Camponotus herculeanus

Camponotus herculeanus is a very common species that has large colonies under rotten bark and in rotten logs and stumps, especially those of conifers and in dead wood of living poplars (Wheeler and Wheeler, 1963; Mackay and Mackay, 2002). It occasionally nests under stones in clay soil, especially incipient nests (Mackay and Mackay, 2002). It is a boreal species generally lives at altitudes above 1200-1300 m in Mongolia. It inhabits different kinds of forest (larch, birch and pine) arid mountain meadows. Nests are built in rotten wood, stumps and tree hollows, and under logs partly in soil. Males and winged queens are seen in July in nests, and some dealated queens were collected outside the nest in late June to July (Aibek & Yamane, 2009, as C. sachalinensis).

Compare with Camponotus chromaiodes, Camponotus modoc, Camponotus novaeboracensis, Camponotus pennsylvanicus, Camponotus sansabeanus.

The largest majors of Camponotus herculeanus have scapes which barely reach, or only slightly surpass the posterior lateral corners of the head. They generally have a dark head and gaster and the mesosoma is at least partially brown. The legs are usually brown. Erect and suberect setae are moderately abundant, specifically on the clypeus (restricted to the margins), on the dorsal surface of the head, ventral surface of the head, dorsal surface of the mesosoma, petiole and gaster, setae are absent on the cheeks, scapes (except at the apex) and tibiae (except for a double row on the flexor surface). Appressed pubescence is sparse and is limited to a few tiny setae on the head, the dorsum of the mesosoma and dorsal surface of the gaster.

The scapes of the intermediate sized workers extend approximately the first two funicular segments past the posterior lateral corner of the head. The head and gaster are generally black, with the mesosoma, petiole and legs deep reddish brown.

The minors are similar except for size, having an oval-shaped head, and having the scapes extend well past the posterior lateral corners of the head.

The females are large (total length 15 mm), mostly dark specimens. The scape extends more than 2 funicular segments past the posterior lateral corners of the head.

The males are relatively small (total length 7 mm) black specimens.

Comparisons
Camponotus herculeanus is relatively easy to recognize among the ants of the subgenus Camponotus, as the scape of the larger majors and female reaches or barely surpasses the posterior lateral corner of the head. The scapes of the majors of most of the other species in the subgenus (except for Camponotus sansabeanus) nearly always extend at least 1 - 2 funicular segments past the posterior lateral corners.

Camponotus herculeanus would most likely be confused with Camponotus sansabeanus (southern US, northern Mexico), with relatively short scapes. In addition, the clypeus of C. sansabeanus has a poorly developed carina, which could result in it being confused with members of the subgenus Camponotus. The species can be separated as the antennal scapes of C. herculeanus are never flattened at the base, whereas they nearly always are in C. sansabeanus. Camponotus herculeanus is also somewhat dull in appearance, whereas C. sansabeanus is usually shiny and little sculptured. The shape of the clypeus is completely different in the 2 species: Camponotus herculeanus has a wide clypeus, which is convex, but without any raised or depressed areas in the medial region. The clypeus of Camponotus sansabeanus is narrower, the central region is usually somewhat elevated and occasionally there are longitudinal oblique depressions which point towards the medial region in C. sansabeanus. Finally, C. herculeanus has no erect and suberect setae on the cheeks, whereas C. sansabeanus normally does.

Camponotus herculeanus could be confused with Camponotus pennsylvanicus (primarily SE Canada, eastern US), Camponotus chromaiodes (eastern US) and Camponotus modoc (primarily western US). The majors of C. herculeanus can be separated by the shorter scapes and the relatively short appressed setae on the dorsum of the gaster (C. modoc has similar short setae). The females and majors of C. herculeanus can be separated as the scapes are shorter than those in the other 3 species, barely extending past the posterior lateral corner (up to the length of about the first funicular segment, extends more than the first funicular segments in the other 3 species). Unfortunately, the amount the scape extends past the corner of the head depends on the size of the major, with smaller majors having relatively longer scapes.

Camponotus herculeanus is often difficult to separate from C. chromaiodes, see the discussion of the latter species on hints to separate them.

It is often difficult to separate C. herculeanus from Camponotus modoc when limited specimens are available. The mesosomata of workers of C. herculeanus are partially red, they are often completely black in workers of C. modoc. The females of both species usually have black mesosomata. Camponotus herculeanus can also be separated from C. modoc by the color (C. modoc usually has red legs and at least part of the mesosoma is usually reddish). There is a considerable amount of variability in characters and these 2 taxa can be nearly indistinguishable.

Camponotus herculeanus is similar to Camponotus novaeboracensis (southern Canada, US) and is genetically similar (Sämi Schär, et al., 2018). The majors of the two species can be separated as the scapes of C. novaeboracensis extend well past the posterior lateral corners. The females of the two species are very difficult to separate, but the head of C. herculeanus is slightly wider (CI ranges from 113 to 114, versus 106 - 107 in C. novaeboracensis). Additionally, the dorsum of the mesosoma of the female of C. herculeanus is nearly always black, whereas it is often partially red in C. novaeboracensis. The mesosoma of the minor worker of C. herculeanus is nearly always dark, whereas it is reddish in C. novaeboracensis, contrasting with the remainder of the ant.

Some females of C. herculeanus from Canada are smaller (TL 10 - 12 mm) but otherwise there are no obvious differences and are considered to be conspecific. Specimens of majors from Europe usually have fewer erect and suberect setae on the mesosoma (0 - 20) as compared to those of North America (20 - 40), but otherwise they appear to be identical.

Distribution
Throughout mountain Europe and extending through Northern Eurasia from Norway to Eastern Siberia to the northernmost tree frontier in Arctic Norway (Collingwood 1979). Also northern North America.

Camponotus herculeanus is found in dense riparian forests, pine, fir and spruce forests, juniper-fir-deciduous forests, spruce birch forests, mixed conifer/hardwood, campground with trees, disturbed coniferous forest, coniferous/hardwood, burned oak scrub slope, birch forest, grassland along seashore, grassland, as well as in a Tamarack bog (Wheeler, 1910a) and deciduous forests, aspen, spruce-fir forests usually in clearings (Mackay and Mackay, 2002). It is the dominant ant in boreal and alpine forests in North America (Wheeler and Wheeler, 1963).

The Reinig Line faunal divide separates East Siberian, Inner Mongolian, Chinese and Tibetan species from those of Central Siberia, West Siberia and the Turanian region (DE LATTIN, 1967). In ants, the Reinig Line is crossed only by a cold resistant species including Camponotus herculeanus, Formica exsecta, Formica gagatoides, Formica lugubris, Formica manchu, Formica picea, Formica pisarskii, Formica uralensis, Lasius flavus, Leptothorax acervorum and Tetramorium sibiricum (DLUSSKY, 1967; FRANCOEUR, 1983; SEIFERT, 2000, 2021a, 2021b).

