A phylogeny of the extant ant subfamilies.
*Cerapachyinae is paraphyletic
‡ The previous dorylomorph subfamilies were synonymized under
Dorylinae by Brady et al. in 2014
Ants are eusocial insects of the family Formicidae and, along with the
related wasps and bees, belong to the order Hymenoptera. Ants evolved
from wasp-like ancestors in the
Cretaceous period, about 140 million
years ago, and diversified after the rise of flowering plants. More
than 12,500 of an estimated total of 22,000 species have been
classified. They are easily identified by their elbowed antennae
and the distinctive node-like structure that forms their slender
Ants form colonies that range in size from a few dozen predatory
individuals living in small natural cavities to highly organised
colonies that may occupy large territories and consist of millions of
individuals. Larger 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" (aner) and
one or more fertile females called "queens" (gynes). The colonies
are described as superorganisms because the ants appear to operate as
a unified entity, collectively working together to support the
(video) Ants gathering food
Ants have colonised almost every landmass on Earth. The only places
lacking indigenous ants are
Antarctica and a few remote or
inhospitable islands. Ants thrive in most ecosystems and may form
15–25% of the terrestrial animal biomass. 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 with other species has led to
mimetic, commensal, parasitic, and mutualistic relationships.
Ant societies have division of labour, communication between
individuals, and an ability to solve complex problems. These
parallels with human societies have long been an inspiration and
subject of study. Many human cultures make use of ants in cuisine,
medication, and rituals. Some species are valued in their role as
biological pest control 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), are regarded as invasive species,
establishing themselves in areas where they have been introduced
2 Taxonomy and evolution
3 Distribution and diversity
5 Life cycle
6 Behaviour and ecology
6.4 Nest construction
6.5 Cultivation of food
6.8 Cooperation and competition
6.9 Relationships with other organisms
7 Relationship with humans
7.1 As food
7.2 As pests
7.3 In science and technology
7.4 As pets
7.5 In culture
8 See also
9.1 Cited texts
10 Further reading
11 External links
The word ant and its chiefly dialectal form emmet come from ante,
emete of Middle English, which come from ǣmette of Old English, and
these are all related to the dialectal Dutch emt and the Old High
German āmeiza, from which comes the modern German Ameise. All of
these words come from West Germanic *ēmaitijǭ, and the original
meaning of the word was "the biter" (from
Proto-Germanic *ai-, "off,
away" + *mait- "cut"). The family name Formicidae is derived
Latin formīca ("ant") from which the words in other
Romance languages, such as the Portuguese formiga, Italian formica,
Spanish hormiga, Romanian furnică, and French fourmi are derived. It
has been hypothesised that a Proto-Indo-European word *morwi- was
Latin formīca, Greek μύρμηξ
Old Church Slavonic
Old Church Slavonic mraviji,
Old Irish moirb, Old Norse
maurr, Dutch mier.
Taxonomy and evolution
Ants fossilised in Baltic amber
Phylogenetic position of the Formicidae.
The family Formicidae belongs to the order Hymenoptera, which also
includes sawflies, bees, and wasps. Ants evolved from a lineage within
the aculeate wasps, and a 2013 study suggests that they are a sister
group of the Apoidea. In 1966,
E. O. Wilson
E. O. Wilson and his colleagues
identified the fossil remains of an ant (Sphecomyrma) that lived in
Cretaceous period. The specimen, trapped in amber dating back to
around 92 million years ago, has features found in some wasps,
but not found in modern ants.
