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Family: Formicidae   Latreille, 1809 

Classification:

Taxonomic History (provided by Barry Bolton, 2020)

Extant: 17 valid subfamilies, 39 valid tribes, 337 valid genera, 13,836 valid species

Fossil: 5 valid subfamilies, 7 valid tribes, 163 valid genera, 787 valid species

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 PDF: 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 PDF: 6; Heer, 1867 PDF: 6 [Formicaria]; Forel, 1870 PDF: 307 [Formicinae]; Forel, 1874 PDF: 19 [Formicariae]; Dalla Torre, 1893 PDF: 1; Forel, 1899b PDF: 1; Ruzsky, 1902d PDF: 5 [Formicarii]; Bingham, 1903 PDF: 1; Ruzsky, 1905b: 91 [Formicariae or Formicidae]; Ashmead, 1905c PDF: 384; all subsequent authors.
Subfamilies (extant) of Formicidae: Agroecomyrmecinae, Amblyoponinae, Aneuretinae, Apomyrminae, Dolichoderinae, Dorylinae, Ectatomminae, Formicinae, Heteroponerinae, Leptanillinae, Martialinae, Myrmeciinae, Myrmicinae, Paraponerinae, Ponerinae, Proceratiinae, Pseudomyrmecinae
Subfamilies (extinct) of Formicidae: †Armaniinae, †Brownimeciinae, †Formiciinae, †Haidomyrmecinae, †Sphecomyrminae
Genera (extinct) incertae sedis in Formicidae: Archaeopone, Baikuris, Calyptites, Camelomecia, Curticorna, Dlusskyidris, Eoaenictites, Eoformica, Fonsecahymen, Klondikia, Kohlsimyrma, Myanmyrma, Petropone, Poneropterus
WORLD SYNOPTIC CLASSIFICATIONS
Forel, 1893b PDF: 161 (Formicidae); Emery, 1895l PDF: 764 (Formicidae); Emery, 1896e PDF: 173 (Formicidae); Ashmead, 1905c PDF: 381 (Formicidae); Wheeler, 1910a PDF: 134 (Formicidae); Emery, 1910b PDF: 3 (Dorylinae); Emery, 1911e PDF: 2 (Ponerinae); Emery, 1913a PDF: 2 (Dolichoderinae); Forel, 1917 PDF: 235 (Formicidae); Emery, 1921c PDF: 3 and Emery, 1922c PDF: 95 (Myrmicinae); Wheeler, 1922: 631 (Formicidae); Emery, 1924f PDF: 207 (Myrmicinae); Emery, 1925d PDF: 2 (Formicinae); Brown, 1973b PDF: 165, 178 (Formicidae); Snelling, 1981: 387 (Formicidae); Wheeler & Wheeler, 1985b PDF: 256 (Formicidae); Dlussky & Fedoseeva, 1988: 77 (Formicidae); Hölldobler & Wilson, 1990: 9 (Formicidae); Bolton, 1994: 12 (Formicidae); Bolton, 2003 PDF: 4 (Formicidae); Engel & Grimaldi, 2005 PDF: 4 (Formicidae); Ward, 2007c PDF: 549 (Formicidae); Borysenko, 2017 PDF: 1
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 PDF: 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 (Formicidae subfamilies, male-based); Cantone, 2017 PDF: 1 (Formicidae male wing venation, key to most genera)
EVOLUTION AND PHYLOGENY OF FORMICIDAE
Wheeler, 1920 PDF: 52; Emery, 1920e: 370; Donisthorpe, 1922b PDF: xlvi; Wheeler, 1923g: 156; Wheeler, 1928h: 124; Stitz, 1935a: 211; Morley, 1938 PDF: 190; Brown & Nutting, 1950 PDF: 113; Bernard, 1951c: 1041; Brown, 1954e PDF: 30; Wilson et al., 1967 PDF: 12; Taylor, 1978a PDF: 982; Rasnitsyn, 1980: 45-46 (foldout page); Lutz, 1986 PDF: 213; Dlussky & Fedoseeva, 1988: 137; Baroni Urbani, 1989 PDF: 137; Brandão et al., 1990 PDF: 201; Hölldobler & Wilson, 1990: 26; Hashimoto, 1991a PDF: 125; Hashimoto, 1991b PDF: 289; Baroni Urbani et al., 1992 PDF: 301; Shattuck, 1992b PDF: 203; Brothers & Carpenter, 1993 PDF: 227; Ward, 1994 PDF: 173; Hashimoto, 1996 PDF: 340; Crozier et al., 1997 PDF: 22; Grimaldi et al., 1997 PDF: 25; Perrault, 1999a PDF: 125; Brothers, 1999: 233; Grimaldi & Agosti, 2000b: 