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{{short description|Family of insects}}
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{{automatic taxobox
| name = Ants
| fossil_range = {{fossilrange|100|0|earliest=140|refs=<ref name=phyl2006 />}} Late [[Albian]]&nbsp;– [[Holocene|Present]]
| image = Fire ants 01.jpg
| image_caption = [[Fire ant]]s
| display_parents = 2
| parent_authority = Latreille, 1809<ref name=Boudinot2020/>
| taxon = Formicidae
| authority = [[Pierre André Latreille|Latreille]], 1809
| type_species = ''[[Formica rufa]]''
| type_species_authority = [[Carl Linnaeus|Linnaeus]], [[Systema Naturae|1761]]
| subdivision_ranks = [[List of ant subfamilies|Subfamilies]]
| subdivision = * [[Agroecomyrmecinae]]
* [[Amblyoponinae]] (incl. "[[Apomyrminae]]")
* [[Aneuretinae]]
* {{extinct}}[[Brownimeciinae]]
* [[Dolichoderinae]]
* [[Dorylinae]]
* [[Ectatomminae]]
* {{extinct}}[[Formiciinae]]
* [[Formicinae]]
* {{extinct}}[[Haidomyrmecinae]]
* [[Heteroponerinae]]
* [[Leptanillinae]]
* [[Martialinae]]
* [[Myrmeciinae]] (incl. "[[Nothomyrmeciinae]]")
* [[Myrmicinae]]
* [[Paraponerinae]]
* [[Ponerinae]]
* [[Proceratiinae]]
* [[Pseudomyrmecinae]]
* {{extinct}}[[Sphecomyrminae]]
* †[[Zigrasimeciinae]]

}}

'''Ants''' are [[Eusociality|eusocial]] [[insect]]s of the [[Family (biology)|family]] '''Formicidae''' and, along with the related [[wasp]]s and [[bee]]s, belong to the [[Taxonomy (biology)|order]] [[Hymenoptera]]. Ants evolved from [[Vespoidea|vespoid wasp]] ancestors in the [[Cretaceous]] period. More than 13,800 of an estimated total of 22,000 [[species]] have been classified. They are easily identified by their geniculate (elbowed) [[Antenna (biology)|antenna]]e and the distinctive node-like structure that forms their slender waists.

Ants form [[Ant colony|colonies]] that range in size from a few dozen individuals often living in small natural cavities to highly organised colonies that may occupy large territories with sizeable nest that consist of millions of individuals or into the hundreds of millions in [[Ant supercolony|super colonies]]. Typical colonies consist of various castes of sterile, wingless females, most of which are workers (ergates), as well as soldiers (dinergates) and other specialised groups. Nearly all ant colonies also have some fertile males called "drones" and one or more fertile females called "[[Queen ant|queens]]" ([[gyne]]s). The colonies are described as [[superorganism]]s because the ants appear to operate as a unified entity, collectively working together to support the colony.

Ants have colonised almost every landmass on [[Earth]]. The only places lacking [[Indigenous (ecology)|indigenous]] ants are [[Antarctica]] and a few remote or inhospitable islands. Ants thrive in moist tropical ecosystems and may exceed the combined [[Biomass (ecology)|biomass]] of wild birds and mammals. Their success in so many environments has been attributed to their social organisation and their ability to modify habitats, tap resources, and defend themselves. Their long [[co-evolution]] with other species has led to [[mimicry|mimetic]], [[commensalism|commensal]], [[parasitism|parasitic]], and [[Mutualism (biology)|mutualistic]] relationships.

Ant societies have [[division of labour]], communication between individuals, and an ability to [[Problem solving|solve complex problems]]. 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 rites. 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'') of South America, are regarded as [[invasive species]] in other parts of the world, establishing themselves in areas where they have been introduced accidentally.

==Etymology==
The word ''ant'' and the archaic word ''emmet''<ref>[https://www.merriam-webster.com/dictionary/emmet emmet]. Merriam-Webster Dictionary</ref> are derived from ''{{lang|enm|ante}}'', ''{{lang|enm|emete}}'' of [[Middle English]], which come from ''{{lang|ang|ǣmette}}'' of [[Old English]]; these are all related to [[Low German|Low Saxon]] ''{{lang|nds|e(e)mt}}'', ''{{lang|nds|empe}}'' and varieties ([[Old Saxon]] ''{{lang|osx|emeta}}'') and to [[German language|German]] ''{{lang|de|Ameise}}'' ([[Old High German]] ''{{lang|goh|āmeiza}}''). All of these words come from West Germanic ''*{{lang|gmw|ǣmaitjōn}}'', and the original meaning of the word was "the biter" (from [[Proto-Germanic]] ''{{lang|gem-x-proto|ai-}}'', "off, away" + ''{{lang|gem-x-proto|mait-}}'' "cut").<ref>{{cite web |url=http://www.merriam-webster.com/dictionary/ant |title=ant|publisher= Merriam-Webster Dictionary |access-date=6 June 2008}}</ref><ref>{{cite web |url=http://www.etymonline.com/index.php?term=ant |title=Ant. Online Etymology Dictionary |access-date=30 May 2009}}</ref>

The family name Formicidae is derived from the [[Latin]] ''{{lang|la|formīca}}'' ("ant")<ref>{{cite book | author = Simpson DP | title = Cassell's Latin Dictionary | publisher = Cassell | year = 1979 | edition = 5th | location = London | isbn=978-0-304-52257-6}}</ref> from which the words in other [[Romance languages]], such as the Portuguese ''{{lang|pt|formiga}}'', Italian ''{{lang|it|formica}}'', Spanish ''{{lang|es|hormiga}}'', Romanian ''{{lang|ro|furnică}}'', and French ''{{lang|fr|fourmi}}'' are derived. It has been hypothesised that a [[Proto-Indo-European language|Proto-Indo-European]] word *morwi- was the root for [[Sanskrit]] ''vamrah'', [[Ancient Greek|Greek]] μύρμηξ ''mýrmēx'', [[Old Church Slavonic]] ''mraviji'', [[Old Irish]] ''moirb'', [[Old Norse]] ''maurr'', [[Dutch language|Dutch]] ''mier'', [[Swedish language|Swedish]] ''myra'', [[Danish language|Danish]] ''myre'', [[Middle Dutch]] ''miere'', and [[Crimean Gothic]]'' miera''.<ref>{{cite web|url=http://www.etymonline.com/index.php?term=formic |title=Formic |publisher=Etymonline.com |access-date=2012-01-30}}</ref><ref>{{cite web|url=https://www.etymonline.com/word/pismire |title=Pismire |publisher=Etymonline.com |access-date=2020-08-27}}</ref>

==Taxonomy and evolution==


The family Formicidae belongs to the order [[Hymenoptera]], which also includes [[sawfly|sawflies]], [[bee]]s, and [[wasp]]s. Ants evolved from a lineage within the [[Aculeata|stinging wasps]], and a 2013 study suggests that they are a sister group of the [[Apoidea]].<ref name=phylo /> However, since Apoidea is a superfamily, ants must be upgraded to the same rank.<ref name=Fernández2021>{{cite journal|author1=Fernando Fernández|author2=Roberto J. Guerrero|author3=Andrés F. Sánchez Restrepo|date=April 2021|url=https://www.researchgate.net/publication/350917357|title=Systematics and diversity of Neotropical ants|journal=Revista Colombiana de Entomología|volume=47|issue=1|page=e11082|doi=10.25100/socolen.v47i1.11082|language=en|hdl=11336/165214|hdl-access=free}}</ref> A more detailed basic taxonomy was proposed in 2020. Three species of the extinct mid-[[Cretaceous]] genera ''[[Camelomecia]]'' and ''[[Camelosphecia]]'' were placed outside of the Formicidae, in a separate [[clade]] within the general superfamily [[Formicoidea]], which, together with Apoidea, forms the higher-ranking group [[Formicapoidina]].<ref name=Boudinot2020>{{cite journal|author1=Brendon E. Boudinot|author2=Vincent Perrichot|author3=Júlio C. M. Chaul|date=December 2020|title=†Camelosphecia gen. nov., lost ant-wasp intermediates from the mid-Cretaceous (Hymenoptera, Formicoidea)|journal=ZooKeys|volume=|issue=1005|pages=21–55 |doi=10.3897/zookeys.1005.57629|doi-access=free|pmid=33390754 |language=en|pmc=7762752|bibcode=2020ZooK.1005...21B }}</ref> Fernández et al. (2021) suggest that the common ancestors of ants and apoids within the Formicapoidina probably existed as early as in the end of the [[Jurassic]] period, before divergence in the Cretaceous.<ref name=Fernández2021/>
{{clade gallery  |width=350px |height=500px
|caption1=Relationship of ants with aculeate wasp families
|footer1=Phylogenetic position of the Formicidae as seen in Johnson et al. (2013)<ref name=phylo>{{cite journal | vauthors = Johnson BR, Borowiec ML, Chiu JC, Lee EK, Atallah J, Ward PS | title = Phylogenomics resolves evolutionary relationships among ants, bees, and wasps | journal = Current Biology | volume = 23 | issue = 20 | pages = 2058–2062 | date = October 2013 | pmid = 24094856 | doi = 10.1016/j.cub.2013.08.050 | doi-access = free | bibcode = 2013CBio...23.2058J }}</ref><ref name=Fernández2021/>
|cladogram1={{clade|style=font-size:100%;line-height:100%
 |label1=[[Aculeata]]
 |1={{clade
    |1=[[Chrysidoidea]]
    |label2=
    |2={{clade
       |label1=
       |1={{clade
          |1=[[Vespidae]]
          |2=[[Rhopalosomatidae]]
          }}
       |label2=
       |2={{clade
          |label1=
          |1={{clade
             |1={{clade
                |1=[[Pompilidae]]
                |2=[[Mutillidae]]
                }}
             |2={{clade
                |1=[[Tiphiidae]]
                |2=[[Chyphotidae]]
                }}
             }}
          |label2=
          |2={{clade
             |1=[[Scolioidea]]
             |label2=
             |2={{clade
                |1=[[Apoidea]]
                |2='''Formicidae'''
                }}
             }}
          }}
       }}
    }}
 }}

|caption2=Relationships of ant subfamilies |cladogram2=
{{clade|style=font-size:100%;line-height:100%
          |label1='''Formicidae'''
          |1={{clade
             |label1=
             |1={{clade
                |label1=Formicoid
                |1={{clade
                   |label1=
                   |1={{clade
                      |label1=
                      |1={{clade
                         |label1=
                         |1={{clade
                            |1=Myrmicinae
                            |label2=
                            |2={{clade
                               |1=Ectatomminae
                               |2=Heteroponerinae
                               }}
                            }}
                         |2=Formicinae
                         }}
                      |label2=
                      |2={{clade
                         |label1=
                         |1={{clade
                            |1=Dolichoderinae
                            |2=Aneuretinae
                            }}
                         |label2=
                         |2={{clade
                            |1=Pseudomyrmecinae
                            |2=Myrmeciinae
                            }}
                         }}
                      }}
                   |2=Dorylinae‡
                   }}
                |label2=Poneroid
                |2={{clade
                   |label1=
                   |1={{clade
                      |1=Ponerinae
                      |label2=
                      |2={{clade
                         |1=Agroecomyrmecinae
                         |2=Paraponerinae
                         }}
                      }}
                   |2=Proceratiinae
                   |label3=
                   |3={{clade
                      |1=Amblyoponinae
                      |2=Apomyrminae
                      }}
                   }}
                }}
             |label2=
             |2={{clade
                |1=Leptanillinae
                |2=Martialinae
                }}
             }}
}}
|footer2=A [[phylogeny]] of the extant ant [[subfamily|subfamilies]].<ref>{{Cite book |last1=Borowiec |first1=Marek L. |chapter=Ants: Phylogeny and Classification |year=2020 |title=Encyclopedia of Social Insects |pages=1–18 |editor-last=Starr |editor-first=Christopher K. |chapter-url=http://link.springer.com/10.1007/978-3-319-90306-4_155-1 |access-date=2024-02-11 |place=Cham |publisher=Springer International Publishing|doi=10.1007/978-3-319-90306-4_155-1 |isbn=978-3-319-90306-4 |last2=Moreau |first2=Corrie S. |last3=Rabeling |first3=Christian|s2cid=219873464 }}</ref><ref name=Ward>{{cite journal | vauthors = Ward PS |title=Phylogeny, classification, and species-level taxonomy of ants (Hymenoptera: Formicidae) | journal = [[Zootaxa]] | volume = 1668 | year=2007 |pages=549–563| url=http://www.mapress.com/zootaxa/2007f/zt01668p563.pdf |archive-url=https://ghostarchive.org/archive/20221009/http://www.mapress.com/zootaxa/2007f/zt01668p563.pdf |archive-date=2022-10-09 |url-status=live | doi = 10.11646/zootaxa.1668.1.26 }}</ref><ref name=martialis>{{cite journal | vauthors = Rabeling C, Brown JM, Verhaagh M | title = Newly discovered sister lineage sheds light on early ant evolution | journal = Proceedings of the National Academy of Sciences of the United States of America | volume = 105 | issue = 39 | pages = 14913–14917 | date = September 2008 | pmid = 18794530 | pmc = 2567467 | doi = 10.1073/pnas.0806187105 | bibcode = 2008PNAS..10514913R | doi-access = free }}</ref>

<nowiki>*</nowiki>Cerapachyinae is [[paraphyletic]]<br />
‡ The previous dorylomorph subfamilies – Ecitoninae, Aenictinae, Aenictogitoninae, Cerapachyinae, Leptanilloidinae – were synonymized under [[Dorylinae]] by Brady ''et al.'' in 2014<ref name="Brady_et_al_2014">{{cite journal | vauthors = Brady SG, Fisher BL, Schultz TR, Ward PS | title = The rise of army ants and their relatives: diversification of specialized predatory doryline ants | journal = BMC Evolutionary Biology | volume = 14 | issue = 93 | date = May 2014 | page = 93 | pmid = 24886136 | pmc = 4021219 | doi = 10.1186/1471-2148-14-93 | doi-access = free | bibcode = 2014BMCEE..14...93B }}</ref>
}}
[[Image:Ants in amber.jpg|thumb|right|Ants fossilised in [[Baltic amber]]]]
In 1966, [[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 [[absolute dating|dating]] back to around 92&nbsp;million years ago, has features found in some wasps, but not found in modern ants.<ref>{{cite journal | vauthors = Wilson EO, Carpenter FM, Brown WL | title = The first mesozoic ants | journal = [[Science (journal)|Science]] | volume = 157 | issue = 3792 | pages = 1038–1040 | date = September 1967 | pmid = 17770424 | doi = 10.1126/science.157.3792.1038 | author-link = E. O. Wilson | bibcode = 1967Sci...157.1038W | s2cid = 43155424 }}</ref> The oldest fossils of ants date to the mid-Cretaceous, around 100 million years ago, which belong to extinct [[stem-group]]s such as the [[Haidomyrmecinae]], [[Sphecomyrminae]] and [[Zigrasimeciinae]], with modern ant subfamilies appearing towards the end of the Cretaceous around 80–70 million years ago.<ref>{{Cite journal |last1=Boudinot |first1=Brendon E |last2=Richter |first2=Adrian |last3=Katzke |first3=Julian |last4=Chaul |first4=Júlio C M |last5=Keller |first5=Roberto A |last6=Economo |first6=Evan P |last7=Beutel |first7=Rolf Georg |last8=Yamamoto |first8=Shûhei |date=2022-07-29 |title=Evidence for the evolution of eusociality in stem ants and a systematic revision of † Gerontoformica (Hymenoptera: Formicidae) |url=https://academic.oup.com/zoolinnean/article/195/4/1355/6523228 |journal=[[Zoological Journal of the Linnean Society]]|volume=195 |issue=4 |pages=1355–1389 |doi=10.1093/zoolinnean/zlab097 |issn=0024-4082|doi-access=free |hdl=10451/55807 |hdl-access=free }}</ref> Ants diversified extensively during the [[Cretaceous Terrestrial Revolution|Angiosperm Terrestrial Revolution]]<ref>{{Cite journal |last1=Jouault |first1=Corentin |last2=Condamine |first2=Fabien L. |last3=Legendre |first3=Frédéric |last4=Perrichot |first4=Vincent |date=11 March 2024 |title=The Angiosperm Terrestrial Revolution buffered ants against extinction |journal=[[Proceedings of the National Academy of Sciences of the United States of America]]|volume=121 |issue=13 |pages=e2317795121 |doi=10.1073/pnas.2317795121 |pmid=38466878 |pmc=10990090 |bibcode=2024PNAS..12117795J |issn=0027-8424 }}</ref> and assumed ecological dominance around 60&nbsp;million years ago.<ref name="grimaldi2001">{{cite journal | vauthors = Grimaldi D, Agosti D | title = A formicine in New Jersey cretaceous amber (Hymenoptera: formicidae) and early evolution of the ants | journal = [[Proceedings of the National Academy of Sciences of the United States of America]] | volume = 97 | issue = 25 | pages = 13678–13683 | date = December 2000 | pmid = 11078527 | pmc = 17635 | doi = 10.1073/pnas.240452097 | bibcode = 2000PNAS...9713678G | doi-access = free }}</ref><ref name=phyl2006>{{cite journal | vauthors = Moreau CS, Bell CD, Vila R, Archibald SB, Pierce NE | title = Phylogeny of the ants: diversification in the age of angiosperms | journal = Science | volume = 312 | issue = 5770 | pages = 101–104 | date = April 2006 | pmid = 16601190 | doi = 10.1126/science.1124891 | bibcode = 2006Sci...312..101M | s2cid = 20729380 }}</ref><ref name=riseofants>{{cite journal | vauthors = Wilson EO, Hölldobler B | title = The rise of the ants: a phylogenetic and ecological explanation | journal = Proceedings of the National Academy of Sciences of the United States of America | volume = 102 | issue = 21 | pages = 7411–7414 | date = May 2005 | pmid = 15899976 | pmc = 1140440 | doi = 10.1073/pnas.0502264102 | bibcode = 2005PNAS..102.7411W | doi-access = free }}</ref><ref>{{cite journal | vauthors = LaPolla JS, Dlussky GM, Perrichot V | s2cid = 40555356 | title = Ants and the fossil record | journal = Annual Review of Entomology | volume = 58 | pages = 609–730 | year = 2013 | pmid = 23317048 | doi = 10.1146/annurev-ento-120710-100600 }}</ref> Some groups, such as the [[Leptanillinae]] and [[Martialinae]], are suggested to have diversified from early primitive ants that were likely to have been predators underneath the surface of the soil.<ref name=martialis/><ref name="Barden2012">{{cite journal |vauthors=Barden P, Grimaldi D |year=2012 |title=Rediscovery of the bizarre Cretaceous ant ''Haidomyrmex'' Dlussky (Hymenoptera: Formicidae), with two new species |journal=American Museum Novitates |pages=1–16 |doi=10.1206/3755.2 |url=http://digitallibrary.amnh.org/dspace/bitstream/handle/2246/6368/N3755.pdf?sequence=1 |issue=3755 |hdl=2246/6368 |s2cid=83598305 |access-date=2013-05-05 |archive-date=2013-04-23 |archive-url=https://web.archive.org/web/20130423110550/http://digitallibrary.amnh.org/dspace/bitstream/handle/2246/6368/N3755.pdf?sequence=1 |url-status=dead }}</ref>