Distribution based on Regional Taxon Lists
Nearctic Region: Canada, United States. Palaearctic Region: Andorra, Armenia, Austria, Azerbaijan, Belarus, Belgium, Bulgaria, China, Croatia, Czech Republic, Denmark, Estonia, Finland, Georgia, Germany, Greece, Hungary, Iberian Peninsula, Italy, Kazakhstan, Kyrgyzstan, Latvia, Liechtenstein, Lithuania, Luxembourg, Netherlands, Norway, Poland, North Macedonia, Republic of Moldova, Romania, Russian Federation, Slovakia, Slovenia, Spain, Sweden, Switzerland, Turkey, Ukraine.

Nests contain multiple dealate females (Mackay and Mackay, unpublished), including unrelated queens (Ellison et al., 2012).

Alates and brood were collected in nests from May - September. In New England sexuals are produced in late summer, cared for throughout the winter and emerge for flights in warm spring days (Ellison et al., 2012). Males and females aggregate on the shores of a lake in Ontario, where they may form swarms elsewhere and are attracted to the lake shore or may be moved there by convectional air movements (Sanders, 1971). Dealate nest founding females were collected under the bark of log and stumps from May to October (Mackay and Mackay, 2002: Mackay and Mackay, unpublished).

Sanders (1972) studied foraging in C. herculeanus and found that the start of seasonal activity was temperature dependent and peaked in mid-summer.

Camponotus herculeanus tends aphids (Jones, 1929; Mackay and Mackay, 2002) and the myrmecophilous lycaenid butterfly Glaucopsyche lygdamus that that feed predominantly on Lupinus bakeri (Fraser et al., 2001). It is the host of the ant cricket Myrmecophilus pergandei (Hebard, 1920), and the secondary host for Nemadus brachyderus (Coleoptera: Leiodidae) (Peck and Cook, 2007), Foragers protect conifer seedlings from feeding damage by the pine weevil Hylobius abietis in Europe (Maňák et al., 2013).

Camponotus herculeanus is infected by the hymenopterous parasite Pseudochalcura gibbosa (Eucharitidae) (Heraty and Barber, 1990; Lachaud, J.-P. and G. Pérez-Lachaud. 2012), as well as by the nematode Oscheius dolichura (Poinar, 2012). Reemer (2012) discusses the parasitism by Microdon sp. flies and it is specifically the host of the syrphid fly parasite Microdon piperi (Duffield, 1981; Akre et al., 1988).

Camponotus herculeanus is a host of the endosymbiotic proteobacterium Wolbachia (Wernegreen et al., 2009) and Candidatus Blochmannia herculeanus, which has close relatives that occur in aphids and the tsetse fly (Sauer et al., 2000). It is also the host of the intracellular endosymbiotic bacterium Blochmannia floridanus (Wolschin et al., 2004). Wolschin et al. (2004) further found that B. floridanus proliferates during pupation and immediately after the eclosion. In older workers the number of bacteria present in the midgut bacteriocytes decreased significantly in C. herculeanus as well as the related C. sericeiventris. Workers have gram-negative prokaryotic endosymbionts in the follicle cells (Peloquin et al., 2001).

Camponotus herculeanus is eaten by bears and woodpeckers (Ellison et al., 2012).

It is the most cold tolerant ant species known, surviving to below -40°C (Ellison et al., 2012).

Hölldobler and Engel-Siegel (1984) discuss the lack of the metapleural gland in this species.

Camponotus herculeanus is a structural pest (Hansen and Klotz, 2005).

Fitzpatrick et al. (2013) used the models MaxEnt and MaxLike to predict the distribution.

Regional Notes
Incipient Colony.

Europe
Collingwood (1979) - This species is a typical denizen of shaded coniferous forest nesting in rotten stumps and occasionally mining in living trees. Fertilised females found nests singly. Alatae are developed in the late summer but overwinter to swarm in June.

New Mexico
Mackay and Mackay (2002) - This very common species normally nests in rotten logs and stumps, but nests are occasionally found under stones, especially incipient nests. This species may form a plesiobiotic relationship with Formica neorufibarbis. Foragers tend several species of aphids on many different plant species. Reproductives and brood were present in the nests from June to August, reproductives until September. Foundress females were found from late June to October.

Hemiptera
This species is associated with the following aphids (Saddiqui et al., 2019 and included references):, , , , , , , , , , , , and.

