List of ant subfamilies

Clades

stem ants †Armaniinae †Sphecomyrminae poneroids Agroecomyrmecinae Amblyoponinae Apomyrminae †Brownimeciinae Paraponerinae Ponerinae Proceratiinae formicoids dorylomorphs Dorylinae myrmeciomorphs Myrmeciinae Pseudomyrmecinae dolichoderomorphs Aneuretinae Dolichoderinae ectaheteromorphs Ectatomminae Heteroponerinae †Formiciinae Formicinae Myrmicinae Leptanillinae Martialinae

Extinct taxa are indicated by a †

Source: Ward (2007), Kück et al. (2011), Brady et al. (2014)

Ants (family Formicidae in the order Hymenoptera) are the most species-rich of all social insects, with more than 12,000 described species and many others awaiting description.^[1] Formicidae is divided into 21 subfamilies, of which 17 contain extant taxa, while four are exclusively fossil.^[2] Ants have come to occupy virtually all major terrestrial habitats, with the exception of tundra and cold ever-wet forests. They display a wide range of social behaviors, foraging habits and associations with other organisms, which has generated scientific and public interest.^[1]

Clades

Beginning in the 1990s, molecular (DNA sequence) data have come to play a central role in attempts to reconstruct the ant "tree of life". Molecular phylogenetic analyses based on multiple nuclear genes have yielded robust results that reinforce some preexisting views but overturn others – and suggest that there has been considerable morphological convergence among some ant lineages. Molecular data provide very strong support for a novel group, the "formicoid clade", not revealed by previous morphological work. This clade comprises 9^[3] of the 16 extant ant subfamilies and about 90% of all described ant species. Formicoids include such widespread and species-rich subfamilies as Myrmicinae, Formicinae and Dolichoderinae, as well as the army ants (Dorylinae). Non-formicoids comprise five "poneroid" subfamilies (Agroecomyrmecinae, Amblyoponinae, Paraponerinae, Ponerinae, and Proceratiinae), Leptanillinae, about which little is known, and Martialinae, the most recently discovered subfamily.^[1][4] Relationships among these remaining seven subfamilies are less well resolved.^[1] A recent study (2011) places Leptanillinae as a sister group to all other ants, with Martialinae, the poneroids and formicoids forming a clade.^[4]

Evolution of ants

Ants first arose during the mid-Cretaceous, more than 100 million years ago, associated with the rise of flowering plants and an increase in forest ground litter.^[5] The earliest known ants evolved from a lineage within the aculeate wasps, and a recent study suggests that they are a sister group of Apoidea.^[6] During the Cretaceous ants were confined to the northern Laurasian supercontinent, with only a few widespread primitive species.^[7] By the middle Eocene, around 50 million years ago, ants had diversified and become ecologically dominant as predators and scavengers. Ant species are less than 2% of the total number of insect species but make up one third of the insect biomass.^[5][8]

History of classification

Formica rufa, one of the seventeen species of ants described by Linnaeus (1758) in his Systema Naturae^[9]

In volume 1 of Systema Naturae, Carl Linnaeus (1758) described seventeen species of ants, all of which he placed in the single genus Formica.^[9] Within a few decades additional genera had been recognized, and this trend continued in the ensuing years, together with the development of a more complex hierarchical classification in which genera were apportioned among subfamilies and tribes. The ant species described by Linnaeus are now dispersed in eleven different genera, belonging to four subfamilies.^[1]

For much of the twentieth century the number of recognized ant subfamilies varied from seven to ten, with the Aneuretinae, Cerapachyinae, Leptanillinae, Myrmeciinae and Pseudomyrmecinae being variously treated as separate subfamilies or (at different times) subsumed within Dolichoderinae, Ponerinae, Dorylinae, Ponerinae, and Myrmicinae, respectively.^[1] In 2014, Brady et al. synonymized the army ant subfamilies and their closest relatives under Dorylinae; this clade, the dorylomorph subfamilies, previously also contained Aenictinae, Aenictogitoninae, Cerapachyinae, Ecitoninae and Leptanilloidinae.^[3]

The last three decades have seen a proliferation of subfamily names, as a result of three factors: (1) the realization that some subfamilies were assemblages of unrelated taxa; (2) abandonment of paraphyletic taxa, and (3) the discovery of novel fossil taxa.^[1] Seventeen extant subfamilies of ants are currently recognized, along with four extinct subfamilies.^[2] One of the fossil taxa, Armaniinae, is often given family rank within the superfamily Formicoidea.^[1][10] About 13 genera are incertae sedis (of uncertain placement), and are not assigned to any subfamily.^[2]

Subfamilies

Extinct taxa are indicated by a †.

