...
that runs the length of the body, with several
ganglia A ganglion (: ganglia) is a group of neuron cell bodies in the peripheral nervous system. In the somatic nervous system, this includes dorsal root ganglia and trigeminal ganglia among a few others. In the autonomic nervous system, there a ...
and branches along the way reaching into the extremities of the appendages.Borror, Triplehorn & Delong (1989), pp. 24–71


Head

An ant's head contains many
sensory organs A sense is a biological system used by an organism for sensation, the process of gathering information about the surroundings through the detection of stimuli. Although, in some cultures, five human senses were traditionally identified as s ...
. Like most insects, ants have
compound eye A compound eye is a Eye, visual organ found in arthropods such as insects and crustaceans. It may consist of thousands of ommatidium, ommatidia, which are tiny independent photoreception units that consist of a cornea, lens (anatomy), lens, and p ...
s made from numerous tiny lenses attached together. Ant eyes are good for acute movement detection, but do not offer a high resolution image. They also have three small
ocelli A simple eye or ocellus (sometimes called a pigment pit) is a form of eye or an optical arrangement which has a single lens without the sort of elaborate retina that occurs in most vertebrates. These eyes are called "simple" to distinguish the ...
(simple eyes) on the top of the head that detect light levels and
polarization Polarization or polarisation may refer to: Mathematics *Polarization of an Abelian variety, in the mathematics of complex manifolds *Polarization of an algebraic form, a technique for expressing a homogeneous polynomial in a simpler fashion by ...
. Compared to
vertebrate Vertebrates () are animals with a vertebral column (backbone or spine), and a cranium, or skull. The vertebral column surrounds and protects the spinal cord, while the cranium protects the brain. The vertebrates make up the subphylum Vertebra ...
s, ants tend to have blurrier eyesight, particularly in smaller species, and a few
subterranean Subterranean(s) or The Subterranean(s) may refer to: * Subterranea (geography), underground structures, both natural and man-made Literature * ''Subterranean'' (novel), a 1998 novel by James Rollins * ''Subterranean Magazine'', an American fa ...
taxa are completely blind. However, some ants, such as Australia's
bulldog ant ''Myrmecia'' is a genus of ants first established by Danish zoologist Johan Christian Fabricius in 1804. The genus is a member of the subfamily Myrmeciinae of the family Formicidae. ''Myrmecia'' is a large genus of ants, comprising at least ...
, have excellent vision and are capable of discriminating the distance and size of objects moving nearly a
meter The metre (or meter in US spelling; symbol: m) is the base unit of length in the International System of Units (SI). Since 2019, the metre has been defined as the length of the path travelled by light in vacuum during a time interval of of ...
away. Based on experiments conducted to test their ability to differentiate between selected wavelengths of light, some ant species such as ''Camponotus blandus, Solenopsis invicta,'' and ''Formica cunicularia'' are thought to possess a degree of colour vision. Two antennae ("feelers") are attached to the head; these organs detect chemicals,
air current In meteorology, air currents are concentrated areas of winds. They are mainly due to differences in atmospheric pressure or temperature. They are divided into horizontal and vertical currents; both are present at Mesoscale meteorology, mesoscale w ...
s, and
vibration Vibration () is a mechanical phenomenon whereby oscillations occur about an equilibrium point. Vibration may be deterministic if the oscillations can be characterised precisely (e.g. the periodic motion of a pendulum), or random if the os ...
s; they also are used to transmit and receive signals through touch. The head has two strong jaws, the
mandibles In jawed vertebrates, the mandible (from the Latin ''mandibula'', 'for chewing'), lower jaw, or jawbone is a bone that makes up the lowerand typically more mobilecomponent of the mouth (the upper jaw being known as the maxilla). The jawbone i ...
, used to carry food, manipulate objects, construct nests, and for defence. In some species, a small pocket (infrabuccal chamber) inside the mouth stores food, so it may be passed to other ants or their larvae.


Mesosoma

Both the
legs A leg is a weight-bearing and locomotive anatomical structure, usually having a columnar shape. During locomotion, legs function as "extensible struts". The combination of movements at all joints can be modeled as a single, linear element cap ...
and wings of the ant are attached to the
mesosoma The mesosoma is the middle part of the body, or tagma, of arthropods whose body is composed of three parts, the other two being the prosoma and the metasoma. It bears the legs, and, in the case of winged insects, the wings. Wasps, bees and a ...
("thorax"). The legs terminate in a hooked
claw A claw is a curved, pointed appendage found at the end of a toe or finger in most amniotes (mammals, reptiles, birds). Some invertebrates such as beetles and spiders have somewhat similar fine, hooked structures at the end of the leg or Arthro ...
which allows them to hook on and climb surfaces. Only reproductive ants (
queens Queens is the largest by area of the Boroughs of New York City, five boroughs of New York City, coextensive with Queens County, in the U.S. state of New York (state), New York. Located near the western end of Long Island, it is bordered by the ...
and males) have wings. Queens shed their wings after the
nuptial flight Nuptial flight is an important phase in the reproduction of most ant, termite, and some bee species. It is also observed in some fly species, such as '' Rhamphomyia longicauda''. During the flight, virgin queens mate with males and then land ...
, leaving visible stubs, a distinguishing feature of queens. In a few species, wingless queens (
ergatoid An ergatoid (from Greek '' ergat-'', "worker" + '' -oid'', "like") is a permanently wingless reproductive adult ant or termite. The similar but somewhat ambiguous term ergatogyne refers to any intermediate form between workers and standard gyne ...
s) and males occur.


Metasoma

The
metasoma The metasoma is the posterior part of the body, or tagma (biology), tagma, of arthropods whose body is composed of three parts, the other two being the prosoma and the mesosoma. In insects, it contains most of the digestive tract, respiratory sy ...
(the "abdomen") of the ant houses important internal organs, including those of the reproductive, respiratory (tracheae), and excretory systems. Workers of many species have their egg-laying structures modified into stings that are used for subduing
prey 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 feeding behaviours that includes parasitism and micropredation (which usually do not ki ...
and defending their nests.


Polymorphism

In the colonies of a few ant species, there are physical castes—workers in distinct size-classes, called minor (micrergates), median, and major ergates (macrergates). Often, the larger ants have disproportionately larger heads, and correspondingly stronger
mandibles In jawed vertebrates, the mandible (from the Latin ''mandibula'', 'for chewing'), lower jaw, or jawbone is a bone that makes up the lowerand typically more mobilecomponent of the mouth (the upper jaw being known as the maxilla). The jawbone i ...
. Although formally known as dinergates, such individuals are sometimes called "soldier" ants because their stronger mandibles make them more effective in fighting, although they still are workers and their "duties" typically do not vary greatly from the minor or median workers. In a few species, the median workers are absent, creating a sharp divide between the minors and majors.
Weaver ant Weaver ants or green ants are eusocial insects of the Hymenoptera family Formicidae belonging to the tribe Oecophyllini. Weaver ants live in trees (they are obligately arboreal) and are known for their unique nest building behaviour where wor ...
s, for example, have a distinct
bimodal In statistics, a multimodal distribution is a probability distribution with more than one mode (i.e., more than one local peak of the distribution). These appear as distinct peaks (local maxima) in the probability density function, as shown ...
size distribution. Some other species show continuous variation in the size of workers. The smallest and largest workers in ''
Carebara diversa ''Carebara'' is a genus of ants in the subfamily Myrmicinae. It is one of the largest myrmicine genera with more than 200 species distributed worldwide in the tropics and the Afrotropical region. Many of them are very tiny cryptic soil and leaf- ...
'' show nearly a 500-fold difference in their dry weights. Workers cannot mate; however, because of the
haplodiploid sex-determination system Haplodiploidy is a sex-determination system in which males develop from unfertilized eggs and are haploid, and females develop from fertilized eggs and are diploid. Haplodiploidy is sometimes called arrhenotoky. Haplodiploidy determines the se ...
in ants, workers of a number of species can lay unfertilised eggs that become fully fertile, haploid males. The role of workers may change with their age and in some species, such as
honeypot ants Honeypot ants, also called honey ants, are ants which have specialized workers—repletes, plerergates or rotunds—that consume large amounts of food to the point that their abdomens swell enormously. This phenomenon of extreme inflation of the ...
, young workers are fed until their gasters are distended, and act as living food storage vessels. These food storage workers are called ''repletes''. For instance, these replete workers develop in the North American honeypot ant ''
Myrmecocystus mexicanus ''Myrmecocystus mexicanus'' is a species of ant in the genus '' Myrmecocystus'', which is one of the six genera that bear the common name " honey ant" or "honeypot ant", due to curious behavior where some of the workers will swell with liquid foo ...
''. Usually the largest workers in the colony develop into repletes; and, if repletes are removed from the colony, other workers become repletes, demonstrating the flexibility of this particular polymorphism. This polymorphism in morphology and behaviour of workers initially was thought to be determined by environmental factors such as nutrition and hormones that led to different developmental paths; however, genetic differences between worker castes have been noted in ''Acromyrmex'' sp. These polymorphisms are caused by relatively small genetic changes; differences in a single gene of ''
Solenopsis invicta ''Solenopsis invicta'', the fire ant, or red imported fire ant (RIFA), is a species of ant native to South America. A member of the genus '' Solenopsis'' in the subfamily Myrmicinae, it was described by Swiss entomologist Felix Santschi as a ...
'' can decide whether the colony will have single or multiple queens. The Australian
jack jumper ant The jack jumper ant (''Myrmecia pilosula''), also known as the jack jumper, jumping jack, hopper ant, or jumper ant, is a species of venomous ant native to Australia. Most frequently found in Tasmania and southeast mainland Australia, it is a ...
(''Myrmecia pilosula'') has only a single pair of chromosomes (with the males having just one chromosome as they are
haploid Ploidy () is the number of complete sets of chromosomes in a cell (biology), cell, and hence the number of possible alleles for Autosome, autosomal and Pseudoautosomal region, pseudoautosomal genes. Here ''sets of chromosomes'' refers to the num ...
), the lowest number known for any animal, making it an interesting subject for studies in the genetics and developmental biology of social insects.


Genome size

Genome size Genome size is the total amount of DNA contained within one copy of a single complete genome. It is typically measured in terms of mass in picograms (trillionths or 10−12 of a gram, abbreviated pg) or less frequently in daltons, or as the tot ...
is a fundamental characteristic of an organism. Ants have been found to have tiny genomes, with the evolution of genome size suggested to occur through loss and accumulation of
non-coding Non-coding DNA (ncDNA) sequences are components of an organism's DNA that do not encode protein sequences. Some non-coding DNA is transcribed into functional non-coding RNA molecules (e.g. transfer RNA, microRNA, piRNA, ribosomal RNA, and regula ...
regions, mainly
transposable element A transposable element (TE), also transposon, or jumping gene, is a type of mobile genetic element, a nucleic acid sequence in DNA that can change its position within a genome. The discovery of mobile genetic elements earned Barbara McClinto ...
s, and occasionally by whole genome duplication. This may be related to
colonisation 475px, Map of the year each country achieved List of sovereign states by date of formation, independence. Colonization (British English: colonisation) is a process of establishing occupation of or control over foreign territories or peoples f ...
processes, but further studies are needed to verify this.


