Ponerinae

Ponerinae is the largest ant subfamily outside the formicoid clade, and is rivaled or exceeded in diversity only by Dolichoderinae, Formicinae and Myrmicinae within that clade. Species of ponerines range from small and cryptic to large and conspicuous.

Identification
The mesosoma is attached to the gaster with a single distinct segment, the petiole. The gaster usually has a slight but distinct impression between the first and second segments. In a few other cases (species of Odontomachus) the gaster is smooth and uniform, but here the mandibles are elongate and straight, with teeth only at the extreme tip, and attached close together along the front margin of the head. The upper surface of the tip of the gaster (the pygidium) is rounded and lacks a row of spines or teeth on its outer and trailing edge. The sting is present (although often retracted and difficult to see).

Most species in this large and diverse subfamily can be identified by the presence of a single-segmented petiole combined with a constriction between the first and second segment of the gaster. In the few cases where the constriction is absent (Odontomachus), the overall shape of the gaster or the shape of the head can be used to identify these ants (see above for details).

Boudinot (2015) - Male Ponerinae share the following characters: antennal toruli situated well-posterad anterior clypeal margin (except Dolioponera); at least 4 closed cells present on forewing (Dolioponera with 3); propodeal lobes usually present; jugal lobe usually present; petiolar tergum and sternum distinct; cinctus between abdominal pre- and posttergites IV usually present; and abdominal sternum IX unpronged and edentate. Three final sets of characters are required for identification: 1) (Platythyreini) mandibles triangular, tibial spur formula 2,2; 2) (Ponerini) mandibles spatulate, linear, or nub-like and mesonotum not anteriorly elongated; and 3) (Ponerini, Dolioponera) forewing with three closed cells, propodeal lobes present, antennal toruli situated at anterior extreme of head, oblique mesopleural sulcus absent, and cinctus present. The eighth abdominal tergum of male Ponerinae may be spiniform, a unique state among the Formicidae, but this character is not present in all genera and may be interspecifically variable. The spur formula of Ponerini is variable.

Australia
They are found throughout Australia from pristine habitats to disturbed sites such as gardens and parks, and can be quite abundant. Workers are predacious, generally forage on the ground, and some specialise on a very limited range of prey. In addition, many species have powerful and painful stings used for subduing prey and as a defensive measure against intruders. Within Australia there are about 200 described species in 10 genera (pre - Schmidt and Shattuck 2012), with numerous species yet to be studied in detail.

Nomenclature

 * [subfamily of Formicidae]
 * Ponerites Lepeletier de Saint-Fargeau, 1835: 185. Type-genus: Ponera Latreille, 1804: 179.