Nomenclature

 * . Formica herculeana Linnaeus, 1758: 579 (q.) (no state data, “Habitat in Europae....; in America septentrionali”).
 * [Note: type-locality Sweden, after Linnaeus, 1761: 426.]
 * [Misspelled as herculanea by Latreille, 1809: 126, Smith, F. 1858b: 10; misspelled as hepculeanus by Kuznetsov-Ugamsky, 1929a: 17; misspelled as helcureanus by Teranishi, 1940: 71, and subsequent pages.]
 * Lepeletier de Saint-Fargeau, 1835: 209 (w.m.); Wheeler, G.C. & Wheeler, J. 1953e: 185 (l.).
 * Combination in Camponotus: Mayr, 1861: 36 (in key);
 * combination in C. (Camponotus): Forel, 1914a: 259.
 * Status as species: Linnaeus, 1761: 426; Scopoli, 1763: 312; Linnaeus, 1767: 962; Fabricius, 1775: 391; Fabricius, 1782: 488; Fabricius, 1787: 307; Gmelin, 1790: 2797; Christ, 1791: 514; Olivier, 1792: 491; Fabricius, 1793: 349; Latreille, 1798: 33; Fabricius, 1804: 395; Jurine, 1807: 272; Gravenhorst, 1807: 286; Latreille, 1809: 126; Leach, 1815: 147; Latreille, 1817d: 96; Billberg, 1820: 104; Brullé, 1833: 326; Losana, 1834: 309; Lepeletier de Saint-Fargeau, 1835: 209; Zetterstedt, 1838: 448; Schilling, 1839: 52; Nylander, 1846a: 894; Nylander, 1846b: 1044; Foerster, 1850a: 9; Schenck, 1852: 123; Mayr, 1855: 308 (redescription); Nylander, 1856b: 56; Gredler, 1858: 3; Smith, F. 1858b: 10, 53; Roger, 1859: 229; Dumeril, 1860: 929; Mayr, 1861: 36 (in key); Meinert, 1861: 309; Mayr, 1863: 399; Roger, 1863b: 1; Cresson, 1865: 426; Dours, 1873: 164; Forel, 1874: 39 (in key); André, 1874: 176 (in key); Emery, 1878b: 44; Emery & Forel, 1879: 447; Forel, 1879a: 56; Provancher, 1881b: 354; André, 1882a: 142 (in key); Provancher, 1883: 597; Mayr, 1886d: 419; Cresson, 1887: 255; Provancher, 1887: 228 (in key); Nasonov, 1889: 9; Forel, 1892i: 306; Dalla Torre, 1893: 233; Emery, 1893i: 674; Ruzsky, 1896: 67; Emery, 1896d: 372 (in list); Ruzsky, 1902d: 5; Forel, 1902i: 699; Ruzsky, 1903c: 205; Forel, 1904b: 381; Ruzsky, 1905b: 214; Wasmann, 1906: 111; Forel, 1907e: 19; Emery, 1908a: 182; Wheeler, W.M. 1908f: 625; Bondroit, 1910: 487; Santschi, 1911d: 7; Karavaiev, 1912b: 592; Forel, 1914a: 266; Stitz, 1914: 95; Bruch, 1914: 228; Ruzsky, 1914b: 100; Bruch, 1915: 535; Forel, 1915d: 68 (in key); Ruzsky, 1915b: 4; Donisthorpe, 1915d: 347; Emery, 1916b: 225; Escherich, 1917: 330 (in key); Bondroit, 1918: 70; Nadig, 1918: 339; Menozzi, 1922c: 142; Soudek, 1922: 95; Kuznetsov-Ugamsky, 1923: 241; Müller, 1923b: 159; Wheeler, W.M. 1923b: 5; Vashkevich, 1924b: 146; Emery, 1925b: 72; Kiseleva, 1925: 73; Ruzsky, 1925a: 287; Essig, 1926: 868; Karavaiev, 1926e: 191; Ruzsky, 1926: 108; Stärcke, 1926: 119 (in key); Karavaiev, 1927a: 295; Donisthorpe, 1927b: 400; Menozzi, 1927b: 92; Kuznetsov-Ugamsky, 1929b: 36; Karavaiev, 1930b: 147; Arnol'di, 1933b: 602 (in key); Stitz, 1934: 4; Grandi, 1935: 102; Karavaiev, 1936: 177 (redescription); Ruzsky, 1936: 89; Kono & Sugihara, 1939: 10; Teranishi, 1940: 71; Novák & Sadil, 1941: 110 (in key); Holgersen, 1942: 10; Holgersen, 1943b: 172 (in key); Holgersen, 1944: 179; Ruzsky, 1946: 69; Creighton, 1950a: 366; Donisthorpe, 1950e: 1066; Röszler, 1950: 210; Yasumatsu & Brown, 1951: 30; Chapman & Capco, 1951: 220; Consani & Zangheri, 1952: 43; Cole, 1954f: 271; Ceballos, 1956: 312; Smith, M.R. 1958c: 142; Pisarski, 1961a: 153; Dlussky, 1962: 181; Collingwood, 1962: 220; Bernard, 1967: 340 (redescription); Arnol'di, 1967: 1819 (redescription); Smith, M.R. 1967: 366; Kutter, 1968a: 60; Collingwood & Yarrow, 1969: 81; Dlussky & Pisarski, 1970: 86; Baroni Urbani, 1971c: 175; Collingwood, 1971: 163; Banert & Pisarski, 1972: 352; Tarbinsky, 1976: 148 (redescription); Aktaç, 1977: 125; van Boven, 1977: 131; Francoeur, 1977b: 207; Kutter, 1977c: 204; Yensen, et al. 1977: 183; Collingwood, 1978: 91 (in key); Collingwood, 1979: 90; Smith, D.R. 1979: 1426; Allred, 1982: 454; Wheeler, G.C. & Wheeler, J. 1986g: 60; Agosti & Collingwood, 1987a: 58; Agosti & Collingwood, 1987b: 283 (in key); Nilsson & Douwes, 1987: 68; Mackay, Lowrie, et al. 1988: 105; Wang, C., Xiao & Wu, 1989a: 224 (in key); Morisita, et al. 1991: 41; Wang, M., 1992: 681; Atanassov & Dlussky, 1992: 210; Arakelian, 1994: 85; Radchenko, 1994b: 116 (in key); Wang, C. & Wu, 1994: 31 (in key); Wheeler, G.C., et al. 1994: 305; Bolton, 1995b: 103; Douwes, 1995: 92; Poldi, et al. 1995: 7; Wu, J. & Wang, 1995: 182; Radchenko, 1996b: 1202 (in key); Espadaler, 1997b: 27; Radchenko, 1997a: 555; Gallé, et al. 1998: 216; Czechowski, et al. 2002: 95; Mackay & Mackay, 2002: 290; Zhang, W. & Zheng, 2002: 218; Coovert, 2005: 166; Hansen & Klotz, 2005: 82; Radchenko, 2005b: 158; Csösz, & Markó, 2005: 228; Karaman, G.S. & Karaman, 2005: 58; Bračko, 2006: 145; Markó, Sipos, et al. 2006: 66; Petrov, 2006: 109 (in key); Schultz, R. et al. 2006: 203; Bračko, 2007: 19; Seifert, 2007: 262; Werner & Wiezik, 2007: 143; Zryanin & Zryanina, 2007: 232; Gratiashvili & Barjadze, 2008: 131; Casevitz-Weulersse & Galkowsky, 2009: 479; Lapeva-Gjonova, et al. 2010: 43; Boer, 2010: 18; Csösz, et al. 2011: 58; Karaman, M.G. 2011b: 70; Legakis, 2011: 30; Ran & Zhou, 2011: 67; Borowiec, L. & Salata, 2012: 476; Czechowski, et al. 2012: 240; Ellison, et al. 2012: 121; Guénard & Dunn, 2012: 28; Kiran & Karaman, 2012: 7; Karaman, C. & Aktaç, 2013: 52 (in key); Borowiec, L. 2014: 32; Lebas, et al. 2016: 126; Radchenko, 2016: 328; Salata & Borowiec, 2018c: 43; Schär, Talavera, et al. 2018: 6; Seifert, 2018: 256; Mackay, 2019: 213 (redescription).
 * [Note: Seifert, 2019a: 1, reports herculeanus × ligniperda worker hybrids.]
 * Senior synonym of altaicus: Schär, Talavera, et al. 2018: 6.
 * Senior synonym of atra: Nylander, 1846a: 894; Foerster, 1850a: 9; Nylander, 1856b: 56; Smith, F. 1858b: 10; Mayr, 1863: 399; Roger, 1863b: 1; Forel, 1874: 96 (in list); Emery & Forel, 1879: 447; Dalla Torre, 1893: 234; Emery, 1925b: 72; Karavaiev, 1936: 178; Bolton, 1995b: 103; Radchenko, 2016: 328.
 * Senior synonym of caucasicus: Arakelian, 1994: 85; Bolton, 1995b: 103; Radchenko, 1997a: 555; Radchenko, 2016: 328.
 * Senior synonym of eudokiae: Radchenko, 1997a: 555; Radchenko, 2016: 328.
 * Senior synonym of intermedia Zetterstedt: Nylander, 1846a: 894; Foerster, 1850a: 9; Nylander, 1856b: 56; Smith, F. 1858b: 10; Mayr, 1863: 399; Roger, 1863b: 1; Emery & Forel, 1879: 447; Emery, 1925b: 72; Karavaiev, 1936: 178; Bolton, 1995b: 103; Radchenko, 1997a: 555; Radchenko, 2016: 328.
 * Senior synonym of jacuticus: Yasumatsu & Brown, 1951: 35; Schär, Talavera, et al. 2018: 6.
