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Family: Formicidae   Latreille, 1809 

Classification:

Taxonomic History (provided by Barry Bolton, 2015)

Extant: 16 subfamilies, 39 tribes, 326 genera, 13,094 species, 2,086 subspecies

Fossil: 4 subfamilies, 8 tribes, 147 genera, 689 species, 2 subspecies

Formicariae Latreille, 1809 PDF: 124 . Type-genus: Formica. AntCat AntWiki

Taxonomic history

Formicidae as family: Latreille, 1809 PDF: 124 [Formicariae]; Leach, 1815: 147 [Formicarides]; Stephens, 1829b: 356 [first spelling as Formicidae]; Haliday, 1836: 331; Westwood, 1839a: 217; Swainson & Shuckard, 1840: 171; Nylander, 1846a PDF: 877; Foerster, 1850a: 1 [Formicariae]; Mayr, 1855 PDF: 275 [Formicina]; Smith, 1857a PDF: 52; Smith, 1858a PDF: 1; Mayr, 1861 PDF: 21; Mayr, 1865: 6; Heer, 1867 PDF: 6 [Formicaria]; Forel, 1870 PDF: 307 [Formicinae]; Forel, 1874 PDF: 19 [Formicariae]; Dalla Torre, 1893 PDF: 1; Forel, 1899B: 1; Ruzsky, 1902d: 5 [Formicarii]; Bingham, 1903 PDF: 1; Ruzsky, 1905b: 91 [Formicariae or Formicidae]; Ashmead, 1905c PDF: 384; all subsequent authors.
WORLD IDENTIFICATION KEYS
Emery, 1896e PDF: 173 (Formicidae subfamilies and genera); Emery, 1910b PDF: 4 (Dorylinae tribes and genera); Emery, 1911e PDF: 4 (Ponerinae tribes and genera); Emery, 1913a PDF: 6 (Dolichoderinae tribes and genera); Escherich, 1917: 2 (Formicidae subfamilies); Emery, 1921c: 9, 12, 16 (Myrmicinae tribes and genera); Wheeler, 1922: 631 (Formicidae subfamilies, tribes and genera); Emery, 1925d PDF: 7 (Formicinae tribes and genera); Clark, 1951 PDF: 14 (Formicidae subfamilies); Wheeler & Wheeler, 1972a PDF: 42 (Formicidae subfamilies); Snelling, 1981: 386 (Formicidae subfamilies); Hölldobler & Wilson, 1990: 33 (Formicidae subfamilies and genera); Bolton, 1990c PDF: 1361 (Formicidae subfamilies); Shattuck, 1992c PDF: 20 (Dolichoderinae genera); Ward, 1990 PDF: 464 (Pseudomyrmecinae genera); Brothers & Finnamore, 1993: 218 (Formicidae subfamilies); Bolton, 1994: 7 (Formicidae subfamilies and genera); Bolton, 2003 PDF: 11 (revision of subfamilies); Boudinot, 2015 PDF: 14 (male-based global subfamily key)
MORPHOLOGY OF FORMICIDAE
Forel, 1878c PDF: 339 (proventriculus); Janet, 1902: 25 (general anatomy); Pietschker, 1910: 1 (brain structure); Wheeler, 1910a: 13 (general morphology, anatomy); Emery, 1913d: 577 (venation); Wheeler, 1918c PDF: 293 (larvae); Janet, 1923: 5 (anatomical drawings synopsis); Bugnion, 1930: 85 (mouthparts); Brown & Nutting, 1950 PDF: 113 (venation); Gregg, 1953c PDF: 328 (promesonotal suture motility); Eisner, 1957 PDF: 439 (proventriculus); Gotwald, 1969: 1 (mouthparts and gaster); Crozier, 1970a PDF: 109 (karyology); Markl, 1973: 258 (stridulitrum); Wheeler & Wheeler, 1976b PDF: 45 (larvae); Hölldobler & Engel, 1979 PDF: 285 and Hölldobler & Engel-Siegel, 1982: 113 (tergal and sternal glands); Imai, Baroni Urbani, Kubota, et al. 1984: 5 (karyology); Imai, Brown, Kubota, et al. 1984: 66 (karyology); Imai, Kubota, Brown, et al. 1985: 46 (karyology); Hölldobler & Engel-Siegel, 1985 PDF: 201 (metapleural gland); Billen, 1986b: 165 (Dufour's gland); Wheeler & Wheeler, 1986e PDF: 684 (larvae); Brown, 1988h: 17 (malpighian tubule numbers); Caetano, 1988: 129 (digestive tract anatomy); Francoeur & Loiselle, 1988a PDF: 333 (strigil, evolution); Hölldobler & Wilson, 1990: 5, 229 (anatomy, exocrine glands); Hashimoto, 1991a PDF: 125 (antennal and labial sensilla); Hashimoto, 1991b PDF: 289 (tibial spurs); Billen, 1993: 4 (exocrine glands); Bolton, 1994: 191 (morphology glossary); Hölldobler, Obermayer & Peeters, 1996: 157 (metatibial gland); Wenseleers, Schoeters, et al. 1998: 121 (cloacal gland); Perrault, 1999A PDF: 125 (thoracic anatomy); Perrault, 2005: 291 (anterior metasomal anatomy); Serna & MacKay, 2010 PDF: 1 (Procryptocerus morphology); Keller, 2011 PDF: 1 (phylogenetic analysis of morphology); Boudinot, 2013 PDF: 29 (male genitalia); Boudinot, 2015 PDF: 1 (family diagnosis, morphology)
PALAEONTOLOGY OF FORMICIDAE
Scudder, 1891: 682 (fossil taxa catalogue); Handlirsch, 1907: 859 (fossil taxa catalogue); Wheeler, 1915i PDF: 1 (Baltic amber fauna synopsis); Burnham, 1979 PDF: 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, in catalogue); Grimaldi, Agosti & Carpenter, 1997 PDF: 1 (Cretaceous genera in amber review); Dlussky, 1997A PDF: 617 (Baltic amber genera key); Bolton, 2003 PDF: 73 (fossil subfamilies synopsis); Dlussky & Rasnitsyn, 2007 PDF: 118 (Palaeontological record); Dlussky & Rasnitsyn, 2009 PDF: 1024 (European Upper Eocene amber, synopsis of species); LaPolla, Dlussky & Perrichot, 2013 PDF (Mesozoic and Cenozoic major ant fossil deposits list, Cretaceous and Cenozoic ant fossils bibliographical review).
OTHER GENERAL REFERENCES
Bolton, 1995a PDF: 1038 (census of extant taxa); Ward, Bolton, Shattuck & Brown, 1996}: 55 (bibliography of systematics); Brown, 2000: 46 (overview of genera); Brandão, 2000: 172 (list of major ant collections); Bolton, 2003 PDF: 15, 77 (diagnosis, synopsis); Bolton, 2007A: 51 (method for taxonomic revisions); Taylor, 2007: 580 (eusociality in ants)
Regional catalogues and checklists
NEARCTIC & PALEARCTIC: Smith, 1951c: 778; Smith, 1958c: 108 (first supplement to previous); Smith, 1967a: 343 (second supplement); Smith, 1979: 1323; Karaman, 2011 PDF (Montenegro catalogue); Borowiec, 2014 PDF (Europe, Mediterranean Basin and adjacent regions)
NEOTROPICAL: Kempf, 1972b PDF: 3; Brandão, 1991 PDF: 319 (supplement to previous)
AFROTROPICAL AND MALAGASY: Wheeler, 1922: 727, 1005
ORIENTAL AND MALESIAN: Chapman & Capco, 1951 PDF: 9
AUSTRAL: Taylor & Brown, 1985: 5; Taylor, 1987a PDF: 1; Taylor, 1987b: 1 (supplement to previous)
Regional and national faunas with keys
PALAEARCTIC
Mayr, 1855 PDF: 299 (Austria); Mayr, 1861 PDF: 25 (Europe); Forel, 1874 PDF: 19 (Switzerland); Saunders, 1880 PDF: 202 (Britain); André, 1882a: 125 (Europe & Algeria); Nasonov, 1889: 50 (Russia); Lameere, 1892: 62 (Belgium); Saunders, 1896: 18 (Britain); Ruzsky, 1905b: 99 (Russian Empire); Wasmann, 1906 PDF: 7 (Luxemburg); Emery, 1908a PDF (Palaearctic); Bondroit, 1910 PDF: 480 (Belgium); Stitz, 1914: 54 (Central Europe); Donisthorpe, 1915f: 65 (Britain); Forel, 1915d: 1 (Switzerland); Emery, 1916a PDF: 92 (Italy); Bondroit, 1918 PDF: 12 (France & Belgium); Kutter, 1920b: 132 (Switzerland); Soudek, 1922b PDF: 17 (Czechoslovakia); Lomnicki, 1925a PDF: 160 (Poland); Stärcke, 1926: 79 (Netherlands); Karavaiev, 1927d: 254 (Ukraine); Donisthorpe, 1927c: 66 (Britain); Arnol'di, 1933a: 596 (Russia); Menozzi, 1933b PDF: 87 (Israel); Karavaiev, 1934: 48 (Ukraine); Stitz, 1939: 56 (Germany); Kratochvíl, 