13680; Ohnishi et al., 2004 PDF: 419; Astruc et al., 2004 PDF: 880; Grimaldi & Engel, 2005 PDF: 440; Wilson & Hölldobler, 2005a PDF: 7411; Brady et al., 2006 PDF: 18172; Moreau et al., 2006 PDF: 101; Ouellette et al., 2006 PDF: 359; Rabeling et al., 2008 PDF: 14914; Keller, 2011 PDF: 1; Moreau & Bell, 2013 PDF: 2245; Johnson et al., 2013 PDF: 2060; Peters et al., 2017 10.1016/j.cub.2017.01.027 PDF: 1015; Branstetter et al., 2017 10.1016/j.cub.2017.03.027 PDF: 1021; Branstetter et al., 2017 10.1111/20141-210X.12742 PDF: 774; Economo et al., 2018 10.1038/s41467-018-04218-4 PDF: 1; Borowiec et al., 2019 10.1016/j.ympev.2019.01.024 PDF: 111
PALAEONTOLOGY OF FORMICIDAE
Scudder, 1891 PDF: 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); Brandão et al., 1990 PDF: 201 (oldest fossil record); Carpenter, 1992: 490 (fossil genera review); Bolton, 1995b: 4 (fossil taxa checklist, in catalogue); Grimaldi et al., 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); Perrichot et al., 2008 PDF: 85 (Cretaceous ants review); Dlussky & Rasnitsyn, 2009 PDF: 1024 (European Upper Eocene amber, synopsis of species); LaPolla et al., 2013 PDF (Mesozoic and Cenozoic major ant fossil deposits list, Cretaceous and Cenozoic ant fossils bibliographical review); Dlussky et al., 2015 PDF (Late Eocene of Sikhote-Alin fossils); Barden & Grimaldi, 2016 10.1016/j.cub.2015.12.060 PDF (Cretaceous stem-group ants and phylogeny); Perrichot et al., 2016 10.1016/j.cub.2016.03.075 PDF: 1468 (Cretaceous ants morphology and ecology); Barden, 2017 PDF (literature review); Zheng et al., 2018 10.1038/s41467-018-05650-2 PDF (subfamily records from Late Cretaceous burmite); Barden & Engel, 2019 10.1007/978-3-319-90306-4_45-1 PDF (broad-perspective review of fossil record); Barden et al., 2020 10.1016/j.cub.2020.06.106 PDF (morphospace)
OTHER GENERAL REFERENCES
Bolton, 1995a PDF: 1038 (census of extant taxa); Ward et al., 1996: 55 (bibliography of systematics); Brown, 2000a PDF: 46 (overview of genera); Brandão, 2000 PDF: 172 (list of major ant collections); Bolton, 2003 PDF: 15, 77 (diagnosis, synopsis); Bolton, 2007a PDF: 51 (method for taxonomic revisions); Taylor, 2007 PDF: 580 (eusociality in ants)
REGIONAL CATALOGUES AND CHECKLISTS
AFROTROPICAL AND MALAGASY
Wheeler, 1922: 727, 1005.
AUSTRALASIAN
Taylor & Brown, 1985: 5; Taylor, 1987a PDF: 1; Taylor, 1987b: 1 (supplement to previous)
NEARCTIC
Smith, 1951c PDF: 778 (North America, north of Mexico, catalogue); Smith, 1958c PDF: 108 (North America, north of Mexico, catalogue, first supplement); Smith, 1967a PDF: 343 (North America, north of Mexico, catalogue, second supplement); Smith, 1979: 1323 (North America, north of Mexico, catalogue, volume 2); Castaño-Meneses et al., 2015 PDF (Mexico partial checklist).
NEOTROPICAL
Kempf, 1972b PDF: 3; Brandão, 1991 PDF: 319 (supplement to previous); Castaño-Meneses et al., 2015 PDF (Mexico partial checklist); Salazar & Donoso, 2013 PDF: 151 (partial checklist)
PALEARCTIC
Karaman, 2011b PDF (Montenegro catalogue); Borowiec, 2014 PDF (Europe, Mediterranean Basin and adjacent regions)
REGIONAL AND NATIONAL FAUNAS WITH 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, 2012a doi:10.1371/journal.pone.0033325 PDF: 11 (Amblyoponinae males); Fisher & Bolton, 2016: 60 (Afrotropical and Malagasy genera key, workers).