Sphecomyrma was possibly a ground
Haidomyrmex and Haidomyrmodes, related genera in
subfamily Sphecomyrminae, are reconstructed as active arboreal
predators. Older ants in the genus Sphecomyrmodes have been found
in 99 million year-old amber from Myanmar. A 2006 study
suggested that ants arose tens of millions of years earlier than
previously thought, up to 168 million years ago. After the rise of
flowering plants about 100 million years ago they diversified and
assumed ecological dominance around 60 million years
ago. Some groups, such as the
Martialinae, are suggested to have diversified from early primitive
ants that were likely to have been predators underneath the surface of
Cretaceous period, a few species of primitive ants ranged
widely on the Laurasian supercontinent (the Northern Hemisphere). They
were scarce in comparison to the populations of other insects,
representing only about 1% of the entire insect population. Ants
became dominant after adaptive radiation at the beginning of the
Paleogene period. By the
Oligocene and Miocene, ants had come to
represent 20–40% of all insects found in major fossil deposits. Of
the species that lived in the
Eocene epoch, around one in 10 genera
survive to the present. Genera surviving today comprise 56% of the
Baltic amber fossils (early Oligocene), and 92% of the
genera in Dominican amber fossils (apparently early Miocene).
Termites, although sometimes called 'white ants', are not ants. They
belong to the sub-order
Isoptera within the order Blattodea. Termites
are more closely related to cockroaches and mantids. Termites are
eusocial, but differ greatly in the genetics of reproduction. The
similarity of their social structure to that of ants is attributed to
convergent evolution. Velvet ants look like large ants, but are
wingless female wasps.
Distribution and diversity
Ants are found on all continents except Antarctica, and only a few
large islands, such as Greenland, Iceland, parts of
Polynesia and the
Hawaiian Islands lack native ant species. Ants occupy a wide
range of ecological niches 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.
Their ecological dominance is demonstrated by their biomass: ants are
estimated to contribute 15–20 % (on average and nearly 25% in
the tropics) of terrestrial animal biomass, exceeding that of the
Ants range in size from 0.75 to 52 millimetres
(0.030–2.0 in), the largest species being the fossil
Titanomyrma giganteum, the queen of which was 6 centimetres
(2.4 in) long with a wingspan of 15 centimetres
(5.9 in). Ants vary in colour; most ants are red or black,
but a few species are green and some tropical species have a metallic
lustre. More than 12,000 species are currently known (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
AntBase 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 ecosystems has made them useful as
indicator species in biodiversity studies.
Ants are distinct in their morphology from other insects in having
elbowed antennae, metapleural glands, and a strong constriction of
their second abdominal segment into a node-like petiole. The head,
mesosoma, and metasoma are the three distinct body segments. The
petiole forms a narrow waist between their mesosoma (thorax 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
Bull ant showing the powerful mandibles and the relatively large
compound eyes that provide excellent vision
Like other insects, ants have an exoskeleton, 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 vertebrates. Insects do not have lungs; oxygen and
other gases, such as carbon dioxide, pass through their exoskeleton
via tiny valves called spiracles. Insects also lack closed blood
vessels; 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 toward the head, thus driving the
circulation of the internal fluids. The nervous system consists of a
ventral nerve cord that runs the length of the body, with several
ganglia and branches along the way reaching into the extremities of
Diagram of a worker ant (Pachycondyla verenae)
An ant's head contains many sensory organs. Like most insects, ants
have compound eyes 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 (simple eyes)
on the top of the head that detect light levels and polarization.
Compared to vertebrates, most ants have poor-to-mediocre eyesight and
a few subterranean species are completely blind. However, some ants,
such as Australia's bulldog ant, have excellent vision and are capable
of discriminating the distance and size of objects moving nearly a
Two antennae ("feelers") are attached to the head; these organs detect
chemicals, air currents, and vibrations; they also are used to
transmit and receive signals through touch. The head has two strong
jaws, the mandibles, 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.
Both the legs and wings of the ant are attached to the mesosoma
("thorax"). The legs terminate in a hooked claw which allows them to
hook on and climb surfaces. Only reproductive ants, queens, and
males, have wings. Queens shed their wings after the nuptial flight,
leaving visible stubs, a distinguishing feature of queens. In a few
species, wingless queens (ergatoids) and males occur.