Taxonomic History

 * Ponerinae as group name: Lepeletier de Saint-Fargeau, 1835: 185 [Ponérites].
 * Ponerinae as family: Smith, F. 1851: 6 [Poneridae]; Smith, F. 1858b: 76 [Poneridae]; Smith, F. 1861b: 44 [Poneridae]; Smith, F. 1871a: 320 [Poneridae]; André, 1882a: 125 [Poneridae]; Cresson, 1887: 93 [Poneridae]; Emery, 1894g: 379 [Poneridae]; Saunders, 1896: 18 [Poneridae]; Ashmead, 1905b: 382 [Poneridae]; Novák & Sadil, 1941: 70 [Poneridae]; Bernard, 1951: 1042 [Poneridae]; Bernard, 1953b: 185 [Poneridae].
 * Ponerinae as tribe of Formicidae: André, 1874: 167 [Poneridae].
 * Ponerinae as subfamily of Poneridae: Ashmead, 1905b: 382.
 * Ponerinae as subfamily of Formicidae: Mayr, 1855: 289, 299 [Poneridae]; Mayr, 1861: 21 [Poneridae]; Mayr, 1862: 712 [Poneridae]; Smith, F. 1857a: 64 [Poneridae]; Smith, F. 1862b: 31 [Poneridae]; Mayr, 1865: 11 [Poneridae]; Mayr, 1868b: 24 [Poneridae]; Forel, 1870: 307 [Poneridae]; Forel, 1874: 21 [Poneridae]; Emery, 1877a: 70 [Poneridae]; Forel, 1878: 366 [Poneridae]; Emery & Forel, 1879a: 455 [Poneridae]; André, 1881b: 64 [Poneridae]; Forel, 1892j: 220 [Poneridae]; Dalla Torre, 1893: 13; Forel, 1893a: 162 [Ponerinae]; Forel, 1895b: 111 [Poneridae]; Nasonov, 1889: 27 [Poneridae]; Emery, 1895j: 766 [subfamily spelled Ponerini]; Emery, 1896e: 176; Forel, 1899c: 2; Emery, 1901a: 36; Bingham, 1903: 23; Ruzsky, 1905b: 174; Wheeler, W.M. 1910g: 134; Emery, 1911d: 2; Wheeler, 1915g: 805 [Ponerides]; Donisthorpe, 1915d: 65; Wheeler, W.M. 1915h: 25; Arnold, 1915: 9; Escherich, 1917: 2 [Ponerini]; Forel, 1917: 235; Bondroit, 1918: 78 [Poneritae]; Wheeler, W.M. 1920: 53; Wheeler, W.M. 1922a: 56, 632, 640; Borgmeier, 1923: 37; Karavaiev, 1934: 49; Clark, 1951: 15; Brown, 1954e: 24; Wheeler, G.C. & Wheeler, J. 1972a: 39; Brown, 1973b: 165; subsequent authors.
 * Ponerinae as poneromorph subfamily of Formicidae: Bolton, 2003: 42, 156.
 * Ponerinae as poneroid subfamily of Formicidae: Ouellette, et al. 2006: 365; Brady, et al. 2006: 18173; Moreau, et al. 2006: 102; Ward, 2007a: 555; Schmidt, C.A. & Shattuck, 2014: 46.

Taxonomic References
Smith, F. 1858b: 76 (diagnosis); Mayr, 1862: 712 (genera key); Mayr, 1865: 11 (Odontomachidae, Poneridae diagnoses), Mayr, 1867a: 79, 81 (Odontomachidae, Poneridae diagnoses); Forel, 1878: 366 (diagnosis); Dalla Torre, 1893: 13 (catalogue); Emery, 1895j: 766 (diagnosis); Emery, 1896e: 176 (genera key); Handlirsch, 1907: 879 (*fossil taxa catalogue); Wheeler, W.M. 1910g: 134 (diagnosis); Emery, 1911d: 3 (diagnosis, tribes key); Forel, 1917: 235 (synoptic classification); Forel, 1921c: 133 (diagnosis); Wheeler, W.M. 1922a: 56, 636 (diagnosis, tribes key); Brown & Nutting, 1950: 124 (venation, phylogeny); Brown, 1954e: 24 (phylogeny); Eisner, 1957: 475 (proventriculus morphology); Bernard, 1967: 79 (diagnosis); Gotwald, 1969: 25 (mouthparts morphology); Wheeler, G.C. & Wheeler, J. 1972a: 39 (diagnosis); Brown, 1973b: 165 (genera, distribution); Wheeler, G.C. & Wheeler, J. 1976b: 48 (larvae, review and synthesis); Snelling, R.R. 1981: 387 (synoptic classification); Wheeler, G.C. & Wheeler, J. 1985: 256 (synoptic classification); Billen, 1986: 168 (Dufour's gland); Fanfani & Dazzini Valcurone, 1986: 115 (exocrine glands synopsis); Dlussky & Fedoseeva, 1988: 78 (synoptic classification); Hölldobler & Wilson, 1990: 9 onward (synoptic classification, genera keys); Baroni Urbani, et al. 1992: 317 (phylogeny); Bolton, 1994: 153 (diagnosis, synoptic classification, genera keys); Bolton, 1995a: 1042 (census); Bolton, 1995b: 14 (catalogue); Grimaldi, et al. 1997: 20 (*Cretaceous genera); Dlussky & Rasnitsyn, 2002: 422 (diagnosis for wingless fossils); Bolton, 2003: 42, 156 (diagnosis, synopsis); Ouellette, et al. 2006: 359 (phylogeny); Brady, et al. 2006: 18173 (phylogeny); Moreau, et al. 2006: 102 (phylogeny); Ward, 2007a: 555 (classification); Keller, 2011: 1 (morphology, phylogeny); Schmidt, C. 2013: 201 (molecular, phylogeny); Schmidt, C.A. & Shattuck, 2014: 18, 24, 36 (revision of world genera, keys to New World, Afrotropical and Malagasy, Eurasian and Australian); Boudinot, 2015: 47 (diagnosis).