During the Cretaceous period, a few species of primitive ants ranged widely on the [[Laurasia]]n supercontinent (the [[Northern Hemisphere]]). Their representation in the fossil record is poor, in comparison to the populations of other insects, representing only about 1% of fossil evidence of insects in the era. Ants became dominant after [[adaptive radiation]] at the beginning of the [[Paleogene period]]. By the [[Oligocene]] and [[Miocene]], ants had come to represent 20–40% of all insects found in major fossil deposits. Of the species that lived in the [[Eocene]] epoch, around one in 10 genera survive to the present. Genera surviving today comprise 56% of the genera in [[Baltic amber]] fossils (early Oligocene), and 92% of the genera in [[Dominican amber]] fossils (apparently early Miocene).<ref name="grimaldi2001"/><ref name=TheAntEvo>Hölldobler & Wilson (1990), pp. 23–24</ref>

[[Termite]]s live in colonies and are sometimes called "white ants", but termites are only distantly related to ants. They are the sub-order [[Isoptera]], and together with [[cockroach]]es, they form the order [[Blattodea]]. Blattodeans are related to [[mantid]]s, [[crickets]], and other winged insects that do not undergo [[Holometabolism|complete metamorphosis]]. Like ants, termites are [[eusociality|eusocial]], with sterile workers, but they differ greatly in the genetics of reproduction. The similarity of their social structure to that of ants is attributed to [[convergent evolution]].<ref>{{cite journal| vauthors = Thorne BL |year=1997 |title=Evolution of eusociality in termites |journal=Annu. Rev. Ecol. Syst. |volume=28 |issue=5 |pages=27–53 |url=http://www.thornelab.umd.edu/Termite_PDFS/EvolutionEusocialityTermites.pdf |doi=10.1146/annurev.ecolsys.28.1.27 |pmc=349550  |bibcode=1997AnRES..28...27T |url-status=dead |archive-url=https://web.archive.org/web/20100530162505/http://www.thornelab.umd.edu/Termite_PDFS/EvolutionEusocialityTermites.pdf |archive-date=2010-05-30 }}</ref> [[Mutillidae|Velvet ants]] look like large ants, but are wingless female [[wasp]]s.<ref>{{cite web |url=http://bugguide.net/node/view/69 |title=Order Isoptera – Termites |access-date=12 June 2008 |publisher=Iowa State University Entomology
|date=16 February 2004| archive-url= https://web.archive.org/web/20080615140449/http://bugguide.net/node/view/69| archive-date= 15 June 2008| url-status= live}}</ref><ref>{{cite web |url=http://bugguide.net/node/view/159/
|title=Family Mutillidae&nbsp;– Velvet ants |access-date=12 June 2008 |publisher=Iowa State University Entomology
|date=16 February 2004| archive-url= https://web.archive.org/web/20080630045231/http://bugguide.net/node/view/159/| archive-date= 30 June 2008| url-status= live}}</ref>

==Distribution and diversity==
{| class="wikitable sortable" style="float:right; margin:10px;"
|-
! [[Region]] !! Number of<br />species&nbsp;<ref name = HolldoblerWilsonAnts>Hölldobler & Wilson (1990), p. 4</ref>
|-
| [[Neotropical realm|Neotropics]] ||align="right"| 2,162
|-
| [[Nearctic realm|Nearctic]] ||align="right"| 580
|-
| [[Europe]] ||align="right"| 180
|-
| [[Africa]] ||align="right"| 2,500
|-
| [[Asia]] ||align="right"| 2,080
|-
| [[Melanesia]] ||align="right"| 275
|-
| [[Australia]] ||align="right"| 985
|-
| [[Polynesia]] ||align="right"| 42
|}
Ants have a [[cosmopolitan distribution]]. They 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?
| work=National Geographic Magazine
| author=Jones, Alice S
| access-date=5 July 2008| archive-url=https://web.archive.org/web/20080730071158/http://ngm.nationalgeographic.com/2007/08/ants/did-you-know-learn| archive-date=30 July 2008 |url-status=dead}}</ref><ref>{{cite web
| url=http://www.hear.org/ants/ |title=Pest Ants in Hawaii
| year=2007
| author=Thomas, Philip
| publisher=Hawaiian Ecosystems at Risk project (HEAR)
| access-date=6 July 2008}}</ref> Ants occupy a wide range of [[ecological niche]]s and exploit many different food resources as direct or indirect herbivores, predators and scavengers. Most ant species are omnivorous [[Generalist and specialist species|generalists]], but a few are specialist feeders. There is considerable variation in ant abundance across habitats, peaking in the moist tropics to nearly six times that found in less suitable habitats.<ref>{{Cite journal |last1=Fayle |first1=Tom M. |last2=Klimes |first2=Petr |date=2022-10-18 |title=Improving estimates of global ant biomass and abundance |journal=Proceedings of the National Academy of Sciences|volume=119 |issue=42 |pages=e2214825119 |doi=10.1073/pnas.2214825119 |doi-access=free |pmid=36197959 |pmc=9586285 |bibcode=2022PNAS..11914825F |issn=0027-8424}}</ref> Their ecological dominance has been examined primarily using estimates of their [[biomass (ecology)|biomass]]: myrmecologist [[E. O. Wilson]] had estimated in 2009 that at any one time the total number of ants was between one and ten [[1,000,000,000,000,000|quadrillion]] ([[short scale]]) (i.e., between 10<sup>15</sup> and 10<sup>16</sup>) and using this estimate he had suggested that the total [[Biomass (ecology)|biomass]] of all the ants in the world was approximately equal to the total biomass of the entire [[human]] race.<ref>{{cite book |url=https://archive.org/details/superorganismbea0000hlld |title=The Superorganism: The Beauty, Elegance, and Strangeness of Insect Societies |vauthors=Holldobler B, Wilson EO |publisher=W.W. Norton |year=2009 |isbn=978-0-393-06704-0 |place=New York |page=[https://archive.org/details/superorganismbea0000hlld/page/5 5] |url-access=registration}}</ref> More careful estimates made in 2022 which take into account regional variations puts the global ant contribution at 12 megatons of dry carbon, which is about 20% of the total human contribution, but greater than that of the wild birds and mammals combined. This study also puts a conservative estimate of the ants at about 20 × 10<sup>15</sup> (20 quadrillion).<ref name="schultz">{{cite journal | vauthors = Schultz TR | title = In search of ant ancestors | journal = Proceedings of the National Academy of Sciences of the United States of America | volume = 97 | issue = 26 | pages = 14028–14029 | date = December 2000 | pmid = 11106367 | pmc = 34089 | doi = 10.1073/pnas.011513798| bibcode = 2000PNAS...9714028S | doi-access = free }}</ref><ref>{{cite web |title=How many ants are there for every one person on earth? |url=http://topics.info.com/How-many-ants-are-there-for-every-one-person-on-earth_452 |url-status=dead |archive-url=https://web.archive.org/web/20130813234547/http://topics.info.com/How-many-ants-are-there-for-every-one-person-on-earth_452 |archive-date=13 August 2013 |access-date=27 July 2013 |publisher=info.com}}</ref><ref>{{Cite journal |last1=Schultheiss |first1=Patrick |last2=Nooten |first2=Sabine S. |last3=Wang |first3=Runxi |last4=Wong |first4=Mark K. L. |last5=Brassard |first5=François |last6=Guénard |first6=Benoit |date=2022-10-04 |title=The abundance, biomass, and distribution of ants on Earth |journal=Proceedings of the National Academy of Sciences|volume=119 |issue=40 |pages=e2201550119 |doi=10.1073/pnas.2201550119 |doi-access=free |pmid=36122199 |pmc=9546634 |bibcode=2022PNAS..11901550S |s2cid=252381912 |issn=0027-8424}}</ref>

Ants range in size from {{convert|0.75|to(-)|52|mm|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 | year = 1999 | publisher = CSIRO | location = Collingwood, Vic | isbn = 978-0-643-06659-5 | page = 149 }}</ref> the largest species being the fossil ''[[Titanomyrma|Titanomyrma giganteum]]'', the queen of which was {{convert|6|cm|in|frac=2|abbr=on}} long with a wingspan of {{convert|15|cm|in|frac=2|abbr=on}}.<ref name="Messel">{{cite journal|journal=Encyclopedia of Life Sciences|author=Schaal, Stephan|date=27 January 2006|doi=10.1038/npg.els.0004143|title=Messel|isbn=978-0-470-01617-6}}</ref> Ants vary in colour; most ants are yellow to red or brown to black, but a few species are green and some tropical species have a metallic [[Lustre (mineralogy)|lustre]]. More than 13,800 species are currently known<ref name=AntWeb>[https://www.antweb.org/statsPage.do AntWeb]</ref> (with upper estimates of the potential existence of about 22,000; see the article [[List of ant genera (alphabetical)|List of ant genera]]), with the greatest diversity in the tropics. Taxonomic studies continue to resolve the classification and systematics of ants. Online databases of ant species, including AntWeb and the Hymenoptera Name Server, help to keep track of the known and newly described species.<ref name=AntWeb/> The relative ease with which ants may be sampled and studied in [[ecosystem]]s has made them useful as [[indicator species]] in [[biodiversity]] studies.<ref>{{cite book | veditors = Agosti D, Majer JD, Alonso JE, Schultz TR | 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 | access-date = 2015-12-13 }}</ref><ref>{{cite web | url = http://atbi.biosci.ohio-state.edu/hymenoptera/nomenclator.home_page | archive-url = https://web.archive.org/web/20160127143848/http://atbi.biosci.ohio-state.edu/hymenoptera/nomenclator.home_page | url-status = dead | archive-date = 27 January 2016 | title = Hymenoptera name server | publisher  = [[Ohio State University]] | vauthors = Johnson NF | year = 2007 | access-date = 6 July 2008 }}</ref>

==Morphology==
[[Image:Scheme ant worker anatomy-en.svg|thumb|450px|left|Diagram of a worker ant (''[[Neoponera verenae]]'')]]

Ants are distinct in their [[morphology (biology)|morphology]] from other insects in having geniculate (elbowed) [[Antenna (biology)|antenna]]e, [[metapleural gland]]s, and a strong constriction of their second [[Abdomen|abdominal]] segment into a node-like [[petiole (insect anatomy)|petiole]]. The head, [[mesosoma]], and [[metasoma]] are the three distinct body segments (formally [[Tagma (biology)|tagmata]]). The petiole forms a narrow waist between their mesosoma ([[Thorax (insect anatomy)|thorax]] plus the first abdominal segment, which is fused to it) and [[gaster (insect anatomy)|gaster]] (abdomen less the abdominal segments in the petiole). The [[Petiole (insect anatomy)|petiole]] may 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> Tergosternal fusion, when the tergite and sternite of a segment fuse together, can occur partly or fully on the second, third and fourth abdominal segment and is used in identification. Fourth abdominal tergosternal fusion was formerly used as character that defined the poneromorph subfamilies, Ponerinae and relatives within their clade, but this is no longer considered a [[synapomorphic]] character.<ref>{{Cite journal |last1=Ouellette |first1=Gary D. |last2=Fisher |first2=Brian L. |last3=Girman |first3=Derek J. |date=2006 |title=Molecular systematics of basal subfamilies of ants using 28S rRNA (Hymenoptera: Formicidae) |url=https://www.sciencedirect.com/science/article/pii/S105579030600100X |journal=Molecular Phylogenetics and Evolution|volume=40 |issue=2 |pages=359–369 |doi=10.1016/j.ympev.2006.03.017 |pmid=16630727 |bibcode=2006MolPE..40..359O |hdl=10211.1/1549 |issn=1055-7903|hdl-access=free }}</ref>

Like other arthropods, 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 [[lung]]s; [[oxygen]] and other gases, such as [[carbon dioxide]], pass through their exoskeleton via tiny valves called [[Spiracle (arthropods)|spiracle]]s. Insects also lack closed [[blood vessel]]s; instead, they have a long, thin, perforated tube along the top of the body (called the "dorsal aorta") that functions like a heart, and pumps [[haemolymph]] 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.<ref name="insectmorph">Borror, Triplehorn & Delong (1989), pp. 24–71</ref>

===Head===
[[File:Bullant head detail.jpg|thumb|[[Bull ant]] showing the powerful [[Mandible (insect mouthpart)|mandibles]] and the relatively large [[compound eye]]s that provide excellent vision]]

An ant's head contains many [[sensory organs]]. Like most insects, ants have [[compound eye]]s made from numerous tiny lenses attached together. Ant eyes are good for acute movement detection, but do not offer a high [[Optical resolution|resolution]] image. They also have three small [[ocellus|ocelli]] (simple eyes) on the top of the head that detect light levels and [[polarization (waves)|polarization]].<ref>{{cite journal | vauthors = Fent K, Wehner R | title = Oceili: a celestial compass in the desert ant cataglyphis | journal = Science | volume = 228 | issue = 4696 | pages = 192–194 | date = April 1985 | pmid = 17779641 | doi = 10.1126/science.228.4696.192 | s2cid = 33242108 | bibcode = 1985Sci...228..192F }}</ref> Compared to [[vertebrate]]s, ants tend to have blurrier eyesight, particularly in smaller species,<ref>{{cite journal | vauthors = Palavalli-Nettimi R, Narendra A | title = Miniaturisation decreases visual navigational competence in ants | journal = The Journal of Experimental Biology | volume = 221 | issue = Pt 7 | page = jeb177238 | date = April 2018 | pmid = 29487158 | doi = 10.1242/jeb.177238 | doi-access = free }}</ref> and a few [[Subterranea (geography)|subterranean]] taxa are completely [[Blindness|blind]].<ref name=Ward /> However, some ants, such as Australia's [[Myrmecia (ant)|bulldog ant]], have excellent vision and are capable of discriminating the distance and size of objects moving nearly a [[Metre|meter]] away.<ref>{{cite journal|journal= Journal of Experimental Biology|volume=119| pages=115–131| year=1985| title=Attack behaviour and distance perception in the Australian bulldog ant ''Myrmecia nigriceps''| vauthors = Eriksson ES |url= http://jeb.biologists.org/content/119/1/115.full.pdf |archive-url=https://ghostarchive.org/archive/20221009/http://jeb.biologists.org/content/119/1/115.full.pdf |archive-date=2022-10-09 |url-status=live | issue=1|doi=10.1242/jeb.119.1.115| doi-access=free}}</ref> Based on experiments conducted to test their ability to differentiate between selected wavelengths of light, some ant species such as ''Camponotus blandus, Solenopsis invicta,'' and ''Formica cunicularia'' are thought to possess a degree of colour vision.<ref>{{Cite journal |last1=Yilmaz |first1=Ayse |last2=Spaethe |first2=Johannes |date=2022 |title=Colour vision in ants (Formicidae, Hymenoptera) |journal=Philosophical Transactions of the Royal Society B: Biological Sciences|volume=377 |issue=1862 |doi=10.1098/rstb.2021.0291 |issn=0962-8436 |pmc=9441231 |pmid=36058251}}</ref>

Two [[antenna (biology)|antennae]] ("feelers") are attached to the head; these organs detect chemicals, [[air current]]s, and [[vibration]]s; they also are 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 may be passed to other ants or their larvae.<ref>{{cite journal |vauthors=Eisner T, Happ GM | title=The infrabuccal pocket of a formicine ant: a social filtration device | journal=Psyche: A Journal of Entomology | volume=69 | pages=107–116 | year=1962 | doi=10.1155/1962/25068 | issue=3| doi-access=free }}</ref>