Formicidae
Subfamily[2]	Extant genera[2]	Fossil genera[2]	Type genus[2]	Comments	Example
Agroecomyrmecinae Carpenter, 1930	2	2	†Agroecomyrmex Wheeler, 1910	Agroecomyrmecines were probably widespread in both hemispheres during the early Tertiary, but are today represented by only two species in two genera, Tatuidris and Ankylomyrma. The two fossil genera, Agroecomyrmex and Eulithomyrmex, are known from Eocene fossils.[11][12]	Tatuidris tatusia
Amblyoponinae Forel, 1893	13	1	Amblyopone Erichson, 1842	Specialized predators found worldwide, but mainly distributed in the tropics.[13][14]	Apomyrma stygia
Aneuretinae Emery, 1913	1	8	Aneuretus Emery, 1893	In addition to 8 fossil genera, the subfamily is represented by one extant genus, Aneuretus. The living genus has a single species, the Sri Lankan relict ant (Aneuretus simoni), endemic to Sri Lanka and only found in a few locations. The species is listed as critically endangered by the IUCN.[15][16]	Aneuretus simoni
Apomyrminae Dlussky & Fedoseeva, 1988	1	1	Apomyrma Brown, Gotwald & Lévieux, 1970	Contains a single described species, the rare subterranean ant Apomyrma stygia, and several undescribed species, all known from the Afrotropics.[17]	Apomyrma stygia
†Armaniinae Dlussky, 1983	0	8	†Armania Dlussky, 1983	Known from Cretaceous fossils, this subfamily is sometimes treated as the family Armaniidae within the superfamily Formicoidea. They seem to lack a metapleural gland, which would exclude them from Formicidae given an apomorphy-based definition requiring such a gland to have evolved. However, the apparent lack of this gland in Armaniinae fossils could be due to poorer preservation.[1][18]	Orapia rayneri
†Brownimeciinae Bolton, 2003	0	1	†Brownimecia Grimaldi, Agosti & Carpenter, 1997	Contains the single species Brownimecia clavata, a fossil ant known from New Jersey amber. Initially placed within Ponerinae, this primitive Cretaceous ant was later moved to its own subfamily.[1][18]	Brownimecia clavata
Dolichoderinae Forel, 1878	28	20	Dolichoderus Lund, 1831	A diverse subfamily distributed worldwide, mainly in the tropics. Most species are generalized scavengers, but some are predacious. They lack a sting and instead rely on chemical defensive compounds produced from the anal gland.[19][20]	Dolichoderus mariae
Dorylinae Leach, 1815	28	1[a][b]	Dorylus Fabricius, 1793	Contains the previous dorylomorph subfamilies (Aenictinae, Aenictogitoninae, Cerapachyinae, Ecitoninae and Leptanilloidinae), including the Old World and New World army ants.[3]	Dorylus helvolus
Ectatomminae Emery, 1895	4	3	Ectatomma Smith, 1858	Found in tropical and warm climates in the New World, Old World and Indo-Australian regions.[20]	Ectatomma ruidum
†Formiciinae Lutz, 1986	0	1[c]	†Formicium[d] Westwood, 1854	Known from Eocene fossils found in Europe and North America, this subfamily contains the genus Titanomyrma and the collective group name Formicium.[24]	Titanomyrma giganteum