Life cycle

The life of an ant starts from an
egg An egg is an organic vessel grown by an animal to carry a possibly fertilized egg cell (a zygote) and to incubate from it an embryo within the egg until the embryo has become an animal fetus that can survive on its own, at which point the ...
; if the egg is fertilised, the progeny will be female
diploid Ploidy () is the number of complete sets of chromosomes in a cell, and hence the number of possible alleles for autosomal and pseudoautosomal genes. Here ''sets of chromosomes'' refers to the number of maternal and paternal chromosome copies, ...
, if not, it will be male
haploid Ploidy () is the number of complete sets of chromosomes in a cell (biology), cell, and hence the number of possible alleles for Autosome, autosomal and Pseudoautosomal region, pseudoautosomal genes. Here ''sets of chromosomes'' refers to the num ...
. Ants develop by
complete metamorphosis Holometabolism, also called complete metamorphosis, is a form of insect development which includes four life stages: egg, larva, pupa, and imago (or adult). Holometabolism is a synapomorphic trait of all insects in the clade Holometabola. Immature ...
with 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 ...
stages passing through a
pupa A pupa (; : pupae) is the life stage of some insects undergoing transformation between immature and mature stages. Insects that go through a pupal stage are holometabolous: they go through four distinct stages in their life cycle, the stages th ...
l stage before emerging as an adult. The larva is largely immobile and is fed and cared for by workers. Food is given to the larvae by
trophallaxis Trophallaxis () is the transfer of food or other fluids among members of a community through mouth-to-mouth (stomodeum, stomodeal) or anus-to-mouth (proctodeum, proctodeal) feeding. Along with nutrients, trophallaxis can involve the transfer of m ...
, a process in which an ant regurgitates liquid food held in its
crop A crop is a plant that can be grown and harvested extensively for profit or subsistence. In other words, a crop is a plant or plant product that is grown for a specific purpose such as food, Fiber, fibre, or fuel. When plants of the same spe ...
. This is also how adults share food, stored in the "social stomach". Larvae, especially in the later stages, may also be provided solid food, such as
trophic egg A trophic egg is an egg (biology), egg whose function is not reproduction but nutrition; in essence, the trophic egg serves as food for offspring hatched from viable eggs. In most species that produce them, a trophic egg is usually an Fertilization, ...
s, pieces of prey, and seeds brought by workers. The larvae grow through a series of four or five
moult In biology, moulting (British English), or molting (American English), also known as sloughing, shedding, or in many invertebrates, ecdysis, is a process by which an animal casts off parts of its body to serve some beneficial purpose, either at ...
s and enter the pupal stage. The pupa has the appendages free and not fused to the body as in a butterfly pupa. The differentiation into queens and workers (which are both female), and different
castes A caste is a fixed social group into which an individual is born within a particular system of social stratification: a caste system. Within such a system, individuals are expected to marry exclusively within the same caste (endogamy), foll ...
of workers, is influenced in some species by the nutrition the larvae obtain. Genetic influences and the control of gene expression by the developmental environment are complex and the determination of caste continues to be a subject of research. Winged male ants, called drones (termed "aner" in old literature), emerge from pupae along with the usually winged breeding females. Some species, such as
army ant The name army ant (or legionary ant or ''marabunta'') is applied to over 200 ant species in different lineages. Because of their aggressive predatory foraging groups, known as "raids", a huge number of ants forage simultaneously over a limited ...
s, have wingless queens. Larvae and pupae need to be kept at fairly constant temperatures to ensure proper development, and so often are moved around among the various brood chambers within the colony. A new ergate spends the first few days of its adult life caring for the queen and young. She then graduates to digging and other nest work, and later to defending the nest and foraging. These changes are sometimes fairly sudden, and define what are called temporal castes. Such age-based task-specialization or polyethism has been suggested as having evolved due to the high casualties involved in foraging and defence, making it an acceptable risk only for ants who are older and likely to die sooner from natural causes. In the Brazilian ant ''
Forelius pusillus ''Forelius pusillus'' is a species of ant in the genus '' Forelius''. Described by Santschi in 1922, the species is endemic to South America South America is a continent entirely in the Western Hemisphere and mostly in the Southern Hem ...
,'' the nest entrance is closed from the outside to protect the colony from predatory ant species at sunset each day. About one to eight workers seal the nest entrance from the outside and they have no chance of returning to the nest and are in effect sacrificed. Whether these seemingly suicidal workers are older workers has not been determined. Ant colonies can be long-lived. The queens can live for up to 30 years, and workers live from 1 to 3 years. Males, however, are more transitory, being quite short-lived and surviving for only a few weeks. Ant queens are estimated to live 100 times as long as solitary insects of a similar size. Ants are active all year long in the tropics; however, in cooler regions, they survive the winter in a state of dormancy known as
hibernation Hibernation is a state of minimal activity and metabolic reduction entered by some animal species. Hibernation is a seasonal heterothermy characterized by low body-temperature, slow breathing and heart-rate, and low metabolic rate. It is mos ...
. The forms of inactivity are varied and some temperate species have larvae going into the inactive state (
diapause In animal dormancy, diapause is the delay in development in response to regular and recurring periods of adverse environmental conditions.Tauber, M.J., Tauber, C.A., Masaki, S. (1986) ''Seasonal Adaptations of Insects''. Oxford University Press It ...
), while in others, the adults alone pass the winter in a state of reduced activity.


Reproduction

A wide range of reproductive strategies have been noted in ant species. Females of many species are known to be capable of reproducing asexually through
thelytokous parthenogenesis Thelytoky (from the Greek θῆλυς ''thēlys'' "female" and τόκος ''tókos'' "birth") is a type of parthenogenesis and is the absence of mating and subsequent production of all female diploid offspring as for example in aphids. Thelytokou ...
. Secretions from the male accessory glands in some species can plug the female genital opening and prevent females from re-mating. Most ant species have a system in which only the queen and breeding females have the ability to mate. Contrary to popular belief, some ant nests have multiple queens, while others may exist without queens. Workers with the ability to reproduce are called " gamergates" and colonies that lack queens are then called gamergate colonies; colonies with queens are said to be queen-right. Drones can also mate with existing queens by entering a foreign colony, such as in
army ant The name army ant (or legionary ant or ''marabunta'') is applied to over 200 ant species in different lineages. Because of their aggressive predatory foraging groups, known as "raids", a huge number of ants forage simultaneously over a limited ...
s. When the drone is initially attacked by the workers, it releases a mating
pheromone A pheromone () is a secreted or excreted chemical factor that triggers a social response in members of the same species. Pheromones are chemicals capable of acting like hormones outside the body of the secreting individual, to affect the behavio ...
. If recognized as a mate, it will be carried to the queen to mate. Males may also patrol the nest and fight others by grabbing them with their mandibles, piercing their
exoskeleton An exoskeleton () . is a skeleton that is on the exterior of an animal in the form of hardened integument, which both supports the body's shape and protects the internal organs, in contrast to an internal endoskeleton (e.g. human skeleton, that ...
and then marking them with a pheromone. The marked male is interpreted as an invader by worker ants and is killed. Most ants are
univoltine Voltinism is a term used in biology to indicate the number of broods or generations of an organism in a year. The term is most often applied to insects, and is particularly in use in sericulture, where silkworm varieties vary in their voltinism. ...
, producing a new generation each year. During the species-specific breeding period, winged females and winged males, known to
entomologists Entomology (from Ancient Greek ἔντομον (''éntomon''), meaning "insect", and -logy from λόγος (''lógos''), meaning "study") is the branch of zoology that focuses on insects. Those who study entomology are known as entomologists. In ...
as
alate Alate (Latin ''ālātus'', from ''āla'' (“wing”)) is an adjective and noun used in entomology and botany to refer to something that has wings or winglike structures. In entomology In entomology, "alate" usually refers to the winged form of ...
s, leave the colony in what is called a
nuptial flight Nuptial flight is an important phase in the reproduction of most ant, termite, and some bee species. It is also observed in some fly species, such as '' Rhamphomyia longicauda''. During the flight, virgin queens mate with males and then land ...
. The nuptial flight usually takes place in the late spring or early summer when the weather is hot and humid. Heat makes flying easier and freshly fallen rain makes the ground softer for mated queens to dig nests. Males typically take flight before the females. Males then use visual cues to find a common mating ground, for example, a landmark such as a
pine tree A pine is any conifer tree or shrub in the genus ''Pinus'' () of the family Pinaceae. ''Pinus'' is the sole genus in the subfamily Pinoideae. ''World Flora Online'' accepts 134 species-rank taxa (119 species and 15 nothospecies) of pines as c ...
to which other males in the area converge. Males secrete a mating pheromone that females follow. Males will mount females in the air, but the actual mating process usually takes place on the ground. Females of some species mate with just one male but in others they may mate with as many as ten or more different males, storing the
sperm Sperm (: sperm or sperms) is the male reproductive Cell (biology), cell, or gamete, in anisogamous forms of sexual reproduction (forms in which there is a larger, female reproductive cell and a smaller, male one). Animals produce motile sperm ...
in their
spermatheca The spermatheca (pronounced : spermathecae ), also called ''receptaculum seminis'' (: ''receptacula seminis''), is an organ of the female reproductive tract in insects, e.g. ants, bees, some molluscs, Oligochaeta worms and certain other in ...
e.Hölldobler & Wilson (1990), pp. 143–179 The genus ''
Cardiocondyla ''Cardiocondyla'' is an Old World genus of ants in the subfamily Myrmicinae. Distribution Approximately 70 species are currently recognized as belonging to this genus, most of which are distributed in the Old World tropics and subtropics, but a ...
'' have species with both winged and wingless males, where the latter will only mate with females living in the same nest. Some species in the genus have lost winged males completely, and only produce wingless males. In '' C. elegans,'' workers may transport newly emerged queens to other conspecific nests where the wingless males from unrelated colonies can mate with them, a behavioural adaptation that may reduce the chances of inbreeding. Mated females then seek a suitable place to begin a colony. There, they break off their wings using their tibial spurs and begin to lay and care for eggs. The females can selectively fertilise future eggs with the sperm stored to produce diploid workers or lay unfertilized haploid eggs to produce drones. The first workers to hatch, known as nanitics, are weaker and smaller than later workers but they begin to serve the colony immediately. They enlarge the nest, forage for food, and care for the other eggs. Species that have multiple queens may have a queen leaving the nest along with some workers to found a colony at a new site, a process akin to
swarming Swarm behaviour, or swarming, is a collective animal behaviour, collective behaviour exhibited by entities, particularly animals, of similar size which aggregate together, perhaps milling about the same spot or perhaps moving ''en masse'' or a ...
in
honeybee A honey bee (also spelled honeybee) is a eusocial flying insect within the genus ''Apis'' of the bee clade, all native to mainland Afro-Eurasia. After bees spread naturally throughout Africa and Eurasia, humans became responsible for the c ...
s.


Nests, colonies, and supercolonies

The typical ant species has a colony occupying a single nest, housing one or more queens, where the brood is raised. There are however more than 150 species of ants in 49 genera that are known to have colonies consisting of multiple spatially separated nests. These polydomous (as opposed to monodomous) colonies have food and workers moving between the nests. Membership to a colony is identified by the response of worker ants which identify whether another individual belongs to their own colony or not. A signature cocktail of body surface chemicals (also known as cuticular hydrocarbons or CHCs) forms the so-called colony odor which other members can recognize. Some ant species appear to be less discriminating and in the Argentine ant ''Linepithema humile,'' workers carried from a colony anywhere in the southern US and Mexico are acceptable within other colonies in the same region. Similarly workers from colonies that have established in Europe are accepted by any other colonies within Europe but not by the colonies in the Americas. The interpretation of these observations has been debated and some have been termed these large populations as supercolonies while others have termed the populations as unicolonial.


Behaviour and ecology


Communication

Ants communicate with each other using
pheromone A pheromone () is a secreted or excreted chemical factor that triggers a social response in members of the same species. Pheromones are chemicals capable of acting like hormones outside the body of the secreting individual, to affect the behavio ...
s, sounds, and touch. Since most ants live on the ground, they use the soil surface to leave pheromone trails that may be followed by other ants. In species that forage in groups, a forager that finds food marks a trail on the way back to the colony; this trail is followed by other ants, these ants then reinforce the trail when they head back with food to the colony. When the food source is exhausted, no new trails are marked by returning ants and the scent slowly dissipates. This behaviour helps ants deal with changes in their environment. For instance, when an established path to a food source is blocked by an obstacle, the foragers leave the path to explore new routes. If an ant is successful, it leaves a new trail marking the shortest route on its return. Successful trails are followed by more ants, reinforcing better routes and gradually identifying the best path. Ants use pheromones for more than just making trails. A crushed ant emits an alarm pheromone that sends nearby ants into an attack frenzy and attracts more ants from farther away. Several ant species even use "
propaganda Propaganda is communication that is primarily used to influence or persuade an audience to further an agenda, which may not be objective and may be selectively presenting facts to encourage a particular synthesis or perception, or using loaded l ...
pheromones" to confuse enemy ants and make them fight among themselves. Pheromones are produced by a wide range of structures including
Dufour's gland Dufour's gland is an abdominal gland of certain insects, part of the anatomy of the ovipositor or sting apparatus in female members of Apocrita. The diversification of Hymenoptera took place in the Cretaceous and the gland may have developed a ...
s, poison glands and glands on the
hindgut The hindgut (or epigaster) is the posterior ( caudal) part of the alimentary canal. In mammals, it includes the distal one third of the transverse colon and the splenic flexure, the descending colon, sigmoid colon and up to the ano-rectal junct ...
,
pygidium The pygidium (: pygidia) is the posterior body part or shield of crustaceans and some other arthropods, such as insects and the extinct trilobites. In groups other than insects, it contains the anus and, in females, the ovipositor. It is compos ...
,
rectum The rectum (: rectums or recta) is the final straight portion of the large intestine in humans and some other mammals, and the gut in others. Before expulsion through the anus or cloaca, the rectum stores the feces temporarily. The adult ...
,
sternum The sternum (: sternums or sterna) or breastbone is a long flat bone located in the central part of the chest. It connects to the ribs via cartilage and forms the front of the rib cage, thus helping to protect the heart, lungs, and major bl ...
, and hind
tibia The tibia (; : tibiae or tibias), also known as the shinbone or shankbone, is the larger, stronger, and anterior (frontal) of the two Leg bones, bones in the leg below the knee in vertebrates (the other being the fibula, behind and to the outsi ...
. Pheromones also are exchanged, mixed with food, and passed by
trophallaxis Trophallaxis () is the transfer of food or other fluids among members of a community through mouth-to-mouth (stomodeum, stomodeal) or anus-to-mouth (proctodeum, proctodeal) feeding. Along with nutrients, trophallaxis can involve the transfer of m ...
, transferring information within the colony. This allows other ants to detect what task group (e.g., foraging or nest maintenance) other colony members belong to. In ant species with queen
caste A caste is a Essentialism, fixed social group into which an individual is born within a particular system of social stratification: a caste system. Within such a system, individuals are expected to marry exclusively within the same caste (en ...
s, when the dominant queen stops producing a specific pheromone, workers begin to raise new queens in the colony.Hölldobler & Wilson (1990), p. 354 Some ants produce sounds by
stridulation Stridulation is the act of producing sound by rubbing together certain body parts. This behavior is mostly associated with insects, but other animals are known to do this as well, such as a number of species of fish, snakes and spiders. The mech ...
, using the gaster segments and their mandibles. Sounds may be used to communicate with colony members or with other species.