AUSTRALASIAN
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 et al., 2013 PDF: 20 (New Caledonia species checklist)
INDOMALAYAN
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 PDF: 190 (Micronesia); Framenau & Thomas, 2008 PDF: 52 (Christmas I.); Terayama, 2009 PDF: 81 (Taiwan); Eguchi et al., 2011 PDF: 7 (Vietnam subfamilies key); Pfeiffer et al., 2011 PDF: 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)
NEARCTIC
Provancher, 1883 PDF: 596 (Canada); Provancher, 1887: 225 (Canada); Cresson, 1887 PDF: 93 (U.S.A.); Wheeler, 1910a PDF: 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 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 PDF: 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 et al., 1996: 353 (Neotropical subfamilies); Fernández & Ospina, 2003: 49 (Neotropical genera synopsis); Palacio & Fernández, 2003: 233 (Neotropical subfamilies and genera keys); Baccaro et al., 2015 10.5281/zenodo.32912 PDF: (Brazil key to genera); Boudinot, 2019 PDF: 487 (Neotropical subfamilies and genera, males key); Fernández et al., 2019c PDF: 459 (Neotropical subfamilies and genera diagnoses).
PALAEARCTIC
Mayr, 1855 PDF: 299 (Austria); Mayr, 1861 PDF: 25 (Europe); Forel, 1874 PDF: 19 (Switzerland); Saunders, 1880 PDF: 202 (Britain); André, 1882a PDF: 125 (Europe & Algeria); Nasonov, 1889: 50 (Russia); Lameere, 1892: 62 (Belgium); Saunders, 1896 PDF: 18 (Britain); Ruzsky, 1905b: 99 (Russian Empire); Wasmann, 1906 PDF: 7 (Luxemburg); Emery, 1908a PDF (Palaearctic); Bondroit, 1910 PDF: 480 (Belgium); Stitz, 1914 PDF: 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, 1926b: 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 PDF: 69 (Central Europe); Novák & Sadil, 1941 PDF: 69 (Central Europe); Ceballos, 1943: 313 (Spain);Holgersen, 1942b PDF: 3 (Northern Norway); Holgersen, 1943c PDF: 166 and Holgersen, 1944a PDF: 198 (Norway); Van Boven, 1947b PDF: 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); Van 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 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 et al., 1990 PDF: 99 (Turkmenistan); Kupyanskaya, 1990a: 83 (Far Eastern Russia); Bolton, in Hölldobler & Wilson, 1990: 34 (Palaearctic genera key); Atanassov & 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 et al., 2002 PDF: 133 (Poland); Aktaç & Radchenko, 2002: 53 (Turkey genera); Yoshimura & Onoyama, 2002b PDF: 424 (Japan genera, males key); Imai et al., 2003 PDF: 8 (Japan); Radchenko, 2005b PDF: 183 (North Korea); Seifert, 2007: 107 (North and Central Europe); Boer, 2010: 8 (Benelux); Lapeva-Gjonova et al., 2010 PDF: 1 (Bulgaria species catalogue); Csosz et al., 2011 PDF: 56 (Hungary species catalogue); Czechowski et al., 2012: 333 (Poland); Douwes et al., 2012: 1 (Sweden).
