Formicidae

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Below is a recent diagnosis of the Family, which is based on Boudinot (2015). For those of you that are not taxonomists or have knowledge about animal classification and diversity - the following is the taxonomic answer to the question - What is an ant?

Remarks
The Formicidae is an unequivocally monophyletic group, previously defined by Bolton (1994, 2003) as eusocial, sexually dimorphic aculeate Hymenoptera bearing metapleural glands and geniculate antennae, among other characters. Several previously unreported synapomorphies exist for the family, including a suite of adaptations for terrestrial locomotion (characters 6 and 7). The “low and lateral” propodeal spiracle placement may also be an adaptation for terrestrial locomotion, as it may reduce the distance oxygen would need to diffuse to leg locomotor muscles. While this does not clarify whether the ancestral ant was hypogaeic or epigaeic, it does indicate that terrestrial locomotion was a crucial transition for the Formicidae, as these apomorphies are present in all adult castes of the family. Previous diagnoses of the family (Brothers 1975; Gauld & Bolton 1988; Goulet & Huber 1993) were significantly improved by Bolton (1994, 2003). Characters indicated in the family diagnosis by Bolton (2003) and above will be valuable to evaluate for critical fossil taxa such as †Armania Dlussky and other fossils assigned to the †Armaniidae whose relationship to the Formicidae is uncertain (see Dlussky 1975:†Archaeopone, †Dolichomyrma, †Poneropterus, †Pseudarmania; Dlussky 1983: †Armaniella; Dlussky 1999: †Khetania; Dlussky et al. 2004: †Orapia; also see discussion in LaPolla et al. 2013).

Diagnosis
Aculeate Hymenoptera with the following apomorphies:

1. Eusocial, wingless worker caste present, colonies perennial (note 1).

2. Sexuals with synchronous nuptial flights (note 2)

3. Head capsule prognathous (worker, gyne) (note 1).

4. Infrabuccal sac present between labium and hypopharynx (note 1).

5. Antenna geniculate between long scape and funiculus (worker, gyne) (notes 1, 3).

6. Disticoxal foramen directed laterally and completely enclosing protrochanteral base, including protrochanteral condyles, such that all disticoxal membrane concealed (all castes, Fig. 3C) (note 4).

7. All meso- and metacoxal cavities small, circular, monocondylic, ventrally-directed, and disticoxae strongly produced laterally (all adult castes, Fig. 3C) (note 5).

8. Metapleural gland present (adult castes, but see note 6).

9. Propodeal spiracle located on lateral propodeal face distant from the anterodorsal propodeal corner, often near propodeum midlength (all adult castes) (note 7).

10. Wings of alate gyne deciduous, being shed after copulation (note 1).

11. Forewing 3rs-m and 2m-cu absent (note 1).

12. Hindwing C not extending along anterior margin, even spectrally (note 8).

13. Hindwing basal/radial cell not produced distally (alate castes) (note 9).

14. Metasoma petiolate (abdominal segment II differentiated from segment II,I which is strongly constricted between the pre- and postsclerites) (all castes), extremely rarely (~ 1 species) abdominal segment III not constricted between pre- and postsclerites (notes 1 and 10).

Additional, non-synapomorphic characters of value for diagnosis and identification include: Antenna with 4–12 antennomeres (female) or 5–13 antennomeres (male) (note 11). Bulbus neck (= radicle) and scape with common axis. Epicnemium extremely reduced, not visible in situ (note 12). Abdominal segment II with sternum and tergum equally sclerotized. Pterostigma present or absent (note 13). Wing venation variable, may be extremely reduced, with at minimum no closed cells (note 14). Jugal lobe present or absent; abdominal sternum IX may be complex and modified apically (including prongs, teeth, and lobes).

Nomenclature

 * [family of Hymenoptera, Aculeata]
 * Formicariae Latreille, 1809: 124. Type-genus: Formica Linnaeus, 1758: 579.