Defence

Ants attack and defend themselves by biting and, in many species, by stinging often injecting or spraying chemicals.
Bullet ant A bullet is a kinetic projectile, a component of firearm ammunition that is shot from a gun barrel. They are made of a variety of materials, such as copper, lead, steel, polymer, rubber and even wax; and are made in various shapes and constru ...
s ('' Paraponera''), located 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 ...
, are considered to have the most painful sting of any insect, although it is usually not fatal to humans. This sting is given the highest rating on the
Schmidt sting pain index The Schmidt sting pain index is a pain scale rating the relative pain caused by different hymenopteran stings. It is mainly the work of Justin O. Schmidt, who was an entomologist at the Carl Hayden Bee Research Center in Arizona. Schmidt's or ...
. The sting of
jack jumper ant The jack jumper ant (''Myrmecia pilosula''), also known as the jack jumper, jumping jack, hopper ant, or jumper ant, is a species of venomous ant native to Australia. Most frequently found in Tasmania and southeast mainland Australia, it is a ...
s can be lethal for humans, and an
antivenom Antivenom, also known as antivenin, venom antiserum, and antivenom immunoglobulin, is a specific treatment for envenomation. It is composed of antibodies and used to treat certain venomous bites and stings. Antivenoms are recommended only if ...
has been developed for it.
Fire ant Fire ants are several species of ants in the genus ''Solenopsis'', which includes over 200 species. ''Solenopsis'' are stinging ants, and most of their common names reflect this, for example, ginger ants and tropical fire ants. Many of the nam ...
s, '' Solenopsis'' spp., are unique in having a venom sac containing
piperidine alkaloids Piperidine alkaloids are naturally occurring chemical compounds from the group of alkaloids, which are chemically derived from piperidine. Alkaloids with a piperidine building block are widespread and are usually further subdivided according to ...
. Their stings are painful and can be dangerous to hypersensitive people. Formicine ants secrete a poison from their glands, made mainly of
formic acid Formic acid (), systematically named methanoic acid, is the simplest carboxylic acid. It has the chemical formula HCOOH and structure . This acid is an important intermediate in chemical synthesis and occurs naturally, most notably in some an ...
. Trap-jaw ants of the genus ''
Odontomachus ''Odontomachus'' is a genus of ants commonly called trap-jaw ants found in the tropics and subtropics throughout the world. Overview Commonly known as trap-jaw ants, species in ''Odontomachus'' have a pair of large, straight mandible (insect) ...
'' are equipped with mandibles called trap-jaws, which snap shut faster than any other
predatory 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 feeding behaviours that includes parasitism and micropredation (which usually do not kill ...
appendage An appendage (or outgrowth) is an external body part or natural prolongation that protrudes from an organism's body such as an arm or a leg. Protrusions from single-celled bacteria and archaea are known as cell-surface appendages or surface app ...
s within the animal kingdom. One study of ''
Odontomachus bauri ''Odontomachus bauri'' is a species of ponerinae ant known as trap jaw ants. The trap jaw consists of mandibles which contain a spring-loaded catch mechanism. This mechanism permits the ants to accumulate energy before striking or releasing the ...
'' recorded peak speeds of between , with the jaws closing within 130
microsecond A microsecond is a unit of time in the International System of Units (SI) equal to one millionth (0.000001 or 10−6 or ) of a second. Its symbol is μs, sometimes simplified to us when Unicode is not available. A microsecond is to one second, ...
s on average. The ants were also observed to use their jaws as a
catapult A catapult is a ballistics, ballistic device used to launch a projectile at a great distance without the aid of gunpowder or other propellants – particularly various types of ancient and medieval siege engines. A catapult uses the sudden rel ...
to eject intruders or fling themselves backward to escape a threat. Before striking, the ant opens its mandibles extremely widely and locks them in this position by an internal mechanism. Energy is stored in a thick band of
muscle Muscle is a soft tissue, one of the four basic types of animal tissue. There are three types of muscle tissue in vertebrates: skeletal muscle, cardiac muscle, and smooth muscle. Muscle tissue gives skeletal muscles the ability to muscle contra ...
and explosively released when triggered by the stimulation of sensory organs resembling hairs on the inside of the mandibles. The mandibles also permit slow and fine movements for other tasks. Trap-jaws also are seen in other ponerines such as ''
Anochetus ''Anochetus'' is a genus of small, carnivorous ants found in the tropics and subtropics The subtropical zones or subtropics are geographical and climate zones immediately to the north and south of the tropics. Geographically part of the ...
'', as well as some genera in the tribe Attini, such as ''
Daceton ''Daceton'' is a Neotropical genus of ants in the subfamily Myrmicinae. The genus contains only two species: '' D. armigerum'', the most studied species, distributed throughout northern South America, and '' D. boltoni'', known from Brazil and Pe ...
'', ''
Orectognathus ''Orectognathus'' is a genus of ants in the subfamily Myrmicinae. Species *''Orectognathus alligator'' Taylor, 1980 *''Orectognathus antennatus'' Smith, 1854 *''Orectognathus biroi'' Szabó, 1926 *''Orectognathus chyzeri'' Emery, 1897 *''Orecto ...
'', and ''
Strumigenys ''Strumigenys'' is a genus of ants in the subfamily Myrmicinae. Biology ''Strumigenys'' form small nests in soil, under or between rocks, or in and under logs or under cattle dung. Some species nest in association with other ants such as '' Both ...
'', which are viewed as examples of
convergent evolution Convergent evolution is the independent evolution of similar features in species of different periods or epochs in time. Convergent evolution creates analogous structures that have similar form or function but were not present in the last comm ...
. A Malaysian species of ant in the ''
Camponotus Carpenter ants (''Camponotus'' spp.) are a genus of large ants (workers ) indigenous to many parts of the world. True carpenter ants build nests inside wood, consisting of galleries chewed out with their mandibles or jaws, preferably in dead, ...
'' ''cylindricus''
group A group is a number of persons or things that are located, gathered, or classed together. Groups of people * Cultural group, a group whose members share the same cultural identity * Ethnic group, a group whose members share the same ethnic iden ...
has enlarged mandibular glands that extend into their gaster. If combat takes a turn for the worse, a worker may perform a final act of suicidal altruism by rupturing the membrane of its gaster, causing the content of its mandibular glands to burst from the
anterior Standard anatomical terms of location are used to describe unambiguously the anatomy of humans and other animals. The terms, typically derived from Latin or Greek roots, describe something in its standard anatomical position. This position pro ...
region of its head, spraying a poisonous, corrosive secretion containing
acetophenone Acetophenone is the organic compound with the formula C6H5C(O)CH3. It is the simplest aromatic ketone. This colorless, viscous liquid is a precursor to useful resins and fragrances. Production Acetophenone is formed as a byproduct of the cumene ...
s and other chemicals that immobilise small insect attackers. The worker subsequently dies. In addition to defence against predators, ants need to protect their colonies from
pathogen In biology, a pathogen (, "suffering", "passion" and , "producer of"), in the oldest and broadest sense, is any organism or agent that can produce disease. A pathogen may also be referred to as an infectious agent, or simply a Germ theory of d ...
s. Secretions from the metapleural gland, unique to the ants, produce a complex range of chemicals including several with antibiotic properties. Some worker ants maintain the hygiene of the colony and their activities include undertaking or ''
necrophoresis Necrophoresis is a sanitation behavior found in social insects – such as ants, bees, wasps, and termites – in which they carry away the dead bodies of members of their colony from the nest or hive area. The term was introduced in 1958 by E. O. ...
'', the disposal of dead nest-mates.
Oleic acid Oleic acid is a fatty acid that occurs naturally in various animal and vegetable fats and oils. It is an odorless, colorless oil, although commercial samples may be yellowish due to the presence of impurities. In chemical terms, oleic acid is cl ...
has been identified as the compound released from dead ants that triggers necrophoric behaviour in ''Atta mexicana'' while workers of ''Linepithema humile'' react to the absence of characteristic chemicals (
dolichodial Dolichodial is a natural chemical compound with two aldehyde groups, which belongs to the group of iridoids. Chemistry It has in its five-membered ring three asymmetric carbon atoms and accordingly exists in four diastereomeric pairs of enantiome ...
and
iridomyrmecin Iridomyrmecin is a defensive chemical, classified as an iridoid Iridoids are a type of monoterpenoids in the general form of cyclopentanopyran, found in a wide variety of plants and some animals. They are biosynthetically derived from 8-oxoge ...
) present on the cuticle of their living nestmates to trigger similar behaviour. In '' Megaponera analis,'' injured ants are treated by nestmastes with secretions from their metapleural glands which protect them from infection. ''Camponotus'' ants do not have a metapleural gland and '' Camponotus maculatus'' as well as '' C. floridanus'' workers have been found to amputate the affected legs of nestmates when the femur is injured. A femur injury carries a greater risk of infection unlike a tibia injury. Nests may be protected from physical threats such as flooding and overheating by elaborate nest architecture. Workers of ''Cataulacus muticus'', an arboreal species that lives in plant hollows, respond to flooding by drinking water inside the nest, and excreting it outside. ''
Camponotus anderseni ''Colobopsis anderseni'', synonym ''Camponotus anderseni'', is a species of mangrove ant found in northern Australia. Taxonomy The species was originally placed in the ant genus ''Camponotus'' when described in 2001 as ''Camponotus (Colobopsis) ...
'', which nests in the cavities of wood in mangrove habitats, deals with submergence under water by switching to
anaerobic respiration Anaerobic respiration is respiration using electron acceptors other than molecular oxygen (O2). Although oxygen is not the final electron acceptor, the process still uses a respiratory electron transport chain. In aerobic organisms undergoing ...
.


Learning

Many animals can learn behaviours by imitation, but ants may be the only group apart from
mammal A mammal () is a vertebrate animal of the Class (biology), class Mammalia (). Mammals are characterised by the presence of milk-producing mammary glands for feeding their young, a broad neocortex region of the brain, fur or hair, and three ...
s where interactive teaching has been observed. A knowledgeable forager of ''
Temnothorax albipennis ''Temnothorax albipennis'', the rock ant is a species of small ant in the subfamily Myrmicinae. It occurs in Europe and builds simple nests in rock crevices. Description This species has the typical ant body pattern of head, mesosoma and metas ...
'' can lead a naïve nest-mate to newly discovered food by the process of
tandem running Tandem running is a pair movement coordination observed in ants and termites. In ants, tandem running is used for social learning, by which one ant leads another native ant from the nest to the food source it has found. Tandem running is also use ...
. The follower obtains knowledge through its leading tutor. The leader is acutely sensitive to the progress of the follower and slows down when the follower lags and speeds up when the follower gets too close. Controlled experiments with colonies of ''
Cerapachys biroi ''Ooceraea biroi'', the clonal raider ant, is a queenless clonal ant in the genus ''Ooceraea'' (recently transferred from the genus ''Cerapachys''). Native to the Asian mainland, this species has become invasive on tropical and subtropical island ...
'' suggest that an individual may choose nest roles based on her previous experience. An entire generation of identical workers was divided into two groups whose outcome in food foraging was controlled. One group was continually rewarded with prey, while it was made certain that the other failed. As a result, members of the successful group intensified their foraging attempts while the unsuccessful group ventured out fewer and fewer times. A month later, the successful foragers continued in their role while the others had moved to specialise in brood care.