 * Senior synonym of montanus Ruzsky: Emery, 1925b: 72; Karavaiev, 1936: 178; Yasumatsu & Brown, 1951: 30; Arnol'di, 1967: 1819; Arakelian, 1994: 85; Bolton, 1995b: 103; Radchenko, 1997a: 555; Radchenko, 2016: 328.
 * Senior synonym of nadigi: Yasumatsu & Brown, 1951: 30; Baroni Urbani, 1971c: 176; Bolton, 1995b: 103; Radchenko, 1997a: 555; Radchenko, 2016: 328.
 * Senior synonym of sachalinensis: Yasumatsu & Brown, 1957: 49; Schär, Talavera, et al. 2018: 6.
 * Senior synonym of shitkowi: Emery, 1925b: 72; Karavaiev, 1936: 178; Arnol'di, 1967: 1819; Bolton, 1995b: 103; Radchenko, 1997a: 555; Radchenko, 2016: 328.
 * Senior synonym of whymperi: Creighton, 1950a: 367; Yasumatsu & Brown, 1951: 35; Smith, M.R. 1958c: 142; Smith, D.R. 1979: 1426; Bolton, 1995b: 103; Mackay, 2019: 214.
 * Material of the unavailable name intermedius Ruzsky referred here by Radchenko, 1997a: 555.
 * altaicus. Camponotus herculeanus subsp. altaicus Ruzsky, 1926: 108.
 * [First available use of Camponotus herculeanus saxatilis var. altaica Ruzsky, 1915b: 6 (w.) RUSSIA; unavailable (infrasubspecific) name.]
 * As unavailable (infrasubspecific) name: Ruzsky, 1925b: 42.
 * Subspecies of saxatilis: Ruzsky, 1936: 90.
 * Junior synonym of sachalinensis: Arnol'di, 1967: 1821; Bolton, 1995b: 85; Radchenko, 1997a: 556.
 * Junior synonym of herculeanus: Schär, Talavera, et al. 2018: 6.
 * atra. Formica atra Zetterstedt, 1838: 450 (m.) SWEDEN.
 * Junior synonym of herculeanus: Nylander, 1846a: 894; Foerster, 1850a: 9; Nylander, 1856b: 56; Smith, F. 1858b: 10; Mayr, 1863: 399; Roger, 1863b: 1; Forel, 1874: 96 (in list); Emery & Forel, 1879: 447; Dalla Torre, 1893: 234; Emery, 1925b: 72; Karavaiev, 1936: 178; Bolton, 1995b: 86; Radchenko, 2016: 328.
 * caucasicus. Camponotus herculeanus subsp. caucasicus Arnol'di, 1967: 1822 (s.w.q.m.) GEORGIA.
 * Junior synonym of herculeanus: Arakelian, 1994: 85; Bolton, 1995b: 91; Radchenko, 1997a: 555; Radchenko, 2016: 328.
 * eudokiae. Camponotus herculeanus var. eudokiae Ruzsky, 1926: 108 (w.) RUSSIA.
 * Subspecies of herculeanus: Ruzsky, 1936: 90; Bolton, 1995b: 98.
 * Junior synonym of herculeanus: Radchenko, 1997a: 555; Radchenko, 2016: 328.
 * intermedia. Formica intermedia Zetterstedt, 1838: 448 (w.) SWEDEN.
 * Junior synonym of herculeanus: Nylander, 1846a: 894; Foerster, 1850a: 9; Nylander, 1856b: 56; Smith, F. 1858b: 10; Mayr, 1863: 399; Roger, 1863b: 1; Emery & Forel, 1879: 447; Emery, 1925b: 72; Karavaiev, 1936: 178; Bolton, 1995b: 105; Radchenko, 2016: 328.
 * jacuticus. Camponotus (Camponotus) herculeanus var. jacuticus Karavaiev, 1929b: 210 (s.w.q.m.) RUSSIA (Sakha (Yakutia)).
 * Subspecies of herculeanus: Karavaiev, 1931b: 30; Karavaiev, 1931c: 107; Chapman & Capco, 1951: 221.
 * Junior synonym of sachalinensis: Arnol'di, 1967: 1821; Radchenko, 1997a: 556; Bolton, 1995b: 106.
 * Junior synonym of herculeanus: Yasumatsu & Brown, 1951: 35; Schär, Talavera, et al. 2018: 6.
 * montanus. Camponotus herculeanus var. montanus Ruzsky, 1904a: 293 (w.) RUSSIA.
 * [Unresolved junior primary homonym of montanus Emery, 1894c: 168 (Bolton, 1995b: 112).]
 * Karavaiev, 1926e: 192 (s.w.q.).
 * Subspecies of herculeanus: Ruzsky, 1904b: 6; Ruzsky, 1905b: 223; Ruzsky, 1915b: 5; Ruzsky, 1916: 4; Ruzsky, 1925b: 42; Ruzsky, 1926: 108; Karavaiev, 1926e: 191; Ruzsky, 1936: 89; Ruzsky, 1946: 69.
 * Junior synonym of herculeanus: Emery, 1925b: 72; Karavaiev, 1936: 178; Yasumatsu & Brown, 1951: 30; Arnol'di, 1967: 1819; Arakelian, 1994: 85; Bolton, 1995b: 112; Radchenko, 1997a: 555; Radchenko, 2016: 328.
 * nadigi. Camponotus herculeanus var. nadigi Menozzi, 1922c: 142 (s.w.q.m.) ITALY.
 * Subspecies of herculeanus: Emery, 1925b: 72; Consani & Zangheri, 1952: 43.
 * Junior synonym of herculeanus: Yasumatsu & Brown, 1951: 30; Baroni Urbani, 1971c: 176; Bolton, 1995b: 112; Radchenko, 1997a: 555; Radchenko, 2016: 328.
 * Material of the nomen nudum vagusherculeanus referred here by Emery, 1925b: 72; Kutter, 1977c: 205.
 * sachalinensis. Camponotus herculeanus var. sachalinensis Forel, 1904b: 38 (q.) MONGOLIA, RUSSIA.
 * Karavaiev, 1912b: 592 (m.).
 * Combination in C. (Camponotus): Emery, 1925b: 72.
 * As unavailable (infrasubspecific) name: Emery, 1908a: 185; Emery, 1925b: 73; Teranishi, 1940: 71; Chapman & Capco, 1951: 221; Azuma, 1955: 80.
 * Status as species: Collingwood, 1981: 29; Bolton, 1995b: 121; Terayama, 1999b: 29 (in key); Collingwood & Heatwole, 2000: 12; Imai, et al. 2003: 39; Guénard & Dunn, 2012: 29.
 * Subspecies of herculeanus: Ruzsky, 1905b: 222; Wheeler, W.M. 1906c: 325; Yano, 1910: 422; Karavaiev, 1912b: 592; Ruzsky, 1925b: 42; Ruzsky, 1926: 108; Karavaiev, 1927d: 344; Teranishi, 1932: 50; Arnol'di, 1967: 1821; Pisarski, 1969a: 230; Pisarski, 1969b: 304; Dlussky & Pisarski, 1970: 86; Collingwood, 1976: 306; Onoyama, 1980: 200; Pisarski & Krzysztofiak, 1981: 159; Kupyanskaya, 1986b: 96; Kupyanskaya, 1990: 166; Morisita, et al. 1991: 41; Radchenko, 1994b: 116 (in key); Radchenko, 1996b: 1203 (in key); Radchenko, 1997a: 555; Radchenko, 2005b: 158.
 * Status as species: Ran & Zhou, 2011: 69.
 * Junior synonym of saxatilis: Kuznetsov-Ugamsky, 1928b: 4; Kuznetsov-Ugamsky, 1929a: 18.
 * Junior synonym of japonicus: Yasumatsu & Brown, 1951: 31.
 * Junior synonym of herculeanus: Yasumatsu & Brown, 1957: 49; Schär, Talavera, et al. 2018: 6.
 * shitkowi. Camponotus herculeanus var. shitkowi Ruzsky, 1904a: 292 (w.q.) RUSSIA.
 * [Misspelled as shitkovi by Ruzsky, 1914b: 100, Ruzsky, 1946: 69, and others.]
 * Subspecies of herculeanus: Ruzsky, 1914b: 100; Ruzsky, 1915b: 5; Ruzsky, 1926: 108; Ruzsky, 1946: 69.
 * Junior synonym of herculeanus: Emery, 1925b: 72; Karavaiev, 1936: 178; Arnol'di, 1967: 1819; Bolton, 1995b: 124; Radchenko, 1997a: 555; Radchenko, 2016: 328.
 * whymperi. Camponotus herculeanus var. whymperi Forel, 1902i: 699 (w.q.) CANADA (Alberta).
 * Wheeler, W.M. 1910d: 330 (s.m.).
 * As unavailable (infrasubspecific) name: Emery, 1908a: 184; Emery, 1925b: 73.
 * Status as species: Ruzsky, 1926: 108.
 * Subspecies of herculeanus: Forel, 1904a: 152; Forel, 1904b: 381; Ruzsky, 1905b: 222; Wheeler, W.M. 1910d: 330 (redescription); Wheeler, W.M. 1910g: 571; Wheeler, W.M. 1917a: 556; Wheeler, W.M. 1917e: 20; Ruzsky, 1920: 79; Kiseleva, 1925: 73; Ruzsky, 1925b: 42; Essig, 1926: 868; Ruzsky, 1936: 90; Cole, 1936a: 39; Wing, 1939: 163; Cole, 1942: 388; Wheeler, G.C. & Wheeler, E.W. 1944: 251; Ruzsky, 1946: 69; Smith, M.R. 1951a: 840.
 * Junior synonym of herculeanus: Creighton, 1950a: 367; Yasumatsu & Brown, 1951: 35; Smith, M.R. 1958c: 142; Smith, D.R. 1979: 1426; Bolton, 1995b: 130; Mackay, 2019: 214.