ANATOMY AND MORPHOLOGY OF FORMICIDAE
GENERAL
Mayr, 1861 PDF: 1 (general morphology); Forel, 1874 PDF: 1 (general anatomy); Lubbock, 1877: 121; Lubbock, 1879: 738; André, 1882 PDF: 5; Janet, 1902: 25 (general anatomy); Janet, 1904 PDF: 3 (general anatomy); Janet, 1905 PDF: 1 (Lasius cranial anatomy); Janet, 1909 1 (general morphology); Wheeler, 1910a PDF: 13 (general morphology, anatomy); Janet, 1923 PDF: 5 (anatomical drawings synopsis); Pavan & Ronchetti, 1955 PDF: 3 (Linepithema humile, anatomy); Whelden, 1957: 271 (Rhytidoponera general anatomy, see also Whelden, 1958a: 80); Whelden, 1958b PDF: 1 (Stigmatomma general anatomy); Whelden, 1960: 93 (Rhytidoponera metallica); Schneider, 1964 PDF: 103 (antennal anatomy review); Bernard, 1967a PDF: 5; Hölldobler & Wilson, 1990: 5 (glossary, glandular anatomy); Bolton, 1994: 191 (morphology glossary); Serna & MacKay, 2010 PDF: 1 (Procryptocerus morphology, see also Serna et al., 2011 PDF: 67); Keller, 2011 PDF: 1 (phylogenetic analysis of morphology); Delsinne et al., 2019 PDF: 387 (morphology glossary)
HEAD (SKELETON AND MUSCULATURE)
Janet, 1899b: 295 (Myrmica tentorium); Bugnion, 1930: 85 (mouthparts); Gotwald, 1969b PDF: 1 (mouthparts and gaster); Masson, 1972b: 343 (Camponotus antennal articulations); Gronenberg & Ehmer, 1996: 153 (Anochetus mandibular muscles); Ehmer & Gronenberg, 1997 PDF: 287 (antennal musculature, physiology); Paul & Gronenberg, 1999 PDF: 797 (cranial muscle attachments, physiology); Paul, 2001 PDF: 7 (mandible musculature, movements); Paul et al., 2002: 39 (cranial, mouthpart musculature); Schofield et al., 2002 PDF: 579 (mandibular tooth hardness); Muscedere et al., 2011 10.1007/s00114-011-0828-6 PDF: 783 (Pheidole cranial muscle maturation); Larabee & Suarez, 2014 PDF: 25 (trap-jaw evolution); Raderschall et al., 2016 10.1242/jeb.134049 PDF: 1449 (head roll control); Lillico-Ouachour et al., 2018 PDF: 383 (Pheidole intercaste cranial anatomy); Khalife et al., 2018 10.1186/s12983-018-0277-6 PDF: 30 (Melissotarsus cranial and leg anatomy); Richter et al., 2019 10.1016/j.asd.2019-02-002 PDF: 26 (cranial and mouthpart anatomy); Kubota et al., 2019 10.1038/s41598-019-43175-w PDF: 1 (Lasius tentorium); Barlow et al., 2019 10.1111/azo.12291 PDF 1 (mandible histology); Richter et al., 2020 10.26049/ASP78-1-2020-06 PDF: 133 (cranium anatomy: Formicinae, Ponerinae).
EYES
Bernstein & Finn, 1971 PDF: 708 (Formica rufa intranidal ommatidium variation); Menzi, 1986 PDF: 11 (nocturnal/diurnal adaptations); Greiner et al., 2007 10.1016/j.cub.2007.08.015 PDF: 879 (Myrmecia photoreceptor variation); Nakanishi et al., 2009 10.1007/s00441-009-0863-1 PDF: e76015 (Polyrhachis sakalava compound eye light-regime adaptations); Narendra et al., 2011 10.1098/rspb.2010.1378 PDF: 1141 (Myrmecia interspecific and inter-caste visual adaptations); Schwarz et al., 2011 10.1016/j.asd.2010.10.003 PDF: 128 (Melophorus bagoti visual system); Narendra et al., 2016 10.1038/srep22331 PDF: 22331 (Camponotus consobrinus inter-caste eye anatomy); Ramirez-Esquivel et al., 2017 10.1016/j.asd.2017.03.005 PDF: 552 (Temnothorax rugatulus setation, compound eye anatomy);
MESOSOMA (SKELETON AND MUSCULATURE)
Lubbock, 1881 PDF: 141 (Lasius w.q. mesosomal skeletomusculature); Janet, 1898f: 393 (Myrmica q. mesosoma); Janet, 1899a: 249 (Myrmica alate mesosoma, flight mechanism); Janet, 1906: 522 (metanotal diaphragm); Emery, 1900a PDF: 103 (mesosomal exoskeleton); Janet, 1907: 1 (Lasius histolyzed flight musculature); Snodgrass, 1910: 81 (mesosomal exoskeleton); Tulloch, 1935 PDF: 93 (mesosomal exoskeleton); Reid, 1941 PDF: 418 (worker thoracic exoskeleton); Gregg, 1953c PDF: 328 (promesonotal suture motility); Markl, 1966: 138 (Formica mesosoma skeletomusculature, innervation); Matsuda, 1970: 353 (review of mesosomal anatomy); Jones et al., 1978: 603 (Solenopsis flight musculature histolysis); Saini et al., 1982: (Camponotus w.q. mesosomal skeletomusculature); Hashimoto, 1991b PDF: 289 (tibial spurs); Menzel & Tautz, 1994 PDF: 735 (subgenual organ); Perrault, 1999b: 125 (thoracic anatomy); Nguyen et al., 2014 10.1016/j.biomech.2013.10.053 PDF: 497 (Formica neck joint); Boudinot, 2015 PDF: 1 (family diagnosis, mesosoma, coxae); Khalife et al., 2018 10.1186/s12983-018-0277-6 PDF: 30 (Melissotarsus cranial and leg anatomy); Liu et al., 2019 10.26049/ASP77-1-2019-01 PDF: 1 (Myrmecia mesosomal anatomy, and apomorphies for family).
