Editing Ant (section)

==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 113-100 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><ref>{{Cite journal |last=Lepeco |first=Anderson |last2=Meira |first2=Odair M. |last3=Matielo |first3=Diego M. |last4=Brandão |first4=Carlos R.F. |last5=Camacho |first5=Gabriela P. |date=April 2025 |title=A hell ant from the Lower Cretaceous of Brazil |url=https://linkinghub.elsevier.com/retrieve/pii/S0960982225003082 |journal=Current Biology |language=en |doi=10.1016/j.cub.2025.03.023}}</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>