Type Material
Type material not examined by Mackay (2019).

Taxonomic Notes
Seifert (2019) found a low-level of hybridisation between C. herculeanus and Camponotus ligniperda. The frequency of hybridization between the two species is estimated for Central Europe as 0.2–1.0%. This low ratio indicates strong reproductive barriers considering syntopic occurrence at about 10% of the observation sites, a nearly complete overlap of swarming times and basically equal meteorological conditions to release swarming.

Schar et al. (2018): Camponotus sachalinensis Forel, 1904 syn. nov: This taxon has long been regarded a synonym of C. herculeanus (Collingwood, 1976; Kupyanskaya, 1990; Radchenko, 1996) but was raised to the rank of species (Bolton, 1995; Collingwood, 1981) without clear justification for this change in status. Our results support the hypothesis of synonymy with C. herculeanus. Camponotus herculeanus and C. sachalinensis form a young clade (~1.8 Ma, Figure 3) with a continuous distribution throughout the Holarctic. Camponotus sachalinensis represents the link between European and North American populations of C. herculeanus (Figures 2 and 3, Supporting Information Appendices S2 and S4). The current view of C. herculeanus occupying a disjunct distribution in the Western Palearctic and North America while being replaced by a distinct species, C. sachalinensis, in the Eastern Palearctic is bio-geographically not realistic. Camponotus sachalinensis is therefore here returned to synonymy with C. herculeanus. Its junior synonyms Camponotus herculeanus altaica Ruzsky, 1915 and Camponotus herculeanus jacuticus Karavaiev, 1929 are also placed in synonymy with C. herculeanus.

Major worker measurements (mm): HL 2.44 - 3.38, HW 2.40 - 3.74, SL 2.42 - 2.76, EL 0.54 - 0.73, CL 0.83 - 1.10, CW 1.08 - 1.40, WL 3.38 - 4.46, FFL 2.08 - 2.70, FFW 0.65 - 0.84. Indices: CI 98 - 111, SI 82 - 99, CLI 127 - 130, FFI 31.

Mandibles with 5 teeth; anterior border of clypeus concave; posterior margin of head weakly concave; eyes failing to reach sides of head by about 1 minimum diameter; scapes reaching or slightly surpassing posterior lateral corner of head; propodeum angulate between faces, 2 faces nearly equal in length, spiracle oblong; petiole narrow in profile, apex strongly convex and even sharp as seen from front.

