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 99
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 waists.
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 mostly of sterile, wingless
females forming castes of "workers", "soldiers", or other specialised
groups. Nearly all ant colonies also have some fertile males called
"drones" and one or more fertile females called "queens ". The
colonies are described as superorganisms because the ants appear to
operate as a unified entity, collectively working together to support
the colony. Play media (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
* 1 Etymology
* 2 Taxonomy and evolution
* 3 Distribution and diversity
* 4 Morphology
* 4.1 Head
* 4.2 Legs
* 4.3 Wings
* 4.5 Polymorphism
* 5 Life cycle
* 5.1 Reproduction
* 6 Behaviour and ecology
* 6.1 Communication
* 6.2 Defence
* 6.3 Learning
* 6.4 Nest construction
* 6.5 Cultivation of food
* 6.6 Navigation
* 6.7 Locomotion
* 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 References
* 9.1 Cited texts
* 10 Further reading
* 11 External links
The word "ant" is derived from ante, emete of
Middle English which
are derived from ǣmette of
Old English , and is related to the
dialectal Dutch emt and the
Old High German
Old High German āmeiza, hence 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 from the
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 used, cf.
Latin formīca, Greek
Old Church Slavonic
Old Church Slavonic mraviji,
Old Irish moirb,
Old Norse maurr, Dutch mier.
TAXONOMY AND EVOLUTION
Ants fossilised in
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 the
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 possibly was a ground
Haidomyrmodes , related genera in
Sphecomyrminae , are reconstructed as active arboreal
predators. Older ants in the genus Sphecomyrmodes have been found in
99 million year-old amber from Myanmar. 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 the soil.
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
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
Iceland , parts of
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
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
the appendages. 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 metre away.
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.
All six legs are attached to the mesosoma ("thorax") and terminate in
a hooked claw .
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.
Leafcutter ant workers of various castes (left) and two
In the colonies of a few ant species, there are physical
castes—workers in distinct size-classes, called minor, median, and
major workers. Often, the larger ants have disproportionately larger
heads, and correspondingly stronger mandibles . 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 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
A new worker 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
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 or inactivity. 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 spermathecae .
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 weak 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.
Insect defences A
Plectroctena sp. attacks another of
its kind to protect its territory
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:
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 it outside.
Camponotus anderseni , which nests in the cavities of wood in mangrove
habitats, deals with submergence under water by switching to anaerobic
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
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
Ant colony Play media Leaf nest of
weaver ants ,
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 other together.
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
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
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. Workers 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 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 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 breathe.
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). A worker
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
Argentine ant , however, does not have this characteristic,
due to lack of genetic diversity, and has become a global pest because
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 father.
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. 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 insect prey.
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 epiphytic
Isotopic labelling studies suggest that plants also obtain
nitrogen from the ants. In return, the ants obtain food from protein-
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
Macaranga tree, 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 (
Melipona ) use chemical defences against ants.
Flies in the Old World genus
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
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
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 southern China
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 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
Paraponera clavata (tocandira) and
(false tocandiras) of
South America and the Myrmecia ants of
Africa , ants are used to help harvest rooibos (Aspalathus
linearis), which are small seeds 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.
Entomophagy Roasted ants in
larvae for sale in
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
Thailand , a paste of
the green weaver ant (
Oecophylla smaragdina ) is served as a condiment
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 (
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:
Biomimetics , and
Ant colony optimization algorithms
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
Ant and 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. In parts of
Africa, ants are considered to be the messengers of the deities. Some
Native American mythology
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
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
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 . Renowned myrmecologist
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
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
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
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Wikiquote has quotations related to: ANT
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* AntWeb from The
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Ant Species Fact Sheets from the National Pest Management
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