Workers of Leptanilla, photographed by Alex Wild.

Belongs within: Vespoidea.
Contains: Amblyoponinae, Proceratiinae, Ponerinae, Dorylinae, Cerapachys, Ectatommini, Myrmicinae, Formicinae, Dolichoderinae, Myrmeciinae.

The Formicidae are the ants, a diverse and familiar clade of social wasps with a caste division between wingless workers and winged reproductives, with the female reproductives (the 'queens') shedding their wings after mating. An elongate scape (first antennal segment) is present in living ants, but was absent in the Cretaceous Sphecomyrminae (Grimaldi & Engel 2005). The morphology of Sphecomyrma indicates that it was an epigean (above-ground) forager, but two of the most basally diverging living ant lineages, the Leptanillinae and Martialis heureka, are hypogean (below-ground) foragers with reduced pigmentation and eyes.

Other small subfamilies of ants include the Agroecomyrmecinae, little-known ants from Africa and South America with eyes placed far back on the head and antennae spaced widely apart.

Synapomorphies (from Grimaldi & Engel 2005): Eusocial habits, with wingless female caste(s) present; metapleural gland opening on propodeum; petiole present.

<==Formicidae [Poneridae]
    |--Sphecomyrminae BS06
    |    |--Cretomyrma WB03
    |    |--Baikuris WB03
    |    |--Dlusskyidris zherichini WB03, GE05
    |    `--Sphecomyrma Wilson & Brown 1967 V10
    |         |--S. canadensis GE05
    |         `--S. freyi RBV08
    `--+--Martialis Rabeling & Verhaagh in Rabeling, Brown & Verhaagh 2008 [Martialinae] RBV08
       |    `--*M. heureka Rabeling & Verhaagh in Rabeling, Brown & Verhaagh 2008 RBV08
       `--+--Leptanillinae RBV08
          |    |--Protanilla RBV08
          |    `--Leptanilla Emery 1870 FT08
          |         |--L. japonica Baroni Urbani 1977 I92
          |         |--L. kubotai Baroni Urbani 1977 I92
          |         |--L. morimotoi Yasumatsu 1960 I92
          |         |--L. oceanica Baroni Urbani 1977 I92
          |         |--L. swani H09
          |         `--L. tanakai Baroni Urbani 1977 I92
          `--+--+--Tatuidris [Agroecomyrmecinae] RBV08
             |  `--Amblyoponinae RBV08
             `--+--+--Proceratiinae RBV08
                |  `--Paraponera [Paraponerinae, Paraponerini] RBV08
                |       `--P. clavata RBV08
                `--+--Ponerinae RBV08
                   `--+--+--Dorylinae BS06
                      |  `--Cerapachyinae WW90
                      |       |--Cerapachys BS06
                      |       `--+--Leptanilloides BS06
                      |          |    |--L. mckennae BS06
                      |          |    `--L. nomada BS06
                      |          `--+--Cylindromyrmex striatus BS06
                      |             `--Acanthostichus BS06
                      |                  |--A. kirbyi BS06
                      |                  `--A. texanus WW90
                      `--+--Dolichoformica helferi GE05
                         `--+--+--Ectatommini BS06
                            |  `--+--Myrmicinae BS06
                            |     `--Formicinae BS06
                            `--+--+--Dolichoderinae BS06
                               |  `--Aneuretinae BS06
                               |       |--Aneuretellus GE05
                               |       `--Aneuretus Emery 1893 B03
                               |            `--*A. simoni Emery 1893 B03
                               `--+--Myrmeciinae BS06
                                  `--Pseudomyrmecinae BS06
                                       |--Tetraponera BS06
                                       |    |--T. extenuata N-ZLI11
                                       |    |--T. punctulata H09
                                       |    `--T. rufonigra BS06
                                       `--+--Myrcidris epicharis BS06
                                          `--Pseudomyrmex BS06
                                               |--P. ferrugineus FJ99
                                               `--P. gracilis BS06

Formicidae incertae sedis:
  Huberia striata (Smith 1876) H82
  Brownimecia clavata GE05
  Haidomyrmex cerberus GE05
  Labauchena A71
  Strongylognathus E89a
    |--S. afer E89a
    |--S. christophi Emery 1889 E89a
    |--S. huberi E89a
    |--S. koreanus Pisarski 1965 I92
    `--S. testaceus A71
  Anergates atratulus A71
  Eoponera berryi GE05
  Gerontoformica cretacica GE05
  Canapone dentata GE05
  Protopone GE05
  Cariridris bipetiolata GE05
  Myrmecocistus albicans G89
  Oxyopomyrmex E89a
    |--O. oculatus E89a
    `--O. saulcyi Emery 1889 E89a
  Harpegnathus rugosus E89b
  Trapeziopelta maligna E89b
  Lordomyrmex caledonica Andre 1889 E14
  Sphecomyrmodes occidentalis Perrichot, Nel et al. 2007 V10

*Type species of generic name indicated


[A71] Askew, R. R. 1971. Parasitic Insects. Heinemann Educational Books: London.