Regional and National Faunas with Keys
Mayr, 1861: 28 (Europe); Mayr, 1868b: 69 (*Baltic Amber); André, 1874: 170 (Europe); Forel, 1874: 29 (Switzerland); André, 1882b: 227 (Europe and Algeria); Cresson, 1887: 97 (U.S.A., genera); Provancher, 1887: 238 (Canada); Nasonov, 1889: 54 (Russia); Forel, 1891b: 9 (Madagascar genera); Forel, 1900c: 52 (India and Sri Lanka); Bingham, 1903: 23 (India, Sri Lanka and Burma); Ruzsky, 1905b: 108 (Russian Empire); Wheeler, W.M. 1910g: 557 (North America genera); Bondroit, 1910: 489 (Belgium); Stitz, 1914: 54 (Central Europe); Gallardo, 1915: 31 (Argentina genera); Forel, 1915d: 6 (Switzerland); Arnold, 1915: 10 (South Africa); Donisthorpe, 1915d: 65 (Britain); Emery, 1916b: 98 (Italy); Wheeler, W.M. 1916m: 580 (U.S.A., Connecticut); Bondroit, 1918: 79 (France and Belgium); Gallardo, 1918b: 4 (Argentina); Soudek, 1922: 18 (Czechoslovakia); Stärcke, 1926: 82 (Netherlands); Donisthorpe, 1927b: 68 (Britain); Menozzi & Russo, 1930: 169 (Dominican Republic); Arnol'di, 1933b: 596 (Russia); Karavaiev, 1934: 50 (Ukraine); Smith, M.R. 1937: 823 (Puerto Rico); Stitz, 1939: 58 (Germany); Smith, M.R. 1943f: 278 (U.S.A. males); Buren, 1944a: 279 (U.S.A., Iowa); Smith, M.R. 1947f: 530 (U.S.A. genera); Creighton, 1950a: 31 (Nearctic); Kusnezov, 1956: 11 (Argentina); Brown, 1958h: 11 (New Zealand); Gregg, 1963: 280 (U.S.A., Colorado); Bernard, 1967: 80 (Western Europe); Wilson & Taylor, 1967: 10 (Polynesia); Kempf, 1972a: 262 (Neotropical synoptic classification); Bolton, 1973a: 323 (West Africa genera); Bolton & Collingwood, 1975: 3 (Britain); van Boven, 1977: 66 (Belgium); Kutter, 1977c: 21 (Switzerland); Arnol'di & Dlussky, 1978: 522 (former European U.S.S.R.); Collingwood, 1978: 74 (Iberian Peninsula); Collingwood, 1979: 29 (Fennoscandia and Denmark); Greenslade, 1979: 14 (South Australia genera); Francoeur, 1979: 30 (Canada, Québec); Schembri & Collingwood, 1981: 417 (Malta); Allred, 1982: 438 (U.S.A., Utah); Baroni Urbani, 1984: 75 (Neotropical genera); Collingwood, 1985: 236 (Saudi Arabia); Wheeler, G.C. & Wheeler, J. 1986g: 17 (U.S.A., Nevada); Agosti & Collingwood, 1987b: 264 (Balkans); Ogata, 1987: 101 (Japan genera); Morisita, et al. 1989: 8 (Japan); Dlussky, et al. 1990: 173 (Turkmenistan); Kupyanskaya, 1990: 85 (Far Eastern Russia); Atanasov & Dlussky, 1992: 51 (Bulgaria); Lattke, in Jaffe, 1993: 166 (Neotropical genera); Arakelian, 1994: 10 (Armenia); Wu, J. & Wang, 1995: 31 (China genera); Collingwood & Agosti, 1996: 308 (Saudi Arabia); Seifert, 1996b: 107 (Central Europe); Collingwood & Prince, 1998: 10 (Portugal); Kim, et al. 1998: 145 (Korea); Shattuck, 1999: 52, 179 (Australia genera synopsis); Andersen, 2000: 19 (northern Australia genera); Zhou, 2001b: 22 (China, Guangxi); Czechowski, et al. 2002: 133 (Poland); Aktaç & Radchenko, 2002: 53 (Turkey genera); Yoshimura & Onoyama, 2002: 436 (Japan genera, males); Mackay & Mackay, 2002: 27 (U.S.A., New Mexico); Csösz, 2003: 147 (Carpathian Basin); Palacio & Fernández, in Fernández, 2003d: 256 (Neotropical genera); Radchenko, 2005b: 184 (North Korea); Coovert, 2005: 23 (U.S.A., Ohio); Yoshimura & Fisher, 2007: 27 (Malagasy genera, males); Clouse, 2007b: 190 (Micronesia); Seifert, 2007: 108 (North and Central Europe); Bolton & Fisher, 2008c: 32 (Afrotropical genera); Fernández & Arias-Penna, 2008: 32 (Neotropical genera); Dlussky, 2009: 1071 (Eocene amber); Terayama, 2009: 101 (Taiwan genera); Heterick, 2009: 34 (south-western Australia genera); Boer, 2010: 68 (Benelux); Czechowski, et al. 2012: 340 (Poland); General & Alpert, 2012: 76 (Philippines genera key) ; Dlussky & Perfilieva, 2014: 433 (British Eocene species key); Baccaro, et al. 2015: 108, 310 (Brazil genera key, text); Lattke, in Delabie, et al. 2015: 55 (Brazil genera, phylogeny, taxonomy); Fisher & Bolton, 2016: 52 (diagnosis).