The metasoma (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 and
defending their nests.
Seven leafcutter ant workers of various castes (left) and two queens
In the colonies of a few ant species, there are physical
castes—workers in distinct size-classes, called minor, median, and
major ergates. Often, the larger ants have disproportionately larger
heads, and correspondingly stronger mandibles. These are known as
macrergates while smaller workers are known as micrergates.
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 ants, for
example, have a distinct bimodal size distribution. Some other
species show continuous variation in the size of workers. The smallest
and largest workers in
Pheidologeton diversus show nearly a 500-fold
difference in their dry-weights. Workers cannot mate; however,
because of the haplodiploid sex-determination system 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, 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
Myrmecocystus mexicanus. 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 can decide whether
the colony will have single or multiple queens. The Australian
jack jumper ant (Myrmecia pilosula) has only a single pair of
chromosomes (with the males having just one chromosome as they are
haploid), the lowest number known for any animal, making it an
interesting subject for studies in the genetics and developmental
biology of social insects.
Meat eater ant
Meat eater ant nest during swarming
The life of an ant starts from an egg. If the egg is fertilised, the
progeny will be female diploid; if not, it will be male haploid. Ants
develop by complete metamorphosis with the larva stages passing
through a pupal 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, a process in which an ant regurgitates
liquid food held in its crop. 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 eggs, pieces
of prey, and seeds brought by workers.
The larvae grow through a series of four or five moults 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 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, emerge
from pupae along with the usually winged breeding females. Some
species, such as army ants, 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.
An explanation for the sequence is suggested by the high casualties
involved in foraging, making it an acceptable risk only for ants who
are older and are likely to die soon of natural causes.
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, but, in cooler regions,
they survive the winter in a state of dormancy known as hibernation.
The forms of inactivity are varied and some temperate species have
larvae going into the inactive state (diapause), while in others, the
adults alone pass the winter in a state of reduced activity.
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.
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. When the drone is initially attacked by the workers, it
releases a mating pheromone. 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 and then marking them with a pheromone. The marked male is
interpreted as an invader by worker ants and is killed.
Fertilised meat-eater ant queen beginning to dig a new colony
Most ants are univoltine, producing a new generation each year.
During the species-specific breeding period, new reproductives,
females, and winged males leave the colony in what is called a nuptial
flight. 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 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 in their
Mated females then seek a suitable place to begin a colony. There,
they break off their wings 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 are known as nanitics, and 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 in honeybees.
Behaviour and ecology
Camponotus sericeus workers communicating through touch and
Ants communicate with each other using pheromones, sounds, and
touch. The use of pheromones as chemical signals is more developed
in ants, such as the red harvester ant, than in other hymenopteran
groups. Like other insects, ants perceive smells with their long,
thin, and mobile antennae. The paired antennae provide information
about the direction and intensity of scents. 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 pheromones" to confuse enemy ants and make them fight
among themselves. Pheromones are produced by a wide range of
structures including Dufour's glands, poison glands and glands on the
hindgut, pygidium, rectum, sternum, and hind tibia. Pheromones
also are exchanged, mixed with food, and passed by trophallaxis,
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 castes, when
the dominant queen stops producing a specific pheromone, workers begin
to raise new queens in the colony.
Some ants produce sounds by stridulation, using the gaster segments
and their mandibles. Sounds may be used to communicate with colony
members or with other species.
Plectroctena sp. attacks another of its kind to protect its
Ants attack and defend themselves by biting and, in many species, by
stinging, often injecting or spraying chemicals, such as formic acid
in the case of formicine ants, alkaloids and piperidines in fire ants,
and a variety of protein components in other ants. Bullet ants
(Paraponera), located in Central and South America, 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 sting of jack jumper ants can be fatal, and an antivenom has
been developed for it.
Fire ants, Solenopsis spp., are unique in having a venom sac
containing piperidine alkaloids. Their stings are painful and can
be dangerous to hypersensitive people.