===Mesosoma===
Both the [[arthropod leg|legs]] and [[insect wing|wings]] of the ant are attached to the [[mesosoma]] ("thorax"). The legs terminate in a hooked [[Chelae|claw]] which allows them to hook on and climb surfaces.<ref>{{cite web|author=Holbrook, Tate|title=Ask a Biologist: Face to Face with Ants|date=22 September 2009|publisher=ASU School of Life Sciences|url=https://askabiologist.asu.edu/explore/ant-anatomy|access-date=2018-01-23}}</ref> Only reproductive ants ([[Queen (insect)|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 ([[ergatoid]]s) and males occur.<ref name="insectmorph"/>

===Metasoma===
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 [[ovipositor|egg-laying structures]] modified into [[stinger (organ)|stings]] that are used for subduing [[predation|prey]] and defending their nests.<ref name="insectmorph"/>

===Polymorphism===
[[Image:Atta.cephalotes.gamut.selection.jpg|thumb|right|Seven [[leafcutter ant]] workers of various castes (left) and two queens (right)]]

In the colonies of a few ant species, there are physical castes—workers in distinct size-classes, called minor (micrergates), median, and major ergates (macrergates). Often, the larger ants have disproportionately larger heads, and correspondingly stronger [[mandible (insect)|mandibles]]. Although formally known as dinergates, such individuals are sometimes called "soldier" ants because their stronger mandibles make them more effective in fighting, although they still are workers and their "duties" typically do not vary greatly from the minor or median workers.<ref name=":1" /> In a few species, the median workers are absent, creating a sharp divide between the minors and majors.<ref>{{cite journal | vauthors = Wilson EO | title = The origin and evolution of polymorphism in ants | journal = The Quarterly Review of Biology | volume = 28 | issue = 2 | pages = 136–156 | date = June 1953 | pmid = 13074471 | doi = 10.1086/399512 | s2cid = 4560071 }}</ref> [[Weaver ant]]s, for example, have a distinct [[bimodal]] size distribution.<ref>{{cite journal|author=Weber, NA|year=1946|title=Dimorphism in the African ''Oecophylla'' worker and an anomaly (Hym.: Formicidae)|journal=Annals of the Entomological Society of America|volume=39|pages=7–10| url=http://antbase.org/ants/publications/10434/10434.pdf |archive-url=https://ghostarchive.org/archive/20221009/http://antbase.org/ants/publications/10434/10434.pdf |archive-date=2022-10-09 |url-status=live|doi=10.1093/aesa/39.1.7}}</ref><ref>{{cite journal | first1 = Edward O. | last1 = Wilson | first2 = Robert W. | last2 = Taylor | name-list-style=vanc|year=1964|title=A Fossil Ant Colony: New Evidence of Social Antiquity |journal=Psyche: A Journal of Entomology|volume=71|pages=93–103|url=http://psyche.entclub.org/pdf/71/71-093.pdf |archive-url=https://ghostarchive.org/archive/20221009/http://psyche.entclub.org/pdf/71/71-093.pdf |archive-date=2022-10-09 |url-status=live |doi=10.1155/1964/17612 |issue=2|doi-access=free }}</ref> Some other species show continuous variation in the size of workers. The smallest and largest workers in ''[[Carebara diversa]]'' show nearly a 500-fold difference in their dry weights.<ref>{{cite journal |vauthors=Moffett MW, Tobin JE |year=1991 |title=Physical castes in ant workers: a problem for ''Daceton armigerum'' and other ants |journal=Psyche: A Journal of Entomology |volume=98 |pages=283–292 |url=http://psyche2.entclub.org/articles/98/98-283.pdf |archive-url=https://web.archive.org/web/20080227015919/http://psyche2.entclub.org/articles/98/98-283.pdf |archive-date=2008-02-27 |doi=10.1155/1991/30265 |issue=4|doi-access=free }}</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=141–146 |doi=10.1007/PL00001692|s2cid=31953751 }}</ref> For instance, these replete workers develop in the North American honeypot ant ''[[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 (biology)|polymorphism]].<ref>{{cite journal | last1=Rissing | first1=Steven W | name-list-style=vanc | year=1984 | title=Replete caste production and allometry of workers in the Honey Ant, ''Myrmecocystus mexicanus'' Wesmael (Hymenoptera: Formicidae) |  journal=Journal of the Kansas Entomological Society | volume=57 | issue=2| pages=347–350}}</ref> 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 [[morphogenesis|developmental paths]]; however, genetic differences between worker castes have been noted in ''Acromyrmex'' sp.<ref>{{cite journal | vauthors = Hughes WO, Sumner S, Van Borm S, Boomsma JJ | title = Worker caste polymorphism has a genetic basis in Acromyrmex leaf-cutting ants | journal = Proceedings of the National Academy of Sciences of the United States of America | volume = 100 | issue = 16 | pages = 9394–9397 | date = August 2003 | pmid = 12878720 | pmc = 170929 | doi = 10.1073/pnas.1633701100 | bibcode = 2003PNAS..100.9394H | doi-access = free }}</ref> These polymorphisms are caused by relatively small genetic changes; differences in a single gene of ''[[Red imported fire ant|Solenopsis invicta]]'' can decide whether the colony will have single or multiple queens.<ref>{{cite journal | vauthors = Ross KG, Krieger MJ, Shoemaker DD | title = Alternative genetic foundations for a key social polymorphism in fire ants | journal = Genetics | volume = 165 | issue = 4 | pages = 1853–1867 | date = December 2003 | doi = 10.1093/genetics/165.4.1853 | pmid = 14704171 | pmc = 1462884 }}</ref> 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.<ref>{{cite journal | vauthors = Crosland MW, Crozier RH | title = ''Myrmecia pilosula'', an ant with only one Pair of chromosomes | journal = Science | volume = 231 | issue = 4743 | pages = 1278 | date = March 1986 | pmid = 17839565 | doi = 10.1126/science.231.4743.1278 | bibcode = 1986Sci...231.1278C | s2cid = 25465053 }}</ref><ref>{{cite journal | vauthors = Tsutsui ND, Suarez AV, Spagna JC, Johnston JS | title = The evolution of genome size in ants | journal = BMC Evolutionary Biology | volume = 8 | issue = 64 | pages = 64 | date = February 2008 | pmid = 18302783 | pmc = 2268675 | doi = 10.1186/1471-2148-8-64 | doi-access = free | bibcode = 2008BMCEE...8...64T }}</ref>

===Genome size===
[[Genome size]] is a fundamental characteristic of an organism. Ants have been found to have tiny genomes, with the evolution of genome size suggested to occur through loss and accumulation of [[non-coding DNA|non-coding]] regions, mainly [[transposable element]]s, and occasionally by whole genome duplication.<ref name="moura">{{cite journal|author1=Moura, Mariana Neves|author2=Cardoso, Danon Clemes|author3=Cristiano, Maykon Passos|title=The tight genome size of ants: diversity and evolution under ancestral state reconstruction and base composition|year=2021|journal=Zoological Journal of the Linnean Society|volume=193|issue=1|pages=124–144|doi=10.1093/zoolinnean/zlaa135|issn=0024-4082|url=https://academic.oup.com/zoolinnean/article/193/1/124/6036549}}</ref> This may be related to [[Colonisation (biology)|colonisation]] processes, but further studies are needed to verify this.<ref name="moura"/>

==Life cycle==
[[Image:Meat eater ant nest swarming02.jpg|thumb|[[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 [[pupa]]l stage before emerging as an adult. The larva is largely immobile and is fed and cared for by workers. Food is given to the larvae by [[trophallaxis]], a process in which an ant [[Regurgitation (digestion)|regurgitates]] liquid food held in its [[Crop (anatomy)|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 egg]]s, pieces of prey, and seeds brought by workers.<ref>{{cite book| vauthors = Hölldobler B, Wilson EO |title=The Ants|url=https://archive.org/details/ants0000hlld |url-access=registration |year=1990 |publisher=Harvard University Press|isbn=978-0-674-04075-5 |page=[https://archive.org/details/ants0000hlld/page/n316 291]}}</ref>

The larvae grow through a series of four or five [[moult]]s and enter the pupal stage. The pupa has the appendages free and not fused to the body as in a [[Chrysalis|butterfly pupa]].<ref>{{cite book|author=Gillott, Cedric |year=1995|title=Entomology|publisher=Springer|isbn=978-0-306-44967-3|page=325}}</ref> The differentiation into queens and workers (which are both female), and different [[caste (biology)|castes]] of workers, is influenced in some species by the nutrition the larvae obtain. Genetic influences and the [[polyphenism|control of gene expression]] by the developmental environment are complex and the determination of caste continues to be a subject of research.<ref>{{cite journal|title=The causes and consequences of genetic caste determination in ants (Hymenoptera: Formicidae) |first1=Kirk E. |last1=Anderson |first2=Timothy A. |last2=Linksvayer |first3=Chris R. |last3=Smith  | name-list-style=vanc |journal=Myrmecol. News |volume=11 |pages=119–132 |year=2008 |url=https://myrmecologicalnews.org/cms/index.php?option=com_content&view=category&id=250&Itemid=73}}</ref> Winged male ants, called drones (termed "aner" in old literature<ref name=":1">{{Cite journal| vauthors = Gaul AT |date=1951|title=A Glossary of Terms and Phrases Used in the Study of Social Insects|url=https://academic.oup.com/aesa/article-lookup/doi/10.1093/aesa/44.3.473|journal=Annals of the Entomological Society of America|volume=44|issue=3|pages=473–484|doi=10.1093/aesa/44.3.473|issn=1938-2901}}</ref>), emerge from pupae along with the usually winged breeding females. Some species, such as [[army ant]]s, have wingless queens. Larvae and pupae need to be kept at fairly constant temperatures to ensure proper development, and so often are moved around among the various brood chambers within the colony.<ref>Hölldobler & Wilson (1990), pp. 351, 372</ref>

A new ergate spends the first few days of its adult life caring for the queen and young. She then graduates to digging and other nest work, and later to defending the nest and foraging. These changes are sometimes fairly sudden, and define what are called temporal castes. Such age-based task-specialization or [[polyethism]] has been suggested as having evolved due to the high casualties involved in foraging and defence, making it an acceptable risk only for ants who are older and likely to die sooner from natural causes.<ref>{{cite journal|journal=Annual Review of Entomology|year=1989|volume=34|pages=191–210|title=Foraging strategies of ants|author=Traniello JFA|doi=10.1146/annurev.en.34.010189.001203}}</ref><ref>{{cite journal|vauthors=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: A Journal of Entomology|volume=91|pages=319–332|year=1984|doi=10.1155/1984/39236|issue=3–4|doi-access=free}}</ref> In the Brazilian ant ''[[Forelius pusillus]],'' the nest entrance is closed from the outside to protect the colony from predatory ant species at sunset each day. About one to eight workers seal the nest entrance from the outside and they have no chance of returning to the nest and are in effect sacrificed.<ref>{{Cite journal |last1=Tofilski |first1=Adam |last2=Couvillon |first2=Margaret J. |last3=Evison |first3=Sophie E. F. |last4=Helanterä |first4=Heikki |last5=Robinson |first5=Elva J. H. |last6=Ratnieks |first6=Francis L. W. |date=2008 |title=Preemptive Defensive Self-Sacrifice by Ant Workers |journal=The American Naturalist|volume=172 |issue=5 |pages=E239–E243 |doi=10.1086/591688 |pmid=18928332 |s2cid=7052340 |issn=0003-0147}}</ref> Whether these seemingly suicidal workers are older workers has not been determined.<ref>{{Cite journal |last1=Shorter |first1=J. R. |last2=Rueppell |first2=O. |date=2012 |title=A review on self-destructive defense behaviors in social insects |url=http://link.springer.com/10.1007/s00040-011-0210-x |journal=Insectes Sociaux|volume=59 |issue=1 |pages=1–10 |doi=10.1007/s00040-011-0210-x |s2cid=253634662 |issn=0020-1812}}</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, being quite short-lived and surviving for 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=235–246 |doi=10.1007/s000400050084 |issue=3|s2cid=24541087 }}</ref> Ant queens are estimated to live 100 times as long as solitary insects of a similar size.<ref name=insencyc>{{cite book|editor-link1=Nigel R. Franks|veditors=Franks NR, Resh VH, Cardé RT|year=2003|title=Encyclopedia of Insects|pages=[https://archive.org/details/encyclopediaofin00bada/page/29 29–32]|isbn=978-0-12-586990-4|publisher=Academic Press|location=San Diego|url=https://archive.org/details/encyclopediaofin00bada/page/29}}</ref>

Ants are active all year long in the tropics; however, in cooler regions, they survive the winter in a state of dormancy known as [[hibernation]]. 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.<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=198–217}}</ref>

===Reproduction===
[[Image:FlyingAnts.jpg|thumb|Honey ant (''[[Prenolepis imparis]]'') mating, the drone is much smaller than the queen]]
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 [[thelytoky|thelytokous parthenogenesis]].<ref>{{cite journal| vauthors = Heinze J, Tsuji K |year=1995|title=Ant reproductive strategies|journal=Res. Popul. Ecol.|volume=37|issue=2|pages=135–149|url=http://meme.biology.tohoku.ac.jp/POPECOL/RP%20PDF/37(2)/pp.135.pdf|doi=10.1007/BF02515814|bibcode=1995PopEc..37..135H |s2cid=21948488|access-date=2009-04-16|archive-url=https://web.archive.org/web/20110527051529/http://meme.biology.tohoku.ac.jp/POPECOL/RP%20PDF/37(2)/pp.135.pdf|archive-date=2011-05-27|url-status=dead}}</ref> Secretions from the male accessory glands in some species can plug the female genital opening and prevent females from re-mating.<ref name="Mikheyev, a. S. 2003. pp.401">{{Cite journal | doi = 10.1007/s00040-003-0697-x | title = Evidence for mating plugs in the fire ant ''Solenopsis invicta'' | journal = Insectes Sociaux | volume = 50 | issue = 4 | pages = 401–402 | year = 2003 | vauthors = Mikheyev AS | s2cid = 43492133 }}</ref> 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 "[[gamergate (ant)|gamergates]]" and colonies that lack queens are then called gamergate colonies; colonies with queens are said to be queen-right.<ref>{{cite journal | vauthors = Peeters C, Hölldobler B | title = Reproductive cooperation between queens and their mated workers: the complex life history of an ant with a valuable nest | journal = Proceedings of the National Academy of Sciences of the United States of America | volume = 92 | issue = 24 | pages = 10977–10979 | date = November 1995 | pmid = 11607589 | pmc = 40553 | doi = 10.1073/pnas.92.24.10977 | bibcode = 1995PNAS...9210977P | doi-access = free }}</ref>

Drones can also mate with existing queens by entering a foreign colony, such as in [[army ant]]s. 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.<ref name="Franks N. 2008">{{Cite journal | vauthors = Franks NR, Hölldobler B | doi = 10.1111/j.1095-8312.1987.tb00298.x | title = Sexual competition during colony reproduction in army ants | journal = Biological Journal of the Linnean Society | volume = 30 | issue = 3 | pages = 229–243 | year = 1987 }}</ref> 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.<ref>{{Cite journal | doi = 10.1007/BF01175395 | title = Pheromonal manipulation of workers by a fighting male to kill his rival males in the ant ''Cardiocondyla wroughtonii'' | journal = Naturwissenschaften | volume = 79 | issue = 6 | pages = 274–276 | year = 1992 | vauthors = Yamauchi K, Kawase N | bibcode = 1992NW.....79..274Y | s2cid = 31191187 }}</ref>

Most ants are [[univoltine]], producing a new generation each year.<ref name="bloodywasp">{{cite book| vauthors = Taylor RW |year=2007 |chapter=Bloody funny wasps! Speculations on the evolution of eusociality in ants|pages=580–609|veditors=Snelling RR, Fisher BL, Ward PS|title=Advances in ant systematics (Hymenoptera: Formicidae): homage to E. O. Wilson&nbsp;– 50 years of contributions. Memoirs of the American Entomological Institute, 80|publisher=American Entomological Institute | chapter-url = http://antbase.org/ants/publications/21292/21292.pdf |access-date=2015-12-13}}</ref> During the species-specific breeding period, winged females and winged males, known to [[entomologists]] as [[alate]]s, 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.<ref name="nuptial flight 2">{{cite journal| vauthors = Wilson EO |year=1957|title=The organization of a nuptial flight of the ant ''Pheidole sitarches'' Wheeler | journal = Psyche: A Journal of Entomology | url = http://www.antwiki.org/wiki/images/7/7f/Wilson_1958d.pdf |volume=64|issue= 2|pages=46–50|doi=10.1155/1957/68319|doi-access=free}}</ref> 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 [[Spermatozoon|sperm]] in their [[spermatheca]]e.<ref name="HolldoblerWilsonAnts2">Hölldobler & Wilson (1990), pp. 143–179</ref> The genus ''[[Cardiocondyla]]'' have species with both winged and wingless males, where the latter will only mate with females living in the same nest. Some species in the genus have lost winged males completely, and only produce wingless males.<ref>{{Cite journal |last=Heinze |first=Jürgen |date=2017 |title=Life-history evolution in ants: the case of Cardiocondyla |journal=Proceedings of the Royal Society B: Biological Sciences |language=en |volume=284 |issue=1850 |pages=20161406 |doi=10.1098/rspb.2016.1406 |issn=0962-8452 |pmc=5360909 |pmid=28298341}}</ref> In ''[[Cardiocondyla elegans|C. elegans]],'' workers may transport newly emerged queens to other conspecific nests where the wingless males from unrelated colonies can mate with them, a behavioural adaptation that may reduce the chances of inbreeding.<ref>{{cite journal | vauthors = Vidal M, Königseder F, Giehr J, Schrempf A, Lucas C, Heinze J | title = Worker ants promote outbreeding by transporting young queens to alien nests | journal = Communications Biology | volume = 4 | issue = 1 | pages = 515 | date = May 2021 | pmid = 33941829 | pmc = 8093424 | doi = 10.1038/s42003-021-02016-1 }}</ref>

[[Image:Meat eater ant qeen excavating hole.jpg|thumb|left|Fertilised meat-eater ant queen beginning to dig a new colony]]

Mated females then seek a suitable place to begin a colony. There, they break off their wings using their tibial spurs and begin to lay and care for eggs. The females can selectively fertilise future eggs with the sperm stored to produce diploid workers or lay unfertilized haploid eggs to produce drones. The first workers to hatch, known as nanitics,<ref name="SuddFranks2013">{{cite book | vauthors = Sudd JH, Franks NR |title=The Behavioural Ecology of Ants|year=2013|publisher=Springer Science & Business Media|isbn=978-9400931237|page=41}}</ref> are weaker and smaller than later workers but they begin to serve the colony immediately. They enlarge the nest, forage for food, and care for the other eggs. Species that have multiple queens may have a queen leaving the nest along with some workers to found a colony at a new site,<ref name="HolldoblerWilsonAnts2"/> a process akin to [[Swarming (honey bee)|swarming]] in [[honeybee]]s.


