Taxonomic history
[Misspelled as herculanea by Latreille, 1809 PDF: 126, Smith, 1858a PDF: 10; misspelled as hepculeanus by Kuznetsov-Ugamsky, 1929a PDF: 17; misspelled as helcureanus by Teranishi, 1940: 71, and subsequent pages.].Lepeletier de Saint-Fargeau, 1835 PDF: 209 (w.m.); Wheeler & Wheeler, 1953e PDF: 185 (l.).Combination in Camponotus: Mayr, 1861 PDF: 36 (in key).Combination in Camponotus (Camponotus): Forel, 1914a PDF: 259.Status as species: Linnaeus, 1761 PDF: 426; Scopoli, 1763 PDF: 312; Linnaeus, 1767 PDF: 962; Fabricius, 1775 PDF: 391; Fabricius, 1782: 488; Fabricius, 1787 PDF: 307; Gmelin, 1790 PDF: 2797; Christ, 1791 PDF: 514; Olivier, 1792: 491; Fabricius, 1793 PDF: 349; Latreille, 1798 PDF: 33; Fabricius, 1804 PDF: 395; Jurine, 1807 PDF: 272; Gravenhorst, 1807 PDF: 286; Latreille, 1809 PDF: 126; Leach, 1815: 147; Latreille, 1817a: 96; Billberg, 1820: 104; Brullé, 1833 PDF: 326; Losana, 1834 PDF: 309; Lepeletier de Saint-Fargeau, 1835 PDF: 209; Zetterstedt, 1838: 448; Schilling, 1839 PDF: 52; Nylander, 1846a PDF: 894; Nylander, 1846b PDF: 1044; Foerster, 1850a: 9; Schenck, 1852 PDF: 123; Mayr, 1855 PDF: 308 (redescription); Nylander, 1856b PDF: 56; Gredler, 1858 PDF: 3; Smith, 1858a PDF: 10, 53; Roger, 1859 PDF: 229; Duméril, 1860: 929; Mayr, 1861 PDF: 36 (in key); Meinert, 1861: 309; Mayr, 1863a PDF: 399; Roger, 1863b PDF: 1; Cresson, 1865b PDF: 426; Dours, 1873 PDF: 164; Forel, 1874 PDF: 39 (in key); André, 1874b: 176 (in key); Emery, 1878: 44; Emery & Forel, 1879 PDF: 447; Forel, 1879a PDF: 56; Provancher, 1881b PDF: 354; André, 1882b PDF: 142 (in key); Provancher, 1883 PDF: 597; Mayr, 1886d PDF: 419; Cresson, 1887 PDF: 255; Provancher, 1887: 228 (in key); Nasonov, 1889: 9; Forel, 1892j PDF: 306; Dalla Torre, 1893 PDF: 233; Emery, 1893k PDF: 674; Ruzsky, 1896 PDF: 67; Emery, 1896j PDF: 372 (in list); Ruzsky, 1902d PDF: 5; Forel, 1902k PDF: 699; Ruzsky, 1903c PDF: 205; Forel, 1904c PDF: 381; Ruzsky, 1905b: 214; Wasmann, 1906 PDF: 111; Forel, 1907h PDF: 19; Emery, 1908a PDF: 182; Wheeler, 1908i PDF: 625; Bondroit, 1910 PDF: 487; Santschi, 1911d PDF: 7; Karavaiev, 1912b PDF: 592; Forel, 1914a PDF: 266; Stitz, 1914 PDF: 95; Bruch, 1914 PDF: 228; Ruzsky, 1914b PDF: 100; Bruch, 1915 PDF: 535; Forel, 1915d: 68 (in key); Ruzsky, 1915b: 4; Donisthorpe, 1915f: 347; Emery, 1916a PDF: 225; Escherich, 1917: 330 (in key); Bondroit, 1918 PDF: 70; Nadig, 1918 PDF: 339; Menozzi, 1922d PDF: 142; Soudek, 1922b PDF: 95; Kuznetsov-Ugamsky, 1923b PDF: 241; Müller, 1923b PDF: 159; Wheeler, 1923c PDF: 5; Vashkevich, 1924b PDF: 146; Emery, 1925d PDF: 72; Kiseleva, 1925 PDF: 73; Ruzsky, 1925a PDF: 287; Essig, 1926 PDF: 868; Karavaiev, 1926e PDF: 191; Ruzsky, 1926 PDF: 108; Stärcke, 1926a PDF: 119 (in key); Karavaiev, 1927a PDF: 295; Donisthorpe, 1927c: 400; Menozzi, 1927b PDF: 92; Kuznetsov-Ugamsky, 1929b PDF: 36; Karavaiev, 1930b PDF: 147; Arnol'di, 1933a: 602 (in key); Stitz, 1934: 4; Grandi, 1935 PDF: 102; Karavaiev, 1936: 177 (redescription); Ruzsky, 1936 PDF: 89; Kôno & Sugihara, 1939 PDF: 10; Teranishi, 1940: 