A weaver ant in fighting position, mandibles wide open
Trap-jaw ants of the genus
Odontomachus are equipped with mandibles
called trap-jaws, which snap shut faster than any other predatory
appendages within the animal kingdom. One study of Odontomachus
bauri recorded peak speeds of between 126 and 230 km/h (78 and
143 mph), with the jaws closing within 130 microseconds on
average. The ants were also observed to use their jaws as a catapult
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 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 the
following genera: Anochetus, Orectognathus, and Strumigenys, plus
some members of the Dacetini tribe, which are viewed as examples
of convergent evolution.
A Malaysian species of ant in the
Camponotus cylindricus group has
enlarged mandibular glands that extend into their gaster. When
disturbed, workers rupture the membrane of the gaster, causing a burst
of secretions containing acetophenones and other chemicals that
immobilise small insect attackers. The worker subsequently dies.
Suicidal defences by workers are also noted in a Brazilian ant,
Forelius pusillus, where a small group of ants leaves the security of
the nest after sealing the entrance from the outside each evening.
Ant mound holes prevent water from entering the nest during rain.
In addition to defence against predators, ants need to protect their
colonies from pathogens. Some worker ants maintain the hygiene of the
colony and their activities include undertaking or necrophory, the
disposal of dead nest-mates.
Oleic acid 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 and iridomyrmecin)
present on the cuticle of their living nestmates to trigger similar
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
Camponotus anderseni, which nests in the cavities of
wood in mangrove habitats, deals with submergence under water by
switching to anaerobic respiration.
Many animals can learn behaviours by imitation, but ants may be the
only group apart from mammals where interactive teaching has been
observed. A knowledgeable forager of
Temnothorax albipennis will lead
a naive nest-mate to newly discovered food by the process of tandem
running. 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
Controlled experiments with colonies of
Cerapachys biroi 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
Leaf nest of weaver ants, Pamalican, Philippines
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 ants of South America, such as the
Eciton burchellii species,
and the driver ants 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
Weaver ant (
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, 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
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
Main article: Ant–fungus mutualism
Myrmecocystus, honeypot ants, store food to prevent colony famine
Most ants are generalist predators, scavengers, and indirect
herbivores, 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 ants (Atta and Acromyrmex)
feed exclusively on a fungus 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. Symbiotic
bacteria on the exterior surface of the ants produce antibiotics that
kill bacteria introduced into the nest that may harm the fungi.
An ant trail
Foraging ants travel distances of up to 200 metres (700 ft) 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, so the ability to find the shortest route back
to the nest reduces that risk. Diurnal desert ants of the genus
Cataglyphis such as the
Sahara desert ant
Sahara desert ant navigate by keeping track of
direction as well as distance travelled. Distances travelled are
measured using an internal pedometer that keeps count of the steps
taken and also by evaluating the movement of objects in their
visual field (optical flow). 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 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 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. The workers may then run
around continuously until they die of exhaustion.
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) is able to jump by synchronising
the action of its mid and hind pairs of legs. There are several
species of gliding ant including Cephalotes atratus; this may be a
common trait among most arboreal ants. Ants with this ability are able
to control the direction of their descent while falling.
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, a species of ant found in Australian mangrove
swamps, can swim and live in underwater nests. Since they lack gills,
they go to trapped pockets of air in the submerged nests to
Cooperation and competition
Meat-eater ants feeding on a cicada: social ants cooperate and
collectively gather food
Not all ants have the same kind of societies. The Australian bulldog
ants are among the biggest and most basal of ants. Like virtually all
ants, they are eusocial, 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 (such as Tetramorium caespitum) attack and take over
neighbouring ant colonies. Others are less expansionist, but just as
aggressive; they invade colonies to steal eggs or larvae, which they
either eat or raise as workers or slaves. Extreme specialists among
these slave-raiding ants, such as the Amazon ants, are incapable of
feeding themselves and need captured workers to survive. Captured
workers of the enslaved species
Temnothorax have evolved a counter
strategy, destroying just the female pupae of the slave-making
Protomognathus americanus, but sparing the males (who don't take part
in slave-raiding as adults).