=== Nests, colonies, and supercolonies ===
The typical ant species has a colony occupying a single nest, housing one or more queens, where the brood is raised. There are however more than 150 species of ants in 49 genera that are known to have colonies consisting of multiple spatially separated nests. These polydomous (as opposed to monodomous) colonies have food and workers moving between the nests.<ref>{{Cite journal |last1=Cook |first1=Zoe |last2=Franks |first2=Daniel W. |last3=Robinson |first3=Elva J.H. |date=2013 |title=Exploration versus exploitation in polydomous ant colonies |url=https://linkinghub.elsevier.com/retrieve/pii/S0022519313000477 |journal=Journal of Theoretical Biology|volume=323 |pages=49–56 |doi=10.1016/j.jtbi.2013.01.022|pmid=23380232 |bibcode=2013JThBi.323...49C }}</ref> Membership to a colony is identified by the response of worker ants which identify whether another individual belongs to their own colony or not. A signature cocktail of body surface chemicals (also known as cuticular hydrocarbons or CHCs) forms the so-called colony odor which other members can recognize.<ref>{{Cite journal |last1=Bos |first1=Nick |last2=d’Ettorre |first2=Patrizia |date=2012 |title=Recognition of Social Identity in Ants |journal=Frontiers in Psychology |volume=3 |page=83 |doi=10.3389/fpsyg.2012.00083 |issn=1664-1078 |pmc=3309994 |pmid=22461777 |doi-access=free }}</ref> Some ant species appear to be less discriminating and in the Argentine ant ''Linepithema humile,'' workers carried from a colony anywhere in the southern US and Mexico are acceptable within other colonies in the same region. Similarly workers from colonies that have established in Europe are accepted by any other colonies within Europe but not by the colonies in the Americas. The interpretation of these observations has been debated and some have been termed these large populations as supercolonies<ref>{{Cite journal |last=Moffett |first=Mark W. |date=2012 |title=Supercolonies of billions in an invasive ant: What is a society? |url=https://academic.oup.com/beheco/article-lookup/doi/10.1093/beheco/ars043 |journal=Behavioral Ecology|volume=23 |issue=5 |pages=925–933 |doi=10.1093/beheco/ars043 |issn=1465-7279}}</ref><ref>{{Cite journal |last1=Van Wilgenburg |first1=Ellen |last2=Torres |first2=Candice W. |last3=Tsutsui |first3=Neil D. |date=2010 |title=The global expansion of a single ant supercolony |journal=Evolutionary Applications|volume=3 |issue=2 |pages=136–143 |doi=10.1111/j.1752-4571.2009.00114.x |issn=1752-4571 |pmc=3352483 |pmid=25567914}}</ref><ref>{{Cite journal |last=Heikki Helanterä |date=2022 |title=Supercolonies of ants (Hymenoptera: Formicidae): ecological patterns, behavioural processes and their implications for social evolution |url=https://myrmecologicalnews.org/cms/index.php?option=com_content&view=category&id=1593&Itemid=441 |journal=Myrmecological News |volume=32 |pages=1–22 |doi=10.25849/MYRMECOL.NEWS_032:001}}</ref> while others have termed the populations as unicolonial.<ref>{{Cite journal |last1=Helanterä |first1=Heikki |last2=Strassmann |first2=Joan E. |last3=Carrillo |first3=Juli |last4=Queller |first4=David C. |date=2009 |title=Unicolonial ants: where do they come from, what are they and where are they going? |url=https://linkinghub.elsevier.com/retrieve/pii/S0169534709000895 |journal=Trends in Ecology & Evolution|volume=24 |issue=6 |pages=341–349 |doi=10.1016/j.tree.2009.01.013|pmid=19328589 |bibcode=2009TEcoE..24..341H }}</ref>

==Behaviour and ecology==

===Communication===
{{See also|Ant communication}}
[[File:Camponotus sericeus Senegal.jpg|thumb|Two ''[[Camponotus sericeus]]'' workers communicating through touch and pheromones]]
[[File:Ants find white cabbage larvae.webm|thumb|right|thumbtime=320|Ants find a dying [[Pieris rapae|white cabbage]] larvae that [[parasitoid wasp]]s larvae exited two days earlier.]]

Ants communicate with each other using [[pheromone]]s, sounds, and touch.<ref name="Jackson2006">{{cite journal | vauthors = Jackson DE, Ratnieks FL | title = Communication in ants | journal = Current Biology | volume = 16 | issue = 15 | pages = R570–R574 | date = August 2006 | pmid = 16890508 | doi = 10.1016/j.cub.2006.07.015 | s2cid = 5835320 | doi-access = free | bibcode = 2006CBio...16.R570J }}</ref> 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.<ref name="Jackson2006"/><ref>{{cite journal| vauthors = Goss S, Aron S, Deneubourg JL, Pasteels JM |year=1989|title=Self-organized shortcuts in the Argentine ant|journal=Naturwissenschaften |volume=76 |pages=579–581 |doi=10.1007/BF00462870 |issue=12|url=http://www.ulb.ac.be/sciences/use/publications/JLD/56.pdf |archive-url=https://ghostarchive.org/archive/20221009/http://www.ulb.ac.be/sciences/use/publications/JLD/56.pdf |archive-date=2022-10-09 |url-status=live|bibcode = 1989NW.....76..579G |s2cid=18506807}}</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 farther away. Several ant species even use "[[propaganda]] pheromones" to confuse enemy ants and make them fight among themselves.<ref>{{cite journal | vauthors = D'Ettorre P, Heinze J | journal = [[Acta Ethologica]] | year = 2001 | volume = 3 | pages = 67–82 | title = Sociobiology of slave-making ants | doi = 10.1007/s102110100038 | issue = 2 | s2cid = 37840769 }}</ref> Pheromones are produced by a wide range of structures including [[Dufour's gland]]s, poison glands and glands on the [[hindgut]], [[pygidium]], [[rectum]], [[sternum]], and hind [[tibia]].<ref name="insencyc"/> Pheromones also are exchanged, mixed with food, and passed by [[trophallaxis]], transferring information within the colony.<ref>{{cite book|vauthors=Detrain C, Deneubourg JL, Pasteels JM |year=1999|title=Information processing in social insects|publisher=Birkhäuser|isbn=978-3-7643-5792-4|pages=224–227}}</ref> This allows other ants to detect what task group (e.g., foraging or nest maintenance) other colony members belong to.<ref>{{cite journal | vauthors = Greene MJ, Gordon DM | title = Structural complexity of chemical recognition cues affects the perception of group membership in the ants Linephithema humile and Aphaenogaster cockerelli | journal = The Journal of Experimental Biology | volume = 210 | issue = Pt 5 | pages = 897–905 | date = March 2007 | pmid = 17297148 | doi = 10.1242/jeb.02706 | doi-access =  | s2cid = 14909476 }}</ref> In ant species with queen [[caste]]s, when the dominant queen stops producing a specific pheromone, workers begin to raise new queens in the colony.<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 | vauthors = Hickling R, Brown RL | title = Analysis of acoustic communication by ants | journal = The Journal of the Acoustical Society of America | volume = 108 | issue = 4 | pages = 1920–1929 | date = October 2000 | pmid = 11051518 | doi = 10.1121/1.1290515 | bibcode = 2000ASAJ..108.1920H }}</ref><ref>{{cite journal | vauthors = Roces F, 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=293–299 |doi=10.1007/s002650050292 |issue=5|bibcode=1996BEcoS..39..293R |s2cid=32884747 }}</ref>

===Defence <!-- Note this spelling is correct in UK English, if you came here to change this to Defense, please note that articles on Wikipedia choose between US and UK spelling based on priority-->===
{{see also|Defense in insects}}
[[File:Plectroctena sp ants.jpg|thumb|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. [[Paraponera clavata|Bullet ant]]s (''[[Paraponera clavata|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]].<ref>{{cite journal|doi=10.1002/arch.940010205|pages=155–160|title=Hemolytic activities of stinging insect venoms|journal=Archives of Insect Biochemistry and Physiology |volume=1|issue=2|year=1983 | vauthors = Schmidt JO, Blum MS, Overal WL }}</ref>

The sting of [[jack jumper ant]]s can be lethal for humans,<ref>{{cite journal | vauthors = Clarke PS | title = The natural history of sensitivity to jack jumper ants (Hymenoptera formicidae Myrmecia pilosula) in Tasmania | journal = The Medical Journal of Australia | volume = 145 | issue = 11–12 | pages = 564–566 | year = 1986 | pmid = 3796365 | doi=10.5694/j.1326-5377.1986.tb139498.x}}</ref> and an [[antivenom]] has been developed for it.<ref>{{cite journal | vauthors = Brown SG, Heddle RJ, Wiese MD, Blackman KE | title = Efficacy of ant venom immunotherapy and whole body extracts | journal = The Journal of Allergy and Clinical Immunology | volume = 116 | issue = 2 | pages = 464–465; author reply 465–466 | date = August 2005 | pmid = 16083810 | doi = 10.1016/j.jaci.2005.04.025 }}</ref> [[Fire ant]]s, ''[[Fire ant|Solenopsis]]'' spp., are unique in having a venom sac containing [[piperidine alkaloids]].<ref>{{cite journal | vauthors = Obin MS, Vander Meer RK | title = Gaster flagging by fire ants (''Solenopsis'' spp.): Functional significance of venom dispersal behavior | journal = Journal of Chemical Ecology | volume = 11 | issue = 12 | pages = 1757–1768 | date = December 1985 | pmid = 24311339 | doi = 10.1007/BF01012125 | bibcode = 1985JCEco..11.1757O | s2cid = 12182722 | url = https://zenodo.org/record/1232476 }}</ref> Their stings are painful and can be dangerous to hypersensitive people.<ref>{{cite journal | vauthors = Stafford CT | title = Hypersensitivity to fire ant venom | journal = Annals of Allergy, Asthma & Immunology | volume = 77 | issue = 2 | pages = 87–95; quiz 96–99 | date = August 1996 | pmid = 8760773 | doi = 10.1016/S1081-1206(10)63493-X }}</ref> [[Formicinae|Formicine ants]] secrete a poison from their glands, made mainly of [[formic acid]].<ref>{{cite journal|vauthors=Lopez LC, Morgan ED, Brand JM|year=1993|title=Hexadecanol and hexadecyl formate in the venom gland of formicine ants|journal=Philosophical Transactions of the Royal Society B|volume=341|issue=1296|pages=177–180|doi=10.1098/rstb.1993.0101|s2cid=85361145 }}</ref>

[[Image:WeaverAntDefense.JPG|thumb|left|A [[weaver ant]] in fighting position, [[mandible (insect)|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]] [[appendage]]s within the [[animalia|animal kingdom]].<ref name="TrapJawPatek">{{cite journal |vauthors=Patek SN, Baio JE, Fisher BL, Suarez AV |title=Multifunctionality and mechanical origins: ballistic jaw propulsion in trap-jaw ants |journal=Proceedings of the National Academy of Sciences of the United States of America |volume=103 |issue=34 |pages=12787–12792 |date=August 2006 |pmid=16924120 |pmc=1568925 |doi=10.1073/pnas.0604290103 |bibcode=2006PNAS..10312787P|doi-access=free }}</ref> One study of ''[[Odontomachus bauri]]'' recorded peak speeds of between {{convert|126|and|230|km/h|mph|abbr=on}}, with the jaws closing within 130 [[microsecond]]s 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.<ref name="TrapJawPatek" /> 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 receptor|sensory]] organs resembling hairs on the inside of the mandibles. The mandibles also permit slow and fine movements for other tasks. Trap-jaws also are seen in other ponerines such as ''[[Anochetus]]'', as well as some genera in the tribe [[Attini]], such as ''[[Daceton]]'', ''[[Orectognathus]]'', and ''[[Strumigenys]]'',<ref name="TrapJawPatek" /><ref>{{cite journal |vauthors=Gronenberg W |title=The trap-jaw mechanism in the dacetine ants ''Daceton armigerum'' and ''Strumigenys'' sp. |journal=The Journal of Experimental Biology |volume=199 |issue=Pt 9 |pages=2021–2033 |year=1996 |doi=10.1242/jeb.199.9.2021 |pmid=9319931 |url=http://jeb.biologists.org/cgi/reprint/199/9/2021.pdf |archive-url=https://ghostarchive.org/archive/20221009/http://jeb.biologists.org/cgi/reprint/199/9/2021.pdf |archive-date=2022-10-09 |url-status=live}}</ref> which are viewed as examples of [[convergent evolution]].

[[Camponotus saundersi|A Malaysian species of ant]] in the ''[[Camponotus]]'' ''cylindricus'' [[superspecies|group]] has enlarged mandibular glands that extend into their gaster. If combat takes a turn for the worse, a worker may perform a final act of [[autothysis|suicidal altruism]] by rupturing the membrane of its gaster, causing the content of its mandibular glands to burst from the [[anterior]] region of its head, spraying a poisonous, corrosive secretion containing [[acetophenone]]s and other chemicals that immobilise small insect attackers. The worker subsequently dies.<ref>{{cite journal | vauthors = Jones TH, Clark DA, Edwards AA, Davidson DW, Spande TF, Snelling RR | title = The chemistry of exploding ants, ''Camponotus'' spp. (''cylindricus'' complex) | journal = Journal of Chemical Ecology |volume=30 |issue=8 |pages=1479–1492 |date=August 2004 |pmid=15537154 |doi=10.1023/B:JOEC.0000042063.01424.28 | bibcode = 2004JCEco..30.1479J |s2cid=23756265}}</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. Secretions from the metapleural gland, unique to the ants, produce a complex range of chemicals including several with antibiotic properties.<ref name=":0">{{Cite journal |last1=Yek |first1=Sze Huei |last2=Mueller |first2=Ulrich G. |date=2011 |title=The metapleural gland of ants |url=http://www.sbs.utexas.edu/Muelleru/pubs/Yek-Mueller2011%20EarlyOnline.pdf |journal=Biological Reviews|volume=86 |issue=4 |pages=774–791 |doi=10.1111/j.1469-185X.2010.00170.x|pmid=21504532 |s2cid=7690884 }}</ref> Some worker ants maintain the hygiene of the colony and their activities include [[wikt:undertaker|undertaking]] or ''[[necrophoresis]]'', the disposal of dead nest-mates.<ref>{{cite journal | vauthors = Julian GE, Cahan S | title = Undertaking specialization in the desert leaf-cutter ant Acromyrmex versicolor | journal = Animal Behaviour | volume = 58 | issue = 2 | pages = 437–442 | date = August 1999 | pmid = 10458895 | doi = 10.1006/anbe.1999.1184 | s2cid = 23845331 }}</ref> [[Oleic acid]] has been identified as the compound released from dead ants that triggers necrophoric behaviour in ''Atta mexicana''<ref>{{cite journal|vauthors=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=353–360| doi = 10.1111/j.1365-3032.2006.00526.x|s2cid=84890901}}</ref> 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 behaviour.<ref>{{cite journal | vauthors = Choe DH, Millar JG, Rust MK | title = Chemical signals associated with life inhibit necrophoresis in Argentine ants | journal = Proceedings of the National Academy of Sciences of the United States of America | volume = 106 | issue = 20 | pages = 8251–8255 | date = May 2009 | pmid = 19416815 | pmc = 2688878 | doi = 10.1073/pnas.0901270106 | bibcode = 2009PNAS..106.8251C | doi-access = free }}</ref> In ''[[Megaponera analis]],'' injured ants are treated by nestmastes with secretions from their metapleural glands which protect them from infection.<ref>{{Cite journal |last1=Frank |first1=Erik. T. |last2=Kesner |first2=Lucie |last3=Liberti |first3=Joanito |last4=Helleu |first4=Quentin |last5=LeBoeuf |first5=Adria C. |last6=Dascalu |first6=Andrei |last7=Sponsler |first7=Douglas B. |last8=Azuma |first8=Fumika |last9=Economo |first9=Evan P. |last10=Waridel |first10=Patrice |last11=Engel |first11=Philipp |last12=Schmitt |first12=Thomas |last13=Keller |first13=Laurent |date=2023-12-29 |title=Targeted treatment of injured nestmates with antimicrobial compounds in an ant society |journal=Nature Communications |language=en |volume=14 |issue=1 |page=8446 |doi=10.1038/s41467-023-43885-w |issn=2041-1723 |pmc=10756881 |pmid=38158416|bibcode=2023NatCo..14.8446F }}</ref> ''Camponotus'' ants do not have a metapleural gland<ref name=":0" /> and ''[[Camponotus maculatus]]'' as well as ''[[Camponotus floridanus|C. floridanus]]'' workers have been found to amputate the affected legs of nestmates when the femur is injured. A femur injury carries a greater risk of infection unlike a tibia injury.<ref>{{Cite journal |last1=Frank |first1=Erik.T. |last2=Buffat |first2=Dany |last3=Liberti |first3=Joanito |last4=Aibekova |first4=Lazzat |last5=Economo |first5=Evan P. |last6=Keller |first6=Laurent |date=2024 |title=Wound-dependent leg amputations to combat infections in an ant society |journal=Current Biology |volume=34 |issue=14 |pages=3273–3278.e3 |language=en |doi=10.1016/j.cub.2024.06.021|pmid=38959879 |doi-access=free |bibcode=2024CBio...34.3273F }}</ref>