1941b: 69 (Central Europe); Novák & Sadil, 1941 PDF: 69 (Central Europe); Ceballos, 1943: 313 (Spain);Holgersen, 1942b PDF: 3 (Northern Norway); Holgersen, 1943c: 166 and Holgersen, 1944a: 198 (Norway); Boven, 1947: 168 (Belgium); Collingwood, 1958b PDF: 69 (Britain); Somfai, 1959: 1 (Hungary); Boven, 1959: 1 (Netherlands); Collingwood, 1964b PDF: 93 (Britain); Bernard, 1967a PDF: 75 (Western Europe); Boven, 1970b: 7 (Netherlands); Tarbinsky, 1976 PDF: 14 (Kyrghyzstan); Bolton & Collingwood, 1975: 1 (Britain); Boven, 1977 PDF: 1 (Belgium); Kutter, 1977c: 19 (Switzerland); Brian, 1977: 33 (Britain); Arnol'di & Dlussky, 1978: 520 (former European U.S.S.R.); Collingwood, 1978 PDF: 65 (Iberian Peninsula); Collingwood, 1979 PDF: 28 (Fennoscandia & Denmark); Schembri & Collingwood, 1981 PDF: 417 (Malta); Verhaeghe, Deligne, et al., 1984: 105 (Belgium genera); Gösswald, 1985: 262 (Germany); Collingwood, 1985 PDF: 230 (Saudi Arabia); Nilsson & Douwes, 1987: 56 (Norway); Agosti & Collingwood, 1987b PDF: 261 (Balkans); Casevitz-Weulersse, 1990b: 135 and Casevitz-Weulersse, 1990c PDF: 415 (Corsica); Dlussky, Soyunov & Zabelin, 1990 PDF: 99 (Turkmenistan); Kupyanskaya, 1990a: 83 (Far Eastern Russia); Bolton, in Hölldobler & Wilson, 1990: 34 (Palaearctic genera key); Atanasov & Dlussky, 1992: 47 (Bulgaria); Arakelian, 1994 PDF: 1 (Armenia); Radchenko, 1994b: 95 (South Siberia); Seifert, 1994a PDF: 1 (Germany, synopsis); Onoyama & Terayama, 1994: 1 (Japan bibliography); Bolton, 1994: 22, 42, 75, 153 (Dolichoderinae, Formicinae, Myrmicinae, Ponerinae genera keys); Douwes, 1995: 83 (Sweden); Kupyanskaya, 1995: 327 (Far Eastern Russia); Wu & Wang, 1995a: 30 (China); Fontenla Rizo, 1995b: 16 (Cuba, tramp species key); Collingwood & Agosti, 1996 PDF: 300 (Saudi Arabia); Collingwood & Van Harten, 1994: 1 (Yemen); Seifert, 1996B: 106 (Central Europe); Skinner & Allen, 1996: 40 (Britain); Dlussky, 1997A PDF: 616 (*Baltic Amber genera); Collingwood & Prince, 1998: 9 (Portugal); Czechowski, Radchenko & Czechowska, 2002 PDF: 133 (Poland); Aktaç & Radchenko, 2002: 53 (Turkey genera); Yoshimura & Onoyama, 2002B PDF: 424 (Japan genera, males key); Imai, Kihara, Kondoh, et al. 2003: 8 (Japan); Radchenko, 2005B PDF: 183 (North Korea); Seifert, 2007: 107 (North and Central Europe); Boer, 2010: 8 (Benelux); Lapeva-Gjonova, et al. 2010: 1 (Bulgaria species catalogue); Cs?sz, Markó & Gallé, 2011 PDF: 56 (Hungary species catalogue); Czechowski, Radchenko, et al. 2012: 333 (Poland); Douwes, Abenius, Cederberg & Wahlstedt, 2012: 1 (Sweden)
NEARCTIC
Provancher, 1883 PDF: 596 (Canada); Provancher, 1887: 225 (Canada); Cresson, 1887 PDF: 93 (U.S.A.); Wheeler, 1910a: 557 (North America genera); Wheeler, 1916r: 579 (U.S.A., Connecticut); Cole, 1942 PDF}: 359 (U.S.A., Utah); Smith, 1943e PDF: 276 (U.S.A. subfamilies & genera, males); Buren, 1944a PDF: 278 (U.S.A., Iowa); Smith, 1947f PDF: 524 (U.S.A. genera); Creighton, 1950a PDF: 29 (North America); Gregg, 1963: 278 (U.S.A., Colorado); Wheeler & Wheeler, 1963: 149 (U.S.A., North Dakota); Francoeur, 1979a PDF: 48 (Canada, Québec, introduction); Allred, 1982: 437 (U.S.A., Utah); Wheeler & Wheeler, 1986g PDF: 16 (U.S.A., Nevada); Mackay, Lowrie, et al., 1988: 81 (U.S.A., New Mexico); Cover, in Hölldobler & Wilson, 1990: 62 (Nearctic genera key); Mackay & MacKay, 2002 PDF: 16 (U.S.A., New Mexico); Dlussky & Rasnitsyn, 2003 PDF: 430 (*Eocene North American genera); Bolton, 1994: 25, 38, 48, 95, 160 (Dolichoderinae, Ecitoninae, Formicinae, Myrmicinae, Ponerinae genera keys); Coovert, 2005: 1 (U.S.A., Ohio); Ward, 2005 PDF: 20 (U.S.A., California genera); Fisher & Cover, 2007: 1 (North American genera key)