71; Novák & Sadil, 1941 PDF: 110 (in key); Holgersen, 1942b PDF: 10; Holgersen, 1943c PDF: 172 (in key); Holgersen, 1944a PDF: 179; Ruzsky, 1946 PDF: 69; Creighton, 1950a PDF: 366; Donisthorpe, 1950e PDF: 1066; Röszler, 1950 PDF: 210; Yasumatsu & Brown, 1951 PDF: 30; Chapman & Capco, 1951 PDF: 220; Consani & Zangheri, 1952 PDF: 43; Cole, 1954f PDF: 271; Ceballos, 1956: 312; Smith, 1958c PDF: 142; Pisarski, 1961 PDF: 153; Dlussky, 1962 PDF: 181; Collingwood, 1962c PDF: 220; Bernard, 1967a PDF: 340 (redescription); Arnol'di, 1967 PDF: 1819 (redescription); Smith, 1967a PDF: 366; Kutter, 1968b: 60; Collingwood & Yarrow, 1969 PDF: 81; Dlussky & Pisarski, 1970 PDF: 86; Baroni Urbani, 1971c PDF: 175; Collingwood, 1971 PDF: 163; Banert & Pisarski, 1972 PDF: 352; Tarbinsky, 1976 PDF: 148 (redescription); Aktaç, 1977 PDF: 125; Van Boven, 1977 PDF: 131; Francoeur, 1977b PDF: 207; Kutter, 1977c: 204; Yensen et al., 1977 PDF: 183; Collingwood, 1978 PDF: 91 (in key); Collingwood, 1979 PDF: 90; Smith, 1979: 1426; Allred, 1982: 454; Wheeler & Wheeler, 1986g PDF: 60; Agosti & Collingwood, 1987a PDF: 58; Agosti & Collingwood, 1987b PDF: 283 (in key); Nilsson & Douwes, 1987: 68; MacKay et al., 1988: 105; Wang et al., 1989a: 224 (in key); Morisita et al., 1991: 41; Wang, 1992: 681; Atanassov & Dlussky, 1992: 210; Arakelian, 1994 PDF: 85; Radchenko, 1994b: 116 (in key); Wang & Wu, 1994 PDF: 31 (in key); Wheeler et al., 1994 PDF: 305; Bolton, 1995b: 103; Douwes, 1995: 92; Poldi et al., 1995: 7; Wu & Wang, 1995a: 182; Radchenko, 1996e: 1202 (in key); Espadaler, 1997g PDF: 27; Radchenko, 1997b PDF: 555; Gallé et al., 1998: 216; Czechowski et al., 2002 PDF: 95; Mackay & Mackay, 2002 PDF: 290; Zhang & Zheng, 2002 PDF: 218; Coovert, 2005 PDF: 166; Hansen & Klotz, 2005: 82; Radchenko, 2005b PDF: 158; Csosz & Markó, 2005 PDF: 228; Karaman & Karaman, 2005 PDF: 58; Bračko, 2006 PDF: 145; Markó et al., 2006 PDF: 66; Petrov, 2006 PDF: 109 (in key); Schultz et al., 2006 PDF: 203; Bračko, 2007 PDF: 19; Seifert, 2007: 262; Werner & Wiezik, 2007 PDF: 143; Zryanin & Zryanina, 2007 PDF: 232; Gratiashvili & Barjadze, 2008 PDF: 131; Casevitz-Weulersse & Galkowski, 2009 PDF: 479; Lapeva-Gjonova et al., 2010 PDF: 43; Boer, 2010: 18; Csosz et al., 2011 PDF: 58; Karaman, 2011b PDF: 70; Legakis, 2011 PDF: 30; Ran & Zhou, 2011: 67; Borowiec & Salata, 2012 PDF: 476; Czechowski et al., 2012: 240; Ellison et al., 2012: 121; Guénard & Dunn, 2012 PDF: 28; Kiran & Karaman, 2012 PDF: 7; Karaman & Aktaç, 2013 PDF: 52 (in key); Borowiec, 2014 PDF: 32; Lebas et al., 2016: 126; Radchenko, 2016: 328; Salata & Borowiec, 2018c 10.5281/zenodo.2199191 PDF: 43; Schär et al., 2018 10.1111/jbi.13380 PDF: 6; Seifert, 2018: 256; Mackay, 2019: 213 (redescription).