[B03] Bingham, C. T. 1903. The Fauna of British India, including Ceylon and Burma. Hymenoptera. Vol. II. Ants and Cuckoo-Wasps. Taylor and Francis: London.

[BS06] Brady, S. G., T. R. Schultz, B. L. Fisher & P. S. Ward. 2006. Evaluating alternative hypotheses for the early evolution and diversification of ants. Proceedings of the National Academy of Sciences of the USA 103 (48): 18172–18177.

[E89a] Emery, C. 1889a. Intorno ad alcune formiche della fauna palearctica. Annali del Museo Civico di Storia Naturale di Genova, Serie 2a 7: 439–443.

[E89b] Emery, C. 1889b. Viaggio di Leonardo Fea in Birmania e regioni vicine. XX.—Formiche di Birmania e del Tenasserim raccolte da Leonardo Fea (1885–87). Annali del Museo Civico di Storia Naturale di Genova, Serie 2a 7: 485–520.

[E14] Emery, C. 1914. Les fourmis de la Nouvelle-Calédonie & des îles Loyalty. In: Sarasin, F. & J. Roux (eds) Nova Caledonia: Forschungen in Neu-Caledonian und auf den Loyalty-Inseln. A. Zoologie vol. 1, Heft IV pp. 393–436, pl. 12. C. W. Kreidels Verlag: Wiesbaden.

[FJ99] Fiala, B., A. Jakob, U. Maschwitz & K. E. Linsenmair. 1999. Diversity, evolutionary specialization and geographic distribution of a mutualistic ant-plant complex: Macaranga and Crematogaster in South East Asia. Biological Journal of the Linnean Society 66: 305–331.

[FT08] Framenau, V. W., & M. L. Thomas. 2008. Ants (Hymenoptera: Formicidae) of Christmas Island (Indian Ocean): identification and distribution. Records of the Western Australian Museum 25 (1): 45–85.

[G89] Gestro, R. 1889. Viaggio ab Assab nel Mar Rosso dei signori G. Doria ed O. Beccari con il R. Avviso «Esploratore» dal 16 Novembre 1879 al 26 Febbraio 1880.—IV. Coleotteri. Annali del Museo Civico di Storia Naturale di Genova, Serie 2a 7: 5–72.

[GE05] Grimaldi, D., & M. S. Engel. 2005. Evolution of the Insects. Cambridge University Press: New York.

[H82] Helmore, D. W. 1982. Drawings of New Zealand insects. Bulletin of the Entomological Society of New Zealand 8: 1–52.

[H09] Heterick, B. E. 2009. A guide to the ants of south-western Australia. Records of the Western Australian Museum Supplement 76: 1–206.

[I92] Iwahashi, J. (ed.) 1992. Reddo Deeta Animaruzu: a pictorial of Japanese fauna facing extinction. JICC: Tokyo.

[N-ZLI11] Nur-Zati A. M., G. T. Lim & A. B. Idris. 2011. Checklist of ants of selected hill dipterocarp forests of peninsular Malaysia. Serangga 16 (1): 91–105.

[RBV08] Rabeling, C., J. M. Brown & M. Verhaagh. 2008. Newly discovered sister lineage sheds light on early ant evolution. Proceedings of the National Academy of Sciences of the USA 105 (39): 14913–14917.

[V10] Vršanský, P. 2010. Cockroach as the earliest eusocial animal. Acta Geologica Sinica (English Edition) 84 (4): 793–808.

[WB03] Ward, P. S., & S. G. Brady. 2003. Phylogeny and biogeography of the ant subfamily Myrmeciinae (Hymenoptera: Formicidae). Invertebrate Systematics 17: 361–386.

[WW90] Wheeler, G. C., & J. Wheeler. 1990. Insecta: Hymenoptera Formicidae. In: Dindal, D. L. (ed.) Soil Biology Guide pp. 1277–1294. John Wiley & Sones: New York.

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