Ant 2594 225956313 2008-07-16T05:09:02Z TXiKiBoT 3171782 robot Adding: [[ur:چیونٹی]] <!--{{pp-semi-protected|small=yes}}--> {{otheruses|Ant (disambiguation)}} {{Taxobox | name = Ants | fossil_range = [[Cretaceous]] - Recent | image = Meat eater ant feeding on honey02.jpg | image_width = 250px | image_caption = [[Meat ant|Meat eater ant]] feeding on honey | regnum = [[Animal]]ia | phylum = [[Arthropod]]a | classis = [[Insect]]a | ordo = [[Hymenoptera]] | subordo = [[Apocrita]] | superfamilia = [[Vespoidea]] | familia = '''Formicidae''' | familia_authority = [[Pierre André Latreille|Latreille]], 1809 | subdivision_ranks = Subfamilies | subdivision = *[[Aenictogitoninae]] *[[Agroecomyrmecinae]] *[[Amblyoponinae]] (incl. "[[Apomyrminae]]") *[[Aneuretinae]] *[[Cerapachyinae]] *[[Dolichoderinae]] *[[Ecitoninae]] (incl. "[[Dorylinae]]" and "[[Aenictinae]]") *[[Ectatomminae]] *[[Formicinae]] *[[Heteroponerinae]] *[[Leptanillinae]] *[[Leptanilloidinae]] *[[Myrmeciinae]] (incl. "[[Nothomyrmeciinae]]") *[[Myrmicinae]] *[[Paraponerinae]] *[[Ponerinae]] *[[Proceratiinae]] *[[Pseudomyrmecinae]] {{hidden|bg1=#D3D3A4|[[Cladogram]] of ant [[subfamily|subfamilies]]| {{clade|style=font-size:75%;line-height:75% |1={{clade| |1=[[Leptanillinae]] |2=[[Amblyoponinae]] |3=[[Paraponerinae]] |4=[[Agroecomyrmecinae]] |5=[[Ponerinae]] |6=[[Proceratiinae]] |7={{clade| |1={{clade| |1={{clade| |1=[[Ecitoninae]] |2=[[Aenictinae]] |3={{clade| |1=[[Dorylini]] |2=[[Aenictogitoninae]] }} }} |2=[[Cerapachyinae]]* |3=[[Leptanilloidinae]] }} |2={{clade| |1={{clade| |1={{clade| |1=[[Dolichoderinae]] |2=[[Aneuretinae]] }} |2={{clade| |1=[[Pseudomyrmecinae]] |2=[[Myrmeciinae]] }} }} |2={{clade| |1={{clade| |1=[[Ectatomminae]] |2=[[Heteroponerinae]] }} |2=[[Myrmicinae]] |3=[[Formicinae]] }} }} }} }} }} <small>A [[phylogeny]] of the extant ant [[subfamily|subfamilies]].<ref>{{cite journal |author=Ward, Philip S |title=Phylogeny, classification, and species-level taxonomy of ants (Hymenoptera: Formicidae)|journal=[[Zootaxa]] |volume=1668 |year=2007|pages=pp. 549&ndash;563 }}</ref><br/> <nowiki>*</nowiki>Cerapachyinae is [[paraphyletic]]</small>}} }} '''Ants''' are [[eusocial|social]] [[insect]]s of the family '''Formicidae''' and, along with the related families of [[wasp]]s and [[bee]]s, belong to the order [[Hymenoptera]]. Ants [[evolution|evolved]] from wasp-like ancestors in the mid-[[Cretaceous]] period between 110 and 130&nbsp;million years ago and [[Evolutionary radiation|diversified]] after the rise of [[flowering plant]]s. Today, more than 14,000 [[List of ant genera (alphabetical)|species]] are classified.<ref>{{cite news |url=http://www.nytimes.com/2008/07/15/science/15wils.html |accessdate=2008-07-15 |title=Taking a Cue From Ants on Evolution of Humans |first=Nicholas |last=Wade |work=New York Times |date=2008-07-15 }}</ref> They are easily identified by their elbowed antennae and a distinctive node-like structure that forms a slender waist. Ants form highly organised [[colony (biology)|colonies]], which may occupy large territories and consist of millions of individuals that are mostly sterile females forming castes of "workers", "soldiers", or other specialised groups. Ant colonies also have some fertile males called "drones" and one or more fertile females called [[Queen ant|"queens"]]. The colonies are sometimes described as [[superorganisms]] because ants appear to operate as a unified entity, collectively working together to support the colony.<ref>{{cite book|author=Oster GF, Wilson EO|year=1978|title=Caste and ecology in the social insects|publisher=Princeton University Press, Princeton|page=pp. 21&ndash;22|isbn=0691023611}}</ref> Ants have colonised almost every landmass on Earth. The only places lacking indigenous ants are remote or inhospitable islands. Ants dominate most ecosystems, and form 15&ndash;20% of the terrestrial animal biomass.<ref name="schultz"/> Their success has been attributed to their social organisation, ability to modify their habitats, tap resources and defend themselves. Their long [[co-evolution]] with other species has led to [[mimicry|mimetic]], [[commensalism|commensal]], [[parasitism|parasitic]] and [[mutualistic]] relationships.<ref name=TheAntsRelations>Hölldobler & Wilson (1990), p. 471</ref> Ant societies have [[division of labour]], communication between individuals, and an ability to [[Problem solving|solve complex problems]].<ref name="SANdisk"/> These parallels with [[Civilization|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.<ref name="HolldoblerWilsonAnts3"/> However, their ability to exploit resources brings ants into conflict with humans, as they can damage crops and invade buildings. Some species, such as the [[red imported fire ant]], are regarded as [[invasive species]], since they can spread rapidly into new areas.<ref name="pests"/> ==Taxonomy and evolution== [[Image:Ants in amber.jpg|thumb|left|Ants fossilised in Baltic [[amber]]]] The family Formicidae belongs to the order [[Hymenoptera]], which also includes [[sawfly|sawflies]], [[bees]] and [[wasps]]. Ants are evolved from a lineage within the [[Vespoidea|vespoid]] wasps. [[Phylogenetic]] analysis suggests that ants arose in the mid-[[Cretaceous]] period about 110 to 130&nbsp;million years ago. After the rise of [[flowering plant]]s about 100&nbsp;million years ago, they diversified and assumed ecological dominance around 60&nbsp;million years ago.<ref name="grimaldi2001">{{cite journal |author=Grimaldi D, Agosti D |year=2001 |title=A formicine in New Jersey Cretaceous amber (Hymenoptera: Formicidae) and early evolution of the ants |journal=[[Proceedings of the National Academy of Sciences]] |volume=97 |pages=pp. 13678&ndash;13683 |url=http://www.pnas.org/cgi/ijlink?linkType=ABST&journalCode=pnas&resid=97/25/13678 | doi = 10.1073/pnas.240452097 |pmid=11078527}}</ref><ref>{{cite journal |journal=[[Science (journal)|Science]] |url=http://www.sciencemag.org/cgi/content/abstract/312/5770/101 | doi = 10.1126/science.1124891|title=Phylogeny of the ants: Diversification in the Age of Angiosperms|author=Moreau CS, Bell CD, Vila R, Archibald SB, Pierce NE |volume=312 |issue=5770 |pages=pp. 101&ndash;104|year=2006|pmid=16601190}}</ref><ref>{{Cite journal | volume = 102| issue = 21 | pages = 7411&ndash;7414| author= Wilson EO, Hölldobler B| title = The rise of the ants: A phylogenetic and ecological explanation| journal = Proceedings of the National Academy of Sciences |url=http://www.pnas.org/cgi/content/full/102/21/7411 | doi = 10.1073/pnas.0502264102| year = 2005| pmid = 15899976}}</ref> In 1966, [[E. O. Wilson]] and his colleagues identified the fossil remains of an ant (''[[Sphecomyrma freyi]]'') that lived in the Cretaceous period. The specimen, trapped in amber [[absolute dating|dating]] back to more than 80&nbsp;million years ago has features of both ants and wasps.<ref>{{cite journal |author=[[E. O. Wilson|Wilson E O]], Carpenter FM, Brown WL |title=The first Mesozoic ants |journal=[[Science (journal)|Science]] |year=1967 |volume=157 |pages=pp. 1038&ndash;1040 | doi = 10.1126/science.157.3792.1038 |pmid=17770424}} </ref> {{userboxtop|toptext=&nbsp;}} {{clade| style=font-size:75%;line-height:75% |label1=[[Vespoidea]] |1={{clade |1=[[Sierolomorphidae]] |2={{clade |1={{clade |1=[[Tiphiidae]] |2={{clade |1=[[Sapygidae]] |2=[[Mutillidae]] }} }} |2={{clade |1={{clade |1=[[Pompilidae]] |2=[[Rhopalosomatidae]] }} |2={{clade |1='''Formicidae''' |2={{clade |1=[[Vespidae]] |2=[[Scoliidae]] }} }} }} }} }} }} <center><small>Phylogenetic position of the Formicidae.<ref>{{cite journal|author=Brothers DJ|year=1999|title=Phylogeny and evolution of wasps, ants and bees (Hymenoptera, Chrysisoidea, Vespoidea, and Apoidea)|journal=Zoologica Scripta|volume=28|pages=pp. 233&ndash;249|doi=10.1046/j.1463-6409.1999.00003.x}}</ref></small></center> {{userboxbottom}} During the Cretaceous period, only a few species of primitive ants ranged widely on the [[Laurasia]]n super-continent (the northern hemisphere). They were scarce in comparison to other insects, and represented about 1% of the insect population. Ants became dominant after [[adaptive radiation]] at the beginning of the [[Tertiary period]]. By the [[Oligocene]] and [[Miocene]] ants had come to represent 20&ndash;40% of all insects found in major fossil deposits. Of the species that lived in the [[Eocene]] epoch, one of approximately ten genera survive to the present. Genera surviving today comprise 56% of the genera in [[Baltic region|Baltic]] amber fossils (early Oligocene), and 92% of the genera in [[Dominican Republic|Dominican]] amber fossils (apparently early Miocene).<ref name=TheAntEvo>Hölldobler & Wilson (1990), pp. 23&ndash;24</ref><ref name="grimaldi2001"/> [[Termite]]s, though sometimes called ''white ants'', are unrelated to ants and belong to the order [[Isoptera]] and resemble ants only in some aspects of their social life. [[Mutillidae|Velvet ant]]s, look like large ants, but are wingless female [[wasp]]s.