Nests may be protected from physical threats such as flooding and overheating by elaborate nest architecture.<ref>{{cite journal | vauthors = Tschinkel WR | title = The nest architecture of the Florida harvester ant, ''Pogonomyrmex badius'' | journal = Journal of Insect Science | volume = 4 | issue = 21 | pages = 21 | year = 2004 | pmid = 15861237 | pmc = 528881 | doi = 10.1093/jis/4.1.21 }}</ref><ref>{{cite journal| vauthors = 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=211–218|doi=10.1007/BF01240479|issue=2|s2cid=41870857}}</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 | vauthors = Maschwitz U, Moog J | title = Communal peeing: a new mode of flood control in ants | journal = Die Naturwissenschaften | volume = 87 | issue = 12 | pages = 563–565 | date = December 2000 | pmid = 11198200 | doi = 10.1007/s001140050780 | bibcode = 2000NW.....87..563M | s2cid = 7482935 }}</ref> ''[[Camponotus anderseni]]'', which nests in the cavities of wood in mangrove habitats, deals with submergence under water by switching to [[anaerobic respiration]].<ref>{{cite journal | vauthors = Nielsen MG, Christian KA | title = The mangrove ant, ''Camponotus anderseni'', switches to anaerobic respiration in response to elevated CO2 levels | journal = Journal of Insect Physiology | volume = 53 | issue = 5 | pages = 505–508 | date = May 2007 | pmid = 17382956 | doi = 10.1016/j.jinsphys.2007.02.002 }}</ref>

===Learning===
[[File:"Follow the leader".jpg|thumb|Two Weaver ants walking in [[Tandem running|tandem]]]]
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]]'' can lead a naïve nest-mate to newly discovered food by the process of [[tandem running]]. The follower obtains knowledge through its leading tutor. The leader is acutely sensitive to the progress of the follower and slows down when the follower lags and speeds up when the follower gets too close.<ref>{{cite journal | vauthors = Franks NR, Richardson T | title = Teaching in tandem-running ants | journal = Nature | volume = 439 | issue = 7073 | pages = 153 | date = January 2006 | pmid = 16407943 | doi = 10.1038/439153a | bibcode = 2006Natur.439..153F | s2cid = 4416276 | doi-access = free }}</ref>

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 brood care.<ref>{{cite journal | vauthors = Ravary F, Lecoutey E, Kaminski G, Châline N, Jaisson P | title = Individual experience alone can generate lasting division of labor in ants | journal = Current Biology | volume = 17 | issue = 15 | pages = 1308–1312 | date = August 2007 | pmid = 17629482 | doi = 10.1016/j.cub.2007.06.047 | s2cid = 13273984 | doi-access = free | bibcode = 2007CBio...17.1308R }}</ref>

===Nest construction===
{{Main|Ant colony}}
[[Image:WeaverAntNest.JPG|thumb|Leaf nest of [[weaver ants]], [[Pamalican]], [[Philippines]]]]

Complex nests are built by many ant species, but other species are nomadic and do not build permanent structures. Ants may form subterranean nests or build them on trees. These nests may be found in the ground, under stones or logs, inside logs, hollow stems, or even acorns. The materials used for construction include soil and plant matter,<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 | vauthors = Franks NR, Hooper J, Webb C, Dornhaus A | title = Tomb evaders: house-hunting hygiene in ants | journal = Biology Letters | volume = 1 | issue = 2 | pages = 190–192 | date = June 2005 | pmid = 17148163 | pmc = 1626204 | doi = 10.1098/rsbl.2005.0302 }}</ref>

The [[army ant]]s of South America, such as the ''[[Eciton burchellii]]'' species, 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|vauthors=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=164–169|doi=10.1111/j.1440-6055.2005.00462.x}}</ref>
[[File:AntBridge Crossing 10.jpg|thumb|left|Ant bridge]]
''[[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 materials.<ref>{{cite journal| vauthors = Frouz J |title=The Effect of Nest Moisture on Daily Temperature Regime in the Nests of ''Formica polyctena'' Wood Ants|journal=Insectes Sociaux| volume=47 |issue=3|year=2000|pages=229–235|doi=10.1007/PL00001708|s2cid=955282}}</ref><ref>{{Cite journal|last1=Kadochová|first1=Štěpánka|last2=Frouz|first2=Jan|date=2013|title=Thermoregulation strategies in ants in comparison to other social insects, with a focus on red wood ants ( Formica rufa group)|journal=F1000Research|volume=2|page=280|doi=10.12688/f1000research.2-280.v2|issn=2046-1402|pmc=3962001|pmid=24715967 |doi-access=free }}</ref>

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.<ref>{{Cite journal|journal= Biological Conservation| volume= 115 |issue= 2| year=2004| pages= 279–289| title= Impact of human dwellings on the distribution of the exotic Argentine ant: a case study in the Doñana National Park, Spain | vauthors = Carpintero S, Reyes-López J, de Reynac LA |doi=10.1016/S0006-3207(03)00147-2| bibcode= 2004BCons.115..279C }}</ref><ref>{{cite journal | vauthors = Friedrich R, Philpott SM | title = Nest-site limitation and nesting resources of ants (Hymenoptera: Formicidae) in urban green spaces | journal = Environmental Entomology | volume = 38 | issue = 3 | pages = 600–607 | date = June 2009 | pmid = 19508768 | doi = 10.1603/022.038.0311 | name-list-style = vanc | s2cid = 20555077 | doi-access = free }}</ref>

===Cultivation of food===
{{Main|Ant–fungus mutualism}}
[[Image:HoneyAnt.jpg|thumb|left|''[[Myrmecocystus]]'', [[honeypot ant]]s, store food to prevent colony famine.]]
Most ants are generalist predators, [[scavenger]]s, and indirect [[herbivore]]s,<ref name=riseofants/> 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<ref>{{cite journal | vauthors = Anderson KE, Russell JA, Moreau CS, Kautz S, Sullam KE, Hu Y, Basinger U, Mott BM, Buck N, Wheeler DE | title = Highly similar microbial communities are shared among related and trophically similar ant species | journal = Molecular Ecology | volume = 21 | issue = 9 | pages = 2282–2296 | date = May 2012 | pmid = 22276952 | doi = 10.1111/j.1365-294x.2011.05464.x | bibcode = 2012MolEc..21.2282A | s2cid = 32534515 }}</ref> to upgrade the nutritional value of the food they collect<ref>{{cite journal | vauthors = Feldhaar H, Straka J, Krischke M, Berthold K, Stoll S, Mueller MJ, Gross R | title = Nutritional upgrading for omnivorous carpenter ants by the endosymbiont Blochmannia | journal = BMC Biology | volume = 5 | pages = 48 | date = October 2007 | pmid = 17971224 | pmc = 2206011 | doi = 10.1186/1741-7007-5-48 | doi-access = free }}</ref> and allow them to survive in nitrogen poor regions, such as rainforest canopies.<ref>{{cite journal | vauthors = Russell JA, Moreau CS, Goldman-Huertas B, Fujiwara M, Lohman DJ, Pierce NE | title = Bacterial gut symbionts are tightly linked with the evolution of herbivory in ants | journal = Proceedings of the National Academy of Sciences of the United States of America | volume = 106 | issue = 50 | pages = 21236–21241 | date = December 2009 | pmid = 19948964 | pmc = 2785723 | doi = 10.1073/pnas.0907926106 | bibcode = 2009PNAS..10621236R | doi-access = free }}</ref> [[Leafcutter ant]]s (''[[Atta (genus)|Atta]]'' and ''[[Acromyrmex]]'') feed exclusively on a [[fungus]] that grows only within their colonies. They continually collect leaves which are taken to the colony, cut into tiny pieces and placed in fungal gardens. Ergates specialise in related tasks according to their sizes. The largest ants cut stalks, smaller workers chew the leaves and the smallest tend the fungus. Leafcutter ants are sensitive enough to recognise the reaction of the fungus to different plant material, apparently detecting chemical signals from the fungus. If a particular type of leaf is found to be toxic to the fungus, the colony will no longer collect it. The ants feed on structures produced by the fungi called ''[[gongylidia]]''. [[Symbiosis|Symbiotic]] bacteria on the exterior surface of the ants produce antibiotics that kill bacteria introduced into the nest that may harm the fungi.<ref>{{cite journal|journal=Nature|year=1999|volume=398|title=Ants, plants and antibiotics|author=Schultz TR|pages=747–748 | doi = 10.1038/19619|issue=6730 |url=http://www.insecta.ufv.br/Entomologia/ent/disciplina/ban%20160/AULAP/aula2/AntsandBacteria.pdf |archive-url=https://ghostarchive.org/archive/20221009/http://www.insecta.ufv.br/Entomologia/ent/disciplina/ban%20160/AULAP/aula2/AntsandBacteria.pdf |archive-date=2022-10-09 |url-status=live|bibcode = 1999Natur.398..747S |s2cid=5167611|doi-access=free}}</ref>

===Navigation===
[[File:Ant trail.jpg|thumb|upright|An ant [[trail]]]]
[[Forage|Foraging]] ants travel distances of up to {{convert|200|m|sigfig=1}} from their nest<ref name=foraging>{{cite journal | title = Ecology of foraging by ants | vauthors = Carrol CR, Janzen DH | journal = Annual Review of Ecology and Systematics | year = 1973 | volume = 4 | issue = 1 | pages = 231–257 | doi = 10.1146/annurev.es.04.110173.001311 | bibcode = 1973AnRES...4..231C }}</ref> 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<ref>{{cite journal | vauthors = Wittlinger M, Wehner R, Wolf H | title = The ant odometer: stepping on stilts and stumps | journal = Science | volume = 312 | issue = 5782 | pages = 1965–1967 | date = June 2006 | pmid = 16809544 | doi = 10.1126/science.1126912 | url = http://sun.menloschool.org/~dspence/biology/pdfs/ant_odometer.pdf | archive-url = https://web.archive.org/web/20110728042723/http://sun.menloschool.org/~dspence/biology/pdfs/ant_odometer.pdf | url-status = dead | bibcode = 2006Sci...312.1965W | s2cid = 15162376 | archive-date = 2011-07-28 }}</ref> and also by evaluating the movement of objects in their visual field ([[optical flow]]).<ref>{{cite journal | title = Desert ants Cataglyphis fortis use self-induced optic flow to measure distances travelled | vauthors = Ronacher B, Werner R | journal = Journal of Comparative Physiology A | year = 1995 | url = http://www.zool.uzh.ch/static/research/nb_wehner/literatur/pdf95/wehner19953.pdf | doi = 10.1007/BF00243395 | volume = 177 | s2cid = 4625001 | access-date = 2011-06-07 | archive-date = 2011-07-27 | archive-url = https://web.archive.org/web/20110727072227/http://www.zool.uzh.ch/static/research/nb_wehner/literatur/pdf95/wehner19953.pdf | url-status = dead }}</ref> Directions are measured using the position of the sun.<ref>{{cite journal | vauthors = Wehner R | title = Desert ant navigation: how miniature brains solve complex tasks | journal = Journal of Comparative Physiology A | volume = 189 | issue = 8 | pages = 579–588 | date = August 2003 | pmid = 12879352 | doi = 10.1007/s00359-003-0431-1 | s2cid = 4571290 | url = http://www.zool.uzh.ch/static/research/nb_wehner/literatur/pdf03/wehner20038.pdf | access-date = 2010-09-07 | archive-date = 2011-07-07 | archive-url = https://web.archive.org/web/20110707004545/http://www.zool.uzh.ch/static/research/nb_wehner/literatur/pdf03/wehner20038.pdf | url-status = dead }}</ref>
They [[path integration|integrate]] this information to find the shortest route back to their nest.<ref>{{cite journal | vauthors = Sommer S, Wehner R | title = The ant's estimation of distance travelled: experiments with desert ants, Cataglyphis fortis | journal = Journal of Comparative Physiology A | volume = 190 | issue = 1 | pages = 1–6 | date = January 2004 | pmid = 14614570 | doi = 10.1007/s00359-003-0465-4 | s2cid = 23280914 | url = https://www.zora.uzh.ch/id/eprint/669/1/ZORA_NL_669.pdf }}</ref>
Like all ants, they can also make use of visual landmarks when available<ref>{{cite journal | title = Visual navigation in desert ants ''Cataglyphis fortis'': are snapshots coupled to a celestial system of reference? | vauthors = Åkesson S, Wehner R | journal = [[Journal of Experimental Biology]] | year = 2002 | volume = 205 | pages = 1971–1978 | url = http://jeb.biologists.org/cgi/reprint/205/14/1971.pdf | issue = 14 | doi = 10.1242/jeb.205.14.1971 | pmid = 12089203 }}</ref> as well as olfactory and tactile cues to navigate.<ref>{{cite journal | vauthors = Steck K, Hansson BS, Knaden M | title = Smells like home: Desert ants, Cataglyphis fortis, use olfactory landmarks to pinpoint the nest | journal = Frontiers in Zoology | volume = 6 | pages = 5 | date = February 2009 | pmid = 19250516 | pmc = 2651142 | doi = 10.1186/1742-9994-6-5 | doi-access = free }}</ref><ref>{{cite journal | vauthors = Seidl T, Wehner R | title = Visual and tactile learning of ground structures in desert ants | journal = The Journal of Experimental Biology | volume = 209 | issue = Pt 17 | pages = 3336–3344 | date = September 2006 | pmid = 16916970 | doi = 10.1242/jeb.02364 | s2cid = 9642888 | doi-access = free }}</ref> Some species of ant are able to use the [[Earth's magnetic field]] for navigation.<ref>{{cite journal
 |title=Orientation by magnetic field in leaf-cutter ants, ''Atta colombica'' (Hymenoptera: Formicidae)
 |vauthors=Banks AN, Srygley RB |journal=Ethology
 |year=2003
 |volume=109
 |pages=835–846
 |doi=10.1046/j.0179-1613.2003.00927.x
 |issue=10
|bibcode=2003Ethol.109..835B }}</ref> The compound eyes of ants have specialised cells that detect polarised light from the Sun, which is used to determine direction.<ref>{{cite journal | vauthors = Fukushi T | title = Homing in wood ants, Formica japonica: use of the skyline panorama | journal = The Journal of Experimental Biology | volume = 204 | issue = Pt 12 | pages = 2063–2072 | date = June 2001 | doi = 10.1242/jeb.204.12.2063 | pmid = 11441048 | url = http://jeb.biologists.org/cgi/content/abstract/204/12/2063 }}</ref><ref>{{cite journal | vauthors = Wehner R, Menzel R | title = Homing in the ant Cataglyphis bicolor | journal = Science | volume = 164 | issue = 3876 | pages = 192–194 | date = April 1969 | pmid = 5774195 | doi = 10.1126/science.164.3876.192 | bibcode = 1969Sci...164..192W | s2cid = 41669795 }}</ref>
These polarization detectors are sensitive in the [[ultraviolet]] region of the light spectrum.<ref>{{cite book | title = The Insects: Structure and Function | url = https://archive.org/details/insectsstructure0000chap | url-access = registration | first = Reginald Frederick | last = Chapman | name-list-style=vanc | year = 1998 | edition = 4th | publisher = Cambridge University Press | isbn = 978-0-521-57890-5 | pages = [https://archive.org/details/insectsstructure0000chap/page/600 600] }}</ref> 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.<ref>{{cite journal | vauthors = Delsuc F | title = Army ants trapped by their evolutionary history | journal = PLOS Biology | volume = 1 | issue = 2 | pages = E37 | date = November 2003 | pmid = 14624241 | pmc = 261877 | doi = 10.1371/journal.pbio.0000037 | doi-access = free }}</ref>