Formicinae Latreille, 1809	51	30	Formica Linnaeus, 1758	Worldwide in distribution, Formicinae is the second largest subfamily; only Myrmicinae is larger.[20]	Formica rufa
Heteroponerinae Bolton, 2003	3	0	Heteroponera Mayr, 1887	Contains the two Neotropical genera, Acanthoponera and Heteroponera, and the genus Aulacopone, which is known from the single species, Aulacopone relicta, from Azerbaijan.[25]	Heteroponera brounii
Leptanillinae Emery, 1910	8	0	Leptanilla Emery, 1870	Small subterranean ants known from Africa, Europe, Asia and a single species from Australia; no species are known from North or South America. Little is known about the biology of these ants.[26][27]	Leptanilla swani
Martialinae Rabeling & Verhaagh, 2008	1	0	Martialis Rabeling & Verhaagh, 2008	The subfamily is only known from a single specimen of the species Martialis heureka, collected in 2003 in the Amazon rainforest. Workers are pale and blind; queens and males are unknown.[28]	Martialis heureka
Myrmeciinae Emery, 1877	2	5[e]	Myrmecia Fabricius, 1804	A formerly widespread subfamily, now only found in the Australasian region distributed in two extant genera. The genus Myrmecia, or "bulldog ants", is known from Australia, New Zealand and New Caledonia, and Nothomyrmecia, with the single species Nothomyrmecia macrops, is known from Australia.[30]	Myrmecia gulosa
Myrmicinae Lepeletier de Saint-Fargeau, 1835	140	35	Myrmica Latreille, 1804	The largest subfamily of ants, distributed worldwide. Contains the seed-harvesting ants and fungus-growing ants.[20]	Myrmica lonae
Paraponerinae Emery, 1901	1	0	Paraponera Smith, 1858	Contains the single genus Paraponera consisting of two species: the extant Paraponera clavata, also known as a bullet ant, found in the Neotropics, and the very small fossil species Paraponera dieteri known from Dominican amber (Early Miocene).[31][32]	Paraponera clavata
Ponerinae Lepeletier de Saint-Fargeau, 1835	47	12	Ponera Latreille, 1804	Predominantly predacious ants distributed in the tropics and subtropics. The subfamily formerly contained all the poneromorph subfamilies (at the time classified as tribes), which have now been divided into six subfamilies: Amblyoponinae, Ectatomminae, Heteroponerinae, Paraponerinae, Ponerinae and Proceratiinae.[20][33]	Ponera coarctata
Proceratiinae Emery, 1895	3	1	Proceratium Roger, 1863	Ants in this subfamily are small to medium in size. Found worldwide, although mainly tropical and subtropical in distribution. Little is known about their biology, but they are thought to be specialized predators.[20][34]	Proceratium google
Pseudomyrmecinae Smith, 1952	3	0	Pseudomyrmex Lund, 1831	A small subfamily consisting of three genera of predominantly arboreal ants found in tropical and subtropical regions. The ants, agile and with large eyes, are characterized by their slender bodies.[35]	Pseudomyrmex pallidus
†Sphecomyrminae Wilson & Brown, 1967	0	9	†Sphecomyrma Wilson & Brown, 1967	Fossil subfamily of stem ants. Most Cretaceous ants are placed in this subfamily.[1][36]	Sphecomyrma freyi