<ref>{{cite web |url=http://bugguide.net/node/view/69 |title=Order Isoptera - Termites |accessdate=2008-06-12 |publisher=Iowa State University Entomology |date=2004-02-16 }} </ref><ref>{{cite web |url=http://bugguide.net/node/view/159/ |title=Family Mutillidae - Velvet ants|accessdate=2008-06-12 |publisher=Iowa State University Entomology |date= 2004-02-16 }}</ref> ===Etymology=== The word ''ant'' is derived from ''ante'' of Middle English which is derived from ''æmette'' and ''emmett'' of Old English and is related to the Old High German ''āmeiza'' from which comes ''ameise'', the German word for ant.<ref>{{cite web|url=http://www.merriam-webster.com/dictionary/ant|title="ant". Merriam-Webster Online Dictionary|accessdate=2008-06-06|publisher=Merriam-Webster}}</ref> The family name ''Formicidae'' is derived from the [[Latin]] ''formīca'' for ant.<ref>{{cite book|author = Simpson DP| title = Cassell's Latin Dictionary | publisher = Cassell Ltd| date = 1979|edition = 5|location = London| isbn=0-304-52257-0}}</ref> ==Distribution and diversity== {| class="wikitable sortable" align="right" style="margin: 10px;" |- ! [[Region]] !! Number of<br />species&nbsp;<ref name = HolldoblerWilsonAnts>Hölldobler & Wilson (1990), p. 4</ref> |- | [[Neotropic ecozone|Neotropics]] ||align="right"| 2162 |- | [[Nearctic ecozone|Nearctic]] ||align="right"| 580 |- | [[Europe]] ||align="right"| 180 |- | [[Africa]] ||align="right"| 2500 |- | [[Asia]] ||align="right"| 2080 |- | [[Melanesia]] ||align="right"| 275 |- | [[Australia]] ||align="right"| 985 |- | [[Polynesia]] ||align="right"| 42 |} Ants are found on all continents except [[Antarctica]] and only a few large islands such as [[Greenland]], [[Iceland]], parts of [[Polynesia]] and the [[Hawaii|Hawaiian Islands]] lack native ant species.<ref>{{cite web|url=http://ngm.nationalgeographic.com/2007/08/ants/did-you-know-learn|title=Fantastic ants - Did you know?|publisher=National Geographic Magazine|author=Jones, Alice S|accessdate=2008-06-05}}</ref><ref>{{Cite web |url=http://www.hear.org/ants/ |title=Pest Ants in Hawaii |date=2007 |author=Thomas, Philip|publisher=Hawaiian Ecosystems at Risk project (HEAR)|accessdate=2008-06-06}}</ref> Ants occupy a wide range of [[ecological niche]]s, and are able to exploit a wide range of food resources either as direct or indirect herbivores, predators and scavengers. Most species are omnivorous [[Generalist and specialist species|generalists]] but a few are specialist feeders. Their ecological dominance may be measured by their [[biomass (ecology)|biomass]], and estimates in different environments suggest that they contribute 15&ndash;20% (on average and nearly 25% in the tropics) of the total terrestrial animal biomass, which exceeds that of the vertebrates.<ref name="schultz">{{cite journal |author=Schultz TR|year=2000 |title=In search of ant ancestors|journal=[[Proceedings of the National Academy of Sciences]] |volume=97 |issue=26 |pages=pp. 14028&ndash;14029 |url=http://www.pnas.org/cgi/content/full/97/26/14028 | doi = 10.1073/pnas.011513798 |pmid=11106367}}</ref> Ants range in size from {{convert|0.75|to|52|mm|abbr=on|lk=on|sigfig=2}}.<ref name = AntsDorylusWilverthaiQueen >Hölldobler & Wilson (1990), p. 589</ref><ref name = OligomyrmexMinor>{{Cite book | author=Shattuck SO| title=Australian ants: their biology and identification | date=1999 | publisher=CSIRO | location=Collingwood, Vic| isbn=0-643-06659-4 |pages=p. 149}}</ref> Their colours vary; most are red or black, green is less common, and some tropical species have a metallic lustre. More than [[List of ant genera (alphabetical)|14,000 species]] are currently recognised, 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.<ref>{{cite web |author=Agosti D, Johnson NF (eds.) |year=2005 |title=Antbase |url=http://www.antbase.org/|publisher=American Museum of Natural History|accessdate=2008-06-06}}</ref> The relative ease with which ants can be sampled and studied in ecosystems has made them useful as [[indicator species]] in [[biodiversity]] studies.<ref>{{cite book |author=Agosti D, Majer JD, Alonso JE, Schultz TR (eds.) |year=2000 |title=Ants: Standard methods for measuring and monitoring biodiversity |publisher=[[Smithsonian Institution|Smithsonian Institution Press]] |url=http://antbase.org/databases/publications_files/publications_20330.htm}}</ref><ref>{{cite web |url=http://atbi.biosci.ohio-state.edu:210/hymenoptera/nomenclator.home_page |title=Hymenoptera name server |publisher=[[Ohio State University]]|author=Johnson NF|year=2007|accessdate=2008-06-06}}</ref> == Morphology == Ants are distinct in their [[morphology (biology)|morphology]] from other insects in having elbowed [[antenna (biology)|antennae]], [[metapleural gland]]s, and a strong constriction of their second [[Abdomen|abdominal]] segment into a node-like [[petiole (insect)|petiole]]. The head, [[mesosoma]] and [[metasoma]] or gaster 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 can be formed by one or two nodes (the second alone, or the second and third abdominal segments).<ref>Borror, Triplehorn & Delong (1989), p. 737</ref> [[Image:Bullant head detail.jpg|thumb|right|[[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 [[vertebrate]]s. Insects do not have lungs; oxygen and other gases like [[carbon dioxide]] pass through their exoskeleton through tiny valves called [[spiracle]]s. 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]] towards 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.<ref name="insectmorph">Borror, Triplehorn & Delong (1989), pp. 24&ndash;71</ref> [[Image:Scheme ant worker anatomy-en.svg|thumb|left|Diagram of a worker ant (''Pachycondyla verenae'')]] An ant's head contains many sensory organs. Like most insects, ants have [[compound eye]]s made from numerous tiny lenses attached together. Ants' eyes are good for acute movement detection but do not give a high [[Optical resolution|resolution]]. They also have three small [[ocellus|ocelli]] (simple eyes) on the top of the head that detect light levels and [[polarization|polarisation]].<ref>{{cite journal | journal=Science | year=1985 |volume=228 |issue=4696 |pages=pp. 192&ndash;194|doi=10.1126/science.228.4696.192|title=Ocelli: A celestial compass in the desert ant ''Cataglyphis''|author=Fent K, Rudiger W|pmid=17779641}}</ref> Compared to vertebrates, most ants have poor-to-mediocre eyesight and a few subterranean species are completely blind. Some ants such as Australia's [[bulldog ant]], however, have exceptional vision. Two [[antenna (biology)|antennae]] ("feelers") are attached to the head; these organs detect chemicals, air currents and vibrations; they are also used to transmit and receive signals through touch. The head has two strong jaws, the [[Mandible (insect)|mandibles]], used to carry food, manipulate objects, construct nests, and for defence.<ref name="insectmorph"/> In some species a small pocket (infrabuccal chamber) inside the mouth stores food, so it can be passed to other ants or their larvae.<ref>{{cite journal|author=Eisner T, Happ GM|title=The infrabuccal pocket of a formicine ant: a social filtration device|journal=Psyche|volume=69|pages=pp. 107&ndash;116|year=1962|url=http://psyche.entclub.org/69/69-107.html|doi=10.1155/1962/25068}}</ref> All six [[arthropod leg|legs]] are attached to the [[mesosoma]] ("thorax"). A hooked [[claw#Arthropods|claw]] at the end of each leg helps ants to climb and hang onto surfaces. Most [[Queen (insect)|queens]] and male ants have [[insect wing|wings]]; queens shed the wings after the [[nuptial flight]], leaving visible stubs, a distinguishing feature of queens. However, wingless queens (ergatoids) and males occur in a few species.<ref name="insectmorph"/> The [[metasoma]] (the "abdomen") of the ant houses important internal organs, including those of the reproductive, respiratory (tracheae) and excretory systems. Many species have [[stinger (organ)|stingers]], used for subduing [[predation|prey]] and defending their nests.<ref name="insectmorph"/> ===Polymorphism=== [[Image:HoneyAnt.jpg|thumb|right|''Myrmecocystus'' (Honeypot) ants store food to prevent colony famine.]] In the colonies of a few ant species, there are physical castes&mdash;workers in distinct size-classes, called minor, median, and major workers. Often the larger ants have disproportionately larger heads, and correspondingly stronger [[mandible (insect)|mandibles]]. Such individuals are sometimes called "soldier" ants because their stronger mandibles make them more effective in fighting, although they are still 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.<ref>{{cite journal |author=[[E. O. Wilson|Wilson EO]] |year=1953 |title=The origin and evolution of polymorphism in ants|journal=[[Quarterly Review of Biology]] |volume=28 |issue=2 |pages=pp. 136&ndash;56 |doi=10.1086/399512}}</ref> Some other species show continuous variation in the size of workers. The smallest and largest workers in ''[[Pheidologeton diversus]]'' show nearly a 500&ndash;fold difference in their dry-weights.