Taxonomic History
Latreille, 1802a: 352 [Formicariae, family-group name]; Latreille, 1809: 124 [Formicariae]; Latreille, 1810: 285; Leach, 1815: 147 [Formicarides]; Stephens, 1829: 356 [first spelling as Formicidae]; Haliday, 1836: 331; Zetterstedt, 1838: 447 [Formicariae]; Westwood, 1839: 217; Swainson & Shuckard, 1840: 171; Nylander, 1846a: 877; Foerster, 1850a: 1 [Formicariae]; Mayr, 1855: 275 [Formicina]; Smith, F. 1857a: 52; Smith, F. 1858b: 1; Mayr, 1861: 21; Meinert, 1861: 305 [Formicae]; Mayr, 1865: 6; Heer, 1867: 6 [Formicaria]; Forel, 1870: 307 [Formicinae]; Forel, 1874: 19, 95 [Formicariae]; Emery, 1877a: 70 [Formicidae]; Dalla Torre, 1893: 1; Forel, 1899c: 1; Ruzsky, 1902d: 5 [Formicarii]; Bingham, 1903: 1; Ruzsky, 1905b: 91 [Formicariae or Formicidae]; Ashmead, 1905b: 384; subsequent authors.

Subfamilies of Formicidae
Agroecomyrmecinae, Amblyoponinae, Aneuretinae, †Brownimeciinae, Dolichoderinae, Dorylinae, Ectatomminae, †Formiciinae, Formicinae, Heteroponerinae, Leptanillinae, Martialinae, Myrmeciinae, Myrmicinae, Paraponerinae, Ponerinae, Proceratiinae, Pseudomyrmecinae, †Sphecomyrminae.

Family-group names unavailable in Formicidae
Alloformicinae, Eucamponotinae, Eudolichoderinae, Eudorylinae, Euformicinae, Eumyrmicinae, Exeuponerinae, Heteroformicinae, Mesocamponotinae, Metadorylinae, Mycetomyrminae, Neoattini, Paleoattini, Paleoponerinae, Procamponotinae, Prodolichoderinae, Prodorylinae, Promyrmicinae, Proponerinae, Pseudoponerinae, Rhagiomyrmicinae, Taraxoponerinae.

Genus-group names incertae sedis in Formicidae
†Archaeopone, †Baikuris, †Calyptites, †Camelomecia, Condylodon, †Dlusskyidris, †Eoformica, Hypochira, †Klondikia, †Kohlsimyrma, †Myanmyrma, †Poneropterus.

Genus-group names unavailable in Formicidae
†Ectatona, Pseudacantholepis, Sphaeromessor, Sulcomyrmex.

Genus-group names nomina nuda in Formicidae
Ancylognathus, †Dolichoformica, Hypopheidole, Leptoxenus, Myrmegis, Pergandea, Salticomorpha, Savignyi, Titusia.

Genus-group names excluded from Formicidae
†Cariridris, †Cretacoformica, Formila, †Myrmicium, †Palaeomyrmex, †Promyrmicium, †Syntaphus.

Introductory references
Hölldobler & Wilson, 1990: 1 (introduction to myrmecology); Bolton, 1995a: 1038 (census of extant taxa); Ward, et al. 1996: 55 (bibliography of systematics); Brown, 2000: 46 (overview of genera); Brandão, 2000: 172 (list of major ant collections); Bolton, 2003: 15, 77 (diagnoses, synopses).

World and zoogeographical regional catalogues
Roger, 1863b: 1 (world Formicidae); Mayr, 1863: 394 (world Formicidae); Dalla Torre, 1893: 1 (world Formicidae); Emery, 1910b: 3 (world Dorylinae); Emery, 1911d: 2 (world Ponerinae); Emery, 1913a: 2 (world Dolichoderinae); Wheeler, W.M. 1922a: 727, 1005 (Afrotropical and Malagasy); Emery, 1921f: 3, Emery, 1922e: 95, and Emery, 1924d: 207 (world Myrmicinae); Emery, 1925b: 2 (world Formicinae); Chapman & Capco, 1951: 9 (Oriental and Malesian); Smith, M.R. 1951a: 778 (Nearctic); Smith, M.R. 1958c: 108 (Nearctic, first supplement to previous); Smith, M.R. 1967: 343 (Nearctic, second supplement); Kempf, 1972a: 3 (Neotropical); Smith, D.R. 1979: 1323 (Nearctic). Taylor & Brown, D.R. 1985: 5 (Austral); Taylor, R.W. 1987a: 1, and Taylor, R.W. 1987b: 1 (Austral); Brandão, 1991: 319 (Neotropical, supplement to Kempf, 1972a); Shattuck, 1994: 1 (world Aneuretinae and Dolichoderinae); Bolton, 1995b: 7 (world Formicidae, extant and fossil).