Nest construction

Complex nests are built by many ant species, but other species are nomadic and do not build permanent structures. Ants may form subterranean nests or build them on trees. These nests may be found in the ground, under stones or logs, inside logs, hollow stems, or even acorns. The materials used for construction include soil and plant matter, and ants carefully select their nest sites; ''Temnothorax albipennis'' will avoid sites with dead ants, as these may indicate the presence of pests or disease. They are quick to abandon established nests at the first sign of threats. The
army ant The name army ant (or legionary ant or ''marabunta'') is applied to over 200 ant species in different lineages. Because of their aggressive predatory foraging groups, known as "raids", a huge number of ants forage simultaneously over a limited ...
s of South America, such as the ''
Eciton burchellii ''Eciton burchellii'' is a species of New World army ant in the genus '' Eciton''. This species performs expansive, organized swarm raids that give it the informal name, ''Eciton'' army ant. This species displays a high degree of worker polymor ...
'' species, and the
driver ant ''Dorylus'', also known as driver ants, safari ants, or siafu, is a large genus of army ants found primarily in central and east Africa, although the range also extends to southern Africa and tropical Asia. The term siafu is a loanword from Swa ...
s of Africa do not build permanent nests, but instead, alternate between nomadism and stages where the workers form a temporary nest ( bivouac) from their own bodies, by holding each other together.
Weaver ant Weaver ants or green ants are eusocial insects of the Hymenoptera family Formicidae belonging to the tribe Oecophyllini. Weaver ants live in trees (they are obligately arboreal) and are known for their unique nest building behaviour where wor ...
(''Oecophylla'' spp.) workers build nests in trees by attaching leaves together, first pulling them together with bridges of workers and then inducing their larvae to produce silk as they are moved along the leaf edges. Similar forms of nest construction are seen in some species of '' Polyrhachis''. ''
Formica polyctena ''Formica polyctena'' is a species of European red wood ant in the genus ''Formica'' and large family Formicidae. The species was first described by Arnold Förster in 1850. The latin species name ''polyctena'' is from Greek and literally mea ...
'', among other ant species, constructs nests that maintain a relatively constant interior temperature that aids in the development of larvae. The ants maintain the nest temperature by choosing the location, nest materials, controlling ventilation and maintaining the heat from solar radiation, worker activity and metabolism, and in some moist nests, microbial activity in the nest materials. Some ant species, such as those that use natural cavities, can be opportunistic and make use of the controlled micro-climate provided inside human dwellings and other artificial structures to house their colonies and nest structures.


Cultivation of food

Most ants are generalist predators,
scavenger Scavengers are animals that consume Corpse decomposition, dead organisms that have died from causes other than predation or have been killed by other predators. While scavenging generally refers to carnivores feeding on carrion, it is also a he ...
s, and indirect
herbivore A herbivore is an animal anatomically and physiologically evolved to feed on plants, especially upon vascular tissues such as foliage, fruits or seeds, as the main component of its diet. These more broadly also encompass animals that eat ...
s, but a few have evolved specialised ways of obtaining nutrition. It is believed that many ant species that engage in indirect herbivory rely on specialized symbiosis with their gut microbes to upgrade the nutritional value of the food they collect and allow them to survive in nitrogen poor regions, such as rainforest canopies.
Leafcutter ant Leafcutter ants are fungus-growing ants that share the behaviour of cutting leaves which they carry back to their nests to farm fungus. Next to humans, leafcutter ants form some of the largest and most complex animal societies on Earth. In a few ...
s ('' Atta'' and ''
Acromyrmex ''Acromyrmex'' is a genus of New World ants of the subfamily Myrmicinae. This genus is found in South America and parts of Central America, México and the Caribbean Islands, and contains 33 known species. Commonly known as " leafcutter ants" th ...
'') feed exclusively on a
fungus A fungus (: fungi , , , or ; or funguses) is any member of the group of eukaryotic organisms that includes microorganisms such as yeasts and mold (fungus), molds, as well as the more familiar mushrooms. These organisms are classified as one ...
that grows only within their colonies. They continually collect leaves which are taken to the colony, cut into tiny pieces and placed in fungal gardens. Ergates specialise in related tasks according to their sizes. The largest ants cut stalks, smaller workers chew the leaves and the smallest tend the fungus. Leafcutter ants are sensitive enough to recognise the reaction of the fungus to different plant material, apparently detecting chemical signals from the fungus. If a particular type of leaf is found to be toxic to the fungus, the colony will no longer collect it. The ants feed on structures produced by the fungi called ''
gongylidia Gongylidia (singular gongylidium) are hyphal swellings of fungus cultivated by higher-attine genera of fungus-growing ants. This fungus no longer exists naturally outside the ant colonies. Developing larvae feed on the gongylidia and distribu ...
''.
Symbiotic Symbiosis (Ancient Greek : living with, companionship < : together; and ''bíōsis'': living) is any type of a close and long-term biolo ...
bacteria on the exterior surface of the ants produce antibiotics that kill bacteria introduced into the nest that may harm the fungi.


Navigation

Foraging Foraging is searching for wild food resources. It affects an animal's fitness because it plays an important role in an animal's ability to survive and reproduce. Foraging theory is a branch of behavioral ecology that studies the foraging behavi ...
ants travel distances of up to from their nest and scent trails allow them to find their way back even in the dark. In hot and arid regions, day-foraging ants face death by
desiccation Desiccation is the state of extreme dryness, or the process of extreme drying. A desiccant is a hygroscopic (attracts and holds water) substance that induces or sustains such a state in its local vicinity in a moderately sealed container. The ...
, so the ability to find the shortest route back to the nest reduces that risk. Diurnal desert ants of the genus ''
Cataglyphis ''Cataglyphis'' is a genus of ant, desert ants, in the subfamily Formicinae. Its most famous species is ''C. bicolor'', the Sahara Desert ant, which runs on hot sand to find insects that died of heat exhaustion, and can, like other several oth ...
'' such as the Sahara desert ant navigate by keeping track of direction as well as distance travelled. Distances travelled are measured using an internal
pedometer A pedometer, or step-counter, is a device, usually portable and electronic or electromechanical, that counts each step a person takes by detecting the motion of the person's hands or hips. Because the distance of each person's step varies, an ...
that keeps count of the steps taken and also by evaluating the movement of objects in their visual field (
optical flow Optical flow or optic flow is the pattern of apparent motion of objects, surfaces, and edges in a visual scene caused by the relative motion between an observer and a scene. Optical flow can also be defined as the distribution of apparent velocit ...
). Directions are measured using the position of the sun. They integrate this information to find the shortest route back to their nest. Like all ants, they can also make use of visual landmarks when available as well as olfactory and tactile cues to navigate. Some species of ant are able to use the
Earth's magnetic field Earth's magnetic field, also known as the geomagnetic field, is the magnetic field that extends from structure of Earth, Earth's interior out into space, where it interacts with the solar wind, a stream of charged particles emanating from ...
for navigation. The compound eyes of ants have specialised cells that detect polarised light from the Sun, which is used to determine direction. These polarization detectors are sensitive in the
ultraviolet Ultraviolet radiation, also known as simply UV, is electromagnetic radiation of wavelengths of 10–400 nanometers, shorter than that of visible light, but longer than X-rays. UV radiation is present in sunlight and constitutes about 10% of ...
region of the light spectrum. In some army ant species, a group of foragers who become separated from the main column may sometimes turn back on themselves and form a circular
ant mill An ant mill is an observed phenomenon in which a group of army ants, separated from the main foraging party, lose the pheromone track and begin to follow one another, forming a continuously rotating circle. This circle is commonly known as a "d ...
. The workers may then run around continuously until they die of exhaustion.


Locomotion

The female worker ants do not have wings and reproductive females lose their wings after their mating flights in order to begin their colonies. Therefore, unlike their wasp ancestors, most ants travel by walking. Some species are capable of leaping. For example, Jerdon's jumping ant (''
Harpegnathos saltator ''Harpegnathos saltator'', sometimes called the Indian jumping ant or Jerdon's jumping ant, is a species of ant found in India. They have long mandibles and have the ability to leap a few inches. They are large-eyed and active Predation, predato ...
'') is able to jump by synchronising the action of its mid and hind pairs of legs. There are several species of
gliding ant Gliding ants are arboreal ants of several different genera that are able to control the direction of their descent when falling from a tree. Living in the rainforest canopy like many other gliders, gliding ants use their gliding to return to the ...
including ''Cephalotes atratus''; this may be a common trait among arboreal ants with small colonies. Ants with this ability are able to control their horizontal movement so as to catch tree trunks when they fall from atop the forest canopy. Other species of ants can form chains to bridge gaps over water, underground, or through spaces in vegetation. Some species also form floating rafts that help them survive floods. These rafts may also have a role in allowing ants to colonise islands. ''
Polyrhachis sokolova '' Polyrhachis sokolova'' is a species of ant from Australia and New Guinea that recently was discovered to be capable of surviving tidal inundations. Researchers at the James Cook University gained attention in 2006 after finding and reportin ...
'', a species of ant found in
Australia Australia, officially the Commonwealth of Australia, is a country comprising mainland Australia, the mainland of the Australia (continent), Australian continent, the island of Tasmania and list of islands of Australia, numerous smaller isl ...
n
mangrove A mangrove is a shrub or tree that grows mainly in coastal saline water, saline or brackish water. Mangroves grow in an equatorial climate, typically along coastlines and tidal rivers. They have particular adaptations to take in extra oxygen a ...
swamps, can swim and live in underwater nests. Since they lack
gill A gill () is a respiration organ, respiratory organ that many aquatic ecosystem, aquatic organisms use to extract dissolved oxygen from water and to excrete carbon dioxide. The gills of some species, such as hermit crabs, have adapted to allow r ...
s, they go to trapped pockets of air in the submerged nests to breathe.


Cooperation and competition

Not all ants have the same kind of societies. The Australian
bulldog ant ''Myrmecia'' is a genus of ants first established by Danish zoologist Johan Christian Fabricius in 1804. The genus is a member of the subfamily Myrmeciinae of the family Formicidae. ''Myrmecia'' is a large genus of ants, comprising at least ...
s are among the biggest and most basal of ants. Like virtually all ants, they are
eusocial Eusociality ( Greek 'good' and social) is the highest level of organization of sociality. It is defined by the following characteristics: cooperative brood care (including care of offspring from other individuals), overlapping generations wit ...
, but their social behaviour is poorly developed compared to other species. Each individual hunts alone, using her large eyes instead of chemical senses to find prey. Some species attack and take over neighbouring ant colonies. Extreme specialists among these
slave-raiding ant Slave-making ants or slaver ants are brood parasites that capture broods of other ant species to increase the worker force of their colony. After emerging in the slave-maker nest, slave workers work as if they were in their own colony, while paras ...
s, such as the Amazon ants, are incapable of feeding themselves and need captured workers to survive. Captured workers of enslaved ''
Temnothorax ''Temnothorax'' is a genus of ants in the subfamily Myrmicinae. It contains more than 380 species. Biology The workers of ''Temnothorax'' species are generally small ( mesosoma length 0.715 mm). Colonies are typically monogynous, although fac ...
'' species have evolved a counter-strategy, destroying just the female pupae of the slave-making ''
Temnothorax americanus ''Temnothorax americanus'' is a species of slave-maker ant in the genus '' Temnothorax''. The ants are 2–3 mm in size, and endemic to the northeastern United States The United States of America (USA), also known as the United State ...
'', but sparing the males (who do not take part in slave-raiding as adults). Ants identify kin and nestmates through their scent, which comes from
hydrocarbon In organic chemistry, a hydrocarbon is an organic compound consisting entirely of hydrogen and carbon. Hydrocarbons are examples of group 14 hydrides. Hydrocarbons are generally colourless and Hydrophobe, hydrophobic; their odor is usually fain ...
-laced secretions that coat their exoskeletons. If an ant is separated from its original colony, it will eventually lose the colony scent. Any ant that enters a colony without a matching scent will be attacked. Parasitic ant species enter the colonies of host ants and establish themselves as social parasites; species such as ''
Strumigenys xenos ''Strumigenys xenos'' is a species of ant in the subfamily Myrmicinae. It is a permanent social parasite, which forms no workers and lives in the nests of its host '' Strumigenys perplexa''. It is found in Australia and New Zealand Ne ...
'' are entirely parasitic and do not have workers, but instead, rely on the food gathered by their ''Strumigenys perplexa'' hosts. This form of parasitism is seen across many ant genera, but the parasitic ant is usually a species that is closely related to its host. A variety of methods are employed to enter the nest of the host ant. A parasitic queen may enter the host nest before the first brood has hatched, establishing herself prior to development of a colony scent. Other species use pheromones to confuse the host ants or to trick them into carrying the parasitic queen into the nest. Some simply fight their way into the nest.Hölldobler & Wilson (1990), pp. 436–448 A conflict between the sexes of a species is seen in some species of ants with these reproducers apparently competing to produce offspring that are as closely related to them as possible. The most extreme form involves the production of clonal offspring. An extreme of sexual conflict is seen in ''
Wasmannia auropunctata The little fire ant (''Wasmannia auropunctata''), also known as the electric ant, is a small (approx long), light to golden brown (ginger) social ant native to Central and South America, now spread to parts of Africa (including Gabon and Camero ...
'', where the queens produce diploid daughters by thelytokous parthenogenesis and males produce clones by a process whereby a diploid egg loses its maternal contribution to produce haploid males who are clones of the father.