<ref>{{cite journal|author=Moffett MW, Tobin JE|year=1991|title=Physical castes in ant workers: a problem for ''Daceton armigerum'' and other ants|journal=Psyche|volume=98|pages=pp. 283&ndash;292|url=http://psyche2.entclub.org/articles/98/98-283.pdf|format=PDF|doi=10.1155/1991/30265}}</ref> 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''.<ref>{{cite journal|author=Børgesen LW|year=2000|title=Nutritional function of replete workers in the pharaoh's ant, ''Monomorium pharaonis'' (L.)|journal=Insectes Sociaux|volume=47|issue=2|pages=pp. 141&ndash;146|doi=10.1007/PL00001692}}</ref> This polymorphism in morphology and behaviour of workers was initially thought to be determined by environmental factors such as nutrition and hormones, which led to different [[morphogenesis|developmental paths]], however genetic differences between worker castes have been noted in ''Acromyrmex'' sp.<ref>{{cite journal|title=Worker caste polymorphism has a genetic basis in ''Acromyrmex'' leaf-cutting ants|author=Hughes WOH, Sumner S, Van Borm S, Boomsma JJ|doi=10.1073/pnas.1633701100|journal=Proceedings of the National Academy of Sciences|year=2003 |volume=100 |issue= 16 |pages=pp. 9394&ndash;9397|pmid=12878720}}</ref> 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.<ref>{{cite journal|title=Alternative genetic foundations for a key social polymorphism in fire ants|author=Rossa KG, Kriegera MJB, Shoemaker DD|journal=Genetics|volume=165|pages=pp. 1853&ndash;1867|year=2003}}</ref> The Australian [[jack jumper ant]] (''Myrmecia pilosula''), has only a single pair of chromosomes (males have 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.<ref>{{cite journal|author=Crosland MWJ, Crozier RH|year=1986|title=''Myrmecia pilosula'', an ant with only one pair of chromosomes|journal=Science|volume=231|pages=p. 1278|doi=10.1126/science.231.4743.1278|pmid=17839565}}</ref><ref>{{cite journal|journal=BMC Evolutionary Biology|year=2008|volume=8|issue=64|url=http://www.biomedcentral.com/1471-2148/8/64|accessdate=2008-06-25|title=The evolution of genome size in ants|author=Tsutsui ND, Suarez AV, Spagna JC, Johnston JS|doi=10.1186/1471-2148-8-64|pages=64}} }}</ref><!--electronic without pagination--> ==Development== [[Image:Meat eater ant nest swarming02.jpg|thumb|Meat eater ant nest during swarming]] The life of an ant starts from an [[egg (biology)|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]]l stages passing through a [[pupa]]l stage before emerging as an adult. The larva is immobile and is fed and cared for by workers. Food is given to the larvae by [[trophallaxis]], a process in which an ant [[Regurgitation (digestion)|regurgitates]] food held in its [[Crop (anatomy)|crop]]. This is also how adults share food, stored in the "social stomach", among themselves. The larvae grow through a series of [[moult]]s and enter the pupal stage. The pupa has the appendages free and not fused to the body as in a [[Pupa#Chrysalis|butterfly pupa]].<ref>{{cite book|author=Gillott, Cedric|year=1995|title=Entomology|publisher=Springer|isbn=0306449676|pages=p. 325}}</ref> The differentiation into queens and workers (which are both female), and different [[caste]]s of workers (when they exist), is determined by the nutrition the larvae obtain. Larvae and pupae need to be kept at fairly constant temperatures to ensure proper development, and so are often moved around the various brood chambers within the colony.<ref>Hölldobler & Wilson (1990), pp. 351, 372</ref> A new worker spends the first few days of its adult life caring for the queen and young. It 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 that are older and are likely to die soon of natural causes.<ref>{{cite journal|journal=Annual Review of Entomology|year=1989|volume=34|pages=pp. 191&ndash;210|title=Foraging strategies of ants|author=Traniello JFA|doi=10.1146/annurev.en.34.010189.001203}}</ref><ref>{{cite journal|author=Sorensen A, Busch TM, Vinson SB|title=Behavioral flexibility of temporal sub-castes in the fire ant, ''Solenopsis invicta'', in response to food|journal=Psyche|volume=91|pages=pp. 319&ndash;332|year=1984|url=http://psyche.entclub.org/91/91-319.html|doi=10.1155/1984/39236}}</ref> [[Image:Meat eater ant qeen excavating hole.jpg|thumb|left|Fertilised queen ant beginning to dig a new colony]] 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 can 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.<ref>{{cite journal|author=Peeters C, Holldobler B|year=1995|journal=Proceedings of the National Academy of Sciences|volume=92|pages=pp. 10977&ndash;10979|url=http://www.pnas.org/cgi/reprint/92/24/10977.pdf|format=PDF|title=Reproductive cooperation between queens and their mated workers: The complex life history of an ant with a valuable nest|doi=10.1073/pnas.92.24.10977|pmid=11607589}}</ref> The winged male ants, called drones, emerge from pupae along with the breeding females (although some species, like [[army ant]]s, have wingless queens), and do nothing in life except eat and mate. During the short breeding period, the reproductives, excluding the colony queen, are carried outside where other colonies of similar species are doing the same. Then, all the winged breeding ants take flight. Mating occurs in flight and the males die shortly afterwards. Females of some species mate with multiple males. 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 store the [[Spermatozoon|sperm]] they obtain during their [[nuptial flight]] to selectively fertilise future eggs. 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. This is how new colonies start in most species. Species that have multiple queens may have a queen leaving the nest along with some workers to found a colony at a new site.<ref name = HolldoblerWilsonAnts2>Hölldobler & Wilson (1990), pp. 143&ndash;179</ref> Ant colonies can be long-lived. The queens can live for up to 30&nbsp;years, and workers live from 1 to 3&nbsp;years. Males, however, are more transitory, and survive only a few weeks.<ref>{{cite journal |author=Keller L|year=1998 |title=Queen lifespan and colony characteristics in ants and termites |journal=[[Insectes Sociaux]] |volume=45 |pages=pp. 235&ndash;246 |doi=10.1007/s000400050084}}</ref> Ant queens are estimated to live 100 times longer than solitary insects of a similar size.<ref name=insencyc>{{cite book|author=Franks NR, Resh VH, Cardé RT (eds)|year=2003|title=Encyclopedia of Insects|publishers=Academic Press|pages=pp. 29&ndash;32|ISBN=0125869908}}</ref> Ants 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. Ants are active all year long in the tropics.<ref>{{cite journal|author=Kipyatkov VE|year=2001|title=Seasonal life cycles and the forms of dormancy in ants (Hymenoptera, Formicoidea)|journal=Acta Societatis Zoologicae Bohemicae|volume=65|issue=2|pages=pp. 198&ndash;217}}</ref> ==Behaviour and ecology== ===Communication=== [[Image:WeaverAntsAgainstRedAnt.JPG|thumb|Weaver ants collaborating to dismember a red ant (the two at the extremities are pulling the red ant, while the middle one cuts the red ant until it snaps)]] Ants communicate with each other using [[pheromone]]s.<ref>{{cite journal |author=Jackson DE, Ratnieks FL |title=Communication in ants |journal=Curr. Biol. |volume=16 |issue=15 |pages=R570–4 |year=2006 |month=August |pmid=16890508 |doi=10.1016/j.cub.2006.07.015}}</ref> These chemical signals are more developed in ants 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 can 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 finding the best path.<ref>{{cite journal|author=Goss S, Aron S, Deneubourg JL, Pasteels JM|year=1989|title=Self-organized shortcuts in the Argentine ant|journal=Naturwissenschaften |volume=76 |pages=pp. 579&ndash;581 |doi=10.1007/BF00462870}}</ref> 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 further away. Several ant species even use "propaganda pheromones" to confuse enemy ants and make them fight among themselves.<ref>{{cite journal |author=D'Ettorre P, Heinze J |journal=[[Acta ethologica]] |year=2001 |volume=3 |pages=pp. 67&ndash;82 |title=Sociobiology of slave-making ants |url=http://www.springerlink.com/content/cj1arl0gqb2amw7h/ | doi = 10.1007/s102110100038}}</ref> 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.<ref name="insencyc"/> Pheromones are also exchanged mixed with food and passed by [[trophallaxis]], transferring information within the colony.<ref>{{cite book|author=Detrain C, Deneubourg JL, Pasteels JM|year=1999|title=Information processing in social insects|publisher=Birkhäuser|isbn=3764357924|pages=pp. 224&ndash;227}}</ref> This allows other ants to detect what task group (e.g., foraging or nest maintenance) other colony members belong to.