Erect and suberect setae present along margins of clypeus, along frontal carinae, extending to posterior margin, present on dorsum of mesosoma, of petiole and all surfaces of gaster; appressed pubescence sparse, present on head, dorsum of mesosoma and dorsum of gaster, where few setae overlap adjacent setae.

Head coriaceous, with scattered punctures, mesosoma coriaceous, gaster finely, transversely striolate, most surfaces dull.

Head and gaster normally black, mesosoma at least partially brown (often mesopleuron, propodeum and petiole reddish brown, remainder black), legs reddish brown.

Minor worker measurements (mm): HL 1.78 - 2.12, HW 1.48 - 2.04, SL 1.70 - 1.96, EL 0.45 - 0.48, CL 0.51 - 0.69, CW 0.70 - 0.88, WL 2.62 - 2.76, FFL 1.50 - 1.76, FFW 0.44 - 0.56. Indices: CI 83 - 96, SI 92 - 96, CLI 127 - 137, FFI 29 - 32.

Minor worker similar to major, except head somewhat oval-shaped, eyes fail to reach sides of head by about ½ minimum diameter, posterior margin weakly convex, scape extends nearly ½ length past posterior lateral corner of head, pilosity, sculpture and color as in major worker.

Female measurements (mm): HL 2.60 - 3.02, HW 2.96 - 3.42, SL 2.18 - 2.46, EL 0.65 - 0.70, CL 0.94 - 1.00, CW 1.15 - 1.23, WL 4.95 - 5.38, FFL 2.16 - 2.56, FFW 0.70 - 0.79. Indices: CI 113 - 114, SI 81 - 84, CLI 123, FFI 31 - 32.

Similar to major worker, except eyes nearly reach sides of head, posterior margin convex, scapes extend 2 - 3 funicular segments past posterior lateral corners of head.

Pilosity, sculpture and color as in major worker.

Male measurements (mm): HL 1.50 - 1.78, HW 1.42 - 1.66, SL 1.72 - 1.98, EL 0.53 - 0.60, CL 0.39 - 0.48, CW 0.65 - 0.79, WL 3.46 - 4.04, FFL 2.26 - 2.52, FFW 0.45 - 0.49. Indices: CI 93 - 95, SI 111 - 115, CLI 166 - 168, FFI 19 - 20.

Anterior border of clypeus slightly concave, carina poorly marked; but longitudinally raised area present; scape relatively long, extending about ½ length past posterior lateral corner of head; apex of petiole sharp, composed of two angulate teeth with concave medial border.