Relationships with other organisms

Ants form
symbiotic Symbiosis (Ancient Greek : living with, companionship < : together; and ''bíōsis'': living) is any type of a close and long-term biolo ...
associations with a range of species, including other ant species, other insects, plants, and fungi. They also are preyed on by many animals and even certain fungi. Some arthropod species spend part of their lives within ant nests, either preying on ants, their larvae, and eggs, consuming the food stores of the ants, or avoiding predators. These
inquiline In zoology, an inquiline (from Latin ''inquilinus'', "lodger" or "tenant") is an animal that lives commensally in the nest, burrow, or dwelling place of an animal of another species. For example, some organisms, such as insects, may live in the ...
s may bear a close resemblance to ants. The nature of this
ant mimicry Ant mimicry or myrmecomorphy is mimicry of ants by other organisms; it has evolved over 70 times. Ants are abundant all over the world, and potential predators that rely on vision to identify their prey, such as birds and wasps, normally avoid t ...
(myrmecomorphy) varies, with some cases involving
Batesian mimicry Batesian mimicry is a form of mimicry where a harmless species has evolved to imitate the warning signals of a harmful species directed at a predator of them both. It is named after the English naturalist Henry Walter Bates, who worked on butt ...
, where the mimic reduces the risk of predation. Others show
Wasmannian mimicry Ant mimicry or myrmecomorphy is mimicry of ants by other organisms; it has evolved over 70 times. Ants are abundant all over the world, and potential predators that rely on vision to identify their prey, such as birds and wasps, normally avoid t ...
, a form of mimicry seen only in inquilines.
Aphid Aphids are small sap-sucking insects in the Taxonomic rank, family Aphididae. Common names include greenfly and blackfly, although individuals within a species can vary widely in color. The group includes the fluffy white Eriosomatinae, woolly ...
s and other
hemiptera Hemiptera (; ) is an order of insects, commonly called true bugs, comprising more than 80,000 species within groups such as the cicadas, aphids, planthoppers, leafhoppers, assassin bugs, bed bugs, and shield bugs. They range in size from ...
n insects secrete a sweet liquid called honeydew, when they feed on
plant sap Sap is a fluid transported in the xylem cells (vessel elements or tracheids) or phloem sieve tube elements of a plant. These cells transport water and nutrients throughout the plant. Sap is distinct from latex, resin, or cell sap; it is a sep ...
. The sugars in honeydew are a high-energy food source, which many ant species collect. In some cases, the aphids secrete the honeydew in response to ants tapping them with their antennae. The ants in turn keep predators away from the aphids and will move them from one feeding location to another. When migrating to a new area, many colonies will take the aphids with them, to ensure a continued supply of honeydew. Ants also tend
mealybug Mealybugs are insects in the family Pseudococcidae, unarmored scale insects found in moist, warm habitats. Of the more than 2,000 described species, many are considered pests as they feed on plant juices of greenhouse plants, house plants and ...
s to harvest their honeydew. Mealybugs may become a serious pest of
pineapple The pineapple (''Ananas comosus'') is a Tropical vegetation, tropical plant with an edible fruit; it is the most economically significant plant in the family Bromeliaceae. The pineapple is indigenous to South America, where it has been culti ...
s if ants are present to protect mealybugs from their natural enemies. Myrmecophilous (ant-loving)
caterpillar Caterpillars ( ) are the larval stage of members of the order Lepidoptera (the insect order comprising butterflies and moths). As with most common names, the application of the word is arbitrary, since the larvae of sawflies (suborder ...
s of the butterfly family
Lycaenidae Lycaenidae is the second-largest family (biology), family of butterflies (behind Nymphalidae, brush-footed butterflies), with over 6,000 species worldwide, whose members are also called gossamer-winged butterflies. They constitute about 30% of ...
(e.g., blues, coppers, or hairstreaks) are herded by the ants, led to feeding areas in the daytime, and brought inside the ants' nest at night. The caterpillars have a gland which secretes honeydew when the ants massage them. The chemicals in the secretions of ''Narathura japonica'' alter the behavior of attendant ''Pristomyrmex punctatus'' workers, making them less aggressive and stationary. The relationship, formerly characterized as "mutualistic", is now considered as possibly a case of the ants being parasitically manipulated by the caterpillars. Some caterpillars produce vibrations and sounds that are perceived by the ants. A similar adaptation can be seen in
Grizzled skipper ''Pyrgus'' is a genus in the skippers butterfly family, Hesperiidae, known as the grizzled skippers. The name "checkered" or "chequered skipper" may also be applied to some species, but also refers to species in the genera '' Burnsius'' and '' C ...
butterflies that emit vibrations by expanding their wings in order to communicate with ants, which are natural predators of these butterflies. Other caterpillars have evolved from ant-loving to ant-eating: these myrmecophagous caterpillars secrete a pheromone that makes the ants act as if the caterpillar is one of their own larvae. The caterpillar is then taken into the ant nest where it feeds on the ant larvae. A number of specialized bacteria have been found as
endosymbiont An endosymbiont or endobiont is an organism that lives within the body or cells of another organism. Typically the two organisms are in a mutualism (biology), mutualistic relationship. Examples are nitrogen-fixing bacteria (called rhizobia), whi ...
s in ant guts. Some of the dominant bacteria belong to the order
Hyphomicrobiales The Hyphomicrobiale' (synonym Rhizobiales) are an order of Gram-negative Alphaproteobacteria. The rhizobia, which fix nitrogen and are symbiotic with plant roots, appear in several different families. The four families '' Nitrobacteraceae'', ' ...
whose members are known for being
nitrogen-fixing Nitrogen fixation is a chemical process by which molecular dinitrogen () is converted into ammonia (). It occurs both biologically and abiological nitrogen fixation, abiologically in chemical industry, chemical industries. Biological nitrogen ...
symbiont Symbiosis (Ancient Greek : living with, companionship < : together; and ''bíōsis'': living) is any type of a close and long-term biological interaction, between two organisms of different species. The two organisms, termed symbionts, can fo ...
s in
legume Legumes are plants in the pea family Fabaceae (or Leguminosae), or the fruit or seeds of such plants. When used as a dry grain for human consumption, the seeds are also called pulses. Legumes are grown agriculturally, primarily for human consum ...
s but the species found in ant lack the ability to fix nitrogen.
Fungus-growing ants Fungus-growing ants (tribe Attini) comprise all the known fungus-growing ant species participating in ant–fungus mutualism. They are known for cutting grasses and leaves, carrying them to their colonies' nests, and using them to grow fungus on ...
that make up the tribe Attini, including
leafcutter ant Leafcutter ants are fungus-growing ants that share the behaviour of cutting leaves which they carry back to their nests to farm fungus. Next to humans, leafcutter ants form some of the largest and most complex animal societies on Earth. In a few ...
s, cultivate certain species of fungus in the genera '' Leucoagaricus'' or ''
Leucocoprinus ''Leucocoprinus'' is a genus of fungi in the family Agaricaceae. Its best-known member is the distinctive yellow mushroom ''Leucocoprinus birnbaumii'', which is found in plant pots and greenhouses worldwide. The type species is ''Leucocoprinus ce ...
'' of the family
Agaricaceae The Agaricaceae are a family of basidiomycete fungi and include the genus ''Agaricus'', as well as basidiomycetes previously classified in the families Tulostomataceae, Lepiotaceae, and Lycoperdaceae. Taxonomy The family Agaricaceae was publishe ...
. In this ant-fungus mutualism, both species depend on each other for survival. The ant ''
Allomerus decemarticulatus ''Allomerus decemarticulatus'' is an Amazonian ant species found in the tropics of South America. This species is most notable for the workers’ complex and extreme predatory behavior, which involves a symbiosis with both a plant and fungal sp ...
'' has evolved a three-way association with the host plant, ''
Hirtella physophora ''Hirtella physophora'' is a species of plant in the family Chrysobalanaceae. It forms an association with the ant species, ''Allomerus decemarticulatus ''Allomerus decemarticulatus'' is an Amazonian ant species found in the tropics of South ...
'' (
Chrysobalanaceae Chrysobalanaceae is a family of flowering plants, consisting of trees and shrubs in 27 genera and about 700 species of pantropical distribution with a centre of diversity in the Amazon. Some of the species contain silica in their bodies for rig ...
), and a sticky fungus which is used to trap their insect prey. Lemon ants make
devil's garden In myrmecology and forest ecology, a devil's garden (Kichwa language, Kichwa: ''Supay chakra''Frederickson, M. E., & Gordon, D. (2007). The devil to pay: the cost of mutualism with ''Myrmelachista schumanni'' ants in 'devil's gardens' is increased ...
s by killing surrounding plants with their stings and leaving a pure patch of lemon ant trees, (''
Duroia hirsuta ''Duroia hirsuta'' is a myrmecophyte tree species from the Amazon Forest. It is one of some 37 species of '' Duroia,'' which are shrubs or canopy trees in the family Rubiaceae, favoring ants ( myrmecophilous), and occurring in Central America as f ...
''). This modification of the forest provides the ants with more nesting sites inside the stems of the ''
Duroia ''Duroia'' is a genus of flowering plants in the family Rubiaceae. The genus is found from Costa Rica to tropical South America. Ecology A number of ''Duroia'' species, and possibly all, are capable of biochemical interactions inhibiting the gr ...
'' trees. Although some ants obtain nectar from flowers, pollination by ants is somewhat rare, one example being of the pollination of the orchid ''
Leporella fimbriata ''Leporella fimbriata'', commonly known as ''hare orchid'' or ''fringed hare orchid'', is the only species in the flowering plant genus ''Leporella'' in the orchid family, Orchidaceae and is endemic to the southern Australia mainland. It is rela ...
'' which induces male ''Myrmecia urens'' to pseudocopulate with the flowers, transferring pollen in the process. One theory that has been proposed for the rarity of pollination is that the secretions of the metapleural gland inactivate and reduce the viability of pollen. Some plants, mostly angiosperms but also some ferns, have special nectar exuding structures,
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, ...
, that provide food for ants, which in turn protect the plant from more damaging
herbivorous A herbivore is an animal anatomically and physiologically evolved to feed on plants, especially upon vascular tissues such as foliage, fruits or seeds, as the main component of its diet. These more broadly also encompass animals that eat n ...
insects. Species such as the bullhorn acacia (''
Acacia cornigera ''Vachellia cornigera'', commonly known as bullhorn acacia (family (biology), family Fabaceae), is a swollen-thorn tree and myrmecophyte native to Mexico and Central America. The common name of "bullhorn" refers to the enlarged, hollowed-out, swo ...
'') in
Central America Central America is a subregion of North America. Its political boundaries are defined as bordering Mexico to the north, Colombia to the southeast, the Caribbean to the east, and the Pacific Ocean to the southwest. Central America is usually ...
have hollow thorns that house colonies of stinging ants (''
Pseudomyrmex ferruginea The acacia ant (''Pseudomyrmex ferruginea'') is a species of ant of the genus ''Pseudomyrmex''. These arboreal, wasp-like ants have an orange-brown body around 3 mm in length and very large eyes. The acacia ant is best known and named for ...
'') who defend the tree against insects, browsing mammals, and
epiphytic An epiphyte is a plant or plant-like organism that grows on the surface of another plant and derives its moisture and nutrients from the air, rain, water (in marine environments) or from debris accumulating around it. The plants on which epiphyt ...
vines.
Isotopic labelling Isotopic labeling (or isotopic labelling) is a technique used to track the passage of an isotope (an atom with a detectable variation in neutron count) through chemical reaction, metabolic pathway, or a biological cell. The reactant is 'labeled' ...
studies suggest that plants also obtain nitrogen from the ants. In return, the ants obtain food from protein- and lipid-rich
Beltian bodies A Beltian body is a detachable tip found on the pinnules of some species of ''Acacia'' and closely related genera. Beltian bodies, named after Thomas Belt, are rich in lipids, sugars and proteins and often red in colour. They are believed to have ...
. In
Fiji Fiji, officially the Republic of Fiji, is an island country in Melanesia, part of Oceania in the South Pacific Ocean. It lies about north-northeast of New Zealand. Fiji consists of an archipelago of more than 330 islands—of which about ...
'' Philidris nagasau'' (Dolichoderinae) are known to selectively grow species of epiphytic ''
Squamellaria ''Squamellaria'' is a genus of myrmecophytic flowering plants in the family Rubiaceae. It includes eight species native to the Solomon Islands, Vanuatu, and Fiji. It is one of five ant-plant genera in the family Rubiaceae, the others being '' ...
'' (Rubiaceae) which produce large domatia inside which the ant colonies nest. The ants plant the seeds and the domatia of young seedling are immediately occupied and the ant faeces in them contribute to rapid growth. Similar dispersal associations are found with other dolichoderines in the region as well. Another example of this type of
ectosymbiosis Ectosymbiosis is a form of symbiotic behavior in which an organism lives on the body surface of another organism (the host), including internal surfaces such as the lining of the digestive tube and the ducts of glands. The ectosymbiotic species, ...
comes from the ''
Macaranga ''Macaranga'' is a large genus of Old World tropical trees of the family Euphorbiaceae and the only genus in the subtribe Macaranginae (tribe Acalypheae). Native to Africa, Australasia, Asia and various islands of the Indian and Pacific Oc ...
'' tree, which has stems adapted to house colonies of ''
Crematogaster ''Crematogaster'' is an ecologically diverse genus of ants found worldwide, which are characterised by a distinctive heart-shaped gaster (insect anatomy), gaster (abdomen), which gives them one of their common names, the Saint Valentine ant. Mem ...