<ref>{{cite journal|title=Structural complexity of chemical recognition cues affects the perception of group membership in the ants ''Linephithema humile'' and ''Aphaenogaster cockerelli''|author=Greene MJ, Gordon DM|year=2007|journal=Journal of Experimental Biology|volume=210|pages=pp. 897&ndash;905 |doi=10.1242/jeb.02706 |url=http://jeb.biologists.org/cgi/content/abstract/210/5/897|pmid=17297148}}</ref> In ant species with queen castes, workers begin to raise new queens in the colony when the dominant queen stops producing a specific pheromone.<ref name=TheAntsPheroCastes>Hölldobler & Wilson (1990), p. 354</ref> 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.<ref>{{cite journal |author=Hickling R, Brown RL |year=2000 |title=Analysis of acoustic communication by ants |journal=[[Journal of the Acoustical Society of America]] |volume=108 |issue=4 |pages=pp. 1920&ndash;1929 |doi=10.1121/1.1290515}}</ref><ref>{{cite journal |author=Roces F, [[Bert Holldöbler|Hölldobler B]] |year=1996 |title=Use of stridulation in foraging leaf-cutting ants: Mechanical support during cutting or short-range recruitment signal?|journal=[[Behavioral Ecology and Sociobiology]] |volume=39 |pages=p. 293 |doi=10.1007/s002650050292}}</ref> ===Defence=== [[Image:WeaverAntDefense.JPG|thumb|left|A [[weaver ant]] in fighting position, [[mandible]]s wide open]] Ants attack and defend themselves by biting and in many species, by stinging, often injecting or spraying chemicals like [[formic acid]]. [[Bullet ant]]s (''[[Paraponera]]''), located in [[Central America|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,<ref>{{cite journal|author=Clarke PS|year=1986|title=The natural history of sensitivity to jack jumper ants (hymenoptera:formicidae:''Myrmecia pilosula'') in Tasmania|journal=Med. J. Aust|volume=145|pages=pp. 564&ndash;566}}</ref> and an [[antivenin]] has been developed.<ref>{{cite journal|title=Efficacy of ant venom immunotherapy and whole body extracts|author=Brown SGA, Heddle RJ, Wiese MD, Blackman KE|journal=Journal of Allergy and Clinical Immunology|volume=116|issue=2|year=2005|pages=pp. 464&ndash;465|doi = 10.1016/j.jaci.2005.04.025}}</ref> [[Fire ant]]s, ''[[Fire ant|Solenopsis]]'' spp., are unique in having a poison sac containing [[piperidine]] alkaloids.<ref>{{cite journal |author=Obin MS, Vander Meer RK |year=1985 |title=Gaster flagging by fire ants (''Solenopsis spp.''): Functional significance of venom dispersal behavior|journal=[[Journal of Chemical Ecology]] |volume=11 |pages=pp. 1757&ndash;1768 |doi=10.1007/BF01012125}}</ref> Their stings are painful and can be dangerous to hypersensitive persons.<ref>{{cite journal|title=Hypersensitivity to fire ant venom|author=Stafford CT|year=1996|journal=Annals of allergy, asthma, & immunology|volume=77|issue=2|pages=pp. 87&ndash;99}}</ref> Ants of the genus ''[[Odontomachus]]'' are equipped with mandibles called trap-jaws. This snap-jaw or catapult mechanism involves a large band of muscles that are released by a "trigger". The movement is incredibly fast, with the long mandibles snapping together within 0.13&nbsp;[[millisecond|ms]] in ''[[Odontomachus bauri]]''. This is far faster than any other predatory movement in the animal kingdom.<ref name="TrapJawPatek"> {{cite journal|title=Multifunctionality and mechanical origins: Ballistic jaw propulsion in trap-jaw ants|journal=Proceedings of the National Academy of Sciences|date=2006-08-22|author=Patek SN, Baio JE, Fisher BL, Suarez AV|volume=103|issue=34|pages=pp. 12787&ndash;12792|doi= 10.1073/pnas.0604290103|url=http://www.life.uiuc.edu/suarez/Patek_etal2006PNAS.pdf|format=PDF|accessdate=2008-06-07|pmid=16924120 }}</ref> Before the strike, the mandibles open wide and are locked in the open position. The release is triggered by stimulation of sensory hairs on the side of the mandibles. The mandibles allow slow and fine movement for other tasks. Trap-jaws are also seen in some ants of the ''[[Dacetini]]'' [[Tribe (biology)|tribe]], an example of [[convergent evolution]].<ref>{{cite journal|title=The trap-jaw mechanism in the Dacetine ant ''Daceton Armigerum'' and ''Strumigenys'' sp.|author=Gronenberg W|year=1996|journal=The Journal of Experimental Biology |volume=199 |issue=9 |pages=pp. 2021&ndash;2033 |url=http://jeb.biologists.org/cgi/reprint/199/9/2021.pdf|format=PDF}}</ref> [[Image:Ant mound.jpg|thumb|Ant mound holes prevent water from entering the nest during rain.]] In addition to defence against predators, ants need to protect their colonies from [[pathogen]]s. Some worker ants maintain the hygiene of the colony and their activities include [[wiktionary:undertaker|undertaking]] or ''necrophory'', the disposal of dead nest-mates.<ref>{{cite journal|author=Julian GE, Cahan S|title=Undertaking specialization in the desert leaf-cutter ant ''Acromyrmex versicolor''|journal=Animal Behaviour|year=1999|volume=58|issue=2|pages=pp. 437&ndash;442|doi=10.1006/anbe.1999.1184}}</ref> [[Oleic acid]] has been identified as the compound released by dead ants that triggers undertaking behaviour in ''Atta mexicana''.<ref>{{cite journal|author=López-Riquelme GO, Malo EA, Cruz-López L, Fanjul-Moles ML|year=2006|title=Antennal olfactory sensitivity in response to task-related odours of three castes of the ant ''Atta mexicana'' (hymenoptera: formicidae)|journal=Physiological Entomology|volume=31|issue=4|pages=pp. 353&ndash;360|url=http://www.blackwell-synergy.com/doi/abs/10.1111/j.1365-3032.2006.00526.x | doi = 10.1111/j.1365-3032.2006.00526.x}}</ref> Nests may be protected from physical threats such as flooding and over-heating by elaborate nest architecture.<ref>{{cite journal|author=Tschinkel WR|year=2004|title=The nest architecture of the Florida harvester ant, ''Pogonomyrmex badius''|journal=Journal of Insect Science|volume=4|issue=21|pages=pp. 1&ndash;19|doi= 10.1672/1536-2442(2004)004[0001:TNAOTF]2.0.CO;2|doi_brokendate=2008-06-27}}</ref><ref>{{cite journal|author=Peeters C, Hölldobler B, Moffett M, Musthak Ali TM|year=1994|title=“Wall-papering” and elaborate nest architecture in the ponerine ant ''Harpegnathos saltator''|journal=Insectes Sociaux|volume=41|pages=pp. 211&ndash;218|doi=10.1007/BF01240479}}</ref> 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.<ref>{{cite journal|title=Communal peeing: a new mode of flood control in ants|journal=Naturwissenschaften|year=2000 |volume=87 |issue=12 |pages=pp. 563&ndash;565 |author=Maschwitz U, Moog J |doi=10.1007/s001140050780}}</ref> ===Learning=== Many animals can learn behaviours by imitation but ants may be the only group apart from [[mammal]]s where interactive teaching has been observed. A knowledgeable forager of ''[[Temnothorax albipennis]]'' leads a naive nest-mate to newly discovered food by the excruciatingly slow process of [[tandem running]]. The follower obtains knowledge through its leading tutor. Both leader and follower are acutely sensitive to the progress of their partner with the leader slowing down when the follower lags, and speeding up when the follower gets too close.<ref>{{cite journal |author=Franks NR, Richardson T| year=2006 |title=Teaching in tandem-running ants|journal=[[Nature (journal)|Nature]] |volume=439 |issue=7073 |pages=p. 153|pmid=16407943 |doi=10.1038/439153a}}</ref> Controlled experiments with colonies of ''Cerapachys biroi'' suggests that individuals may choose nest roles based on their 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 less and less. A month later, the successful foragers continued in their role while the others moved to specialise in brood care.<ref>{{cite journal |author=Ravary F, Lecoutey E, Kaminski G, Châline N, Jaisson P| year=2007 |title=Individual experience alone can generate lasting division of labor in ants|journal=[[Current Biology]] |volume=17 |issue=15 |pages=pp. 1308&ndash;1312 |doi=10.1016/j.cub.2007.06.047}} </ref> ===Nest construction=== {{main|Ant colony|Ant hill}} [[Image:WeaverAntNest.JPG|thumb|Leaf nest of [[weaver ants]], [[Pamalican]], [[Philippines]]]] Complex nests are built by many ants, but other species are nomadic and do not build permanent structures. Ants may form subterranean nests or build them on trees. These nests can 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,<ref name = HolldoblerWilsonAnts2/> 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.<ref>{{cite journal|author=Franks NR, Hooper J, Webb C, Dornhaus A|year= 2005|title=Tomb evaders: house-hunting hygiene in ants|journal=[[Biology Letters]]|volume=1|issue=2|pages=pp. 190&ndash;192|doi=10.1098/rsbl.2005.0302}}</ref> The [[army ant]]s of South America and the [[driver ant]]s of Africa do not build permanent nests, but instead alternate between nomadism and stages where the workers form a temporary nest ([[bivouac (ants)|bivouac]]) from their own bodies, by holding each other together.<ref>Hölldobler & Wilson (1990), p. 573</ref> [[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''.