References based on Global Ant Biodiversity Informatics

 * Agosti, D. and C.A. Collingwood. 1987. A provisional list of the Balkan ants (Hym. Formicidae) and a key to the worker caste. I. Synonymic list. Mitteilungen der Schweizerischen Entomologischen Gesellschaft, 60: 51-62
 * Aibek U., and S. Yamane. 2009. Taxonomic review of the genus Camponotus (Hymenoptera, Formicidae, Formicinae) from Mongolia. Biogeography 11: 97-108.
 * Allred D. M. 1982. Ants of Utah. The Great Basin Naturalist 42: 415-511.
 * Allred, D.M. 1982. The ants of Utah. Great Basin Naturalist 42:415-511.
 * Andoni V. 1977. Kontribut mbi Himenopteret e familjes Formicidae te vendit tone. Buletini I Shkencave te Natyres 31(2): 93-101.
 * AntArea. Accessed on February 5th 2014 at http://antarea.fr/fourmi/
 * Antarea (Personal Communication - Rumsais Blatrix- 27 April 2018)
 * Arnan X., M. Gracia, L. Comas, and J. Retana. 2009. Forest management conditioning ground ant community structure and composition in temperate conifer forests in the Pyrenees Mountains. Forest Ecology and Management 258(2): 5159.
 * ArtDatabanken Bugs (via GBIG)
 * Assing V. 1989. Die Ameisenfauna (Hym.: Formicidae) nordwestdeutscher Calluna-Heiden. Drosera 89: 49-62.
 * Azuma M. 1955. A list of ants (Formicidae) from Hokkaido Is. Hyogo Biology 3:79-80.
 * Banschbach V. S., and E. Ogilvy. 2014. Long-term Impacts of Controlled Burns on the Ant Community (Hymenoptera: Formicidae) of a Sandplain Forest in Vermont. Northeastern Naturalist 21(1): 1-12.
 * Baroni Urbani C., and C. A. Collingwood. 1977. The zoogeography of ants (Hymenoptera, Formicidae) in Northern Europe. Acta Zoologica Fennica 152: 1-34.
 * Bayartogtokh B., U. Aibek, S. Yamane, and M. Pfeiffer. 2014. Diversity and biogeography of ants in Mongolia (Hymenoptera: Formicidae). Asian Myrmecology 6: 1-20.
 * Belcher A. K., M. R. Berenbaum, and A. V. Suarez. 2016. Urbana House Ants 2.0.: revisiting M. R. Smith's 1926 survey of house-infesting ants in central Illinois after 87 years. American Entomologist 62(3): 182-193.
 * Belskaya E., A. Gilev, and E. Belskii. 2017. Ant (Hymenoptera, Formicidae) diversity along a pollution gradient near the Middle Ural Copper Smelter, Russia. Environ Sci Pollut Res DOI 10.1007/s11356-017-8736-8
 * Bernadou A., G. Latil, V. Fourcassié, and X. Espadaler. 2006. Etude des communautés de fourmis d'une vallée andorrane. Union International pour l'Etude des Insectes Sociaux, Colloque annuel de la section francaise, 4pp.
 * Bernadou A., V. Fourcassié, and X. Espadaler. 2013. A preliminary checklist of the ants (Hymenoptera, Formicidae) of Andorra. Zookeys 277: 13-23.
 * Bernadou A., X. Espadaler, A. Le Goff, and V. Fourcassie. 2015. Ant community organization along elevational gradients in a temperate ecosystem. Insect. Soc. 62:5971
 * Bernadou, A., G. Latil, V. Fourcassié, and X. Espadaler. "Les formigues de la Vall del Madriu-Perafita-Claror : diversitat i distribució." Hàbitats, 13 (2006): 10-21.
 * Bernard F. 1976. Écologie des fourmis des grès d'Annot, comparées à celles de la Provence calcaire. Annales du Muséum d'Histoire Naturelle de Nice 3: 33-54.
 * Bezdecka P. 1996. The ants of Slovakia (Hymenoptera: Formicidae). Entomofauna carpathica 8: 108-114.
 * Bezdeckova K., and P. Bezdecka. 2008. The ants of the Bohemian-Moravian Highlands. Acta rerum naturalium 5: 243252.
 * Blades, D.C.A. and S.A. Marshall. Terrestrial arthropods of Canadian Peatlands: Synopsis of pan trap collections at four southern Ontario peatlands. Memoirs of the Entomological Society of Canada 169:221-284
 * Blatrix R., C. Lebas, C. Galkowski, P. Wegnez, P. Pimenta, and D. Morichon. 2016. Vegetation cover and elevation drive diversity and composition of ant communities (Hymenoptera: Formicidae) in a Mediterranean ecosystem. – Myrmecological News 22: 119-127.
 * Boer P., W. Dekoninck, A. J. van Loon, and F. Vankerkhoven. 2003. List of ants (Hymenoptera: Formicidae) of Belgium and The Netherlands, their status and Dutch vernacular names. Entomologische Berichten 63 (3): 54-58.
 * Borchert, H.F. and N.L. Anderson. 1973. The Ants of the Bearpaw Mountains of Montana (Hymenoptera: Formicidae). Journal of the Kansas Entomological Society 46(2):200-224
 * Borowiec L. 2014. Catalogue of ants of Europe, the Mediterranean Basin and adjacent regions (Hymenoptera: Formicidae). Genus (Wroclaw) 25(1-2): 1-340.
 * Borowiec L., and S. Salata. 2012. Ants of Greece - Checklist, comments and new faunistic data (Hymenoptera: Formicidae). Genus 23(4): 461-563.
 * Boucher P., C. Hebert, A. Francoeur, and L. Sirois. 2015. Postfire succession of ants (Hymenoptera: Formicidae) nesting in dead wood of northern boreal forest. Environ. Entomol. 44(5): 1316-1327: DOI: 10.1093/ee/nvv109
 * Boven J. van 1949. Notes sur la faune des Hautes-Fagnes en Belgique. Bulletin et Annales de la Société Entomologique de Belgique 85: 135-143.
 * Bracko G. 2007. Checklist of the ants of Slovenia (Hymenoptera: Formicidae). Natura Sloveniae 9: 15-24
 * Bracko, G. 2006. Review of the ant fauna (Hymenoptera:Formicidae) of Croatia. Acta Entomologica Slovenica 14(2): 131-156.
 * Bracko, G.. "Review of the ant fauna (Hymenoptera: Formicidae) of Croatia." Acta Entomologica Slovenica Vol 14 st (2006): 131-156.
 * Bruch C. 1914. Catálogo sistemático de los formícidos argentinos. Revista del Museo de La Plata 19: 211-234.
 * Carniel A. 1998. Ricerche sulla mirmecofauna delle Prealpi Orobiche (Lombardia) (Insecta, Hymenoptera, Formicidae). Atti. Mus. Civ. Stor. Nat. Morbegno 9: 29-39.
 * Carroll T. M. 2011. The ants of Indiana (Hymenoptera: Formicidae). Master's Thesis Purdue university, 385 pages.
 * Casevitz-Weulersse J., and C. Galkowski. 2009. Liste actualisee des Fourmis de France (Hymenoptera, Formicidae). Bull. Soc. Entomol. Fr. 114: 475-510.
 * Casevitz-Weulersse J., and M. Prost. 1991. Fourmis de la Côte-d'Or présentes dans les collections du Muséum d'Histoire Naturelle de Dijon. Bulletin Scientifique de Bourgogne 44: 53-72.
 * Chen Ying-Wu et al. 2008. Preliminary list of Formicidae in Gansu Province. Journal of Anhui Agri. Sci. 36(32): 14133-14134
 * Chen Z. L., S. Y. Zhou, D. D. Ye, Y. Chen, and C. W. Lu. 2013. Moleular phylogeny of the ant subfamily Formicinae (Hymenoptera, Formicidae) from China based on Mitochondrial genes. Sociobiology 60(2): 135-144.
 * Cherix D., and S. Higashi. 1979. Distribution verticale des fourmis dans le Jura vaudois et recensement prelimaire des bourdons (Hymenoptera, Formicidae et Apidae). Bull. Soc. Vaud. Sc. Nat. 356(74): 315-324.
 * Cole A. C., Jr. 1942. The ants of Utah. American Midland Naturalist 28: 358-388.
 * Cole, A.C. 1936. An annotated list of the ants of Idaho (Hymenoptera; Formicidae). Canadian Entomologist 68(2):34-39
 * Collingwood C. A. 1956. Ant hunting in France. Entomologist 89: 106-108.
 * Collingwood C. A. 1971. A synopsis of the Formicidae of north Europe. Entomologist 104: 150-176