World synoptic classifications
Forel, 1893a: 161 (Formicidae); Emery, 1895j: 764 (Formicidae); Emery, 1896e: 173 (Formicidae); Ashmead, 1905b: 381 (Formicidae); Wheeler, W.M. 1910g: 134 (Formicidae); Emery, 1910b: 3 (Dorylinae); Emery, 1911d: 2 (Ponerinae); Emery, 1913a: 2 (Dolichoderinae); Forel, 1917: 235 (Formicidae); Emery, 1921f: 3, and Emery, 1922e: 95 (Myrmicinae); Wheeler, W.M. 1922a: 631 (Formicidae); Emery, 1924d: 207 (Myrmicinae); Emery, 1925b: 2 (Formicinae); Brown, 1973b: 165, 178 (Formicidae); Snelling, R.R. 1981: 387 (Formicidae); Wheeler, G.C. & Wheeler, J. 1985: 256 (Formicidae); Dlussky & Fedoseeva, 1988: 77 (Formicidae); Hölldobler & Wilson, 1990: 9 (Formicidae); Bolton, 1994: 12 (Formicidae); Bolton, 2003: 77 (Formicidae); Engel & Grimaldi, 2005: 4 (Formicidae); Ward, 2007a: 549 (Formicidae); Boudinot, 2015: 12 (Formicidae).

World identification keys to higher taxa
Emery, 1896e: 173 (Formicidae subfamilies and genera); Emery, 1910b: 4 (Dorylinae tribes and genera); Emery, 1911d: 4 (Ponerinae tribes and genera); Emery, 1913a: 6 (Dolichoderinae tribes and genera); Escherich, 1917: 2 (Formicidae subfamilies); Emery, 1921f: 9, 12, 16 (Myrmicinae tribes and genera); Wheeler, W.M. 1922a: 631 (Formicidae subfamilies, tribes and genera); Emery, 1925b: 7 (Formicinae tribes and genera); Clark, 1951: 14 (Formicidae subfamilies); Wheeler, G.C. & Wheeler, J. 1972a: 42 (Formicidae subfamilies); Snelling, R.R. 1981: 386 (Formicidae subfamilies); Hölldobler & Wilson, 1990: 33 (Formicidae subfamilies and genera); Bolton, 1990c: 1361 (Formicidae subfamilies); Shattuck, 1992c: 20 (Dolichoderinae genera); Ward, 1990: 464 (Pseudomyrmecinae genera); Brothers & Finnamore, 1993: 218 (Formicidae subfamilies); Bolton, 1994: 7 (Formicidae subfamilies and genera); Bolton, 2003: 11 (Formicidae subfamilies); Boudinot, 2015: 14 (Formicidae subfamilies, males); Fisher & Bolton, 2016: 28, 60, 128 (Afrotropical and Malagasy subfamilies key, Afrotropical genera key, Malagsy genera key).

Evolution and phylogeny
Wheeler, W.M. 1920: 52; Emery, 1920d: 370; Donisthorpe, 1922: xlvi; Wheeler, W.M. 1923f: 156; Wheeler, W.M. 1928f: 124; Stitz, 1936: 211; Morley, 1938: 190; Brown & Nutting, 1950: 113; Bernard, 1951: 1041; Brown, 1954e: 30; Wilson, et al. 1967: 12; Taylor, 1978a: 982; Rasnitsyn, 1980: 45-46 (foldout page); Lutz, 1986: 213; Dlussky & Fedoseeva, 1988: 137; Baroni Urbani, 1989: 137; Hölldobler & Wilson, 1990: 26; Hashimoto, 1991a: 125; Hashimoto, 1991b: 289; Baroni Urbani, et al. 1992: 301; Shattuck, 1992b: 203; Brothers & Carpenter, 1993: 227; Ward, 1994: 173; Hashimoto, 1996: 340; Crozier, et al. 1997: 22; Grimaldi, et al. 1997: 25; Perrault, 1999: 125; Brothers, 1999: 233; Grimaldi & Agosti, 2000: 13680; Ward & Brady, 2003: 361; Ohnishi, et al. 2003: 419; Astruc, et al. 2004: 880; Ward & Downie, 2005: 310; Grimaldi & Engel, 2005: 440; Wilson & Hölldobler, 2005: 7411; Brady, et al. 2006: 18172; Moreau, et al. 2006: 101; Ouellette, et al. 2006: 359; Ward, 2007a: 549; Rabeling, et al. 2008: 14914; Ward, et al. 2010: 342; Keller, 2011: 1; Moreau & Bell, 2013; Schmidt, 2013: 201; Ward, et al. 2015: 61.