NEOTROPICAL
Gallardo, 1915 PDF: 30 (Argentina genera); Menozzi & Russo, 1930 PDF: 169 (Dominican Republic); Smith, 1937 PDF: 822 (Puerto Rico); Kusnezov, 1956a PDF: 7 (Argentina); Kusnezov, 1960b PDF: 330 (West Patagonia); Snelling & Hunt, 1975 PDF: 63 (Chile); Kempf, 1978a: 33 (Neotropical subfamilies); Baroni Urbani, 1984 PDF: 73 (Neotropical genera); Mackay & Mackay, 1989: 1 (Mexico genera); Cover, in Hölldobler & Wilson, 1990: 69 (Neotropical genera key); Lattke, in Jaffe, 1993: 145 (Neotropical subfamilies & genera); Bolton, 1994: 26, 38, 49, 98, 162 (Dolichoderinae, Ecitoninae, Formicinae, Myrmicinae, Ponerinae genera keys); Fernández, Palacio, MacKay & MacKay, 1996: 353 (Neotropical subfamilies); Fernández & Ospina, in {ref 133005}: 49 (Neotropical genera synopsis); Palacio & Fernández, in {ref 133005}: 233 (Neotropical subfamilies and genera keys)
AFROTROPICAL AND MALAGASY
Forel, 1891c PDF: 8 (Madagascar genera); Arnold, 1915: 6 (South Africa); Arnold, 1916 PDF: 159 (South Africa); Bolton, 1973a PDF: 321 (West Africa subfamilies & genera); Bolton, in Hölldobler & Wilson, 1990: 39 (Afrotropical and Malagasy genera key); Bolton, 1994: 23, 44, 79, 155 (Dolichoderinae, Formicinae, Myrmicinae, Ponerinae genera keys); Yoshimura & Fisher, 2007 PDF: 27 (Malagasy subfamilies, males key); Yoshimura & Fisher, 2009 PDF: 8 (Proceratiinae males); Yoshimura & Fisher, 2011 PDF: 7 (Dolichoderinae males); Yoshimura & Fisher, 2012 PDF: 11 (Amblyoponinae males)
ORIENTAL AND MALESIAN
Forel, 1892k PDF: 219 (India & Sri Lanka); Bingham, 1903 PDF: 1 (India, Sri Lanka & Burma); Wilson & Taylor, 1967b PDF: 10 (Polynesia); Bolton, in Hölldobler & Wilson, 1990: 46 (Oriental and Indo-Australian genera key); Bolton, 1994: 23, 45, 84, 158 (Dolichoderinae, Formicinae, Myrmicinae, Ponerinae genera keys); Wu & Wang, 1995a: 30 (China); Zhou, 2001A PDF: 21 (China, Guangxi subfamilies); Clouse, 2007B: 190 (Micronesia); Framenau & Thomas, 2008 PDF: 52 (Christmas I.); Terayama, 2009 (Taiwan); Eguchi, Bui & Yamane, 2011 PDF: 7 (Vietnam subfamilies key); Pfeiffer, et al. 2011: 32 (ants of Borneo, checklist); General & Alpert, 2012 PDF: 68 (Philippines genera key); Sarnat & Economo, 2012 PDF: 1 (Fiji); Yamane, 2013: 1 (Krakatau Islands, species keys)
AUSTRAL
Forel, 1905d PDF: 353 (New Zealand); Brown, 1958h PDF}: 1 (New Zealand); Brown & Taylor, 1970}: 958 (Australia subfamilies); Greenslade, 1979: 10 (South Australia subfamilies); Taylor, R.W., in Hölldobler & Wilson, 1990: 55 (Australia genera key); Taylor, 1991d: 987 (Australia subfamilies); Bolton, 1994: 24, 47, 91, 159 (Dolichoderinae, Formicinae, Myrmicinae, Ponerinae genera keys); Shattuck, 1999: 21 (Australia subfamilies & genera); Andersen, 2000: 14 (northern Australia subfamilies & genera); Don, 2007: 39 (New Zealand); Heterick, 2009 PDF: 16, 28 (south-western Australia subfamilies & genera); Jennings, Krogmann & Burwell, 2013 PDF: 20 (New Caledonia species checklist)