<ref>{{cite journal|author=Robson SK, Kohout RJ|year=2005|title=Evolution of nest-weaving behaviour in arboreal nesting ants of the genus ''Polyrhachis'' Fr. Smith (Hymenoptera: Formicidae)|journal=Australian Journal of Entomology|volume=44|issue=2|pages=pp. 164&ndash;169|doi=10.1111/j.1440-6055.2005.00462.x}}</ref> ===Food cultivation=== {{main|Ant-fungus mutualism}} [[Image:Atta.cephalotes.gamut.selection.jpg|thumb|left|Seven [[Leafcutter ant]] workers of various castes (left) and two Queens (right)]] [[Leafcutter ant]]s (''[[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 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 toxic to the fungus the colony will no longer collect it. The ants feed on structures produced by the fungi called ''[[gongylidia]]''. [[Symbiosis|Symbiotic]] bacteria on the exterior surface of the ants produce antibiotics that kill bacteria that may harm the fungi.<ref>{{cite journal|journal=Nature|year=1999|volume=398|title=Ants, plants and antibiotics|author=Schultz TR|pages=pp. 747&ndash;748 | doi = 10.1038/19619}}</ref> ===Navigation=== [[Forage|Foraging]] ants travel distances of up to {{convert|200|m|ft}} from their nest<ref>{{cite journal|journal=Annual Review of Ecology and Systematics|volume=4|pages= pp. 231&ndash;257|year=1973|doi=10.1146/annurev.es.04.110173.001311|title=Ecology of foraging by ants|author=Carrol CR, Janzen DH}}</ref> and usually find their way back using scent trails. Some ants forage at night. Day foraging ants in hot and arid regions face death by desiccation, so the ability to find the shortest route back to the nest reduces that risk. Diurnal desert ants (''Cataglyphis fortis'') use visual landmarks in combination with other cues to navigate.<ref>{{cite journal |author=Åkesson S, Wehner R|year=2002 |title=Visual navigation in desert ants ''Cataglyphis fortis'': are snapshots coupled to a celestial system of reference? |journal=[[Journal of Experimental Biology]] |volume=205 |pages=pp. 1971&ndash;1978 |url=http://jeb.biologists.org/cgi/reprint/205/14/1971.pdf|format=PDF}}</ref> In the absence of visual landmarks, the closely related [[Sahara desert ant]] (''Cataglyphis bicolor'') navigates by keeping track of direction as well as distance travelled, like an internal [[pedometer]] that counts how many steps they take in each direction. They integrate this information to find the shortest route back to their nest.<ref>{{cite journal |author=Sommer S, Wehner R|year=2004 |title=The ant's estimation of distance travelled: experiments with desert ants, ''Cataglyphis fortis''|journal=Journal of Comparative Physiology |volume=190 |issue=1 |pages=pp. 1&ndash;6 |url=http://www.springerlink.com/content/bywx5wqjchmh85t2/ | doi = 10.1007/s00359-003-0465-4 }}</ref> Several species of ants are able to use the Earth's magnetic field.<ref>{{cite journal|title=Orientation by magnetic field in leaf-cutter ants, ''Atta colombica (Hymenoptera: Formicidae)|author=Banks AN, Srygley RB|year=2003|journal=Ethology|volume=109|pages=pp. 835&ndash;46|doi=10.1046/j.0179-1613.2003.00927.x}}</ref> Ants' compound eyes have specialised cells that detect polarised light from the Sun, which is used to determine direction.<ref>{{cite journal |author=Fukushi T |year=2001 |title=Homing in wood ants, ''Formica japonica'': use of the skyline panorama|journal=[[Journal of Experimental Biology]] |volume=204 |pages=pp. 2063&ndash;2072 |url=http://jeb.biologists.org/cgi/content/abstract/204/12/2063 |pmid=11441048}}</ref><ref>{{cite journal |author=Wehner R, Menzel R|year=1969 |title=Homing in the ant ''Cataglyphis bicolor'' |journal=[[Science (journal)|Science]] |volume=164 |issue=3876 |pages=pp. 192&ndash;194 |doi=10.1126/science.164.3876.192 |pmid=5774195}}</ref> ===Locomotion=== [[Image:Harpegnathos saltator fight.jpg|thumb|''[[Harpegnathos saltator]]'', a jumping ant]] 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.<ref>{{cite journal |author=Baroni-Urbani C, Boyan GS, Blarer A, Billen J, Musthak Ali TM|year=1994 |title=A novel mechanism for jumping in the Indian ant ''Harpegnathos saltator'' (Jerdon) (Formicidae, Ponerinae) |journal=[[Experientia]] |volume=50 |pages=pp. 63&ndash;71 |doi=10.1007/BF01992052}}</ref> 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.<ref>{{cite journal |author=Yanoviak SP, Dudley R, Kaspari M|year=2005 |title=Directed aerial descent in canopy ants|journal=[[Nature (journal)|Nature]] |volume=433 |pages=pp. 624&ndash;626 |url=http://www.canopyants.com/Nature05.pdf|format=PDF|doi=10.1038/nature03254}}</ref> 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.<ref>{{cite journal|author=Morrison LW|year=1998|title=A review of Bahamian ant (Hymenoptera: Formicidae) biogeography|journal=Journal of Biogeography|volume=25|issue=3|pages=pp. 561&ndash;571|doi=10.1046/j.1365-2699.1998.2530561.x}}</ref> ''[[Polyrhachis sokolova]]'', a species of ant found in [[Australia]]n [[mangrove]] swamps, can swim and these ants live in nests that are underwater. Since they lack [[gill]]s, they breathe in trapped pockets of air in the submerged nests.<ref>{{cite journal |author=Clay RE, Andersen AN|year=1996 |title=Ant fauna of a mangrove community in the Australian seasonal tropics, with particular reference to zonation |journal=[[Australian Journal of Zoology]] |volume=44 |pages=pp. 521&ndash;533 |doi=10.1071/ZO9960521}}</ref> ===Cooperation and competition=== [[Image:Meat eater ants feeding on honey.jpg|thumb|left|These meat eater ants are feeding on honey. Social ants cooperate and collectively gather food.]] Not all ants have the same kind of societies. The Australian [[bulldog ant]]s are among the biggest and most [[basal (phylogenetics)|basal]] (primitive) of ants. Like all ants they are [[eusocial]], but their social behaviour is poorly developed compared to other species. Each individual hunts alone, using its large eyes instead of its chemical senses to find prey.<ref name=Crosland1988>{{cite journal | doi = 10.1111/j.1440-6055.1988.tb01179.x | title = Aspects of the biology of the primative ant genus Myrmecia F. (Hymenoptera: Formicidae) | year = 1988 | journal = Australian Journal of Entomology | volume = 27 | pages =pp. 305&ndash;309 | author = Crosland MWJ, Crozier RH, Jefferson E | url = http://www.blackwell-synergy.com/action/showPdf?doi=10.1111%2Fj.1440-6055.1988.tb01179.x }}</ref><ref>{{cite Web|url=http://ngm.nationalgeographic.com/ngm/0705/feature6/index.html|title=Ant, Bulldog Ants|accessdate=2008-06-12 |publisher=National Geographic |author=Moffett MW}}</ref> 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/slaves. Extreme specialists among these slave-raiding ants, such as the [[Amazon ant]]s, are incapable of feeding themselves and need captured workers to survive.<ref>{{cite journal |author=Diehl E, Junqueira LK, Berti-Filho E|year=2005 |title=Ant and termite mound coinhabitants in the wetlands of Santo Antonio da Patrulha, Rio Grande do Sul, Brazil|journal=[[Brazilian Journal of Biology]] |volume=65 |issue=3 |pages=pp. 431&ndash;437 |url=http://www.scielo.br/scielo.php?script=sci_arttext&pid=S1519-69842005000300008&lng=en&nrm=iso |doi=10.1590/S1519-69842005000300008}}</ref> 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.<ref>{{cite journal |author=Henderson G, Andersen JF, Phillips JK, Jeanne RL|year=2005 |title=Internest aggression and identification of possible nestmate discrimination pheromones in polygynous ant ''Formica montana'' |journal=[[Journal of Chemical Ecology]] |volume=16 |issue=7 |pages=pp. 2217&ndash;2228 |doi=10.1007/BF01026932}}</ref> Parasitic ant species enter the colonies of host ants and establish themselves as social parasites; species like ''Strumigenys xenos'' are entirely parasitic and do not have workers, but instead rely on the food gathered by their ''Strumigenys perplexa'' hosts.<ref>{{cite journal|author=Ward PS|year=1996|title=A new workerless social parasite in the ant genus ''Pseudomyrmex'' (Hymenoptera: Formicidae), with a discussion of the origin of social parasitism in ants|journal=Systematic Entomology|volume= 21|pages=pp. 253&ndash;263|url=http://www.archive.org/details/ants_08424|doi=10.1046/j.1365-3113.1996.d01-12.x}}</ref><ref>{{cite journal|author=Taylor RW|year=1968|title=The Australian workerless inquiline ant, ''Strumigenys xenos'' Brown (Hymenoptera-Formicidae) recorded from New Zealand|journal= New Zealand Entomologist|volume=4|issue=1|pages=pp. 47&ndash;49|url=http://www.archive.org/details/ants_10687}}</ref> 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 can 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.<ref name=TheAntParasites>Hölldobler & Wilson (1990), pp. 436&mdash;448</ref> ===Relationships with other organisms=== Ants form [[symbiotic]] associations with a range of species, including other ant species, insects, plants, and fungi. They 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 ants' food stores, or avoiding predators. These [[inquiline]]s can 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 [[Mimicry#Wasmannian|Wasmannian mimicry]], a form of mimicry seen only in inquilines.