===Locomotion===
The female worker ants do not have wings and reproductive females lose their wings after their mating flights in order to begin their colonies. Therefore, unlike their wasp ancestors, most ants travel by walking. Some species are capable of leaping. For example, Jerdon's jumping ant (''[[Harpegnathos saltator]]'') is able to jump by synchronising the action of its mid and hind pairs of legs.<ref>{{cite journal |vauthors=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=63–71 |doi=10.1007/BF01992052|s2cid=42304237 }}</ref> There are several species of [[gliding ant]] including ''Cephalotes atratus''; this may be a common trait among arboreal ants with small colonies. Ants with this ability are able to control their horizontal movement so as to catch tree trunks when they fall from atop the forest canopy.<ref>{{cite journal | vauthors = Yanoviak SP, Dudley R, Kaspari M | title = Directed aerial descent in canopy ants | journal = Nature | volume = 433 | issue = 7026 | pages = 624–626 | date = February 2005 | pmid = 15703745 | doi = 10.1038/nature03254 | url = http://www.canopyants.com/Nature05.pdf | url-status = dead | bibcode = 2005Natur.433..624Y | s2cid = 4368995 | archive-url = https://wayback.archive-it.org/all/20070616090223/http://www.canopyants.com/Nature05.pdf | archive-date = 2007-06-16 }}</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.<ref name="Mlot">{{cite journal | vauthors = Mlot NJ, Tovey CA, Hu DL | title = Fire ants self-assemble into waterproof rafts to survive floods | journal = Proceedings of the National Academy of Sciences of the United States of America | volume = 108 | issue = 19 | pages = 7669–7673 | date = May 2011 | pmid = 21518911 | pmc = 3093451 | doi = 10.1073/pnas.1016658108 | bibcode = 2011PNAS..108.7669M | doi-access = free }}</ref> 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=561–571|doi=10.1046/j.1365-2699.1998.2530561.x|bibcode=1998JBiog..25..561M |s2cid=84923599 }}</ref> ''[[Polyrhachis sokolova]]'', a species of ant found in [[Australia]]n [[mangrove]] swamps, can swim and live in underwater nests. Since they lack [[gill]]s, they go to trapped pockets of air in the submerged nests to breathe.<ref>{{cite journal |vauthors=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=521–533 |doi=10.1071/ZO9960521 |issue=5}}</ref>

===Cooperation and competition===
[[File:Ants eating cicada, jjron 22.11.2009.jpg|thumb|right|[[Meat ant|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 ant]]s are among the biggest and most [[basal (phylogenetics)|basal]] of ants. Like virtually all ants, they are [[Eusociality|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.<ref name=Crosland1988>{{cite journal | doi = 10.1111/j.1440-6055.1988.tb01179.x | title = Aspects of the biology of the primitive ant genus Myrmecia F. (Hymenoptera: Formicidae) | year = 1988 | journal = Australian Journal of Entomology | volume = 27 | pages = 305–309 | vauthors = Crosland MW, Crozier RH, Jefferson E | issue = 4 | doi-access = free }}</ref>

Some species attack and take over neighbouring ant colonies. Extreme specialists among these [[slave-raiding ant]]s, such as the [[Polyergus|Amazon ant]]s, are incapable of feeding themselves and need captured workers to survive.<ref>{{cite journal | vauthors = Diehl E, Junqueira LK, Berti-Filho E | 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 = 431–437 | date = August 2005 | pmid = 16341421 | doi = 10.1590/S1519-69842005000300008 | doi-access = free }}</ref> Captured workers of enslaved ''[[Temnothorax]]'' species have evolved a counter-strategy, destroying just the female pupae of the slave-making ''[[Temnothorax americanus]]'', but sparing the males (who do not take part in slave-raiding as adults).<ref>{{cite journal | vauthors = Achenbach A, Foitzik S | title = First evidence for slave rebellion: enslaved ant workers systematically kill the brood of their social parasite protomognathus americanus | journal = Evolution; International Journal of Organic Evolution | volume = 63 | issue = 4 | pages = 1068–1075 | date = April 2009 | pmid = 19243573 | doi = 10.1111/j.1558-5646.2009.00591.x | s2cid = 9546342 | doi-access = free }} See also ''[[New Scientist]]'', 9 April 2009</ref>

[[Image:Harpegnathos saltator fight.jpg|left|upright|thumb|A worker ''[[Harpegnathos saltator]]'' (a jumping ant) engaged in battle with a rival colony's queen (on top)]]
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 | vauthors = Henderson G, Andersen JF, Phillips JK, Jeanne RL | title = Internest aggression and identification of possible nestmate discrimination pheromones in polygynous antFormica montana | journal = Journal of Chemical Ecology | volume = 16 | issue = 7 | pages = 2217–2228 | date = July 1990 | pmid = 24264088 | doi = 10.1007/BF01026932 | bibcode = 1990JCEco..16.2217H | s2cid = 22878651 }}</ref>

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.<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=253–263|url=https://archive.org/details/ants_08424|doi=10.1046/j.1365-3113.1996.d01-12.x|issue=3|bibcode=1996SysEn..21..253W |s2cid=84198690}}</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=47–49|url=https://archive.org/details/ants_10687|doi=10.1080/00779962.1968.9722888|bibcode=1968NZEnt...4...47T |s2cid=83791596 }}</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 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.<ref name=TheAntParasites>Hölldobler & Wilson (1990), pp. 436–448</ref>

A [[sexual conflict|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.<ref>{{cite journal | vauthors = Fournier D, Estoup A, Orivel J, Foucaud J, Jourdan H, Le Breton J, Keller L | title = Clonal reproduction by males and females in the little fire ant | journal = Nature | volume = 435 | issue = 7046 | pages = 1230–1234 | date = June 2005 | pmid = 15988525 | doi = 10.1038/nature03705 | bibcode = 2005Natur.435.1230F | s2cid = 1188960 | url = https://serval.unil.ch/resource/serval:BIB_4B6DEB46C264.P001/REF.pdf }}</ref>

===Relationships with other organisms===
[[File:Myrmarachne plataleoides female thailand.jpg|thumb|The spider ''[[Myrmarachne plataleoides]]'' (female shown) mimics [[weaver ant]]s 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 [[inquiline]]s 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.<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=1–8|doi=10.2307/2387854|jstor=2387854|bibcode=1977Biotr...9....1R }}</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=165–193|doi=10.2307/3495552|url=http://www.fcla.edu/FlaEnt/fe80p165.pdf |archive-url=https://ghostarchive.org/archive/20221009/http://www.fcla.edu/FlaEnt/fe80p165.pdf |archive-date=2022-10-09 |url-status=live|jstor=3495552|doi-access=free}}</ref>

[[Image:Ant Receives Honeydew from Aphid.jpg|thumb|right|An ant collects [[Honeydew (secretion)|honeydew]] from an [[aphid]]]]
[[File:Calico scale.webm|thumb|thumbtime=32|[[Formicidae|Ant]]s collecting [[Honeydew (secretion)|honeydew]] from Calico scales ([[Eulecanium cerasorum]] then played at 30 times speed to show the pumping action of the scale.]]

[[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 | vauthors = Styrsky JD, Eubanks MD | title = Ecological consequences of interactions between ants and honeydew-producing insects | journal = Proceedings. Biological Sciences | volume = 274 | issue = 1607 | pages = 151–164 | date = January 2007 | pmid = 17148245 | pmc = 1685857 | doi = 10.1098/rspb.2006.3701 }}</ref> In some cases, the aphids secrete the honeydew in response to ants tapping them with their antennae. The ants in turn keep predators away from the aphids and will move them from one feeding location to another. When migrating to a new area, many colonies will take the aphids with them, to ensure a continued supply of honeydew. Ants also tend [[mealybug]]s to harvest their honeydew. Mealybugs may become a serious pest of [[pineapple]]s if ants are present to protect mealybugs from their natural enemies.<ref>{{cite journal |vauthors=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 |issue=5 |pages=1124–1129|doi=10.1093/jee/89.5.1124 |doi-access=free }}</ref>

[[Myrmecophile|Myrmecophilous]] (ant-loving) [[caterpillar]]s 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. The chemicals in the secretions of ''Narathura japonica'' alter the behavior of attendant ''Pristomyrmex punctatus'' workers, making them less aggressive and stationary. The relationship, formerly characterized as "mutualistic", is now considered as possibly a case of the ants being parasitically manipulated by the caterpillars.<ref>{{Cite journal |last1=Hojo|first1=Masaru K.|last2=Pierce|first2=Naomi E.|last3=Tsuji |first3=Kazuki|year=2015 |title=Lycaenid Caterpillar secretions manipulate attendant ant behavior|url= |journal=Current Biology |language=en |volume=25 |issue=17 |pages=2260–2264 |doi=10.1016/j.cub.2015.07.016|bibcode=2015CBio...25.2260H }}</ref> 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=76–82 |doi=10.1038/scientificamerican1092-76 |issue=4|bibcode=1992SciAm.267d..76D }}</ref>  A similar adaptation can be seen in [[Grizzled skipper]] butterflies that emit vibrations by expanding their wings in order to communicate with ants, which are natural predators of these butterflies.<ref>{{Cite journal|last=Elfferich|first=Nico W. | name-list-style=vanc |date=1998|title=Is the larval and imaginal signalling of Lycaenidae and other Lepidoptera related to communication with ants|url=http://natuurtijdschriften.nl/search?identifier=538588|journal=Deinsea|volume=4|issue=1}}</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 ant nest where it feeds on the ant larvae.<ref>{{cite journal | vauthors = Pierce NE, Braby MF, Heath A, Lohman DJ, Mathew J, Rand DB, Travassos MA | title = The ecology and evolution of ant association in the Lycaenidae (Lepidoptera) | journal = Annual Review of Entomology | volume = 47 | pages = 733–771 | year = 2002 | pmid = 11729090 | doi = 10.1146/annurev.ento.47.091201.145257 }}</ref> A number of specialized bacteria have been found as [[endosymbiont]]s in ant guts. Some of the dominant bacteria belong to the order [[Hyphomicrobiales]] whose members are known for being [[Nitrogen fixation|nitrogen-fixing]] [[symbiont]]s in [[legume]]s but the species found in ant lack the ability to fix nitrogen.<ref>{{cite journal | vauthors = Kautz S, Rubin BE, Russell JA, Moreau CS | title = Surveying the microbiome of ants: comparing 454 pyrosequencing with traditional methods to uncover bacterial diversity | journal = Applied and Environmental Microbiology | volume = 79 | issue = 2 | pages = 525–534 | date = January 2013 | pmid = 23124239 | pmc = 3553759 | doi = 10.1128/AEM.03107-12 | bibcode = 2013ApEnM..79..525K }}</ref><ref>{{cite journal | vauthors = Neuvonen MM, Tamarit D, Näslund K, Liebig J, Feldhaar H, Moran NA, Guy L, Andersson SG|author8-link=Siv G. E. Andersson| title = The genome of Rhizobiales bacteria in predatory ants reveals urease gene functions but no genes for nitrogen fixation | language = En | journal = Scientific Reports | volume = 6 | issue = 1 | pages = 39197 | date = December 2016 | pmid = 27976703 | pmc = 5156944 | doi = 10.1038/srep39197| bibcode = 2016NatSR...639197N }}</ref> [[Fungus-growing ants]] that make up the tribe [[Attini]], including [[leafcutter ant]]s, cultivate certain species of fungus in the genera ''[[Leucoagaricus]]'' or ''[[Leucocoprinus]]'' of the family [[Agaricaceae]]. 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 | vauthors = Dejean A, Solano PJ, Ayroles J, Corbara B, Orivel J | title = Insect behaviour: arboreal ants build traps to capture prey | journal = Nature | volume = 434 | issue = 7036 | pages = 973 | date = April 2005 | pmid = 15846335 | doi = 10.1038/434973a | bibcode = 2005Natur.434..973D | s2cid = 4428574 | doi-access = free }}</ref>
[[File:Ants on a dandelion.jpg|thumb|Ants may obtain nectar from flowers such as the [[dandelion]], but are only rarely known to pollinate flowers.]]
[[File:Aphids and ants.webm|thumb|thumbtime=35|Ants tending [[aphid]]s and collecting [[Honeydew (secretion)|honeydew]] secreted. A [[Podabrus rugosulus|wrinkled solder beetle]] flies in and eats an aphid before being chased away by the ants.]][[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 | vauthors = Frederickson ME, Gordon DM | 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. Biological Sciences | volume = 274 | issue = 1613 | pages = 1117–23 | date = April 2007 | pmid = 17301016 | pmc = 2124481 | doi = 10.1098/rspb.2006.0415 }}</ref> Although some ants obtain nectar from flowers, pollination by ants is somewhat rare, one example being of the pollination of the orchid ''[[Leporella fimbriata]]'' which induces male ''Myrmecia urens'' to [[Pseudocopulation|pseudocopulate]] with the flowers, transferring pollen in the process.<ref>{{cite journal | vauthors = Peakall R, Beattie AJ, James SH | title = Pseudocopulation of an orchid by male ants: a test of two hypotheses accounting for the rarity of ant pollination | journal = Oecologia | volume = 73 | issue = 4 | pages = 522–524 | date = October 1987 | pmid = 28311968 | doi = 10.1007/BF00379410 | s2cid = 3195610 | bibcode = 1987Oecol..73..522P }}</ref> One theory that has been proposed for the rarity of pollination is that the secretions of the metapleural gland inactivate and reduce the viability of pollen.<ref>{{cite journal|title=Ant Inhibition of Pollen Function: A Possible Reason Why Ant Pollination is Rare|first1=Andrew J.|last1= Beattie| last2 = Turnbull | first2 = Christine | first3 = Knox | last3 =  R. B. | first4 = Williams | last4 = E. G. | name-list-style=vanc |journal=American Journal of Botany|volume=71| issue=3| year=1984| pages= 421–426|doi=10.2307/2443499|jstor=2443499}}</ref><ref>{{cite book|last=New|first=Tim R.|chapter=Classic Themes: Ants, Plants and Fungi|date=2017|pages=63–103|publisher=Springer International Publishing|doi=10.1007/978-3-319-58292-4_4|isbn=9783319582917|title=Mutualisms and Insect Conservation}}</ref> Some plants, mostly angiosperms but also some ferns,<ref>{{Cite journal |last1=Suissa |first1=Jacob S. |last2=Li |first2=Fay-Wei |last3=Moreau |first3=Corrie S. |date=2024 |title=Convergent evolution of fern nectaries facilitated independent recruitment of ant-bodyguards from flowering plants |journal=Nature Communications |language=en |volume=15 |issue=1 |page=4392 |doi=10.1038/s41467-024-48646-x |issn=2041-1723 |pmc=11126701 |pmid=38789437|bibcode=2024NatCo..15.4392S }}</ref> have special nectar exuding structures, [[Extrafloral nectary|extrafloral nectaries]], that provide food for ants, which in turn [[plant defense against herbivory|protect]] the plant from more damaging [[herbivorous]] insects.<ref>{{cite journal | vauthors = 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 = 119–124 | volume = 7 | issue = 2 | doi = 10.1111/j.1479-8298.2004.00057.x | s2cid = 85428729 | url = https://barrel.repo.nii.ac.jp/?action=repository_uri&item_id=5267 | hdl = 10252/00005880 | hdl-access = free }}</ref> 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 [[Epiphyte|epiphytic]] vines. [[Isotopic labelling]] studies suggest that plants also obtain nitrogen from the 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''|vauthors=Fischer RC, Wanek W, Richter A, Mayer V |year=2003|journal=Journal of Ecology|volume=91|issue=1 |pages=126–134|doi=10.1046/j.1365-2745.2003.00747.x  |doi-access=free|bibcode=2003JEcol..91..126F }}</ref> In return, the ants obtain food from protein- and lipid-rich [[Beltian bodies]]. In [[Fiji]] ''[[Philidris nagasau]]'' (Dolichoderinae) are known to selectively grow species of epiphytic ''[[Squamellaria]]'' (Rubiaceae) which produce large domatia inside which the ant colonies nest. The ants plant the seeds and the domatia of young seedling are immediately occupied and the ant faeces in them contribute to rapid growth.<ref>{{cite journal | vauthors = Chomicki G, Renner SS | title = Obligate plant farming by a specialized ant | journal = Nature Plants | volume = 2 | issue = 12 | pages = 16181 | date = November 2016 | pmid = 27869787 | doi = 10.1038/nplants.2016.181 | bibcode = 2016NatPl...216181C | s2cid = 23748032 }}</ref> Similar dispersal associations are found with other dolichoderines in the region as well.<ref>{{cite journal | vauthors = Chomicki G, Janda M, Renner SS | title = The assembly of ant-farmed gardens: mutualism specialization following host broadening | journal = Proceedings. Biological Sciences | volume = 284 | issue = 1850 | pages = 20161759 | date = March 2017 | pmid = 28298344 | pmc = 5360912 | doi = 10.1098/rspb.2016.1759 }}</ref> Another example of this type of [[ectosymbiosis]] comes from the ''[[Macaranga]]'' tree, which has stems adapted to house colonies of ''[[Crematogaster]]'' ants.<ref>{{cite journal | vauthors = Fiala B, Maschwitz U, Pong TY, Helbig AJ | title = Studies of a South East Asian ant-plant association: protection of Macaranga trees by Crematogaster borneensis | journal = Oecologia | volume = 79 | issue = 4 | pages = 463–470 | date = June 1989 | pmid = 28313479 | doi = 10.1007/bf00378662 | s2cid = 21112371 | bibcode = 1989Oecol..79..463F | url = https://opus.bibliothek.uni-wuerzburg.de/files/3903/Fiala_Macaranga.pdf }}</ref>