Harpegnathos saltator (a jumping ant) engaged in battle with
a rival colony's queen
Ants identify kin and nestmates through their scent, which comes from
hydrocarbon-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. Also, the reason why two separate colonies of
ants will attack each other even if they are of the same species is
because the genes responsible for pheromone production are different
between them. The Argentine ant, however, does not have this
characteristic, due to lack of genetic diversity, and has become a
global pest because of it.
Parasitic ant species enter the colonies of host ants and establish
themselves as social parasites; species such as
Strumigenys xenos 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.
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, 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
Relationships with other organisms
Myrmarachne plataleoides (female shown) mimics weaver ants
to avoid predators.
Ants form symbiotic 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 inquilines may bear a close resemblance
to ants. The nature of this ant mimicry (myrmecomorphy) varies, with
some cases involving Batesian mimicry, where the mimic reduces the
risk of predation. Others show Wasmannian mimicry, a form of mimicry
seen only in inquilines.
An ant collects honeydew from an aphid
Aphids and other hemipteran insects secrete a sweet liquid called
honeydew, when they feed on plant sap. 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
mealybugs to harvest their honeydew. Mealybugs may become a serious
pest of pineapples if ants are present to protect mealybugs from their
Myrmecophilous (ant-loving) caterpillars of the butterfly family
Lycaenidae (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. Some caterpillars produce
vibrations and sounds that are perceived by the ants. A similar
adaptation can be seen in
Grizzled skipper 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.
Fungus-growing ants that make up the tribe Attini, including
leafcutter ants, cultivate certain species of fungus in the
Leucocoprinus genera of the
Agaricaceae family. In
this ant-fungus mutualism, both species depend on each other for
survival. The ant
Allomerus decemarticulatus has evolved a three-way
association with the host plant, Hirtella physophora
(Chrysobalanaceae), and a sticky fungus which is used to trap their
Ants may obtain nectar from flowers such as the dandelion but are only
rarely known to pollinate flowers.
Lemon ants make devil's gardens by killing surrounding plants with
their stings and leaving a pure patch of lemon ant trees, (Duroia
hirsuta). This modification of the forest provides the ants with more
nesting sites inside the stems of the
Duroia trees. Although some
ants obtain nectar from flowers, pollination by ants is somewhat
rare. Some plants have special nectar exuding structures,
extrafloral nectaries, that provide food for ants, which in turn
protect the plant from more damaging herbivorous insects. Species
such as the bullhorn acacia (Acacia cornigera) in
Central America have
hollow thorns that house colonies of stinging ants (Pseudomyrmex
ferruginea) who defend the tree against insects, browsing mammals, and
Isotopic labelling studies suggest that plants also
obtain nitrogen from the ants. In return, the ants obtain food
from protein- and lipid-rich Beltian bodies. In Fiji Philidris nagasau
(Dolichoderinae) are known to selectively grow species of epiphytic
Squamellaria (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 comes from the
which has stems adapted to house colonies of
Many tropical tree species have seeds that are dispersed by ants.
Seed dispersal by ants or myrmecochory is widespread and new estimates
suggest that nearly 9% of all plant species may have such ant
associations. Some plants in fire-prone grassland 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, elaiosomes, that
are sought after by ants as food.
A convergence, possibly a form of mimicry, is seen in the eggs of
stick insects. They have an edible elaiosome-like structure and are
taken into the ant nest where the young hatch.
A meat ant tending a common leafhopper nymph
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 and Termitopone) while a few
Cerapachyinae prey on other ants. Some termites, including
Nasutitermes corniger, form associations with certain ant species to
keep away predatory ant species. The tropical wasp Mischocyttarus
drewseni 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 and Melipona) use chemical defences against ants.