<ref>{{cite journal|title=Ant-mimicry in Panamanian clubionid and salticid spiders (Araneae: Clubionidae, Salticidae)|author=Reiskind J|journal=Biotropica|volume=9|issue=1|year=1977|pages=pp. 1&ndash;8|doi=10.2307/2387854}}</ref><ref>{{cite journal|title=Myrmecomorphy and myrmecophily in spiders: A Review|author=Cushing PE|journal=The Florida Entomologist|volume=80|issue=2|year=1997|pages=pp. 165&ndash;193|doi=10.2307/3495552|url=http://www.fcla.edu/FlaEnt/fe80p165.pdf|format=PDF}}</ref> [[Image:Myrmarachne-plataleoides1.jpg|thumb|The spider ''[[Myrmarachne plataleoides]]'' mimics [[weaver ant]]s to avoid predators, this male appears to be one ant carrying another.]] [[Aphid]]s and other [[hemiptera]]n insects secrete a sweet liquid called [[Honeydew (secretion)|honeydew]] when they feed on [[plant sap]]. The sugars in honeydew are a high-energy food source, which many ant species collect.<ref>{{cite journal |author=Styrsky JD, Eubanks MD |title=Ecological consequences of interactions between ants and honeydew-producing insects |journal=Proc. Biol. Sci. |volume=274 |issue=1607 |pages=151–64 |year=2007 |month=January |pmid=17148245 |pmc=1685857 |doi=10.1098/rspb.2006.3701 |url=http://journals.royalsociety.org/openurl.asp?genre=article&id=doi:10.1098/rspb.2006.3701}}</ref> In some cases the aphids secrete the honeydew in response to the ants' tapping them with their antennae. The ants in turn keep predators away and will move the aphids between feeding locations. On migrating to a new area, many colonies will take the [[aphid]]s with them, to ensure a continued supply of honeydew. Ants also tend [[mealybug]]s to harvest their honeydew. Mealybugs can become a serious pest of pineapples if ants are present to protect mealybugs from their natural enemies.<ref>{{cite journal |author=Jahn GC, Beardsley JW |year=1996 |title=Effects of ''Pheidole megacephala'' (Hymenoptera: Formicidae) on survival and dispersal of ''Dysmicoccus neobrevipes'' (Homoptera: Pseudococcidae) |journal=[[Journal of Economic Entomology]] |volume=89 |pages=pp. 1124&ndash;1129}}</ref> [[Myrmecophile|Myrmecophilous]] (ant-loving) [[caterpillar]]s of the 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.<ref>{{cite journal |author=DeVries PJ|year=1992 |title=Singing caterpillars, ants and symbiosis |journal=[[Scientific American]] |volume=267 |pages=p. 76}}</ref> 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 ants' nest where it feeds on the ant larvae.<ref>{{cite journal|journal=Annual Review of Entomology|year=2002|volume=47|pages=pp. 733&ndash;771|title=The ecology and evolution of ant association in the Lycaenidae (Lepidoptera)|author=Pierce NE, Braby MF, Heath A, et al|doi=10.1146/annurev.ento.47.091201.145257}}</ref> [[Fungus-growing ants]] that make up the tribe [[Attini]], including [[leafcutter ant]]s, cultivate certain species of fungus in the ''[[Leucoagaricus]]'' or ''[[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.<ref>{{cite journal |author=Dejean A, Solano PJ, Ayroles J, Corbara B, Orivel J |year=2005 |title=Arboreal ants build traps to capture prey |journal=[[Nature (journal)|Nature]] |volume=434|pages=p. 973 |doi=10.1038/434973a}}</ref> [[Image:Ant Receives Honeydew from Aphid.jpg|thumb|An ant collects [[honeydew (secretion)|honeydew]] from an [[aphid]].]] [[Myrmelachista schumanni|Lemon ants]] make [[devil's garden]]s 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.<ref>{{cite journal|author= Frederickson ME, Gordon DM|year=2007|title = The devil to pay: a cost of mutualism with ''Myrmelachista schumanni'' ants in ‘devil’s gardens’ is increased herbivory on ''Duroia hirsuta'' trees|journal=Proceedings of the Royal Society B|volume=274 |pages=pp. 1117&ndash;1123|url=http://www.stanford.edu/~dmgordon/frederickson_gordon2007.pdf|format=PDF | doi = 10.1111/j.1461-0248.2005.00741.x}}</ref> Some trees have extrafloral [[nectary|nectaries]] that provide food for ants, which in turn [[plant defense against herbivory|protect]] the plant from [[herbivorous]] insects.<ref>{{cite journal|author=Katayama N, Suzuki N|title=Role of extrafloral nectaries of ''Vicia faba'' in attraction of ants and herbivore exclusion by ants|year=2004|journal=Entomological Science|pages=pp. 119&ndash;124 |volume=7|issue=2|url=http://www.blackwell-synergy.com/doi/abs/10.1111/j.1479-8298.2004.00057.x | doi = 10.1111/j.1479-8298.2004.00057.x }}</ref> Species like the bullhorn acacia (''[[Acacia cornigera]]'') in Central America have hollow thorns that house colonies of stinging ants (''[[Pseudomyrmex ferruginea]]'') that defend the tree against insects, browsing mammals, and [[Epiphyte|epiphytic]] vines. [[Isotopic labeling]] studies suggest that plants also obtain nitrogen from the symbiotic ants.<ref>{{cite journal|title=Do ants feed plants? A <sup>15</sup>N labelling study of nitrogen fluxes from ants to plants in the mutualism of ''Pheidole'' and ''Piper''|author= Fischer RC, Wanek W, Richter A, Mayer V|year=2003|journal=Journal of Ecology|volume=91|pages=pp. 126&ndash;134|doi=10.1046/j.1365-2745.2003.00747.x }}</ref> In return, the ants obtain food from protein-lipid [[Beltian bodies]]. Another example of this type of [[ectosymbiosis]] comes from the ''[[Macaranga]]'' tree, which has stems adapted to house colonies of ''[[Crematogaster]]'' ants. Many tropical tree species have seeds that are dispersed by ants.<ref>{{cite journal |author=Hanzawa FM, Beattie AJ, Culver DC |year=1988 |title=Directed dispersal: demographic analysis of an ant-seed mutualism |journal=[[American Naturalist]] |volume=131 |issue=1 |pages=pp. 1&ndash;13 |doi=10.1086/284769}}</ref> [[Seed#Seed_dispersal|Seed dispersal]] by ants or [[myrmecochory]] is widespread particularly in Africa and Australia.<ref>{{cite journal|author=Giladi I|title= Choosing benefits or partners: a review of the evidence for the evolution of myrmecochory|journal=Oikos |volume=112|issue=3|year=2006|pages=pp. 481&ndash;492|doi=10.1111/j.0030-1299.2006.14258.x}}</ref> Some plants in fire-prone grassland systems are particularly dependent on ants for their survival and dispersal. Many ant-dispersed seeds have special external structures, [[elaiosome]]s, that are sought after by ants as food.<ref>{{cite journal|author=Fischer RC, Ölzant SM, Wanek W, Mayer V|title=The fate of ''Corydalis cava'' elaiosomes within an ant colony of ''Myrmica rubra'': elaiosomes are preferentially fed to larvae|journal=Insectes sociaux|year=2005|volume=52|issue=1|pages=pp. 55&ndash;62 |doi=10.1007/s00040-004-0773-x}}</ref> A [[convergent evolution|convergence]], possibly a form of [[mimicry]], is seen in the eggs of [[stick insect]]s. They have an edible elaiosome-like structure and are taken into the ant nest where the young hatch.<ref>{{cite journal|author=Hughes L, Westoby M|year=1992|title=Capitula on stick insect eggs and elaiosomes on seeds: convergent adaptations for burial by ants|journal=Functional Ecology|volume=6|pages=pp. 642&ndash;648|doi=10.2307/2389958}}</ref> [[Image:Common jassid nymph and ant.jpg|thumb|left|A [[Meat ant]] tending a common [[leafhopper]] nymph]] Flies in the Old World genus ''[[Bengalia]]'' ([[Calliphoridae]]) [[predator|prey]] on ants and are [[kleptoparasite]]s, snatching prey or brood from the mandibles of adult ants.<ref name="sivinski">{{cite journal |author=Sivinski J, Marshall S, Petersson E|year=1999 |title=Kleptoparasitism and phoresy in the Diptera |journal=[[Florida Entomologist]] |volume=82 |issue=2 |pages=pp. 179&ndash;197 |url=http://www.fcla.edu/FlaEnt/fe82p179.pdf|format=PDF|doi=10.2307/3496570}}</ref> Wingless and legless females of the [[Malaysia]]n [[phoridae|phorid]] fly (''[[Vestigipoda myrmolarvoidea]]'') live in the nests of ants of the genus ''[[Aenictus]]'' and are cared for by the ants.<ref name="sivinski"/> The fungus ''[[Cordyceps]]'' infects ants, causing them to climb up plants and sink their mandibles into plant tissue. The fungus kills the ant, grows on its remains, and produces a [[fruiting body]]. It appears that the fungus alters the behaviour of the ant to help disperse its spores.<ref>{{Cite journal |author=Schaechter E |year=2000 |title=Some weird and wonderful fungi |journal=[[Microbiology Today]] |volume=27 |issue=3 |pages=pp. 116&ndash;117}}</ref> [[Strepsiptera]]n parasites also manipulate their ant host to climb grass stems, to help the parasite find mates.<ref>{{cite journal|author=Wojcik DP|year=1989|title=Behavioral interactions between ants and their parasites|journal=The Florida Entomologist|volume=72|issue=1|pages=pp. 43&ndash;51|doi=10.2307/3494966}}</ref> 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, and makes them 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 the further spread of the nematode.