Taxonomic Notes
Schmidt and Shattuck (2012) - The following formal diagnosis for Ponerinae is adapted from Bolton (2003): Torulus fused to frontal lobe. Antenna with 12 segments (13 in males). Lateral margins of frontal lobes form short semicircles or blunt triangles, with a pinched-in appearance posteriorly. Promesonotal suture flexible. Metapleural gland orifice without a dorsal cuticular flange or flap. Propodeal lobes present. Petiole (A2) distinctly separated posteriorly from A3 and with only a narrow attachment to it. Petiole without tergosternal fusion. A3 continuous with the remainder of the gaster. A3 and A4 with tergosternal fusion. A4 with presclerites and usually a girdling constriction between pre- and postsclerites. Spiracles of A5–A7 concealed by posterior margins of preceding tergites. Sting present and strongly developed.

Ponerines are most readily identified by the following combination of traits: toruli fused to frontal lobes, frontal lobes prominent and with a pinched-in appearance posteriorly, waist formed of a single segment (petiole, A3) which attaches narrowly to the undifferentiated postpetiole (A4), petiole without tergosternal fusion, and sting present and well-developed. The identity of ponerine synapomorphies is uncertain. Bolton (2003) gave the complete fusion of the toruli to the frontal lobes as an autapomorphy of Ponerinae, but noted the presence of similar fusion (to various degrees) among some members of Amblyoponinae. Bolton also listed the characteristic shape of the frontal lobes in Ponerinae as synapomorphic for the subfamily, but similar frontal lobe structure occurs in many amblyoponines (pers. observation). Given the close but incompletely resolved relationship between Ponerinae and Amblyoponinae, we consider the ancestral condition of these characteristics (torular fusion and frontal lobe shape) to be ambiguous within the poneroid clade and are therefore hesitant to treat them as apomorphies of Ponerinae.

We tentatively recognize the loss of tergosternal fusion of the petiole as a possible synapomorphy of Ponerinae. Bolton (2003) treated the absence of petiolar tergosternal fusion as plesiomorphic within Formicidae, but recent molecular phylogenies (Moreau et al., 2006; Brady et al., 2006; Rabeling et al., 2008; Schmidt, 2013) suggest that the ancestral ant may have had a fused petiole. The phylogenetic distribution of this character implies that Ponerinae secondarily lost tergosternal fusion of the petiole, though this interpretation depends on the phylogenetic rooting of Formicidae and could conceivably be symplesiomorphic, with repeated evolution of a fused petiole in other poneroid lineages.