LEGS
Janet, 1895a: 691 (Myrmica strigil); Gennerich, 1922 PDF: 26 (strigil); Dashman, 1953 PDF: 56 (general pretarsal terminology); Freeland et al., 1982 PDF: 257 (arolium presence across Formicidae); Schönitzer & Lawitzky, 1986: 189 (antenna cleaning apparatus ultrastructure); Schönitzer & Lawitzky, 1987: (antenna cleaning apparatus Aculeata comparison); Francoeur & Loiselle, 1988a PDF: 333 (strigil, evolution); Basibuyuk & Quicke, 1994 PDF: 205 (Dorylus strigil); Basibuyuk & Quicke, 1995: 171, 176 (antenna cleaning apparatus); Basibuyuk & Quickle, 1999 10.1111/j.1096-3642.1999.tb00597.x PDF: 349 (grooming, tarsus, tibial spur morphology); Basibuyuk et al., 2000 10.1603/0013-8746(2000)093[0625:MASOTO]2.0.CO;2 PDF: 625 (manubrium, manubrial sensillae); Federle et al., 2000 PDF: 505 (comparative attachment capacities); Federle et al., 2001 10.1073/pnas.111139298 PDF: 6215 (pretarsal anatomy, functional morphology); Orivel et al., 2001: 449 (Ponerinae pretarsal morphology); Federle et al., 2002 PDF: 1100 (Oecophylla smaragdina aroliar wet adhesion); Schulmeister, 2003 PDF: 156 (Cataglyphis tarsal plantulae); Böröczky et al., 2013 10.1073/pnas.1212466110 PDF: 3616 (Camponotus pennsylvanicus grooming); Federle et al., 2004 10.1242/jeb.00716 PDF: 67 (Oecophylla smaragdina arolium biomechanics); Federle & Endlein, 2004 10.1016/j.asd.2003.11.001 PDF: 67 (Oecophylla smaragdina arolium control); Endlein & Federle, 2015 10.1371/journal.pone.0141269 PDF: e0141269 (Oecophylla smaragdina arolium, distal tarsal brush); Hackmann et al., 2015 10.1098/rsos.150129 PDF: (functional morphology antenna cleaner); Billen et al., 2017 10.1016/j.sjbs.2016.06.007 PDF: 24 (Brachyponera sennaarensis, Daceton armigerum pretarsus, climbing ability); Beutel et al., 2020 10.1002/jmor.21133 PDF: 1 (Formica tarsal anatomy, grooming and attachment structures).
WINGS
Emery, 1913d PDF: 577 (wing venation); Brown & Nutting, 1950 PDF: 113 (wing venation); Ogata, 1984: 4; Ogata, 1991b PDF: 64 (forewing venation); Basibuyuk & Quicke, 1997: 1563 (hamuli); Perfilieva, 2010 PDF: 965 (wing venation); Cantone & Von Zuben, 2019 10.1155/2019/7929717 PDF: 1 (hindwing venation); Cantone, 2019 PDF: 1 (wing venation).
METASOMA (SKELETON AND MUSCULATURE)
Janet, 1893: 159 (stridulum); Janet, 1894a: 109 (Myrmica stridulum); Janet, 1894e: 591 (Myrmica metasoma); Janet, 1894f: 185 Myrmica petiole); Janet, 1902: 1 (Myrmica metasoma}; Short, 1959 PDF: 194 (Dorylus anterior metasomal skeletomusculature); Gotwald, 1969b PDF: 1 (mouthparts and gaster); Markl, 1973: 258 (stridulum); Bolton, 1990a PDF: (abdominal morphology); Hashimoto, 1996 PDF: 340 (anterior metasomal skeletomusculature); Perrault, 2005 https://doi.org/10.1080/00379271.2004.10697428 PDF: 291 (anterior metasomal anatomy).
VENOM APPARATUS
Janet, 1898c: 638 (sting function, glands); Janet, 1898e: 1 (Myrmica sting function, gland); Kugler, 1978a: 413 (sting morphology, Myrmicinae); Kugler, 1979c: 117 (sting evolutionary morphology [for treatment of other groups, search for C. Kugler references]); Hermann & Chao, 1983: 329 (sting comparative anatomy, emphasis on furcula).
