Workers of Leptanilla, photographed by Alex Wild.

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

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. Leptanillinae are found in the tropics of the Old World and are characterised by a two-segmented petiole and twelve-segmented antennae without frontal lobes concealing the base.

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. The shape of the gaster in Agroecomyrmecinae is also distinctive, with abdominal sternite 4 reduced while the corresponding tergite is much larger and vaulted.

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

<==Formicidae [Formicariae, Formiciformes, Formicoidea, Paleosminthuridae, Poneridae]
    |--Sphecomyrminae [Sphecomyrmidae] BS06
    |    |--Cretomyrma WB03
    |    |--Baikuris mandibularis Dlussky 1987 RJ93
    |    |--Dlusskyidris zherichini WB03, GE05
    |    |--Paleomyrmex Dlussky 1975 H02
    |    |    `--P. zherichini H02
    |    `--Sphecomyrma Wilson & Brown 1967 RBV08, 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. revelierii Emery 1870 TB85
          |         |--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 Wheeler 1932 TB85
          |         `--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 [Cerapachyini, Cerapachysii, Prodorylinae] 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
                            `--+--+--Myrmeciinae BS06
                               |  `--Pseudomyrmecinae BS06
                               `--+--Dolichoderinae BS06
                                  `--Aneuretinae BS06
                                       |--Aneuretellus GE05
                                       `--Aneuretus Emery 1893 B03
                                            `--*A. simoni Emery 1893 B03

Formicidae incertae sedis:
  Huberia striata (Smith 1876) H82
  Brownimecia clavata GE05
  Haidomyrmex cerberus GE05
  Strongylognathus E89a
    |--S. afer E89a
    |--S. caeciliae ES12
    |--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
  Oxyopomyrmex E89a
    |--O. oculatus E89a
    `--O. saulcyi Emery 1889 E89a
  Trapeziopelta maligna E89b
  Lordomyrmex caledonica Andre 1889 E14
  Sphecomyrmodes occidentalis Perrichot, Nel et al. 2007 V10
  Labidus praedator B10
  Eusphinctus Emery 1893 TB85
  Chalepoxenus muellerianus ES12
  Cretacoformica explicata Jell & Duncan 1986 Z01
  Formicium giganteum S12
  Eomyrmex Hong 1974 H02
    `--E. guchengziensis Z01
  Aeromyrma sophiae Emery 1890 H02

*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. 2. 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.

[B10] Brown, B. V. 2010. Phoridae (hump-backed flies, scuttle flies). In: Brown, B. V., A. Borkent, J. M. Cumming, D. M. Wood, N. E. Woodley & M. A. Zumbado (eds) Manual of Central American Diptera vol. 2 pp. 725–761. NRC Research Press: Ottawa.

[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 pt 4 pp. 393–436, pl. 12. C. W. Kreidels Verlag: Wiesbaden.

[ES12] Espadaler, X., & S. Santamaria. 2012. Ecto- and endoparasitic fungi on ants from the Holarctic region. Psyche 2012: 168478.

[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.

[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.

[H02] Hong Y. 2002. Amber Insect of China. Beijing Scientific and Technological Publishing House.

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

[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.

[RJ93] Ross, A. J., & E. A. Jarzembowski. 1993. Arthropoda (Hexapoda; Insecta). In: Benton, M. J. (ed.) The Fossil Record 2 pp. 363–426. Chapman & Hall: London.

[S12] Schaal, S. 2012. Messel Pit fossil site—the legacy of the environment and life of the Eocene. In: Talent, J. A. (ed.) Earth and Life: Global biodiversity, extinction intervals and biogeographic perturbations through time pp. 225–236. Springer.

[TB85] Taylor, R. W., & D. R. Brown. 1985. Formicoidea. In: Walton, D. W. (ed.) Zoological Catalogue of Australia vol. 2. Hymenoptera: Formicoidea, Vespoidea and Sphecoidea pp. 1–149. Australian Government Publishing Service: Canberra.

[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.

[Z01] Zhou, S. 2001. Ants of Guangxi. Guangxi Normal University Press: Guilin (China).

Last updated: 14 June 2020.

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