See also

• List of ant genera
• Poneromorph subfamilies

Notes

1. The fossil genus †Curticorna was placed in Dorylinae by Hong (2002), but this genus is "definitely not Dorylinae" according to Barry Bolton.^[21] Dlussky & Wedmann (2012) suggest that †Curticorna may belong to Ectatomminae.^[13] It is listed as incertae sedis within Formicidae in this list.^[21]
2. The fossil genus †Eoaenictites was placed in Aenictinae (now Dorylinae) by Hong (2002), but this genus is "definitely not Aenictinae" according to Barry Bolton. It is listed as incertae sedis within Formicidae in this list.^[22]
3. The subfamily also contains the collective group name †Formicium.^[23]
4. The type genus of the subfamily, †Formicium, was previously treated as a nominal genus, but was later reclassified as a collective group name by Archibald et al. (2011). However, according to the International Code of Zoological Nomenclature (ICZN Code), the genus remains the type genus of Formiciinae.^[24]
5. The subfamily also contains the collective group name †Myrmeciites.^[29]

References

1. Ward, Philip S. (2007), "Phylogeny, classification, and species-level taxonomy of ants (Hymenoptera: Formicidae)." (PDF), Zootaxa, 1668: 549–563, doi:10.11646/zootaxa.1668.1.26
2. Bolton, B. (2015). "Formicidae". AntCat. Retrieved 30 August 2015.
3. Brady, Seán G; Fisher, Brian L; Schultz, Ted R; Ward, Philip S (2014). "The rise of army ants and their relatives: diversification of specialized predatory doryline ants". BMC Evolutionary Biology. 14 (1): 2–14. Bibcode:2014BMCEE..14...93B. doi:10.1186/1471-2148-14-93. PMC 4021219. PMID 24886136.
4. Kück, P.; Hita Garcia, F.; Misof, B.; Meusemann, K. (2011). "Improved Phylogenetic Analyses Corroborate a Plausible Position of Martialis heureka in the Ant Tree of Life". PLOS ONE. 6 (6): e21031. Bibcode:2011PLoSO...621031K. doi:10.1371/journal.pone.0021031. PMC 3123331. PMID 21731644.
5. Wilson, E. O.; Hölldobler, B. (2005). "The rise of the ants: A phylogenetic and ecological explanation". Proceedings of the National Academy of Sciences. 102 (21): 7411–7414. Bibcode:2005PNAS..102.7411W. doi:10.1073/pnas.0502264102. PMC 1140440. PMID 15899976.
6. Johnson, B.R.; Borowiec, M.L.; Chiu, J.C.; Lee, E.K.; Atallah, J.; Ward, P.S. (2013). "Phylogenomics resolves evolutionary relationships among ants, bees, and wasps". Current Biology. 23 (20): 2058–2062. Bibcode:2013CBio...23.2058J. doi:10.1016/j.cub.2013.08.050. PMID 24094856.
7. Hölldobler, Bert; Wilson, Edward O. (1990). The Ants. Cambridge, Massachusetts: Belknap Press of Harvard University Press. pp. 23–24. ISBN 978-0-674-04075-5.
8. Grimaldi, D.; Agosti, D. (2000). "A formicine in New Jersey Cretaceous amber (Hymenoptera: Formicidae) and early evolution of the ants". Proceedings of the National Academy of Sciences. 97 (25): 13678–13683. Bibcode:2000PNAS...9713678G. doi:10.1073/pnas.240452097. PMC 17635. PMID 11078527.
9. Linnaeus, C. (1758). Systema naturae per regna tria naturae, secundum classes, ordines, genera, species, cum characteribus, differentiis, synonymis, locis. Tomus I (10th ed.). Holmiae. (Laurentii Salvii). pp. 579–582.
10. Engel, M. S.; Grimaldi, David A. (2005). "Primitive New Ants in Cretaceous Amber from Myanmar, New Jersey, and Canada (Hymenoptera: Formicidae)" (PDF). American Museum Novitates (3485): 1–23. doi:10.1206/0003-0082(2005)485[0001:pnaica]2.0.co;2. S2CID 84043939.
11. Brown, W. L. Jr.; Kempf, W. W. (1968), "Tatuidris, a remarkable new genus of Formicidae (Hymenoptera).", Psyche: A Journal of Entomology, 74 (3) (published 1967): 183–190, doi:10.1155/1967/868759
12. Donoso, D.A. (2012), "Additions to the taxonomy of the armadillo ants (Hymenoptera, Formicidae, Tatuidris)." (PDF), Zootaxa, 3503: 61–81, doi:10.11646/zootaxa.3503.1.5
13. Dlussky, G. M.; Wedmann, S. (2012). "The poneromorph ants (Hymenoptera, Formicidae: Amblyoponinae, Ectatomminae, Ponerinae) of Grube Messel, Germany: High biodiversity in the Eocene". Journal of Systematic Palaeontology. 10 (4): 725–753. Bibcode:2012JSPal..10..725D. doi:10.1080/14772019.2011.628341. S2CID 83928415.
14. Saux, Corrie; Fisher, Brian L.; Spicer, Greg S. (2004). "Dracula ant phylogeny as inferred by nuclear 28S rDNA sequences and implications for ant systematics (Hymenoptera: Formicidae: Amblyoponinae)". Molecular Phylogenetics and Evolution. 33 (2): 457–468. doi:10.1016/j.ympev.2004.06.017. PMID 15336679.