GENITALIA
Kempf, 1956 PDF: 101 (male genitalia); Ogata, 1983a: 2 (male genitalia), Ogata, 1991b PDF: 61 (male genitalia); Boudinot, 2013 PDF: 29 (male genitalia).
SKELETON AND MUSCULATURE: MISCELLANEOUS
Janet, 1895b: 610 (muscles); Janet, 1895c: 1 (articulatory membrane, tendon, muscle); Janet, 1897b: 1 (Myrmica post-cephalic segmentation, musculature); Janet, 1898a: 435 (post-cephalic segmentation, membranes); Harris, 1979 10.5281/zenodo.26215 PDF: 1 (surface sculpture); Gronenberg & Ehmer, 1995: 68 (tubular muscle fibers).
ALIMENTARY CANAL
Forel, 1878c PDF: 339 (proventriculus); Bugnion, 1926 33 (anterior alimentary canal, see also: Bugnion, 1927a: 59; Bugnion, 1927c: 105; Bugnion, 1929: 55; Bugnion, 1930: 85); Haskins & Whelden, 1954: 33 (Myrmecia food transfer); Eisner, 1957 PDF: 439 (proventriculus); Brown, 1988h: 17 (malpighian tubule numbers); Caetano, 1988: 129 (digestive tract anatomy); Casadei-Ferreira et al., 2020 10.1016/j.asd.2020.100977 PDF: (thoracic crop, Pheidole)
TRACHEAL SYSTEM
Keister, 1963: 336 (Camponotus tracheal system anatomy); Délye, 1965: 285 (spiracle anatomy, function); Fedoseeva, 2017 PDF: 49 (metathoracic spiracles).
GLANDULAR SYSTEM
Janet, 1894c: 989 (general); Janet, 1894d: 45 (Formica pharyngeal glands); Janet, 1898b: 1168 (Myrmicinae); Janet, 1898d: 1 ({tMyrmica}); Whelden, 1963a: 13 (Eciton w., q. comparison, see also: Whelden, 1963b: 90); Hölldobler & Engel, 1979 PDF: 285 and Hölldobler & Engel-Siegel, 1982: 113 (tergal and sternal glands); Hölldobler & Engel-Siegel, 1985 PDF: 201 (metapleural gland); Billen, 1986b: 165 (Dufour's gland); Hölldobler & Wilson, 1990: 5, 229 (anatomy, exocrine glands); Billen, 1993: 4 (exocrine glands); Hölldobler et al., 1996: 157 (metatibial gland); Schönitzer et al., 1996: 107 (Messor rufitarsis antenna cleaner gland); Wenseleers et al., 1998: 121 (cloacal gland); Jensen & Borgesen, 2000: 171 (Monomorium fat body); Gusmão et al., 2001 PDF: 31 (Atta ultramorphology metapleural gland); Lommelen et al., 2003 10.1016/S1467-8039(03)00052-5 PDF: 209 (Neoponera labial gland development); Billen, 2019 PDF: 165 (synopsis of exocrine glands).
NEUROANATOMY
Janet, 1899b: 295 (Myrmica cranial nerves, corpora allata); Pietschker, 1910 PDF: 1 (brain structure); Thompson, 1913: 515 (mushroom body comparative anatomy); Brun, 1952 http://doi.org/10.5169/seals-401148 PDF: 74 (Tetramorium schneideri neuroanatomy); Steiger, 1967 PDF: 511 (mushroom body, "corpus pedunculatum"); Goll, 1967: 143 (Formica neuroanatomy); Masson, 1969: 171 (Camponotus deutocerebrum); Bressac & Rabitsch, 1969: 135 (Aphaenogaster senilis cranial nervous system); Masson, 1972a: 31 (deutocerebrum); Masson, 1972b: 343 (Camponotus, see also Masson et al., 1972); Schoeters & Billen, 1999 PDF: 139 (terminal abdominal ganglia); Julian & Gronenberg, 2002: 152 (comparison worker, queens); Paul & Gronenberg, 2002 PDF: 255 (mandible closer muscle motor control); Seid et al., 2005 10.1002/cne.20545 PDF: 269 (Pheidole age-related change mushroom bodies); Kühn-Bühlmann & Wehner, 2006 10.1002/neu.20235 PDF: 511 (Cataglyphis mushroom body variation); Gronenberg, 2008 PDF: 25 (family-level neuroanatomy review); Wehner, 2009 PDF: 85 (navigational architecture); Kelber et al., 2009 PDF: 273 (Attini/Attina antennal lobe anatomy); Stieb et al., 2010 10.1002/dneu.20785 PDF: 408 (Cataglyphis mushroom body variation); Bressan et al., 2015 10.3389/fnana.2014.00166 PDF 1 (Cardiocondyla brain map); Ramirez-Esquivel et al., 2017 10.1016/j.asd.2017.03.005 PDF: 552 (Temnothorax rugatulus setation, compound eye anatomy); Habenstein et al., 2020 10.1002/cne.24934 PDF 1 (Cataglyphis brain map)
GAMETES
Blochmann, 1886 PDF: 143 (oogenesis); Blochmann, 1884 PDF: 243 (oogenesis); Lams, 1908: 528 (Camponotus herculeanus spermatogenesis); Buchner, 1918: 1 (oogenesis); Wheeler et al., 1990: 343 (sperm comparative ultrastructure); Moya et al., 2007 PDF: 181 (Pseudomyrmex sperm ultrastructure).