'' ants. Many plant species have seeds that are adapted for dispersal by ants.
Seed dispersal In spermatophyte plants, seed dispersal is the movement, spread or transport of seeds away from the parent plant. Plants have limited mobility and rely upon a variety of dispersal vectors to transport their seeds, including both abiotic vectors, ...
by ants or
myrmecochory Myrmecochory ( (sometimes myrmechory); from ("ant") and ''khoreíā'' ("circular dance") is seed dispersal by ants, an ecologically significant Myrmecophily, ant–plant Biological interaction, interaction with worldwide distribution. Most ...
is widespread, and new estimates suggest that nearly 9% of all plant species may have such ant associations. Often, seed-dispersing ants perform directed dispersal, depositing the seeds in locations that increase the likelihood of seed survival to reproduction. Some plants in arid, fire-prone systems are particularly dependent on ants for their survival and dispersal as the seeds are transported to safety below the ground. Many ant-dispersed seeds have special external structures,
elaiosome Elaiosomes ( ''élaion'' "oil" + ''sóma'' "body") are fleshy structures that are attached to the seeds of many plant species. The elaiosome is rich in lipids and proteins, and may be variously shaped. Many plants have elaiosomes that attract ...
s, that are sought after by ants as food. Ants can substantially alter rate of decomposition and nutrient cycling in their nest. By myrmecochory and modification of soil conditions they substantially alter vegetation and nutrient cycling in surrounding ecosystem. A
convergence Convergence may refer to: Arts and media Literature *''Convergence'' (book series), edited by Ruth Nanda Anshen *Convergence (comics), "Convergence" (comics), two separate story lines published by DC Comics: **A four-part crossover storyline that ...
, possibly a form of
mimicry In evolutionary biology, mimicry is an evolved resemblance between an organism and another object, often an organism of another species. Mimicry may evolve between different species, or between individuals of the same species. In the simples ...
, is seen in the eggs of
stick insect The Phasmatodea (also known as Phasmida or Phasmatoptera) are an order of insects whose members are variously known as stick insects, stick bugs, walkingsticks, stick animals, or bug sticks. They are also occasionally referred to as Devil's da ...
s. They have an edible elaiosome-like structure and are taken into the ant nest where the young hatch. Most ants are predatory and some prey on and obtain food from other social insects including other ants. Some species specialise in preying on termites (''
Megaponera ''Megaponera analis'' is the sole species of the genus ''Megaponera''. They are a strictly termite-eating (termitophagous) Ponerinae, ponerine ant species widely distributed in Sub-Saharan Africa and most commonly known for their column-like raid ...
'' and ''Termitopone'') while a few Cerapachyinae prey on other ants. Some termites, including ''
Nasutitermes corniger ''Nasutitermes corniger'' is a species of arboreal termite that is endemic to the neotropics. It is very closely related to '' Nasutitermes ephratae''. The species has been studied relatively intensively, particularly on Barro Colorado Island, P ...
'', form associations with certain ant species to keep away predatory ant species. The tropical wasp ''
Mischocyttarus drewseni ''Mischocyttarus drewseni'', which is sometimes spelled "drewsenii", is a social wasp in the family Vespidae. It is commonly found in Neotropical regions of South America, including Brazil, Peru, Venezuela, Colombia and Paraguay. This mid-sized ...
'' coats the pedicel of its nest with an ant-repellent chemical. It is suggested that many tropical wasps may build their nests in trees and cover them to protect themselves from ants. Other wasps, such as '' A. multipicta'', defend against ants by blasting them off the nest with bursts of wing buzzing. Stingless bees (''
Trigona ''Trigona'' is one of the largest genera of stingless bees, comprising about 32 species, exclusively occurring in the New World, and formerly including many more subgenera than the present assemblage; many of these former subgenera have been ele ...
'' and ''
Melipona ''Melipona'' is a genus of stingless bees, widespread in warm areas of the Neotropics, from Sinaloa and Tamaulipas (México) to Tucumán Province, Tucumán and Misiones Province, Misiones (Argentina). About 70 species are known.Grüter, C. 2020. ...
'') use chemical defences against ants. Flies in the Old World genus ''
Bengalia ''Bengalia'' is a genus of blow flies in the family Calliphoridae with one authority considering the genus to belong to a separate family Bengaliidae.Lehrer, A.Z., 2003, Bengaliidae n. fam. Une nouvelle famille de Diptera Cyclorrhapha. Entom. Croa ...
'' (
Calliphoridae The Calliphoridae (commonly known as blowflies, blow flies, blow-flies, carrion flies, bluebottles, or greenbottles) are a family of insects in the order Diptera, with almost 1,900 known species. The maggot larvae, often used as fishing bait, ...
)
prey 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 feeding behaviours that includes parasitism and micropredation (which usually do not ki ...
on ants and are
kleptoparasite Kleptoparasitism (originally spelt clepto-parasitism, meaning "parasitism by theft") is a form of feeding in which one animal deliberately takes food from another. The strategy is Evolutionarily stable strategy, evolutionarily stable when stealin ...
s, snatching prey or brood from the mandibles of adult ants. Wingless and legless females of the
Malaysia Malaysia is a country in Southeast Asia. Featuring the Tanjung Piai, southernmost point of continental Eurasia, it is a federation, federal constitutional monarchy consisting of States and federal territories of Malaysia, 13 states and thre ...
n phorid fly (''Vestigipoda myrmolarvoidea'') live in the nests of ants of the genus '' Aenictus'' and are cared for by the ants. Fungi in the genera ''
Cordyceps ''Cordyceps'' is a genus of ascomycete fungi (sac fungi) that includes over 260 species worldwide, many of which are parasitic. Diverse variants of cordyceps have had more than 1,500 years of use in Chinese medicine. Most ''Cordyceps'' specie ...
'' and '' Ophiocordyceps'' infect ants. Ants react to their infection by climbing up plants and sinking their mandibles into plant tissue. The fungus kills the ants, grows on their remains, and produces a
fruiting body The sporocarp (also known as fruiting body, fruit body or fruitbody) of fungi is a multicellular structure on which spore-producing structures, such as basidia or asci, are borne. The fruitbody is part of the sexual phase of a fungal life cyc ...
. It appears that the fungus alters the behaviour of the ant to help disperse its spores in a microhabitat that best suits the fungus.
Strepsiptera The Strepsiptera () are an order of insects with eleven extant families that include about 600 described species. They are endoparasites of other insects, such as bees, wasps, leafhoppers, Zygentoma, silverfish, and cockroaches. Females of most s ...
n parasites also manipulate their ant host to climb grass stems, to help the parasite find mates. A
nematode The nematodes ( or ; ; ), roundworms or eelworms constitute the phylum Nematoda. Species in the phylum inhabit a broad range of environments. Most species are free-living, feeding on microorganisms, but many are parasitic. Parasitic worms (h ...
(''Myrmeconema neotropicum'') that infects canopy ants (''
Cephalotes atratus ''Cephalotes'' is a genus (biology), genus of arboreal, tree-dwelling ant species from the Americas, commonly known as turtle ants. All appear to be gliding ants, with the ability to "parachute" and steer their fall so as to land back on the tre ...
'') causes the black-coloured gasters of workers to turn red. The parasite also alters the behaviour of the ant, causing them to carry their gasters high. The conspicuous red gasters are mistaken by birds for ripe fruits, such as ''Hyeronima alchorneoides'', and eaten. The droppings of the bird are collected by other ants and fed to their young, leading to further spread of the nematode. A study of '' Temnothorax nylanderi'' colonies in Germany found that workers parasitized by the tapeworm '' Anomotaenia brevis'' (ants are intermediate hosts, the definitive hosts are
woodpecker Woodpeckers are part of the bird family (biology), family Picidae, which also includes the piculets, wrynecks and sapsuckers. Members of this family are found worldwide, except for Australia, New Guinea, New Zealand, Madagascar and the extreme ...
s) lived much longer than unparasitized workers and had a reduced mortality rate, comparable to that of the queens of the same species, which live for as long as two decades. South American
poison dart frog Poison dart frog (also known as dart-poison frog, poison frog or formerly known as poison arrow frog) is the common name of a group of frogs in the family Dendrobatidae which are native to tropical Central and South America. These species are ...
s in the genus '' Dendrobates'' feed mainly on ants, and the toxins in the skin of some species come from the ants. Formicine ants in the genera '' Brachymyrmex'' and '' Paratrechina'' have been found to contain pumiliotoxin found in ''
Dendrobates pumilio The strawberry poison frog, strawberry poison dart frog or blue jeans poison frog (''Oophaga pumilio'', formerly ''Dendrobates pumilio'') is a species of small poison dart frog found in Central America. It is common throughout its range, which ex ...
''. The West African frog ''
Phrynomantis microps ''Phrynomantis microps'' is an African frog with many interesting adaptations to the savannah. These frogs are also called the Accra snake-necked frog. They are found across Western and Central Africa. One of its distinguishing features is the pe ...
'' is able to move within the nests of '' Paltothyreus tarsatus'' ants, producing peptides on its skin that prevent the ants from stinging them.
Army ant The name army ant (or legionary ant or ''marabunta'') is applied to over 200 ant species in different lineages. Because of their aggressive predatory foraging groups, known as "raids", a huge number of ants forage simultaneously over a limited ...
s which is the toxin found in forage in a wide roving column, attacking any animals in that path that are unable to escape. In Central and South America, ''
Eciton burchellii ''Eciton burchellii'' is a species of New World army ant in the genus '' Eciton''. This species performs expansive, organized swarm raids that give it the informal name, ''Eciton'' army ant. This species displays a high degree of worker polymor ...
'' is the swarming ant most commonly attended by " ant-following" birds such as
antbird The antbirds are a large passerine bird family, Thamnophilidae, found across subtropical and tropical Central and South America, from Mexico to Argentina. There are more than 230 species, known variously as antshrikes, antwrens, antvireos, fire ...
s and
woodcreeper The woodcreepers (Dendrocolaptinae) comprise a subfamily of suboscine passerine birds endemic to the Neotropics. They have traditionally been considered a distinct family Dendrocolaptidae, but most authorities now place them as a subfamily of th ...
s. This behaviour was once considered mutualistic, but later studies found the birds to be
parasitic Parasitism is a close relationship between species, where one organism, the parasite, lives (at least some of the time) on or inside another organism, the host, causing it some harm, and is adapted structurally to this way of life. The ent ...
. Direct
kleptoparasitism Kleptoparasitism (originally spelt clepto-parasitism, meaning "parasitism by theft") is a form of feeding in which one animal deliberately takes food from another. The strategy is evolutionarily stable when stealing is less costly than direct f ...
(birds stealing food from the ants' grasp) is rare and has been noted in Inca doves which pick seeds at nest entrances as they are being transported by species of ''
Pogonomyrmex ''Pogonomyrmex'' is a genus of harvester ants, occurring primarily in the deserts of North, Central, and South America, with three endemic species from Haiti.Cole, A.C. 1968. Pogonomyrmex Harvester Ants: a study of the genus in North America. Un ...
''. Birds that follow ants eat many prey insects and thus decrease the foraging success of ants. Birds indulge in a peculiar behaviour called
anting Anting () is a town in Jiading District, Shanghai, bordering Kunshan, Jiangsu to the west. It has 96,000 inhabitants and, after the July 2009 merger of Huangdu (), an area of .
that, as yet, is not fully understood. Here birds rest on ant nests, or pick and drop ants onto their wings and feathers; this may be a means to remove
ectoparasite 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 ...
s from the birds.
Anteater Anteaters are the four extant mammal species in the suborder Vermilingua (meaning "worm tongue"), commonly known for eating ants and termites. The individual species have other names in English and other languages. Together with sloths, they ar ...
s,
aardvark Aardvarks ( ; ''Orycteropus afer'') are medium-sized, burrowing, nocturnal mammals native to Africa. Aardvarks are the only living species of the family Orycteropodidae and the order Tubulidentata. They have a long proboscis, similar to a pi ...
s,
pangolin Pangolins, sometimes known as scaly anteaters, are mammals of the order Pholidota (). The one extant family, the Manidae, has three genera: '' Manis'', '' Phataginus'', and '' Smutsia''. ''Manis'' comprises four species found in Asia, while ' ...
s,
echidna Echidnas (), sometimes known as spiny anteaters, are quill-covered monotremes (egg-laying mammals) belonging to the Family (biology), family Tachyglossidae , living in Australia and New Guinea. The four Extant taxon, extant species of echidnas ...
s and
numbat The numbat (''Myrmecobius fasciatus''), also known as the noombat or walpurti, is an insectivorous marsupial. It is diurnal and its diet consists almost exclusively of termites. The species was once widespread across southern Australia, but i ...
s have special
adaptation In biology, adaptation has three related meanings. Firstly, it is the dynamic evolutionary process of natural selection that fits organisms to their environment, enhancing their evolutionary fitness. Secondly, it is a state reached by the p ...
s for living on a diet of ants. These adaptations include long, sticky tongues to capture ants and strong claws to break into ant nests.
Brown bear The brown bear (''Ursus arctos'') is a large bear native to Eurasia and North America. Of the land carnivorans, it is rivaled in size only by its closest relative, the polar bear, which is much less variable in size and slightly bigger on av ...
s (''Ursus arctos'') have been found to feed on ants. About 12%, 16%, and 4% of their faecal volume in spring, summer and autumn, respectively, is composed of ants.