Many plant species have seeds that are adapted for dispersal by ants.<ref name="Aaron D 2009">{{cite journal|title=Convergent evolution of seed dispersal by ants, and phylogeny and biogeography in flowering plants: A global survey|first1=Szabolcs |last1=Lengyel| first2 = Aaron D. | last2 = Gove | first3 = Andrew M. | last3 = Latimer | first4 = Jonathan D. | last4 = Majer | first5 = Robert R. | last5 = Dunn |name-list-style=vanc|year=2010|journal=Perspectives in Plant Ecology, Evolution and Systematics |volume=12|issue=1 | pages=43–55|doi=10.1016/j.ppees.2009.08.001|bibcode=2010PPEES..12...43L }}</ref> [[Seed#Seed dispersal|Seed dispersal]] by ants or [[myrmecochory]] is widespread, and new estimates suggest that nearly 9% of all plant species may have such ant associations.<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=481–492|doi=10.1111/j.0030-1299.2006.14258.x|bibcode= 2006Oikos.112..481G|citeseerx=10.1.1.530.1306}}</ref><ref name="Aaron D 2009"/> Often, seed-dispersing ants perform directed dispersal, depositing the seeds in locations that increase the likelihood of seed survival to reproduction.<ref>{{cite journal |vauthors=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=1–13 |doi=10.1086/284769|s2cid=85317649 }}</ref> Some plants in arid, fire-prone systems are particularly dependent on ants for their survival and dispersal as the seeds are transported to safety below the ground.<ref>{{cite journal |vauthors=Auld TD |date=1996|title=Ecology of the Fabaceae in the Sydney region: fire, ants and the soil seedbank |journal=Cunninghamia|volume=4|issue=22}}</ref> Many ant-dispersed seeds have special external structures, [[elaiosome]]s, that are sought after by ants as food.<ref>{{cite journal|vauthors=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=55–62 |doi=10.1007/s00040-004-0773-x|s2cid=21974767}}</ref> Ants can substantially alter rate of decomposition and nutrient cycling in their nest.<ref>{{Cite journal|last=Frouz|first=Jan|date=1997|title=The effect of wood ants (Formica polyctena Foerst) on the transformation of phosphorus in a spruce plantation|url=https://ci.nii.ac.jp/naid/10027223277/|journal=Pedobiologia|volume=41|issue=5 |pages=437–447|doi=10.1016/S0031-4056(24)00314-7 |doi-access=free|bibcode=1997Pedob..41..437F }}</ref><ref>{{Cite journal|last1=Frouz|first1=J|last2=Jílková|first2=V|date=2008|title=The effect of ants on soil properties and processes (Hymenoptera: Formicidae)|url=https://myrmecologicalnews.org/cms/index.php?filename=volume11/mn11_191-199_printable.pdf&format=raw&option=com_download&view=download|journal=Myrmecological News|volume=1|pages=191–199}}</ref>&nbsp;By myrmecochory and modification of soil conditions they substantially alter vegetation and nutrient cycling in surrounding ecosystem.<ref>{{cite book|last1=Frouz|first1=Jan|chapter=Contribution of wood ants to nutrient cycling and ecosystem function|date=2016|chapter-url=https://www.cambridge.org/core/books/wood-ant-ecology-and-conservation/contribution-of-wood-ants-to-nutrient-cycling-and-ecosystem-function/83392472CE3BE1547CDD95120D4C9E12|title=Wood Ant Ecology and Conservation|pages=207–220|editor-last=Robinson|editor-first=Elva J. H.|series=Ecology, Biodiversity and Conservation|place=Cambridge|publisher=Cambridge University Press|isbn=978-1-107-04833-1|access-date=2021-07-12|last2=Jílková|first2=Veronika|last3=Sorvari|first3=Jouni|editor2-last=Stockan|editor2-first=Jenni A.}}</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|vauthors=Hughes L, Westoby M |date=1992|title=Capitula on stick insect eggs and elaiosomes on seeds: convergent adaptations for burial by ants|journal=Functional Ecology|volume=6|pages=642–648|doi=10.2307/2389958|issue=6|jstor=2389958|bibcode=1992FuEco...6..642. }}</ref>
[[File:Common jassid nymph and ant02.jpg|thumb|right|A [[meat ant]] tending a common [[leafhopper]] nymph]]
[[File:PhidippusWithCutwormAnts.webm|thumb|thumbtime=1|right|Bold Jumping Spider ([[Phidippus audax]]) with a cutworm (tribe [[Noctuini]]) and then lost to ants (Family Formicidae)]]
[[File:Long jawed spider ants.webm|thumb|thumbtime=2|right|[[Formicidae|Ant]]s from different colonies [[Kleptoparasitism|steal]] the [[cranefly]] that a pair of [[Tetragnathidae|Long-jawed orb weaver spider]]s were consuming.]]
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.<ref name=foraging/> Some termites, including ''[[Nasutitermes corniger]]'', form associations with certain ant species to keep away predatory ant species.<ref>{{cite journal|journal= Journal of Insect Behavior |title= Behavioural Interactions Between ''Crematogaster brevispinosa rochai'' Forel (Hymenoptera: Formicidae) and Two Nasutitermes Species (Isoptera: Termitidae)|volume=18|issue= 1|pages=1–17|year= 2005|vauthors=Quinet Y, Tekule N, de Biseau JC |doi=10.1007/s10905-005-9343-y|bibcode= 2005JIBeh..18....1Q|s2cid= 33487814}}</ref> The tropical wasp ''[[Mischocyttarus drewseni]]'' coats the pedicel of its nest with an ant-repellent chemical.<ref>{{cite journal | vauthors = Jeanne RL |year=1972|title=Social biology of the neotropical wasp ''Mischocyttarus drewseni''|journal=Bull. Mus. Comp. Zool.|volume=144|pages=63–150|url=http://biostor.org/reference/692}}</ref> 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 ''[[Agelaia multipicta|A. multipicta]]'', defend against ants by blasting them off the nest with bursts of wing buzzing.<ref>{{Cite journal|url = https://www.researchgate.net/publication/233927144|title = Foraging in Social Wasps: Agelaia lacks recruitment to food (Hymenoptera: Vespidae)|last = Jeanne|first = Robert | name-list-style=vanc |date = July 1995|journal = Journal of the Kansas Entomological Society}}</ref> Stingless bees (''[[Trigona]]'' and ''[[Melipona]]'') use chemical defences against ants.<ref name=foraging/>

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 |vauthors=Sivinski J, Marshall S, Petersson E |year=1999 |title=Kleptoparasitism and phoresy in the Diptera |journal=[[Florida Entomologist]] |volume=82 |issue=2 |pages=179–197 |url=http://www.fcla.edu/FlaEnt/fe82p179.pdf |archive-url=https://ghostarchive.org/archive/20221009/http://www.fcla.edu/FlaEnt/fe82p179.pdf |archive-date=2022-10-09 |url-status=live|doi=10.2307/3496570 |jstor=3496570|doi-access=free }}</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"/>
[[File:Oecophylla_fungus.jpg|thumb|''Oecophylla smaragdina'' killed by a fungus]]
Fungi in the genera ''[[Cordyceps]]'' and ''[[Ophiocordyceps]]'' infect ants. Ants react to their infection by climbing up plants and sinking their mandibles into plant tissue. The fungus kills the ants, grows on their remains, and produces a [[fruiting body]]. 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=116–117}}</ref> in a microhabitat that best suits the fungus.<ref>{{cite journal | vauthors = Andersen SB, Gerritsma S, Yusah KM, Mayntz D, Hywel-Jones NL, Billen J, Boomsma JJ, Hughes DP | title = The life of a dead ant: the expression of an adaptive extended phenotype | journal = The American Naturalist | volume = 174 | issue = 3 | pages = 424–433 | date = September 2009 | pmid = 19627240 | doi = 10.1086/603640 | hdl = 11370/e6374602-b2a0-496c-b78e-774b34fb152b | s2cid = 31283817 | url = https://pure.rug.nl/ws/files/67295795/603640.pdf | hdl-access = free }}</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=43–451|doi=10.2307/3494966|jstor=3494966|url=https://naldc-legacy.nal.usda.gov/naldc/download.xhtml?id=23159&content=PDF|access-date=2017-10-25|archive-date=2021-03-09|archive-url=https://web.archive.org/web/20210309052432/https://naldc-legacy.nal.usda.gov/naldc/download.xhtml?id=23159&content=PDF|url-status=dead}}</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, 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.<ref>{{cite journal | vauthors = Poinar G, Yanoviak SP | 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 | issue = 2 | pages = 145–153 | date =  2008 | pmid = 18038201 | doi = 10.1007/s11230-007-9125-3| s2cid = 8473071 }}</ref>

[[File:Jumping spider with prey.jpg|thumb|Spiders (Like this ''[[Menemerus bivittatus|Menemerus]]'' jumping spider) sometimes feed on ants|right]]
A study of ''[[Temnothorax nylanderi]]'' colonies in Germany found that workers parasitized by the tapeworm ''[[Anomotaenia brevis]]'' (ants are intermediate hosts, the [[Host (biology)|definitive hosts]] are [[woodpecker]]s) lived much longer than unparasitized workers and had a reduced mortality rate, comparable to that of the queens of the same species, which live for as long as two decades.<ref>{{cite journal | vauthors = Stoldt M, Klein L, Beros S, Butter F, Jongepier E, Feldmeyer B, Foitzik S | title = Parasite Presence Induces Gene Expression Changes in an Ant Host Related to Immunity and Longevity | journal = Genes | volume = 12 | issue = 1 | pages = 202118 | date = January 2021 | pmc = 8131941 | doi = 10.1098/rsos.202118 | pmid = 34017599 }}</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=75–101 |doi=10.1111/j.1469-7998.1996.tb05487.x }}</ref>

[[Army ant]]s 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-follower|ant-following]]" birds such as [[antbird]]s and [[woodcreeper]]s.<ref name = "Willis">{{cite journal|doi=10.1146/annurev.es.09.110178.001331 | vauthors = Willis E, Oniki Y |year=1978|title=Birds and Army Ants|journal=Annual Review of Ecology and Systematics|volume=9| issue = 1 |pages=243–263| bibcode = 1978AnRES...9..243W }}</ref><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=271–275|url=http://www.ibiologia.unam.mx/pdf/links/neo/rev12/vol_12_3/orni_12_3_%20271-276.pdf |archive-url=https://ghostarchive.org/archive/20221009/http://www.ibiologia.unam.mx/pdf/links/neo/rev12/vol_12_3/orni_12_3_%20271-276.pdf |archive-date=2022-10-09 |url-status=live|access-date=8 June 2008}}
</ref> This behaviour was once considered [[Mutualism (biology)|mutualistic]], but later studies found the birds to be [[parasitism|parasitic]]. Direct [[kleptoparasitism]] (birds stealing food from the ants' grasp) is rare and has been noted in [[Inca dove]]s which pick seeds at nest entrances as they are being transported by species of ''[[Pogonomyrmex]]''.<ref>{{Cite journal|last1=Inzunza|first1=Ernesto Ruelas|last2=Martínez-Leyva|first2=J. Eduardo|last3=Valenzuela-González|first3=Jorge E. | name-list-style=vanc |title=Doves kleptoparasitize ants|journal=The Southwestern Naturalist|volume=60|issue=1|pages=103–106|doi=10.1894/msh-03.1|year=2015|s2cid=85633598}}</ref> Birds that follow ants eat many prey insects and thus decrease the foraging success of ants.<ref>{{cite journal|author=Wrege PH|title=Antbirds parasitize foraging army ants|journal=Ecology|volume=86|year=2005| pages=555–559|doi=10.1890/04-1133|last2=Wikelski|first2=Martin|last3=Mandel|first3=James T.|last4=Rassweiler|first4=Thomas|last5=Couzin|first5=Iain D. | name-list-style=vanc |issue=3|bibcode=2005Ecol...86..555W }}</ref> Birds indulge in a peculiar behaviour called [[Anting (bird activity)|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 [[ectoparasite]]s from the birds.

[[Anteater]]s, [[aardvark]]s, [[pangolin]]s, [[echidna]]s and [[numbat]]s 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. About 12%, 16%, and 4% of their faecal volume in spring, summer and autumn, respectively, is composed of ants.<ref>{{cite journal|vauthors=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=551–561 |doi=10.1139/z99-004|bibcode=1999CaJZ...77..551S }}</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. 619–629</ref> On the other hand, ants may become nuisances when they invade buildings or cause economic losses.

In some parts of the world (mainly Africa and South America), large ants, especially [[army ant]]s, are used as [[surgical 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 |vauthors=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 |archive-url=https://web.archive.org/web/20070616090223/http://www.ewma.org/pdf/fall04/Historical_Aspects.pdf |archive-date=2007-06-16|pages=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=1861–1864|doi=10.1001/jama.1925.02660500069048|issue=24}}</ref><ref>{{cite book|first=Robert M.|last=Sapolsky | name-list-style=vanc |author-link=Robert Sapolsky|year=2001|title=A Primate's Memoir: A Neuroscientist's Unconventional Life Among the Baboons|url=https://archive.org/details/primatesmemoir00robe|url-access=registration|pages=[https://archive.org/details/primatesmemoir00robe/page/156 156]|publisher=Simon and Schuster|isbn=978-0-7432-0241-1}}</ref> The large heads of the dinergates (soldiers) of the leafcutting ant ''[[Atta cephalotes]]'' are also used by native surgeons in closing wounds.<ref>{{cite book|last1=Wheeler|first1=William M. | name-list-style=vanc |author-link=William Morton Wheeler|title=Ants: Their Structure, Development and Behavior|year=1910|series=[[Columbia University Biological Series]]|volume=9|publisher=[[Columbia University Press]]|isbn=978-0-231-00121-2|pages=10|doi=10.5962/bhl.title.1937|lccn=10008253//r88|oclc=560205|title-link=s:Index:Ants, Wheeler (1910).djvu }}</ref>

Some ants have [[ant venom|toxic venom]] and are of [[Ants of medical importance|medical importance]]. The species include ''[[Paraponera clavata]]'' (tocandira) and ''[[Dinoponera]]'' spp. (false tocandiras) of South America<ref>{{cite journal | vauthors = Haddad Junior V, Cardoso JL, Moraes RH | 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 Sao Paulo | volume = 47 | issue = 4 | pages = 235–238 | year = 2005 | pmid = 16138209 | doi = 10.1590/S0036-46652005000400012 | doi-access = free | hdl = 11449/30504 | hdl-access = free }}</ref> and the ''Myrmecia'' ants of Australia.<ref>{{cite journal | vauthors = McGain F, Winkel KD | title = Ant sting mortality in Australia | journal = Toxicon | volume = 40 | issue = 8 | pages = 1095–1100 | date = August 2002 | pmid = 12165310 | doi = 10.1016/S0041-0101(02)00097-1 | bibcode = 2002Txcn...40.1095M }}</ref>

In [[South Africa]], ants are used to help harvest the seeds of [[rooibos]] (''Aspalathus linearis''), a plant used to make a herbal tea. The plant disperses its seeds widely, making manual collection difficult. Black ants collect and store these and other seeds in their nest, where humans can gather them ''en masse''. Up to half a pound (200&nbsp;g) of seeds may be collected from one ant-heap.<ref>{{cite web|url=http://www.ciel.org/Publications/InnovativeMechanisms.pdf|title=Innovative mechanisms for sharing benefits of biodiversity and related knowledge|vauthors=Downes D, Laird SA |year=1999|publisher=The Center for International Environmental Law|access-date=8 June 2008| archive-url= https://web.archive.org/web/20080423211123/http://www.ciel.org/Publications/InnovativeMechanisms.pdf| archive-date= 23 April 2008 | url-status= live}}</ref><ref>{{cite journal|journal=Economic Botany|title=Rooibos tea, a South African contribution to world beverages|volume=17|issue=3|pages=186–194|year=1963|vauthors=Cheney RH, Scholtz E |doi=10.1007/BF02859435|bibcode=1963EcBot..17..186C |s2cid=37728834}}</ref>

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.<ref>{{cite journal|title=The influence of sociality on the conservation biology of social insects | vauthors = Chapman RE, Bourke AF |journal= Ecology Letters|volume=4|issue=6|year=2001|pages=650–662|doi=10.1046/j.1461-0248.2001.00253.x|doi-access=| bibcode = 2001EcolL...4..650C | s2cid = 86796899 }}</ref>