KARYOLOGY
Whelden & Haskins, 1954: 579 (family-level comparisons); Crozier, 1970a PDF: 109 (karyology); Imai et al., 1984 PDF: 5 (karyology); Imai et al., 1984 PDF: 66 (karyology); Imai et al., 1985 PDF: 46 (karyology).
LARVAE
Wheeler, 1918c PDF: 293 (larvae); Athias-Henriot, 1947 PDF: 247 (larval anatomy); Wheeler & Wheeler, 1976b PDF: 45 (larvae); Petralia & Vinson, 1979 PDF: 472 (Solenopsis invicta development egg, larva); Petralia & Vinson, 1980: 89 (Solenopsis invicta fourth instar internal anatomy); Petralia & Vinson, 1980 PDF: 375 (larval ventral region); Wheeler & Wheeler, 1986e PDF: 684 (larvae); Petralia & Haut, 1986: 16 ({TCamponotus} larval labial gland)
SENSORY STRUCTURE AND NEUROANATOMY
Janet, 1894b: 814 (antennal nerves, chordotonal organ); Debaisieux, 1934: 139 (scolopidial organs)}; Jaisson, 1969 PDF: 279 (Myrmicinae antennal sensilla distribution); Jaisson, 1970: 1192 (Atta laevigata sensilla sexual dimorphism); Masson, 1970: 346 (receptor anatomy, function); Masson & Gabouriaut, 1973: 39 (Camponotus Johnston's organ); Masson & Friggi, 1971 PDF: 618 (Camponotus sensilla coeloconica); Dumpert, 1972b PDF: 95 (Lasius fuliginosus antennal sensilla); Walther, 1979a PDF: 30 (antennal sensilla, comparison to other Aculeata); Walther, 1979b: 313 (Formica rufa antennal sensilla); Walther, 1981 PDF: 146 (cuticular sense organs, phylogeny); Walther, 1983 PDF: 373 (antennal sensilla comparative morphology); Walther, 1985 PDF: 173 (antennal sensilla review); Hashimoto, 1990 PDF: 491 (antennal sensilla, comparison to other Aculeata); Hashimoto, 1991a PDF: 125 (antennal and labial sensilla); Walther, 1994: 360 (Nothomyrmecia macrops antennal sensilla); Gronenberg & Hölldobler, 1999 PDF: 229 (optical and antennal neuroanatomy); Kleineidam et al., 2000 PDF: 43 (Atta sexdens sensillum ampullaceum ultrastructure, physiology); Renthal et al., 2003 10.1016/S0968-4238(03)00050-7 PDF: 405 (Solenopsis invicta antennal sensilla distribution); Marques-Silva et al., 2006 PDF: 885 (Dinoponera lucida antennal sensilla, secretory glands); Nakanishi et al., 2009 10.1007/s00441-009-0863-1 PDF: 79 (Camponotus japonicus sexual dimorphism antennal sensilla); Barsagade et al., 2010 PDF: 53 (Camponotus compressus: mouthpart sensilla); Mysore et al., 2010 10.1016/j.asd.2010.04.003 PDF: 310 (Camponotus compressus antennal sensilla, glomeruli, anatomy, development); Siddiqui et al., 2010 PDF: 6956 (Brachyponera sennaarensis antennal sensilla ultrastructure); Barsagade et al., 2013 10.20362/am.005012 PDF: 113 (Camponotus compressus: antennal sensilla); Gill et al., 2013 10.1086/673712 PDF: 834 (antennal sensilla density-dependent function); Ramirez-Esquivel et al., 2014 10.1016/j.asd.2014.07.004 PDF: 543 (Myrmecia pyriformis antennal sensilla); Barsagade et al., 2017 PDF: 99 (Leptogenys dentilobis: mouthpart sensilla); Gathalkar & Barsagade, 2018 10.7324/JABB.2017.60101 PDF: 1 (Myrmicaria brunnea head, antennal sensilla); Barsagade et al., 2018 PDF: 848 (Tetraponera rufonigra leg sensilla); Barsagade et al., 2019 10.7324/JABB.2019.70201 PDF: 1 (Leptogenys chinensis, antennal sensilla)