<ref>{{cite journal|author=Poinar G Jr., Yanoviak SP|year=2008|title=''Myrmeconema neotropicum'' n. g., n. sp., a new tetradonematid nematode parasitising South American populations of ''Cephalotes atratus'' (Hymenoptera: Formicidae), with the discovery of an apparent parasite-induced host morph|journal=Systematic Parasitology|volume=69|pages=pp. 145&ndash;153|url=http://www.canopyants.com/2008_SystParasit.pdf|format=PDF|doi=10.1007/s11230-007-9125-3}}</ref> South American [[poison dart frog]]s in the genus ''[[Dendrobates]]'' feed mainly on ants, and the toxins in their skin may come from the ants.<ref>{{cite journal |author=Caldwell JP|year=1996 |title=The evolution of myrmecophagy and its correlates in poison frogs (Family Dendrobatidae) |journal=[[Journal of Zoology]] |volume=240 |issue=1 |pages=pp. 75&ndash;101}}</ref> Several South American [[antbird]]s follow army ants to feed on the insects that are flushed from cover by the foraging ants.<ref>{{cite journal|author=Vellely AC|title=Foraging at army ant swarms by fifty bird species in the highlands of Costa Rica|journal=Ornitologia Neotropical|volume=12|year=2001|pages=pp 271&ndash;275|url=http://www.ibiologia.unam.mx/pdf/links/neo/rev12/vol_12_3/orni_12_3_%20271-276.pdf|format=PDF|accessdate=2008-06-08}} </ref> This behaviour was once considered [[mutualism|mutualistic]], but later studies show that it is instead [[Kleptoparasitism|kleptoparastic]], with the birds stealing prey.<ref>{{cite journal|author=Wrege PH| coauthors = Wikelski M, Mandel JT, Rassweiler T, Couzin ID|title=Antbirds parasitize foraging army ants|journal=Ecology|volume=86|year=2005|pages=pp. 555&ndash;559|url=|doi=10.1890/04-1133}} </ref> Birds indulge in a peculiar behaviour called [[Anting (bird activity)|anting]] that is as yet not fully understood. Here birds rest on ant nests, or pick and drop ants onto their wings and feathers; this may remove [[Parasitism#Types of parasitism|ectoparasites]]. [[Anteater]]s, [[pangolin]]s and several [[marsupial]] species in Australia have special [[adaptation]]s for living on a diet of ants. These adaptations include long, sticky tongues to capture ants and strong claws to break into ant nests. [[Brown bear]]s (''Ursus arctos'') have been found to feed on ants, and about 12%, 16%, and 4% of their faecal volume in spring, summer, and autumn, respectively, is composed of ants.<ref>{{cite journal |author=Swenson JE, Jansson A, Riig R, Sandegren R |year=1999 |title=Bears and ants: myrmecophagy by brown bears in central Scandinavia |journal=[[Canadian Journal of Zoology]] |volume=77 |issue=4 |pages=pp. 551&ndash;561 |url=http://rparticle.web-p.cisti.nrc.ca/rparticle/AbstractTemplateServlet?journal=cjz&volume=77&msno=z99-004&calyLang=eng | doi = 10.1139/cjz-77-4-551 }}</ref> ==Relationship with humans== [[Image:AntsStitchingLeave.jpg|thumb|upright|[[Weaver ant]]s 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 (organism)|pest]] populations and aeration of the [[soil]]. The use of [[weaver ant]]s in citrus cultivation in southern China is considered one of the oldest known applications of [[biological control]].<ref name = HolldoblerWilsonAnts3>Hölldobler & Wilson (1990), pp. 619&ndash;629</ref> On the other hand, ants can become nuisances when they invade buildings, or cause economic losses. In some parts of the world, large ants, especially [[army ant]]s, are used as [[suture]]s. The wound is pressed together and ants are applied along it. The ant seizes the edges of the wound in its mandibles and locks in place. The body is then cut off and the head and mandibles remain in place to close the wound.<ref>{{cite journal |author=Gottrup F, Leaper D|year=2004 |title=Wound healing: Historical aspects |journal=[[EWMA Journal]] |volume=4 |issue=2 |url=http://www.ewma.org/pdf/fall04/Historical_Aspects.pdf|format=PDF|pages=p. 5}}</ref><ref>{{cite journal|author=Gudger EW|year=1925|title=Stitching wounds with the mandibles of ants and beetles|journal=Journal of the American Medical Association|volume=84|pages=pp. 1861&ndash;1864}}</ref> Some ants of the family Ponerinae have toxic venom and are of medical importance. The species include ''Paraponera clavata'' (''Tocandira'') and ''Dinoponera'' spp. (false ''Tocandira''s) of South America<ref>{{cite journal|author=Haddad Jr. V, Cardoso JLC, Moraes RHP|year=2005|title=Description of an injury in a human caused by a false tocandira (''Dinoponera gigantea'', Perty, 1833) with a revision on folkloric, pharmacological and clinical aspects of the giant ants of the genera Paraponera and Dinoponera (sub-family Ponerinae)|journal=Revista do Instituto de Medicina Tropical de São Paulo|volume=47|issue=4 |pages=pp. 235&ndash;238|url=http://www.scielo.br/pdf/rimtsp/v47n4/25664.pdf|format=PDF|doi=10.1590/S0036-46652005000400012}}</ref> and the ''Myrmecia'' ants of Australia.<ref>{{cite journal|journal=Toxicon|volume=40|issue=8|pages=pp. 1095&ndash;1100|title=Ant sting mortality in Australia|author=McGain F, Winkel KD|year=2002|doi=10.1016/S0041-0101(02)00097-1}}</ref> In [[South 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 of seeds can be collected from one ant-heap.<ref>{{cite web|url=http://www.ciel.org/Publications/InnovativeMechanisms.pdf|format=PDF|title=Innovative mechanisms for sharing benefits of biodiversity and related knowledge|author=Downes D, Laird SA|year=1999|publisher=The Center for International Environmental Law|accessdate=2008-06-08}}</ref><ref>{{cite journal|journal=Economic Botany|title=Rooibos tea, a South African contribution to world beverages|volume=17|issue=3|pages=pp. 186&ndash;194|year=1963|author=Cheney RH, Scholtz E|doi=10.1007/BF02859435|doi_brokendate=2008-06-27}}</ref> === As food === {{main|Entomophagy}} [[Image:Ants Eggs Market Thailand.jpg|thumb|left|Ant larvae on sale in [[Isaan]], Thailand]] Ants and their larvae are eaten in different parts of the world. The eggs of two species of ants are the basis for the dish in Mexico known as ''[[escamoles]]''. They are considered a form of insect caviar and can sell for as much as 40&nbsp;[[USD]] per pound because they are seasonal and hard to find. In the [[Colombia]]n department of [[Santander Department|Santander]], ''hormigas culonas'' (Spanish for "fatass ants") ''[[Atta laevigata]]'' are toasted alive and eaten.<ref>{{cite journal|author=DeFoliart GR|title=Insects as food: Why the western attitude is important|journal=Annual Review of Entomology|volume=44|pages=pp. 21&ndash;50|year=1999|doi=10.1146/annurev.ento.44.1.21}}</ref> In areas of [[India]], and throughout [[Burma]] and [[Thailand]], a paste of the green weaver ant (''[[Oecophylla smaragdina]]'') is served as a condiment with [[curry]].<ref>{{cite book|author=Bingham CT|year=1903|title=Fauna of British India. Hymenoptera Volume 3|pages=p. 311}}</ref> [[Oecophylla|Weaver ant]] eggs and larvae as well as the ants themselves may be used in a Thai salad, ''yum'' (ยำ), in a dish called ''yum khai mod daeng'' (ยำไข่มดแดง) or red ant egg salad, a dish that comes from the Issan or north-eastern region of Thailand. [[William Saville-Kent|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".<ref name="beq">{{cite journal |author=Bequaert J|year=1921 |title=Insects as food: How they have augmented the food supply of mankind in early and recent times|journal=Natural History Journal|volume=21|pages=pp. 191&ndash;200}}</ref> In his ''First Summer in the Sierra'', [[John Muir]] notes that the [[Paiute|Digger Indians]] of [[California]] ate the tickly 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]]'').<ref name="beq"/> === As pests === [[Image:Pharaoh Ant close up.JPG|thumb|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 pests,<ref name="pests">{{cite web|url=http://www.ipm.ucdavis.edu/PMG/PESTNOTES/pn7411.html|title=Pest Notes: Ants (Publication 7411)|publisher=University of California Agriculture and Natural Resources|year=2007|accessdate=2008-06-05}}</ref> and because of the adaptive nature of ant colonies, eliminating the entire colony is nearly impossible. Pest management is therefore a matter of controlling local populations, instead of eliminating an entire colony, and most attempts at control are temporary solutions. Ants classified as pests include [[pavement ant]]s, [[yellow crazy ant]]s, [[sugar ant]]s, [[Pharaoh ant]]s, [[carpenter ants]], [[Argentine ant]]s, and [[red imported fire ant]]s. Populations are controlled using insecticide baits, either in granule or liquid formulations. Bait is gathered by the ants as food and brought back to the nest where the poison is inadvertently spread to other colony members through trophallaxis. [[Boric acid]] and [[borax]] are often used as [[insecticide]]s that are relatively safe for humans. Bait may be broadcast over a large area to control species like the red fire ant that occupy large areas. Nests of red fire ants may be destroyed by following the ants' trails back to the nest and then pouring boiling water into it to kill the queen. This works in about 60% of the mounds and requires about {{convert|14|l|impgal|lk=on|sigfig=1}} per mound.