15. Karunarathna, D. A. G. N. B.; Karunaratne, W. A. I. P. (2013). "Two new localities of Sri Lankan Relict Ant Aneuretus simoni Emery, 1893 (Formicidae: Aneuretinae) with the very first record in the intermediate zone". Journal of Threatened Taxa. 5 (11): 4604–4607. doi:10.11609/JoTT.o3334.4604-7.
16. Social Insects Specialist Group (1996). "Aneuretus simoni". IUCN Red List of Threatened Species. 1996. Retrieved 30 August 2015.{{cite iucn}}: old-form url (help)
17. Ward, P. S. (2014). "The phylogeny and evolution of ants" (PDF). Annual Review of Ecology, Evolution, and Systematics. 45: 23–43. doi:10.1146/annurev-ecolsys-120213-091824. S2CID 56577465.
18. Grimaldi, D.; Agosti, D.; Carpenter, J.M. (1997), "New and rediscovered primitive ants (Hymenoptera: Formicidae) in Cretaceous amber from New Jersey, and their phylogenetic relationships.", American Museum Novitates (3208): 1–43
19. Cardoso, D.; Cristiano, M. P.; Barros, L. S.; Lopes, D.; Pompolo, S. (2012). "First cytogenetic characterization of a species of the arboreal ant genus Azteca Forel, 1978 (Dolichoderinae, Formicidae)". Comparative Cytogenetics. 6 (2): 107–114. doi:10.3897/CompCytogen.v6i2.2397. PMC 3833797. PMID 24260655.
20. Fisher, Brian L.; Cover, Stefan P. (2007). Ants of North America: A Guide to the Genera. University of California Press. pp. 54–58. ISBN 978-0-520-93455-9.
21. Bolton, B. (2015). "†Curticorna". AntCat. Retrieved 30 August 2015.
22. Bolton, B. (2015). "†Eoaenictites". AntCat. Retrieved 30 August 2015.
23. Bolton, B. (2015). "†Formicium". AntCat. Retrieved 30 August 2015.
24. Archibald, S. B.; Johnson, K. R.; Mathewes, R. W.; Greenwood, D. R. (2011). "Intercontinental dispersal of giant thermophilic ants across the Arctic during early Eocene hyperthermals". Proceedings of the Royal Society B. 278 (1725): 3679–3686. doi:10.1098/rspb.2011.0729. PMC 3203508. PMID 21543354.
25. Taylor, Robert. W. (2011). "Australasian ants of the subfamily Heteroponerinae (Hymenoptera: Formicidae): (1) General introduction and review of the Heteroponera leae (Wheeler, 1923) species group, with descriptions of two new species". Myrmecological News. 15: 117–123.
26. Borowiec, Marek L.; Schultz, Andreas; Alpert, Gary D.; Baňař, Petr (2011). "Discovery of the worker caste and descriptions of two new species of Anomalomyrma (Hymenoptera: Formicidae: Leptanillinae) with unique abdominal morphology". Zootaxa. 2810: 1–14. doi:10.11646/zootaxa.2810.1.1.
27. Shattuck, S. (2000). Australian Ants: Their Biology and Identification. Csiro Publishing. p. 148. ISBN 978-0-643-06659-5.
28. Rabeling, Christian; Brown, Jeremy M.; Verhaagh, Manfred (2008). "Newly discovered sister lineage sheds light on early ant evolution". Proceedings of the National Academy of Sciences of the United States of America. 105 (39): 14913–14917. Bibcode:2008PNAS..10514913R. doi:10.1073/pnas.0806187105. PMC 2567467. PMID 18794530.
29. Bolton, B. (2015). "†Myrmeciites". AntCat. Retrieved 30 August 2015.
30. Ward, Philip S.; Brady, Seán G. (2003). "Phylogeny and biogeography of the ant subfamily Myrmeciinae (Hymenoptera: Formicidae)". Invertebrate Systematics. 17 (3): 361–386. doi:10.1071/IS02046.
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32. Baroni Urbani, C. (1994). "The identity of the Dominican Paraponera (Amber Collection Stuttgart: Hymenoptera, Formicidae. V: Ponerinae, partim)". Stuttgarter Beiträge zur Naturkunde. Serie B (Geologie und Paläontologie): 1–9.
33. Bolton, B. (2003). "Synopsis and classification of Formicidae". Memoirs of the American Entomological Institute. 71: 370. ISBN 1-887988-15-7.
34. Sosa-Calvo, Jeffrey (2007). Studies in Neotropical Ant Diversity. pp. 84–109. ISBN 978-0-549-26277-0.
35. Ward, P. S. (1990). "The ant subfamily Pseudomyrmecinae (Hymenoptera: Formicidae): Generic revision and relationship to other formicids". Systematic Entomology. 15 (4): 449–489. Bibcode:1990SysEn..15..449W. doi:10.1111/j.1365-3113.1990.tb00077.x. S2CID 86012514.
36. Barden, P.; Grimaldi, D. (2013). "A New Genus of Highly Specialized Ants in Cretaceous Burmese Amber (Hymenoptera: Formicidae)" (PDF). Zootaxa. 3681 (4): 405–412. doi:10.11646/zootaxa.3681.4.5. PMID 25232618.

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