===As food===
{{See also|Entomophagy}}
[[File:Ants For Food SG.jpg|thumb|left|Roasted ants in Colombia]]
[[Image:Ants Eggs Market Thailand.jpg|thumb|left|Ant larvae for sale in [[Isaan]], Thailand]]
Ants and their larvae are eaten in different parts of the world. The eggs of two species of ants are used in Mexican ''[[escamoles]]''. They are considered a form of insect [[caviar]] and can sell for as much as US$50 per kg going up to US$200 per kg (as of 2006) because they are seasonal and hard to find.<ref>{{Cite journal |last1=Cruz-Labana |first1=J. D. |last2=Tarango-Arámbula |first2=L. A. |last3=Alcántara-Carbajal |first3=J. L. |last4=Pimentel-López |first4=J. |last5=Ugalde-Lezama |first5=S. |last6=Ramírez-Valverde |first6=G. |last7=Méndez-Gallegos |first7=S. J. |date=2014 |title=Habitat use by the "Escamolera" ant (Liometopum apiculatum Mayr) in central Mexico |url=http://www.scielo.org.mx/scielo.php?script=sci_abstract&pid=S1405-31952014000600001&lng=es&nrm=iso&tlng=en |journal=Agrociencia|volume=48 |issue=6 |pages=569–582 |issn=1405-3195}}</ref> In the [[Colombia]]n department of [[Santander Department|Santander]], ''hormigas culonas'' (roughly interpreted as "large-bottomed ants") ''[[Atta laevigata]]'' are toasted alive and eaten.<ref>{{cite journal | vauthors = DeFoliart GR | title = Insects as food: why the western attitude is important | journal = Annual Review of Entomology | volume = 44 | pages = 21–50 | year = 1999 | pmid = 9990715 | 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|last=Bingham|first= C.T.|year=1903|title=Fauna of British India. Hymenoptera. Volume 2|page=311|url=https://archive.org/details/hymenoptera02bing/page/311/mode/1up| publisher=Taylor and Francis|place=London}}</ref> [[Oecophylla|Weaver ant]] eggs and larvae, as well as the ants, may be used in a [[Thai salad]], ''yam'' ({{langx|th|ยำ}}), in a dish called ''yam khai mot daeng'' ({{langx|th|ยำไข่มดแดง}}) or red [[ant eggs|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=191–200|url=http://www.naturalhistorymag.com/editors_pick/1921_03-04_pick.html}}</ref> Ants or their pupae are used as starters for yogurt making in parts of Bulgaria and Turkey.<ref>{{Cite journal |last=Mutlu Sirakova |first=Sevgi |date=2023 |title=Forgotten Stories of Yogurt: Cultivating Multispecies Wisdom |url=https://journals.sagepub.com/doi/10.1177/02780771231194779 |journal=Journal of Ethnobiology |language=en |volume=43 |issue=3 |pages=250–261 |doi=10.1177/02780771231194779 |issn=0278-0771}}</ref>

In his ''First Summer in the Sierra'', [[John Muir]] notes that the [[Mono people|Digger Indians]] of [[California]] ate the tickling, acid gasters of the large jet-black [[carpenter ant]]s. The Mexican Indians eat the [[Honeypot ant|repletes]], or living honey-pots, of the [[honey ant]] (''[[Myrmecocystus]]'').<ref name="beq"/>

===As pests===
{{See also|Ants of medical importance}}
[[File:Monomorium pharaonis.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 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]], [[Tetramorium immigrans|immigrant pavement ant]], [[yellow crazy ant]], [[banded sugar ant]], [[pharaoh ant]], [[Formica rufa|red wood ant]], [[black carpenter ant]], [[Tapinoma sessile|odorous house ant]], [[red imported fire ant]], and [[Myrmica rubra|European fire ant]]. Some ants will raid stored food, some will seek water sources, others may damage indoor structures, some may damage agricultural crops directly or by aiding sucking pests. Some will sting or bite.<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|access-date=5 June 2008}}</ref> The adaptive nature of ant colonies make it nearly impossible to eliminate entire colonies and most pest management practices aim to control local populations and tend to be temporary solutions. Ant populations are managed by a combination of approaches that make use of chemical, biological, and physical methods. Chemical methods include the use of insecticidal bait which is gathered by ants as food and brought back to the nest where the poison is inadvertently spread to other colony members through [[trophallaxis]]. Management is based on the species and techniques may vary according to the location and circumstance.<ref name="pests" />

===In science and technology===
{{See also|Myrmecology|Biomimetics|Ant colony optimization algorithms}}
[[File:Formicariums.jpg|''[[Camponotus nearcticus]]'' workers travelling between two formicaria through connector tubing|thumb|right]]
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, [[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. [[Ultraviolet vision]] was first discovered in ants by [[John Lubbock, 1st Baron Avebury|Sir John Lubbock]] in 1881.<ref>{{cite journal| vauthors = Lubbock J | year=1881|title=Observations on ants, bees, and wasps. IX. Color of flowers as an attraction to bees: Experiments and considerations thereon|journal=J. Linn. Soc. Lond. (Zool.)|volume=16|pages=110–112|doi=10.1111/j.1096-3642.1882.tb02275.x|issue=90|doi-access=| url=https://www.biodiversitylibrary.org/part/377086}}</ref> 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 strategy|evolutionarily stable strategies]].<ref>{{cite book|vauthors=Stadler B, Dixon AF|year=2008|title=Mutualism: Ants and their insect partners|publisher=Cambridge University Press|isbn=978-0-521-86035-2}}</ref> Ant colonies may 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=17–20|hdl=1811/3802}}</ref> Individuals may be tracked for study by marking them with dots of colours.<ref>{{cite journal|vauthors=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=74–78|doi=10.2307/3496231|jstor=3496231|doi-access=free}}</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, for example [[Ant colony optimization]] and [[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.<ref name="SANdisk">{{cite journal |vauthors=Dicke E, Byde A, Cliff D, Layzell P |year=2004 |title=An ant-inspired technique for storage area network design |journal=Proceedings of Biologically Inspired Approaches to Advanced Information Technology: First International Workshop, BioADIT 2004 LNCS 3141 |pages=364–379 | veditors = Ispeert AJ, Murata M, Wakamiya N }}</ref>

===As pets===
{{main|Ant-keeping}}
From the late 1950s through the late 1970s, [[formicarium|ant farms]] were popular educational children's toys in the United States. Some later commercial versions use transparent gel instead of soil, allowing greater visibility at the cost of stressing the ants with unnatural light.<ref>{{cite patent |country= US |number= 5803014 |status= granted |title= Habitat media for ants and other invertebrates|pubdate= |gdate= 8 September 1998 |inventor=  Guri A |assign1= Plant Cell Technology Inc }}</ref>

===In culture===
[[Image:The Ant and the Grasshopper - Project Gutenberg etext 19994.jpg|thumb|left|[[Aesop]]'s ants: illustration by [[Milo Winter]], 1888–1956 ]]
[[Anthropomorphism|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.<ref>{{cite book |title=Quran |chapter= The Ant, The Ants | volume = Surah 27 | pages = 18–19 |chapter-url=http://www.wright-house.com/religions/islam/Quran/27-ant.html |url-status=dead |archive-url=https://web.archive.org/web/20070101181115/http://www.wright-house.com/religions/islam/Quran/27-ant.html |archive-date=2007-01-01 }}</ref><ref>{{cite book | first = Sahih | last = Bukhari | name-list-style=vanc | title = Sunnah | volume = 4 Book 54 | issue = 536 | chapter = Beginning of Creation | chapter-url=http://www.usc.edu/dept/MSA/fundamentals/hadithsunnah/bukhari/054.sbt.html|archive-url=https://web.archive.org/web/20000818092520/http://www.usc.edu/dept/MSA/fundamentals/hadithsunnah/bukhari/054.sbt.html|url-status=dead|archive-date=2000-08-18}}</ref> In the [[Book of Proverbs]] in the [[Bible]], ants are held up as a good example of hard work and cooperation.<ref>{{Bibleverse|Proverbs|30:25|web|Proverbs 30:25|nobook=|wlink=wikisource:Bible (World English)/Proverbs#30:25|multi=}}</ref> [[Aesop]] did the same in his fable [[The Ant and the Grasshopper]]. In the [[Quran]], [[Solomon|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.{{CiteQuran Ayah|27|18|s=y|b=yl}},<ref>Mentioned once in the{{Quran ref|a=ASA|597|i=1|h=1}}, [[Muhammad Asad]] translates the verse as following: till, when they came upon a valley [full] of ants, and an ant exclaimed: "O you ants! Get into your dwellings, lest Solomon and his hosts crush you without [even] being aware [of you]! (27:18)"</ref><ref>{{cite book|first=Mawil Y. Izzi|last=Deen | name-list-style=vanc |chapter=Islamic Environmental Ethics, Law, and Society|title=Ethics of Environment and Development| veditors = Engel JR, Engel JG |year=1990|publisher=Bellhaven Press, London|chapter-url=http://www.mbcru.com/Texas%20Tech%20Mypage/Conservation%20Biology/Assignment%202/IzziDeenIslamicEcol.pdf|archive-url=https://web.archive.org/web/20110714055330/http://www.mbcru.com/Texas%20Tech%20Mypage/Conservation%20Biology/Assignment%202/IzziDeenIslamicEcol.pdf|url-status=dead|archive-date=2011-07-14}}</ref> In parts of Africa, ants are considered to be the messengers of the deities. Some [[Native American mythology]], such as the [[Hopi mythology]], considers ants as the first animals. 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 (journal)|Ethnohistory]] |volume=47 |issue=2 |year=2000 |pages=399–422|doi=10.1215/00141801-47-2-399|s2cid=162813070 }}</ref> Ant bites are used in the [[initiation]] ceremonies of some Amazon Indian cultures as a test of endurance.<ref>{{cite book |vauthors=Cesard N, Deturche J, Erikson P |year=2003 |chapter=Les Insectes dans les pratiques médicinales et rituelles d'Amazonie indigène | veditors = Motte-Florac E, Thomas JM |title=Les insectes dans la tradition orale|publisher=Peeters-Selaf, Paris|pages=395–406|language=fr}}</ref><ref>{{cite journal | vauthors = Schmidt RJ | title = The super-nettles. A dermatologist's guide to ants in the plants | journal = International Journal of Dermatology | volume = 24 | issue = 4 | pages = 204–210 | date = May 1985 | pmid = 3891647 | doi = 10.1111/j.1365-4362.1985.tb05760.x | s2cid = 73875767 | url = http://www.botanical-dermatology-database.info/BotDermReviews/Myrmecophytes.html }}</ref> In [[Greek mythology]], the goddess [[Athena]] turned the maiden [[Myrmex (Attic woman)|Myrmex]] into an ant when the latter claimed to have invented the plough, when in fact it was Athena's own invention.<ref>[[Maurus Servius Honoratus|Servius]], ''Commentary on Virgil's Aeneid'' [https://topostext.org/work/548#4.402 4.402]; [[William Smith (lexicographer)|Smith]] 1873, s.v. [https://www.perseus.tufts.edu/hopper/text?doc=Perseus%3Atext%3A1999.04.0104%3Aalphabetic+letter%3DM%3Aentry+group%3D32%3Aentry%3Dmyrmex-bio-1 Myrmex]</ref>

[[File:Multia vaakuna.svg|thumb|upright|An ant pictured in the coat of arms of [[Multia, Finland|Multia]], a town in Finland]]

Ant society has always fascinated humans and has been written about both humorously and seriously. [[Mark Twain]] wrote about ants in his 1880 book ''[[A Tramp Abroad]]''.<ref>{{cite book|chapter=22 The Black Forest and Its Treasures|title=A Tramp Abroad|last=Twain|first=Mark|name-list-style=vanc|year=1880|chapter-url=http://www.gutenberg.org/files/119/|isbn=978-0-19-510137-9|publisher=Oxford University Press|location=New York|access-date=2015-12-13|url-access=registration|url=https://archive.org/details/trampabroad0000twai_z6u7}}</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 ''[[Les Fourmis]]'' science-fiction trilogy is divided between the worlds of ants and humans; ants and their behaviour are described using contemporary scientific knowledge. [[H. G. Wells]] wrote about intelligent ants destroying human settlements in Brazil and threatening human civilization in his 1905 science-fiction short story, ''[[Empire of the Ants|The Empire of the Ants]].'' A similar German story involving army ants, ''[[Leiningen Versus the Ants]]'', was written in 1937 and recreated in movie form as [[The Naked Jungle]] in 1954.<ref>{{Cite journal |last=Winthrop-Young |first=Geoffrey |date=2021 |title=A Green Hell Makes Better Germans: Carl Stephenson's "Leiningen" and the Almost Aryan Countertextual Army Ants |url=https://www.tandfonline.com/doi/full/10.1080/00168890.2021.1977228 |journal=The Germanic Review: Literature, Culture, Theory|volume=96 |issue=4 |pages=339–356 |doi=10.1080/00168890.2021.1977228 |s2cid=245125375 |issn=0016-8890}}</ref> In more recent times, animated cartoons and 3-D animated films featuring ants have been produced including ''[[Antz]]'', ''[[A Bug's Life]]'', ''[[The Ant Bully (film)|The Ant Bully]]'', ''[[The Ant and the Aardvark]]'', ''[[Ferdy the Ant (TV series)|Ferdy the Ant]]'' and ''[[Atom Ant]].'' Renowned [[Myrmecology|myrmecologist]] [[E. O. Wilson]] wrote a short story, "Trailhead" in 2010 for ''[[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.<ref name="Trailhead">{{cite magazine|author=Wilson, EO|url=http://www.newyorker.com/fiction/features/2010/01/25/100125fi_fiction_wilson?currentPage=all |title=Trailhead|magazine=The New Yorker|date=25 January 2010|pages=56–62}}</ref> The French neuroanatomist, psychiatrist and [[Eugenics|eugenicist]] [[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.<ref>{{cite journal | vauthors = Parent A | title = Auguste Forel on ants and neurology | journal = The Canadian Journal of Neurological Sciences | volume = 30 | issue = 3 | pages = 284–291 | date = August 2003 | pmid = 12945958 | doi = 10.1017/s0317167100002754 | doi-access = free }}</ref>

In the early 1990s, the video game ''[[SimAnt]]'', which simulated an ant colony, won the 1992 [[Software and Information Industry Association#CODiE Awards|Codie award]] for "Best Simulation Program".<ref>{{cite web |url=http://www.siia.net/codies/2009/pw_1992.asp |archive-url=https://web.archive.org/web/20090611002722/http://www.siia.net/codies/2009/pw_1992.asp |archive-date=2009-06-11 |title= 1992 Excellence in Software Awards Winners |access-date=3 April 2008|publisher=Software & Information Industry Association}}</ref>

Ants also are quite popular inspiration for many [[science-fiction]] [[Insectoids in science fiction|insectoid]]s, 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 (film)|Empire of the Ants]],'' [[Marvel Comics]]' super hero [[Ant-Man]], and ants mutated into super-intelligence in ''[[Phase IV (1974 film)|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 [[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=255–268|s2cid=321326}}</ref>

== See also ==
{{Main|Outline of ants}}
* [[Glossary of ant terms]]
* [[International Union for the Study of Social Insects]]
* ''[[Myrmecological News]]'' (journal)
* [[Task allocation and partitioning in social insects]]

== References ==
{{Reflist}}

===Cited texts===
* {{cite book|vauthors=Borror DJ, Triplehorn CA, Delong DM |title=Introduction to the Study of Insects|edition=6th|publisher=Saunders College Publishing|year=1989|isbn=978-0-03-025397-3|ref=none}}
* {{cite book |vauthors=Hölldobler B, Wilson EO |title=The Ants |publisher=[[Harvard University Press]]|year=1990|isbn=978-0-674-04075-5|title-link=The Ants |ref=none}}

== Further reading ==
{{refbegin}}
* {{cite book|author=Bolton, Barry|publisher=Harvard University Press|year=1995|isbn=978-0-674-61514-4|title=A New General Catalogue of the Ants of the World|ref=none}}
* {{cite book|vauthors=Hölldobler B, Wilson EO|year=1998|title=Journey to the Ants: A Story of Scientific Exploration|publisher=Belknap Press|isbn=978-0-674-48526-6|url=https://archive.org/details/journeytoants00holl|ref=none}}	
* {{cite book|vauthors=Hölldobler B, Wilson EO|year=2009|title=The Superorganism: The Beauty, Elegance and Strangeness of Insect Societies|publisher=Norton & Co.|isbn=978-0-393-06704-0|url-access=registration|url=https://archive.org/details/superorganismbea0000hlld|ref=none}}
{{refend}}

== External links ==
{{Wikiquote}}
{{Commons category|Formicidae}}
{{Wikispecies|Formicidae}}
* {{cite EB9 |wstitle= Ant |volume= II |last= Wilson |first= Andrew |author-link= Andrew Wilson (zoologist) |pages=94-100 |short=1}}
* [http://www.antweb.org/ AntWeb from The California Academy of Sciences]
* [http://www.antwiki.org/wiki/Welcome_to_AntWiki AntWiki&nbsp;– Bringing Ants to the World]
* [http://www.pestworld.org/pest-guide/ants/ Ant Species Fact Sheets] from the [[National Pest Management Association]] on Argentine, Carpenter, Pharaoh, Odorous, and other ant species
* [http://www.antwiki.org/wiki/Category:Genus_Distribution_Map Ant Genera of the World&nbsp;– distribution maps]
* [http://www.botanical-dermatology-database.info/BotDermReviews/Myrmecophytes.html The super-nettles. A dermatologist's guide to ants-in-the-plants]

{{Hymenoptera|2}}
{{Ant taxonomy}}
{{Eusociality}}
{{Taxonbar|from=Q7386}}
{{Authority control}}

[[Category:Ants| ]]
[[Category:Symbiosis]]
[[Category:Extant Albian first appearances]]
[[Category:Articles containing video clips]]
[[Category:Insects in culture]]