// Distribution

Distribution:

  Geographic regions (According to curated Geolocale/Taxon lists):
    Africa: Algeria, Angola, Benin, Botswana, Burkina Faso, Burundi, Cabo Verde, Cameroon, Cape Verde, Central African Republic, Chad, Chagos Islands, Comoros, Congo, Democratic Republic of Congo, Djibouti, Egypt, Equatorial Guinea, Eritrea, Ethiopia, Europa Island, Gabon, Gambia, Ghana, Grande Glorieuse, Guinea, Ile du Lys, Ivory Coast, Juan de Nova Island, Kenya, Lesotho, Liberia, Libya, Macaronesia, Madagascar, Madeira, Malawi, Mali, Mauritania, Mauritius, Mayotte, Morocco, Mozambique, Namibia, Niger, Nigeria, Reunion, Rwanda, Saint Helena, Sao Tome and Principe, Senegal, Seychelles, Sierra Leone, Somalia, South Africa, South Sudan, Sudan, Swaziland, Tanzania, Togo, Tromelin Island, Tunisia, Uganda, Zambia, Zimbabwe
    Americas: Antigua and Barbuda, Argentina, Aruba, Bahamas, Barbados, Belize, Bermuda, Bolivia, Bonaire, Sint Eustatius and Saba, Brazil, British Virgin Islands, Canada, Cayman Islands, Chile, Colombia, Costa Rica, Cuba, Curaçao, Dominica, Dominican Republic, Ecuador, El Salvador, French Guiana, Galapagos Islands, Grenada, Guadeloupe, Guatemala, Guyana, Haiti, Honduras, Jamaica, Martinique, Mexico, Montserrat, Nicaragua, Panama, Paraguay, Peru, Puerto Rico, Saint Kitts and Nevis, Saint Lucia, Saint Vincent and the Grenadines, Suriname, Trinidad and Tobago, Turks and Caicos Islands, United States, United States Virgin Islands, Uruguay, Venezuela
    Antarctica_region: Kerguelen Islands
    Asia: Afghanistan, Armenia, Azerbaijan, Bangladesh, Bhutan, Borneo, Brunei, Cambodia, China, Christmas Island, Cyprus, Georgia, India, Indonesia, Iran, Iraq, Israel, Japan, Jordan, Kazakhstan, Kuwait, Kyrgyzstan, Laos, Lebanon, Malaysia, Mongolia, Myanmar, Nepal, North Korea, Oman, Pakistan, Palestine, Philippines, Qatar, Saudi Arabia, Singapore, South Korea, Sri Lanka, Syria, Taiwan, Tajikistan, Thailand, Timor-Leste, Turkey, Turkmenistan, United Arab Emirates, Uzbekistan, Vietnam, Yemen
    Europe: Albania, Andorra, Austria, Belarus, Belgium, Bosnia and Herzegovina, Bulgaria, Croatia, Czech Republic, Denmark, Estonia, Finland, France, Germany, Gibraltar, Greece, Hungary, Ireland, Italy, Latvia, Liechtenstein, Lithuania, Luxembourg, Macedonia, Malta, Moldova, Monaco, Montenegro, Netherlands, Norway, Poland, Portugal, Romania, Russia, Serbia, Slovakia, Slovenia, Spain, Sweden, Switzerland, Ukraine, United Kingdom
    Oceania: American Samoa, Australia, Cook Islands, Fiji, French Polynesia, Guam, Hawaii, Kiribati, Marshall Islands, Micronesia, Nauru, New Caledonia, New Zealand, Niue, Norfolk Island, Northern Mariana Islands, Palau, Palmyra Atoll, Papua New Guinea, Samoa, Solomon Islands, Tonga, Tuvalu, Vanuatu, Wallis and Futuna
  Biogeographic regions (According to curated Bioregion/Taxon lists):
    Afrotropical, Antarctica, Australasia, Indomalaya, Malagasy, Nearctic, Neotropical, Oceania, Palearctic

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.



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