Flies in the Old World genus
Bengalia (Calliphoridae) prey on ants and
are kleptoparasites, snatching prey or brood from the mandibles of
adult ants. Wingless and legless females of the Malaysian phorid
fly (Vestigipoda myrmolarvoidea) live in the nests of ants of the
Aenictus and are cared for by the ants.
Fungi in the genera
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. It appears that the fungus
alters the behaviour of the ant to help disperse its spores  in a
microhabitat that best suits the fungus. Strepsipteran parasites
also manipulate their ant host to climb grass stems, to help the
parasite find mates.
A nematode (Myrmeconema neotropicum) that infects canopy ants
(Cephalotes atratus) 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.
Spiders sometimes feed on ants
South American poison dart frogs in the genus
Dendrobates feed mainly
on ants, and the toxins in their skin may come from the ants.
Army ants forage in a wide roving column, attacking any animals in
that path that are unable to escape. In Central and South America,
Eciton burchellii is the swarming ant most commonly attended by
"ant-following" birds such as antbirds and woodcreepers.
This behaviour was once considered mutualistic, but later studies
found the birds to be parasitic. Direct kleptoparasitism (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. Birds that follow ants eat many prey
insects and thus decrease the foraging success of ants. Birds
indulge in a peculiar behaviour called anting 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
ectoparasites from the birds.
Anteaters, aardvarks, pangolins, echidnas and numbats have special
adaptations 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 bears (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
Weaver ants are used as a biological control for citrus cultivation in
Ants perform many ecological roles that are beneficial to humans,
including the suppression of pest populations and aeration of the
soil. The use of weaver ants in citrus cultivation in southern China
is considered one of the oldest known applications of biological
control. 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 ants, are used as surgical sutures. 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
Paraponera clavata (tocandira) and
Dinoponera spp. (false
South America  and the Myrmecia ants of
In South Africa, ants are used to help harvest the seeds of rooibos
(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.
It has been estimated by
E.O. Wilson that the total number of
individual ants alive in the world at any one time is between one and
ten quadrillion (short scale) (i.e. between 1015 and 1016). According
to this estimate, the total biomass of all the ants in the world is
approximately equal to the total biomass of the entire human
race. Also, according to this estimate, there are approximately 1
million ants for every human on Earth.
See also: Entomophagy
Roasted ants in Colombia
Ant larvae for sale in Isaan, Thailand
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 and can sell for as much as US$40
per pound ($90/kg) because they are seasonal and hard to find. In the
Colombian department of Santander, hormigas culonas (roughly
interpreted as "large-bottomed ants")
Atta laevigata are toasted alive
In areas of India, and throughout
Burma and Thailand, a paste of the
green weaver ant (
Oecophylla smaragdina) is served as a condiment with
Weaver ant eggs and larvae, as well as the ants, may be
used in a Thai salad, yam (Thai: ยำ), in a dish called yam khai
mot daeng (Thai: ยำไข่มดแดง) 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".
In his First Summer in the Sierra,
John Muir notes that the Digger
California ate the tickling, acid gasters of the large
jet-black carpenter ants. The Mexican Indians eat the replete workers,
or living honey-pots, of the honey ant (Myrmecocystus).
See also: Ants of medical importance
The tiny pharaoh ant is a major pest in hospitals and office blocks;
it can make nests between sheets of paper
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, pavement ant, yellow
crazy ant, banded sugar ant, Pharaoh ant, carpenter ants, odorous
house ant, red imported fire ant, and European fire ant. Some ants
will raid stored food, others may damage indoor structures, some can
damage agricultural crops directly (or by aiding sucking pests), and
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
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.
Management is based on the species and techniques can vary according
to the location and circumstance.
In science and technology
Camponotus nearcticus workers travelling between two formicaria
through connector tubing
See also: Myrmecology, Biomimetics, and
Ant colony optimization
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, study ants in the laboratory and in their natural
conditions. Their complex and variable social structures have made
ants ideal model organisms.