Relationship with humans

Ants perform many ecological roles that are beneficial to humans, including the suppression of pest populations and aeration of the
soil Soil, also commonly referred to as earth, is a mixture of organic matter, minerals, gases, water, and organisms that together support the life of plants and soil organisms. Some scientific definitions distinguish dirt from ''soil'' by re ...
. The use of
weaver ant Weaver ants or green ants are eusocial insects of the Hymenoptera family Formicidae belonging to the tribe Oecophyllini. Weaver ants live in trees (they are obligately arboreal) and are known for their unique nest building behaviour where wor ...
s in
citrus ''Citrus'' is a genus of flowering trees and shrubs in the family Rutaceae. Plants in the genus produce citrus fruits, including important crops such as oranges, mandarins, lemons, grapefruits, pomelos, and limes. ''Citrus'' is nativ ...
cultivation in southern China is considered one of the oldest known applications of
biological control Biological control or biocontrol is a method of controlling pests, whether pest animals such as insects and mites, weeds, or pathogens affecting animals or plants by using other organisms. It relies on predation, parasitism, herbivory, or o ...
.Hölldobler & Wilson (1990), pp. 619–629 On the other hand, ants may become nuisances when they invade buildings or cause economic losses. In some parts of the world (mainly Africa and South America), large ants, especially
army ant The name army ant (or legionary ant or ''marabunta'') is applied to over 200 ant species in different lineages. Because of their aggressive predatory foraging groups, known as "raids", a huge number of ants forage simultaneously over a limited ...
s, are used as
surgical suture A surgical suture, also known as a stitch or stitches, is a medical device used to hold Tissue (biology), body tissues together and approximate wound edges after an injury or surgery. Application generally involves using a Sewing needle, needle w ...
s. The wound is pressed together and ants are applied along it. The ant seizes the edges of the wound in its mandibles and locks in place. The body is then cut off and the head and mandibles remain in place to close the wound. The large heads of the dinergates (soldiers) of the leafcutting ant '' Atta cephalotes'' are also used by native surgeons in closing wounds. Some ants have toxic venom and are of medical importance. The species include ''
Paraponera clavata ''Paraponera clavata'', commonly known as the bullet ant, is a species of ant known for its extremely painful Insect bites and stings, sting. It inhabits humid lowland rainforests in Central America, Central and South America. Etymology The Spe ...
'' (tocandira) and '' Dinoponera'' spp. (false tocandiras) of South America and the ''Myrmecia'' ants of Australia. In
South Africa South Africa, officially the Republic of South Africa (RSA), is the Southern Africa, southernmost country in Africa. Its Provinces of South Africa, nine provinces are bounded to the south by of coastline that stretches along the Atlantic O ...
, ants are used to help harvest the seeds of
rooibos Rooibos ( ; , ), or , is a broom (shrub), broom-like member of the plant family Fabaceae that grows in South Africa's Fynbos biome. The leaves are used to make a caffeine-free herbal tea, herbal infusion that has been popular in Southern Afri ...
(''Aspalathus linearis''), a plant used to make a herbal tea. The plant disperses its seeds widely, making manual collection difficult. Black ants collect and store these and other seeds in their nest, where humans can gather them ''en masse''. Up to half a pound (200 g) of seeds may be collected from one ant-heap. Although most ants survive attempts by humans to eradicate them, a few are highly endangered. These tend to be island species that have evolved specialized traits and risk being displaced by introduced ant species. Examples include the critically endangered Sri Lankan relict ant (''Aneuretus simoni'') and '' Adetomyrma venatrix'' of Madagascar.


As food

Ants and their larvae are eaten in different parts of the world. The eggs of two species of ants are used in Mexican '' escamoles''. They are considered a form of insect
caviar Caviar or caviare is a food consisting of salt-cured roe of the family Acipenseridae. Caviar is considered a delicacy and is eaten as a garnish or spread. Traditionally, the term caviar refers only to roe from wild sturgeon in the Caspi ...
and can sell for as much as US$50 per kg going up to US$200 per kg (as of 2006) because they are seasonal and hard to find. In the
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 ...
n department of Santander, ''hormigas culonas'' (roughly interpreted as "large-bottomed ants") ''
Atta laevigata ''Atta laevigata'' ( Smith, 1858) is one of about a dozen species of leafcutter ants in the genus '' Atta'', found from Venezuela and south to Paraguay. This species is one of the largest leafcutter species, and can be recognized by the smooth ...
'' are toasted alive and eaten. In areas of
India India, officially the Republic of India, is a country in South Asia. It is the List of countries and dependencies by area, seventh-largest country by area; the List of countries by population (United Nations), most populous country since ...
, and throughout
Burma Myanmar, officially the Republic of the Union of Myanmar; and also referred to as Burma (the official English name until 1989), is a country in northwest Southeast Asia. It is the largest country by area in Mainland Southeast Asia and ha ...
and
Thailand Thailand, officially the Kingdom of Thailand and historically known as Siam (the official name until 1939), is a country in Southeast Asia on the Mainland Southeast Asia, Indochinese Peninsula. With a population of almost 66 million, it spa ...
, a paste of the green weaver ant (''
Oecophylla smaragdina ''Oecophylla smaragdina'' (common names include Asian weaver ant, weaver ant, green ant, green tree ant, and orange gaster) is a species of tree, arboreal ant found in tropical Asia and Australia. These ants form colonies with multiple nests in ...
'') is served as a condiment with curry.
Weaver ant Weaver ants or green ants are eusocial insects of the Hymenoptera family Formicidae belonging to the tribe Oecophyllini. Weaver ants live in trees (they are obligately arboreal) and are known for their unique nest building behaviour where wor ...
eggs and larvae, as well as the ants, may be used in a
Thai salad Salads that are internationally known as Thai salads with a few exceptions fall into four main preparation methods. In Thai cuisine these are called ''yam, tam, lap'' and ''phla''. A few other dishes can also be regarded as being a salad. Overvi ...
, ''yam'' (), in a dish called ''yam khai mot daeng'' () or red ant egg salad, a dish that comes from the Issan or north-eastern region of Thailand. Saville-Kent, in the ''Naturalist in Australia'' wrote "Beauty, in the case of the green ant, is more than skin-deep. Their attractive, almost sweetmeat-like translucency possibly invited the first essays at their consumption by the human species". Mashed up in water, after the manner of lemon squash, "these ants form a pleasant acid drink which is held in high favor by the natives of North Queensland, and is even appreciated by many European palates". Ants or their pupae are used as starters for yogurt making in parts of Bulgaria and Turkey. In his ''First Summer in the Sierra'',
John Muir John Muir ( ; April 21, 1838December 24, 1914), also known as "John of the Mountains" and "Father of the national park, National Parks", was a Scottish-born American naturalist, author, environmental philosopher, botanist, zoologist, glaciologi ...
notes that the Digger Indians of
California California () is a U.S. state, state in the Western United States that lies on the West Coast of the United States, Pacific Coast. It borders Oregon to the north, Nevada and Arizona to the east, and shares Mexico–United States border, an ...
ate the tickling, acid gasters of the large jet-black
carpenter ant Carpenter ants (''Camponotus'' spp.) are a genus of large ants (workers ) indigenous to many parts of the world. True carpenter ants build nests inside wood, consisting of galleries chewed out with their mandibles or jaws, preferably in dead, ...
s. The Mexican Indians eat the repletes, or living honey-pots, of the honey ant ('' Myrmecocystus'').


As pests

Some ant species are considered as pests, primarily those that occur in human habitations, where their presence is often problematic. For example, the presence of ants would be undesirable in sterile places such as hospitals or kitchens. Some species or genera commonly categorized as pests include the
Argentine ant The Argentine ant (''Linepithema humile'', formerly ''Iridomyrmex humilis'') is an ant native to northern Argentina, Uruguay, Paraguay, Bolivia and southern Brazil. This invasive species was inadvertently introduced by humans on a global scale a ...
, immigrant pavement ant, yellow crazy ant, banded sugar ant,
pharaoh ant The pharaoh ant (''Monomorium pharaonis'') is a small (2 mm) yellow or light brown, almost transparent ant notorious for being a major indoor nuisance pest, especially in hospitals. A cryptogenic species, it has now been introduced to vir ...
, red wood ant, black carpenter ant, odorous house ant,
red imported fire ant ''Solenopsis invicta'', the fire ant, or red imported fire ant (RIFA), is a species of ant native to South America. A member of the genus ''Fire ant, Solenopsis'' in the subfamily Myrmicinae, it was Species description, described by Swiss ento ...
, and European fire ant. Some ants will raid stored food, some will seek water sources, others may damage indoor structures, some may damage agricultural crops directly or by aiding sucking pests. Some will sting or bite. The adaptive nature of ant colonies make it nearly impossible to eliminate entire colonies and most pest management practices aim to control local populations and tend to be temporary solutions. Ant populations are managed by a combination of approaches that make use of chemical, biological, and physical methods. Chemical methods include the use of insecticidal bait which is gathered by ants as food and brought back to the nest where the poison is inadvertently spread to other colony members through
trophallaxis Trophallaxis () is the transfer of food or other fluids among members of a community through mouth-to-mouth (stomodeum, stomodeal) or anus-to-mouth (proctodeum, proctodeal) feeding. Along with nutrients, trophallaxis can involve the transfer of m ...
. Management is based on the species and techniques may vary according to the location and circumstance.