Taxonomic Treatment (provided by Plazi)

Treatment Citation: Brown, W. L., 1952, Heteroponera Mayr reinstated (Hymenoptera: Formicidae)., Psyche 59, pp. 70-70

Heteroponera Mayr ReinstatedHNS (Hymenoptera: FormicidaeHNS) .

- Heteroponera MayrHNS (1887, Verh. zool.-bot. Ges. Wien, 37: 533) has remained suppressed as a synonym of Acanthoponera MayrHNS for many years. Wheeler (1923, loc. cit.) has shown, however that AcanthoponeraHNS can be split into two groups; one group { AcanthoponeraHNS s. str.) having the tarsal claws with an extra, strong tooth and a basal lobe (or tooth), while the second group (Anacanthoponera Wheeler) has the tarsal claws at most with a single, weak median tooth. In this second group, Wheeler included Heteroponera carinifrons MayrHNS, and since HeteroponeraHNS, with the genotype H. carinifronsHNS, has precedence, it must be reinstated as a good genus. Anacanthoponera Wheeler (1923, Psyche, 30: 176, as a subgenus of AcanthoponeraHNS) is a new synonym of HeteroponeraHNS, since the genotype, Ponera dolo RogerHNS, is congeneric with H. carinifronsHNS.

AcanthoponeraHNS is neotropical, and has well developed propodeal teeth and the petiolar apex produced as a long tooth or spine; Kusnezov (in litt.) finds A. mucronataHNS to have 6, 4 palpal segmentation. HeteroponeraHNS is neotropical and Australasian , and the propodeal teeth and dentiform petiolar apex are absent or feebly developed; H. imbellis EmeryHNS has 3, 3 palpal segmentation (my dissection). Until more species can be critically examined, AcanthoponeraHNS and HeteroponeraHNS should be considered as distinct genera. AcanthoponeraHNS appears to be the most generalized living member of the EctatomminiHNS, and is probably close to the stem from which the proceratiines, the myrmicines and ParaponeraHNS arose. - W. L. Brown, Jr., Museum of Comparative Zoology , Harvard University.

(-1 examples)



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