[Note: Seifert, 2019a 10.1007/s00040-019-00693-0 PDF: 1, reports Camponotus herculeanus × Camponotus ligniperda worker hybrids.].Senior synonym of Camponotus herculeanus altaicus: Schär et al., 2018 10.1111/jbi.13380 PDF: 6.Senior synonym of Camponotus atra: Nylander, 1846a PDF: 894; Foerster, 1850a: 9; Nylander, 1856b PDF: 56; Smith, 1858a PDF: 10; Mayr, 1863a PDF: 399; Roger, 1863b PDF: 1; Forel, 1874 PDF: 96 (in list); Emery & Forel, 1879 PDF: 447; Dalla Torre, 1893 PDF: 234; Emery, 1925d PDF: 72; Karavaiev, 1936: 178; Bolton, 1995b: 103; Radchenko, 2016: 328.Senior synonym of Camponotus herculeanus caucasicus: Arakelian, 1994 PDF: 85; Bolton, 1995b: 103; Radchenko, 1997b PDF: 555; Radchenko, 2016: 328.Senior synonym of Camponotus herculeanus eudokiae: Radchenko, 1997b PDF: 555; Radchenko, 2016: 328.Senior synonym of Camponotus intermedia: Nylander, 1846a PDF: 894; Foerster, 1850a: 9; Nylander, 1856b PDF: 56; Smith, 1858a PDF: 10; Mayr, 1863a PDF: 399; Roger, 1863b PDF: 1; Emery & Forel, 1879 PDF: 447; Emery, 1925d PDF: 72; Karavaiev, 1936: 178; Bolton, 1995b: 103; Radchenko, 1997b PDF: 555; Radchenko, 2016: 328.Senior synonym of Camponotus herculeanus jacuticus: Yasumatsu & Brown, 1951 PDF: 35; Schär et al., 2018 10.1111/jbi.13380 PDF: 6.Senior synonym of Camponotus herculeanus montanus: Emery, 1925d PDF: 72; Karavaiev, 1936: 178; Yasumatsu & Brown, 1951 PDF: 30; Arnol'di, 1967: 1819; Arakelian, 1994 PDF: 85; Bolton, 1995b: 103; Radchenko, 1997b PDF: 555; Radchenko, 2016: 328.Senior synonym of Camponotus herculeanus nadigi: Yasumatsu & Brown, 1951 PDF: 30; Baroni Urbani, 1971c PDF: 176; Bolton, 1995b: 103; Radchenko, 1997b PDF: 555; Radchenko, 2016: 328.Senior synonym of Camponotus herculeanus sachalinensis: Yasumatsu & Brown, 1957 PDF: 49; Schär et al., 2018 10.1111/jbi.13380 PDF: 6.Senior synonym of Camponotus herculeanus shitkowi: Emery, 1925d PDF: 72; Karavaiev, 1936: 178; Arnol'di, 1967 PDF: 1819; Bolton, 1995b: 103; Radchenko, 1997b PDF: 555; Radchenko, 2016: 328.Senior synonym of Camponotus whymperi: Creighton, 1950a PDF: 367; Yasumatsu & Brown, 1951 PDF: 35; Smith, 1958c PDF: 142; Smith, 1979: 1426; Bolton, 1995b: 103; Mackay, 2019: 214.Material of the unavailable name Camponotus herculeanus shitkowi intermedius referred here by Radchenko, 1997b PDF: 555.Records
(Map 59): Bulgaria ( Agosti and Collingwood 1987a , Atanassov and Dlusskij 1992 ); Stara Planina Mts ( Seifert 1989 ); Western Stara Planina Mts: Vilya-glava mine ( Atanassov 1934 ); Sofia Basin: Sofia ( Atanassov 1934 ); Vitosha Mt. ( Atanassov 1936 , 1952 ); Plana Mt.: above Dyavolski most bridge (Kokalyane vill.), Bukov dol loc. (Pasarel vill.), Peyova buka hut (Pasarel vill.) ( Vagalinski and Lapeva-Gjonova in press ); Ihtimanska Sredna Gora Mts: Benkovski peak ( Atanassov 1934 ); Strandzha Mt.: Papia peak ( Atanassov 1934 ); Belasitsa Mt. ( Atanassov 1964 ); Krupnik-Sandanski-Petrich Valley: along Luda Mara river ( Atanassov 1964 ); Rila Mt. ( Seifert 1989 ): Elenin peak, Rilska river valley ( Forel 1892 ), Borovets ( Atanassov 1934 , 1936 ), Kostenets ( Atanassov 1934 ); Pirin Mt. ( Seifert 1989 ): Bansko-Banderitsa ( Atanassov 1934 ); Rhodopi Mts ( Seifert 1989 ); Western Rhodopi Mts: Karlak peak (Atanasov 1934), Perelik peak ( Atanassov 1936 ), Rakitovo, Batak ( Lapeva-Gjonova in press (a) ); Southern Black Sea coast: Maslen nos ( Atanassov 1934 ).