In science and technology

Observed by humans since the dawn of history, the behaviour of ants has been documented and the subject of early writings and fables passed from one century to another. Those using scientific methods,
myrmecologists Myrmecology (; from Greek: μύρμηξ, ''myrmex'', "ant" and λόγος, ''logos'', "study") is a branch of entomology focusing on the study of ants. Ants continue to be a model of choice for the study of questions on the evolution of social ...
, study ants in the laboratory and in their natural conditions. Their complex and variable social structures have made ants ideal
model organism A model organism is a non-human species that is extensively studied to understand particular biological phenomena, with the expectation that discoveries made in the model organism will provide insight into the workings of other organisms. Mo ...
s.
Ultraviolet vision Tetrachromacy (from Ancient Greek ''tetra'', meaning "four" and ''chroma'', meaning "color") is the condition of possessing four independent channels for conveying color information, or possessing four types of cone cell in the eye. Organisms w ...
was first discovered in ants by
Sir John Lubbock John Lubbock, 1st Baron Avebury (30 April 1834 – 28 May 1913), known as Sir John Lubbock, 4th Baronet, from 1865 until 1900, was an English banker, Liberal politician, philanthropist, scientist and polymath. Lubbock worked in his family comp ...
in 1881. Studies on ants have tested hypotheses in
ecology Ecology () is the natural science of the relationships among living organisms and their Natural environment, environment. Ecology considers organisms at the individual, population, community (ecology), community, ecosystem, and biosphere lev ...
and
sociobiology Sociobiology is a field of biology that aims to explain social behavior in terms of evolution. It draws from disciplines including psychology, ethology, anthropology, evolution, zoology, archaeology, and population genetics. Within the study of ...
, and have been particularly important in examining the predictions of theories of
kin selection Kin selection is a process whereby natural selection favours a trait due to its positive effects on the reproductive success of an organism's relatives, even when at a cost to the organism's own survival and reproduction. Kin selection can lead ...
and evolutionarily stable strategies. Ant colonies may be studied by rearing or temporarily maintaining them in '' formicaria'', specially constructed glass framed enclosures. Individuals may be tracked for study by marking them with dots of colours. The successful techniques used by ant colonies have been studied in computer science and robotics to produce distributed and fault-tolerant systems for solving problems, for example
Ant colony optimization In computer science and operations research, the ant colony optimization algorithm (ACO) is a probabilistic technique for solving computational problems that can be reduced to finding good paths through graphs. Artificial ants represent multi ...
and Ant robotics. This area of
biomimetics Biomimetics or biomimicry is the emulation of the models, systems, and elements of nature for the purpose of solving complex human problems. The terms "biomimetics" and "biomimicry" are derived from (''bios''), life, and μίμησις (''mimes ...
has led to studies of ant locomotion, search engines that make use of "foraging trails", fault-tolerant storage, and networking algorithms.


As pets

From the late 1950s through the late 1970s, ant farms were popular educational children's toys in the United States. Some later commercial versions use transparent gel instead of soil, allowing greater visibility at the cost of stressing the ants with unnatural light.


In culture

Anthropomorphised Anthropomorphism is the attribution of human traits, emotions, or intentions to non-human entities. It is considered to be an innate tendency of human psychology. Personification is the related attribution of human form and characteristics to ...
ants have often been used in fables, children's stories, and religious texts to represent industriousness and cooperative effort, such as in the
Aesop Aesop ( ; , ; c. 620–564 BCE; formerly rendered as Æsop) was a Greeks, Greek wikt:fabulist, fabulist and Oral storytelling, storyteller credited with a number of fables now collectively known as ''Aesop's Fables''. Although his existence re ...
fable
The Ant and the Grasshopper The Ant and the Grasshopper, alternatively titled The Grasshopper and the Ant (or Ants), is one of Aesop's Fables, numbered 373 in the Perry Index. The fable describes how a hungry grasshopper begs for food from an ant when winter comes and is ...
. In the
Quran The Quran, also Romanization, romanized Qur'an or Koran, is the central religious text of Islam, believed by Muslims to be a Waḥy, revelation directly from God in Islam, God (''Allah, Allāh''). It is organized in 114 chapters (, ) which ...
, Sulayman is said to have heard and understood an ant warning other ants to return home to avoid being accidentally crushed by Sulayman and his marching army., In parts of Africa, ants are considered to be the messengers of the deities. Some Native American mythology, such as the
Hopi mythology The Hopi maintain a complex religious and mythological tradition stretching back over centuries. However, it is difficult to definitively state what all Hopis as a group believe. Like the oral traditions of many other societies, Hopi mythology i ...
, considers ants as the first animals. Ant bites are often said to have curative properties. The sting of some species of '' Pseudomyrmex'' is claimed to give fever relief. Ant bites are used in the
initiation Initiation is a rite of passage marking entrance or acceptance into a group or society. It could also be a formal admission to adulthood in a community or one of its formal components. In an extended sense, it can also signify a transformatio ...
ceremonies of some Amazon Indian cultures as a test of endurance. In
Greek mythology Greek mythology is the body of myths originally told by the Ancient Greece, ancient Greeks, and a genre of ancient Greek folklore, today absorbed alongside Roman mythology into the broader designation of classical mythology. These stories conc ...
, the goddess
Athena Athena or Athene, often given the epithet Pallas, is an ancient Greek religion, ancient Greek goddess associated with wisdom, warfare, and handicraft who was later syncretism, syncretized with the Roman goddess Minerva. Athena was regarde ...
turned the maiden Myrmex into an ant when the latter claimed to have invented the plough, when in fact it was Athena's own invention. Ant society has always fascinated humans and has been written about both humorously and seriously.
Mark Twain Samuel Langhorne Clemens (November 30, 1835 – April 21, 1910), known by the pen name Mark Twain, was an American writer, humorist, and essayist. He was praised as the "greatest humorist the United States has produced," with William Fau ...
wrote about ants in his 1880 book ''
A Tramp Abroad ''A Tramp Abroad'' is a work of travel literature, including a mixture of autobiography and fictional events, by American author Mark Twain, published in 1880. The book details a journey by the author, with his friend Harris (a character created ...
''. Some modern authors have used the example of the ants to comment on the relationship between society and the individual. Examples are
Robert Frost Robert Lee Frost (March26, 1874January29, 1963) was an American poet. Known for his realistic depictions of rural life and his command of American Colloquialism, colloquial speech, Frost frequently wrote about settings from rural life in New E ...
in his poem "Departmental" and T. H. White in his fantasy novel ''
The Once and Future King ''The Once and Future King'' is a collection of fantasy novels by T. H. White about the legend of King Arthur. It is loosely based upon the 1485 work ''Le Morte d'Arthur'' by Sir Thomas Malory. It was first published in 1958 as a collection o ...
''. The plot in French entomologist and writer Bernard Werber's '' Les Fourmis'' science-fiction trilogy is divided between the worlds of ants and humans; ants and their behaviour are described using contemporary scientific knowledge.
H. G. Wells Herbert George Wells (21 September 1866 – 13 August 1946) was an English writer, prolific in many genres. He wrote more than fifty novels and dozens of short stories. His non-fiction output included works of social commentary, politics, hist ...
wrote about intelligent ants destroying human settlements in Brazil and threatening human civilization in his 1905 science-fiction short story, '' The Empire of the Ants.'' A similar German story involving army ants, '' Leiningen Versus the Ants'', was written in 1937 and recreated in movie form as
The Naked Jungle ''The Naked Jungle'' is a 1954 American adventure film directed by Byron Haskin, and starring Charlton Heston and Eleanor Parker. Telling the story of an attack of army ants on a Brazilian cocoa plantation, it was based on the 1937 short st ...
in 1954. In more recent times, animated cartoons and 3-D animated films featuring ants have been produced including ''
Antz ''Antz'' is a 1998 American animated adventure comedy film directed by Eric Darnell and Tim Johnson from a screenplay written by Todd Alcott and the writing team of Chris and Paul Weitz. It was produced by DreamWorks Pictures, DreamWorks An ...
'', ''
A Bug's Life ''A Bug's Life'' (stylized in all lowercase) is a 1998 American animated comedy film directed by John Lasseter from a screenplay written by Andrew Stanton, Donald McEnery, and Bob Shaw, and a story conceived by Lasseter, Stanton, and Joe Ran ...
'', '' The Ant Bully'', '' The Ant and the Aardvark'', '' Ferdy the Ant'' and ''
Atom Ant Atom Ant is a cartoon ant and superhero, created by Hanna-Barbera in 1965. Atom costarred in '' The Atom Ant/Secret Squirrel Show'' (sharing top billing with Secret Squirrel) on Saturday mornings. In syndication, Atom Ant aired alongside '' The ...
.'' Renowned
myrmecologist Myrmecology (; from Greek: μύρμηξ, ''myrmex'', "ant" and λόγος, ''logos'', "study") is a branch of entomology focusing on the study of ants. Ants continue to be a model of choice for the study of questions on the evolution of social ...
E. O. Wilson Edward Osborne Wilson (June 10, 1929 – December 26, 2021) was an American biologist, naturalist, ecologist, and entomologist known for developing the field of sociobiology. Born in Alabama, Wilson found an early interest in nature and frequ ...
wrote a short story, "Trailhead" in 2010 for ''
The New Yorker ''The New Yorker'' is an American magazine featuring journalism, commentary, criticism, essays, fiction, satire, cartoons, and poetry. It was founded on February 21, 1925, by Harold Ross and his wife Jane Grant, a reporter for ''The New York T ...
'' magazine, which describes the life and death of an ant-queen and the rise and fall of her colony, from an ants' point of view. Ants also are quite popular inspiration for many
science-fiction Science fiction (often shortened to sci-fi or abbreviated SF) is a genre of speculative fiction that deals with imaginative and futuristic concepts. These concepts may include information technology and robotics, biological manipulations, sp ...
insectoids, such as the Formics of ''
Ender's Game ''Ender's Game'' is a 1985 military science fiction novel by American author Orson Scott Card. Set at an unspecified date in Earth's future, the novel presents an imperiled humankind after two conflicts with an insectoid alien species they ...
'', the Bugs of ''
Starship Troopers ''Starship Troopers'' is a military science fiction novel by American writer Robert A. Heinlein. Written in a few weeks in reaction to the US suspending nuclear tests, the story was first published as a two-part serial in ''The Magazine of ...
'', the giant ants in the films ''
Them! ''Them!'' is a 1954 black-and-white science fiction giant monster film starring James Whitmore, Edmund Gwenn, Joan Weldon, and James Arness. Produced by David Weisbart, the film was directed by Gordon Douglas, based on an original story by ...
'' and '' Empire of the Ants,''
Marvel Comics Marvel Comics is a New York City–based comic book publishing, publisher, a property of the Walt Disney Company since December 31, 2009, and a subsidiary of Disney Publishing Worldwide since March 2023. Marvel was founded in 1939 by Martin G ...
' super hero
Ant-Man Ant-Man is the name of several superheroes appearing in books published by Marvel Comics. Publication history Created by Stan Lee, Larry Lieber and Jack Kirby, his first appearance was in '' Tales to Astonish'' #27 (January 1962) as Dr. Henry ...
, and ants mutated into super-intelligence in '' Phase IV''. In computer
strategy games A strategy game or strategic game is a game in which the players' uncoerced, and often autonomous, decision-making skills have a high significance in determining the outcome. Almost all strategy games require internal decision tree-style thinki ...
, ant-based species often benefit from increased production rates due to their single-minded focus, such as the Klackons in the ''
Master of Orion ''Master of Orion'' (abbreviated as MoO) is a turn-based, 4X science fiction strategy game in which the Player character, player leads one of ten races to dominate the galaxy through a combination of diplomacy and conquest while developing techn ...
'' series of games or the ChCht in '' Deadlock II''. These characters are often credited with a hive mind, a common misconception about ant colonies. In the early 1990s, the video game '' SimAnt'', which simulated an ant colony, won the 1992 Codie award for "Best Simulation Program".


See also

* Glossary of ant terms *
International Union for the Study of Social Insects The International Union for the Study of Social Insects has at its purpose to promote and encourage the study of social insects and other social organisms in the broadest sense. Both research and the dissemination of knowledge about social insects ...
* '' Myrmecological News'' (journal) * Task allocation and partitioning in social insects


References


Cited texts

* *


Further reading

* * *


External links


AntWiki – Bringing Ants to the World
*
AntWeb from The California Academy of Sciences

Ant Species Fact Sheets
from the
National Pest Management Association The National Pest Management Association (NPMA), is a non-profit trade association founded in 1933 that aims to represent the interests of the professional pest management and pest control industries in the United States.Ant Genera of the World – distribution maps


{{Authority control Symbiosis Extant Albian first appearances Articles containing video clips Insects in culture