<ref>{{cite web|title=Two step method for fire ant control|url=http://www.ento.okstate.edu/fireants/twostep.htm|publisher =Oklahoma State University|accessdate=2008-06-05}}</ref> === In science and technology === {{See also|Myrmecology|Biomimetics|Ant colony optimisation}} [[Myrmecology|Myrmecologists]] study ants in the laboratory and in their natural conditions. Their complex and variable social structures have made ants ideal [[model organism]]s. Studies on ants have tested hypotheses in [[ecology]], [[sociobiology]] and have been particularly important in examining the predictions of theories of [[kin selection]] and [[Evolutionarily stable strategy|evolutionarily stable strategies]]. Ant colonies can be studied by rearing or temporarily maintaining them in [[formicarium|formicaria]], specially constructed glass framed enclosures.<ref name=myrmtech>{{cite journal|title=Myrmecological technique. IV. Collecting ants by rearing pupae|author=Kennedy CH|journal=The Ohio Journal of Science|volume=51|issue=1|year=1951|pages=pp. 17&ndash;20|url=http://hdl.handle.net/1811/3802}}</ref> Individuals may be tracked for study by marking them with colours.<ref>{{cite journal|author=Wojcik DP, Burges RJ, Blanton CM, Focks DA|year=2000|title=An improved and quantified technique for marking individual fire ants (Hymenoptera: Formicidae)|journal=The Florida Entomologist|volume=83|issue=1|pages=pp. 74&ndash;78|doi=10.2307/3496231|url=http://www.fcla.edu/FlaEnt/fe83p74.pdf|format=PDF}}</ref> 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. 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.<ref name="SANdisk">{{cite journal |author=Dicke E, Byde A, Cliff D, Layzell P |year=2004 |chapter=An ant-inspired technique for storage area network design |title=Proceedings of Biologically Inspired Approaches to Advanced Information Technology: First International Workshop, BioADIT 2004 LNCS 3141 |pages=pp. 364&ndash;379 |editor=A. J. Ispeert, M. Murata & N. Wakamiya}}</ref> === In culture === [[Image:The Ant and the Grasshopper - Project Gutenberg etext 19994.jpg|thumb|left|[[Aesop]]'s ants: picture by [[Milo Winter]], 1888&ndash;1956]] Ants have often been used in fables and children's stories to represent industriousness and cooperative effort. They are also mentioned in religious texts.<ref>{{cite book |title=[[Quran]] 27:18–19 |url=http://www.wright-house.com/religions/islam/Quran/27-ant.html}}</ref><ref>{{cite book |title=[[Sahih Bukhari]] Vol 4, Book 54, Number 536 |url=http://www.usc.edu/dept/MSA/fundamentals/hadithsunnah/bukhari/054.sbt.html}}</ref><!-- This citation needs to be reformatted, see [[WP:CITE/ES]] --> In the [[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 Grasshopper and the Ants". In parts of Africa, ants are considered to be the messengers of the gods. Ant bites are often said to have curative properties. The sting of some species of ''[[Pseudomyrmex]]'' is claimed to give fever relief.<ref>{{cite journal |author=Balee WL|title=Antiquity of traditional ethnobiological knowledge in Amazonia: The Tupi-Guarani family and time |journal=[[Ethnohistory]] |volume=47 |issue=2 |year=2000 |pages=pp. 399&ndash;422|doi=10.1215/00141801-47-2-399}}</ref> Some [[Native American mythology]], such as the [[Hopi mythology]], consider ants as the very first animals. Others use ant bites in [[initiation]] ceremonies as a test of endurance.<ref>{{fr icon}} {{cite book |author=Cesard N, Deturche J, Erikson P |year=2003 |chapter=Les Insectes dans les pratiques médicinales et rituelles d’Amazonie indigène |editor=Motte-Florac, E. & J. M. C. Thomas |title=Les insectes dans la tradition orale|publisher=Peeters-Selaf, Paris|pages=pp. 395&ndash;406}}</ref><ref>{{cite journal|author=Schmidt RJ|year=1985|title=The super-nettles: a dermatologist's guide to ants in the plants|journal=International Journal of Dermatology|volume=24|issue=4|pages=pp. 204&ndash;210|doi= 10.1111/j.1365-4362.1985.tb05760.x}}</ref> The Japanese word for ant, ''ari'', is represented by an ideograph formed of the character for ''insect'' combined with the character signifying ''moral rectitude'', ''propriety'' (''giri''). So the Japanese character could possibly be read as ''The Propriety-Insect''.<ref>{{cite book |author=Hearn L|year=1904 |title=Kwaidan: Stories and studies Of strange things|publisher=Tuttle publishing (2005 reprint)|isbn=0804836620|pages=p. 223}}</ref> Ant society has always fascinated humans and has been written about both humorously and seriously. [[Mark Twain]] wrote about ants in his ''[[A Tramp Abroad]]''.<!--<ref>{{cite book|chapter=22 The Black Forest and Its Treasures|title=A Tramp Abroad|last=Twain|first=Mark|year=1880|url=http://www.gutenberg.org/files/119/}}</ref>--> 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]] in his fantasy novel ''[[The Once and Future King]]''. The plot in French entomologist and writer [[Bernard Werber]]'s science-fiction novel ''[[Les Fourmis]]'' is divided between the worlds of ants and humans, ants and their behaviour is described using contemporary scientific knowledge. In more recent times, animated cartoons and 3D animated movies featuring ants have been produced include ''[[Antz]]'', ''[[A Bug's Life]]'', ''[[The Ant Bully]]'', ''[[The Ant and the Aardvark]]'', ''[[Atom Ant]]'', and there is a [[comic book]] superhero called [[Ant-Man]]. From the late 1950s through the late 1970s, [[formicarium|ant farms]] were popular educational children's toys in the United States. In the early 1990s, the video game [[SimAnt]], which simulated an ant colony, won the 1992 [[Codie awards|Codie award]] for "Best Simulation Program".<ref>{{cite web |url=http://www.siia.net/codies/2007/history_1992.asp |title= 1992 Excellence in Software Awards Winners |accessdate=2008-04-03|publisher=Software & Information Industry Association}}</ref> Ants are also quite popular inspiration for many [[science-fiction]] creatures, such as the Formics of ''[[Ender's Game]]'', the Bugs of ''[[Starship Troopers]]'', and the giant ants in the film ''[[Them!]]''. In [[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 [[Group mind (science fiction)|hive mind]], a common misconception about ant colonies.<ref>{{cite journal|title=Robots, insects and swarm intelligence|journal=Artificial Intelligence Review|volume=26|issue=4|year=2006|author=Sharkey AJC|doi=10.1007/s10462-007-9057-y|pages=pp. 255&ndash;268}}</ref> ==Footnotes== {{reflist|2}} ==References== * {{cite book|author=Borror DJ, Triplehorn CA, Delong DM|title=Introduction to the Study of Insects, 6th Edition|publisher=Saunders College Publishing|year=1989|isbn=0030253977}} * {{cite book |author=Hölldobler B, Wilson EO|title=The Ants |publisher=[[Harvard University Press]]|year=1990|isbn=0674040759}} == Further reading == * {{cite book|author=Bolton, Barry|publisher=Harvard University Press|year=1995|isbn=9780674615144|title=A New General Catalogue of the Ants of the World}} * {{cite book|author=Hölldobler B, Wilson EO|year=1998|title=Journey to the Ants: A Story of Scientific Exploration|publisher=Belknap Press|isbn=0674485262}} ==External links== {{commonscat|Formicidae}} {{Wikispecies|Formicidae}} * [http://www.antweb.org/ Antweb from The California Academy of Sciences] * [http://antbase.org/ AntBase - a taxonomic database with literature sources] * [http://www.discoverlife.org/20/q?search=Formicidae Discover Life&mdash; images, information and links] * [http://bugguide.net/node/view/165 BugGuide] {{featured article}} [[Category:Ants| ]] [[Category:Vespoidea]] [[Category:Hymenoptera]] [[Category:Symbiosis]] {{Link FA|ca}} [[ang:Ǣmette]] [[ar:نمل]] [[an:Formicidae]] [[as:পৰুৱা]] [[ast:Formiga]] [[gn:Tahýi]] [[ay:K'isimira]] [[zh-min-nan:Káu-hiā]] [[bs:Mrav]] [[br:Merien]] [[bg:Мравки]] [[ca:Formiga]] [[cs:Mravencovití]] [[cy:Morgrugyn]] [[da:Myre]] [[de:Ameisen]] [[et:Sipelglased]] [[el:Μυρμήγκι]] [[es:Formicidae]] [[eo:Formiko]] [[eu:Inurri]] [[fr:Formicidae]] [[gl:Formiga]] [[hak:Ngie-kûng]] [[ko:개미]] [[hr:Mravi]] [[io:Formiko]] [[id:Semut]] [[ia:Formica]] [[is:Maurar]] [[it:Formicidae]] [[he:נמליים]] [[jv:Semut]] [[pam:Panas]] [[kn:ಇರುವೆ]] [[ht:Foumi]] [[la:Formica]] [[lv:Skudru dzimta]] [[lt:Skruzdėlės]] [[hu:Hangyák]] [[mg:Vitsika]] [[ml:ഉറുമ്പ്]] [[nah:Āzcatl]] [[nl:Mieren]] [[ja:アリ]] [[no:Maur]] [[nn:Maur]] [[oc:Formiga]] [[nds:Miechhammeke]] [[pl:Mrówkowate]] [[pt:Formiga]] [[ro:Furnică]] [[qu:Sisi]] [[ru:Муравьи]] [[scn:Furmìcula]] [[simple:Ant]] [[sk:Mravcovité]] [[sl:Mravlje]] [[sr:Мрав]] [[sh:Mrav]] [[su:Sireum]] [[fi:Muurahaiset]] [[sv:Myror]] [[tl:Langgam]] [[ta:எறும்பு]] [[te:చీమ]] [[th:มด]] [[vi:Kiến]] [[tg:Мӯрча]] [[chr:ᏙᏒᏓᎵ]] [[tr:Karınca]] [[uk:Мурахи]] [[ur:چیونٹی]] [[yi:מוראשקע]] [[zh:蚂蚁]]