Notes:
Camponotus herculeanus is aboreo-montane species and its record from "Maslen nos" seems doubtful.
Caucase septentrional, district Maikop, mont Bambak, 2 [[ queen ]], 30. VIII. 1894 (S. Prichodko!); Transcaucasie, Gouv. Kutais, Artvin, 1 [[ queen ]], 23. VI. 1888 (Derjugin!); Turkestan, Ferghana, fl. Kugart, 7000 ' h., 2 [[ queen ]], 5. VIII. 1895
(KoRZINSKIj!).
Camponotus herculeanus (or Hercules ant)[1] is a species of ant in the genus Camponotus, the carpenter ants,[2] occurring in Northern Eurasia, from Norway to Eastern Siberia, and North America. First described as Formica herculeana by Linnaeus in 1758,[3] the species was moved to Camponotus by Mayr in 1861.[4]
The colony of Camponotus herculeanus consists of one or several wingless females (queens), some fertile males, and three castes of sterile workers, known as majors, intermediates, and minors, in decreasing order of size. The queens are large, about 15 mm (0.6 in) in length, and are blackish in colour. The males are a similar colour but about half the size of the queens. The workers usually have blackish heads and gasters, and dark reddish-brown mesosomas, petioles and legs. In majors, the scapes (the long segments of the antenna, before the elbow) are shorter than the length of the head; in intermediates they are about the same length, and in minors, they extend well beyond the back of the head. The head and the dorsal surfaces of the mesosoma and gaster of the largest majors are bristly.[5]
Camponotus herculeanus has a widespread distribution in the Northern Hemisphere, being present in most of Europe, Central and Northern Asia, Canada and the United States. It is common in mountainous regions and is the dominant ant species in mountainous and northerly parts of North America. It occupies a range of habitats including various types of conifer and hardwood forests, clearings, oak scrubland, disturbed areas, pastures and seashore grassland.[5]
Nests of Camponotus herculeanus are built in timber, living or rotting trees, stumps, fallen logs and occasionally the structural timbers of buildings. The ants use their strong jaws to excavate galleries and chambers under the bark or in the wood, with a preference for damp wood or timber with fungal decay. In standing trees, their tunnels sometimes extend for 10 m (30 ft) above the ground.[1] Satellite colonies, linked to the original nest by underground tunnels, may develop nearby, often in warmer, drier locations. These house older larvae, pupae, winged reproductives and workers, with the eggs and younger larvae remaining in the main nest.[6]
A colony of Camponotus herculeanus contains several wingless females, which may be unrelated. Winged reproductives are produced in late summer and overwinter in the colony, emerging to fly in swarms on warm spring days. The workers become active in spring and forage in the vicinity of the nest.[5] They tend aphids, and the larvae of the silvery blue butterfly (Glaucopsyche lygdamus), which often feeds on the lupine Lupinus bakeri.[5] The diet consists of the honeydew produced by sap-sucking insects and the ants also consume any insect larvae that they encounter.[7] The ant cricket Myrmecophilus pergandei sometimes lives in the colony, where it is tolerated by the ants.[5]
Camponotus herculeanus (or Hercules ant) is a species of ant in the genus Camponotus, the carpenter ants, occurring in Northern Eurasia, from Norway to Eastern Siberia, and North America. First described as Formica herculeana by Linnaeus in 1758, the species was moved to Camponotus by Mayr in 1861.
Head of a major worker