Morphology
Forel, 1878: 339 (proventriculus); Janet, 1902: 25 (general anatomy); Pietschker, 1911: 43 (brain structure); Wheeler, W.M. 1910g: 13 (general morphology); Emery, 1913c: 577 (venation); Wheeler, W.M. 1918c: 293 (larvae); Janet, 1923: 5 (anatomical drawings synopsis); Bugnion, 1930: 85 (mouthparts); Brown & Nutting, 1950: 113 (venation); Gregg, 1953a: 328 (promesonotal suture motility); Eisner, 1957: 439 (proventriculus); Gotwald, 1969: 1 (mouthparts and gaster); Crozier, 1970: 109 (karyology); Markl, 1973: 258 (stridulitrum); Wheeler, G.C. & Wheeler, J. 1976b: 45 (larvae); Hölldobler & Engel, 1979: 285 and Hölldobler & Engel-Siegel, 1982: 113 (tergal and sternal glands); Imai, Baroni Urbani, Kubota et al. 1984: 5 (karyology); Imai, et al. 1984: 66 (karyology); Imai, et al. 1985: 46 (karyology); Hölldobler & Engel-Siegel, 1985: 201 (metapleural gland); Billen, 1986: 165 (Dufour's gland); Wheeler, G.C. & Wheeler, J. 1986e: 684 (larvae); Brown, 1988h: 17 (malpighian tubule numbers); Caetano, 1988: 129 (digestive tract anatomy); Francoeur & Loiselle, 1988a: 333 (strigil, evolution); Hölldobler & Wilson, 1990: 5, 229 (anatomy, exocrine glands); Hashimoto, 1991a: 125 (antennal and labial sensilla); Hashimoto, 1991b: 289 (tibial spurs); Billen, 1993: 4 (exocrine glands); Bolton, 1994: 191 (morphology glossary); Hölldobler, et al. 1996: 157 (metatibial gland); Wenseleers, et al. 1998: 121 (cloacal gland); Perrault, 1999: 125 (thoracic anatomy); Perrault, 2004: 291 (anterior metasomal anatomy); Keller, 2011: 1 (phylogenetic analysis of morphology); Keller, et al. 2014: 1 (thorax architecture); Boudinot, 2015: 12 (males); Fisher & Bolton, 2016: 459 (morphology glossary).

Palaeontology
Scudder, 1891: 682 (fossil taxa catalogue); Handlirsch, 1907: 859 (fossil taxa catalogue); Wheeler, W.M. 1915h: 1 (Baltic amber fauna synopsis); Burnham, 1979: 108 (fossils in amber checklist); Keilbach, 1982: 272 (fossils in amber checklist); Spahr, 1987: 41 (fossils in amber checklist); Carpenter, 1992: 490 (fossil genera review); Bolton, 1995b: 4 (fossil taxa checklist); Grimaldi, et al. 1997: 1 (Cretaceous genera in amber); Dlussky, 1997: 617 (Baltic amber genera key); Bolton, 2003: 73 (fossil subfamilies synopsis); Dlussky & Rasnitsyn, 2007: 118 (Palaeontological record); Dlussky & Rasnitsyn, 2009: 1024 (European Upper Eocene amber, synopsis of species); Dlussky & Perfilieva, 2014: 433 (Bembridge Marls (Eocene, U.K.) species, key); Perkovsky, 2016: 113 (list of ants in Late Eocene ambers).