Ultraviolet vision was first discovered in
ants by Sir John Lubbock in 1881. Studies on ants have tested
hypotheses in ecology and sociobiology, and have been particularly
important in examining the predictions of theories of kin selection
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 robotics. This area of biomimetics has led to
studies of ant locomotion, search engines that make use of "foraging
trails", fault-tolerant storage, and networking algorithms.
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
Aesop's ants: picture by Milo Winter, 1888–1956
Anthropomorphised ants have often been used in fables and children's
stories to represent industriousness and cooperative effort. They also
are mentioned in religious texts. In the
Book of Proverbs
Book of Proverbs in
the Bible, ants are held up as a good example for humans for their
hard work and cooperation.
Aesop did the same in his fable The
the Grasshopper. In the Quran, 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.[Quran 27:18] In parts of Africa, ants are considered
to be the messengers of the deities. Some Native American mythology,
such as the Hopi mythology, considers ants as the very 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 ceremonies of some Amazon Indian
cultures as a test of endurance.
Ant society has always fascinated humans and has been written about
both humorously and seriously.
Mark Twain wrote about ants in his 1880
book A Tramp Abroad. Some modern authors have used the example of
the ants to comment on the relationship between society and the
individual. Examples are
Robert Frost in his poem "Departmental" and
T. H. White
T. H. White in his fantasy novel The Once and Future King. 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 is described using contemporary
H.G. Wells 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. In more recent times, animated cartoons and 3-D animated
films featuring ants have been produced including Antz, A Bug's Life,
Ant Bully, The
Ant and the Aardvark, Ferdy the
Ant and Atom Ant.
E. O. Wilson
E. O. Wilson wrote a short story, "Trailhead"
in 2010 for
The New Yorker
The New Yorker 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. The French neuroanatomist, psychiatrist and
Auguste Forel believed that ant societies were models for
human society. He published a five volume work from 1921 to 1923 that
examined ant biology and society.
In the early 1990s, the video game SimAnt, which simulated an ant
colony, won the 1992 Codie award for "Best Simulation Program".
Ants also are quite popular inspiration for many science-fiction
insectoids, such as the Formics of Ender's Game, the Bugs of Starship
Troopers, the giant ants in the films
Them! and Empire of the Ants,
Marvel Comics' super hero Ant-Man, and ants mutated into
super-intelligence in Phase IV. In computer strategy games, ant-based
species often benefit from increased production rates due to their
single-minded focus, such as the Klackons in the Master of Orion
series of games or the ChCht in Deadlock II. These characters are
often credited with a hive mind, a common misconception about ant
Main article: Outline of ants
Glossary of ant terms
International Union for the Study of Social Insects
Myrmecological News (journal)
Task allocation and partitioning of social insects
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Wikiquote has quotations related to: Ant
Wikimedia Commons has media related to Formicidae.
Wikispecies has information related to Formicidae
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Sierolomorphidae (sierolomorphid wasps)
Mutillidae (velvet ants)
Pompilidae (spider wasps)
Sapygidae (sapygid, or club-horned wasps)
Scoliidae (scoliid wasps)
Ampulicidae (cockroach wasps)
Crabronidae (sand wasps, bee wolves)
Sphecidae (thread-waisted wasps)
Andrenidae (mason bees)
Apidae (honey bees, bumblebees, cuckoo bees, carpenter bees, orchid
bees, stingless bees)
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Megachilidae (mason bees, leafcutter bees)
Italic are paraphyletic groups
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List of ant genera
Evolution of eusociality
Identity in social insects
Sexual selection in social insects
Karl von Frisch
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Charles Duncan Michener
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E. O. Wilson
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Sociobiology: The New Synthesis 1975
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Living things in culture
Fauna Europaea: 11356
BNF: cb11931991b (d