 * Collingwood C. A. 1976. Ants (Hymenoptera: Formicidae) from North Korea. Annales Historico-Naturales Musei Nationalis Hungarici 68:
 * Collingwood C., and H. Heatwole. 2000. Ants from Northwestern China (Hymenoptera, Formicidae). Psyche 103 (1-2): 1-24.
 * Collingwood C.A. 1959. Scandinavian Ants. Entomol. Rec. 71: 78-83
 * Collingwood, C. A. 1974. A revised list of Norwegian ants (Hymenoptera: Formicidae). Norsk Entomologisk Tidsskrift 21: 31-35.
 * Collingwood, C. A.. "The Formicidae (Hymenoptera) of Fennoscandia and Denmark." Fauna Entomologica Scandinavica 8 (1979): 1-174.
 * Consani M. 1947. Reperti corologici sulle formiche italiane. Redia 32: 179-182.
 * Consani M., and P. Zangheri. 1952. Fauna di Romagna. Imenotteri - Formicidi. Memorie della Societa Entomologica Italiana 31: 38-48.
 * Coovert, G.A. 2005. The Ants of Ohio (Hymenoptera: Formicidae) Ohio Biological Survey Bulletin New Series Volume 15(2):1-196
 * Csosz S., B. Marko, K. Kiss, A. Tartally, and L. Galle. 2002. The ant fauna of the Ferto-Hansag National Park (Hymenoptera: Formicidae). In: Mahunka, S. (Ed.): The fauna of the Fert?-Hanság National Park. Hungarian Natural History Museum, Budapest, pp. 617-629.
 * Csősz S. and Markó, B. 2005. European ant species (Hymenoptera: Formicidae) in the ant collection of the Natural History Museum of Sibiu (Hermannstadt/Nagyszeben), Romania II. Subfamily Formicinae. Annales Historico-Naturales Musei Nationalis Hungarici 97: 225-240.
 * Csősz S., B. Markó, and L. Gallé. 2011. The myrmecofauna (Hymenoptera: Formicidae) of Hungary: an updated checklist. North-Western Journal of Zoology 7: 55-62.
 * Cuní, M.. "Excursión entomológica a varias localidades de la provincia de Gerona (Cataluña)." Anales de la Sociedad española de Historia Natural (1885): 51.
 * Cuní, M.. "Insectos observados en los alrededores de Barcelona." Anales de la Sociedad española de Historia Natural XVII (1888): 133.
 * Czechowski W., A. Radchenko, W. Czechowska and K. Vepsäläinen. 2012. The ants of Poland with reference to the myrmecofauna of Europe. Fauna Poloniae 4. Warsaw: Natura Optima Dux Foundation, 1-496 pp
 * Dahms H., L. Lenoir, R. Lindborg, V. Wolters, and J. Dauber. 2008. Restoration of seminatural grasslands: what is the impact on ants? Restoration Ecology 18(3): 330-337.
 * Dekoninck W., and A. Pauly. 2002. Camponotus vagus Scopoli, 1763 (Hymenoptera Formicidae) a new ant species for Belgium? Bull. Soc. R. Belge Entomol. 138: 29-30.
 * Del Toro, I. 2010. PERSONAL COMMUNICATION. MUSEUM RECORDS COLLATED BY ISRAEL DEL TORO
 * Della Santa E. 1994. Guide pour l'identification des principales espèces de fourmis de Suisse. Miscellanea Faunistica Helvetiae 3: 1-124.
 * Dlussky G. M., and B. Pisarski. 1970. Formicidae aus der Mongolei. Ergebnisse der Mongolisch-Deutschen Biologischen Expeditionen seit 1962, Nr. 46. Mitteilungen aus dem Zoologischen Museum in Berlin 46: 85-90.
 * Donisthorpe, H.. "A first instalment of the ants of Turkey." Annals and Magazine of Natural History (12)3 (1950): 1057-1067.
 * Downing H., and J. Clark. 2018. Ant biodiversity in the Northern Black Hills, South Dakota (Hymenoptera, Formicidae). Journal of the Kansas Entomological Society 91(2): 119-132.
 * Drummond F. A., A. M. llison, E. Groden, and G. D. Ouellette. 2012. The ants (Formicidae). In Biodiversity of the Schoodic Peninsula: Results of the Insect and Arachnid Bioblitzes at the Schoodic District of Acadia National Park, Maine. Maine Agricultural and forest experiment station, The University of Maine, Technical Bulletin 206. 217 pages
 * Dubovikoff D. A., and Z. M. Yusupov. 2018. Family Formicidae - Ants. In Belokobylskij S. A. and A. S. Lelej: Annotated catalogue of the Hymenoptera of Russia. Proceedingss of the Zoological Institute of the Russian Academy of Sciences 6: 197-210.
 * Dusmet, J. M.. "Linneo y los himenópteros de España." Linneo en España. Homenaje a su segundo centenario Zaragoza (1907): 475.
 * Eidmann V. H., and H. Münden. 1933. Zur Kenntnis der Ameisenfauna von Sudlabrador. Zoologischer Anzeiger 101(7): 202-221.
 * Ellison A. M., E. J. Farnsworth, and N. J. Gotelli. 2002. Ant diversity in pitcher-plant bogs of Massachussetts. Northeastern Naturalist 9(3): 267-284.
 * Espadaler X., X. Roig, K. Gómez, and F. García. 2011. Formigues de les Planes de Son i mata de València (Hymenoptera, Formicidae) Treballs de la Institució Catalana d'Història Natural 16: 609-627.
 * Espadaler, X., X. Roig, and K. Gómez. "Cuatro nuevas citas de hormigas (Hymenopera, Formicidae) y actualización del listado para Cataluña (Península Ibérica)." Boletín de la Sociedad Entomológica Aragonesa 40 (2007): 313-316.
 * Espadaler, X.. "Contribución al conocimiento de los formícidos (Hymenoptera, Formicidae) del Pirineo catalán." Tesis Universida (1979): 285 pp.
 * Fiedler, K., F. Kuhlmann, B. C. Schlick-Steiner, F. M. Steiner and G. Gebauer. 2007. Stable N-isotope signatures of central European ants  assessing positions in a trophic gradient. Insectes Sociaux 54(4):393-402.
 * Field Museum Collection, Chicago, Illinois (C. Moreau)
 * Forel A. 1904. Note sur les fourmis du Musée Zoologique de l'Académie Impériale des Sciences à St. Pétersbourg. Ezheg. Zool. Muz. 8: 368-388.
 * Franch, J., and X. Espadaler. "Ants as colonizing agents of pine stumps in San Juan de la Peña (Huesca, Spain)." Vie et Milieu 38 (1988): 149-154.
 * Francoeur A. 2001. Les fourmis de la forêt boréale (Formicidae, Hymenoptera). Le naturaliste canadien 125(3): 108-114.
 * Francoeur, A. 1983. The ant fauna near the tree-line in Northern Quebec (Formicidae, Hymenoptera). Nordicana 47: 177-180
 * Francoeur, A. 1997. Ants (Hymenoptera: Formicidae) of the Yukon. Pages 901 910 in H.V. Danks and J.A. Downes (Eds.), Insects of the Yukon. Biological Survey of Canada (Terrestrial Arthropods), Ottawa.
 * Gadeau de Kerville H. 1922. Materiaux pour la Faune des Hymenopteres de la Normandie. Bull. Soc. Amis Sc. Nat. Rouen 1916-1921, 1922: 217-225.
 * Galle L. 1993. Data to the ant fauna of the Bukk (Hymenoptera: Formicidae). Natural history of the national parks of Hungary 7: 445-448.
 * Gallé L., B. Markó, K. Kiss, E. Kovács, H. Dürgő, K. Kőváry, and S. Csősz. 2005. Ant fauna of Tisza river basin (Hymenoptera: Formicidae).  In: Gallé, L. (szerk.): Vegetation and Fauna of Tisza River Basin I. Tiscia Monograph Series 7; Szeged, pp. 149-197.
 * Garcia Garcia F. 2017. Camponotus herculeanus (Linnaeus, 1758) and Camponotus ligniperdus (Latreille, 1802) (Hymenoptera: Formicidae) in Galicia (NW Iberia). Iberomyrmex 9: 45-47.
 * Garcia Garcia F., and A. D. Cuesta-Esgura. 2017. First catalogue of the ants of Burgos province, Spain (Hymenoptera: Formicidae). Boletín de la Sociedad Entomológica Aragonesa 60: 245–258.
 * Gibb H., and T. Johansson. 2011. Field tests of interspecific competition in ant assemblages: revisiting the dominant red wood ants. Journal of Animal Ecology 80: 548-557.
 * Gilev A. V., I. V. Kuzmin, V. A. Stolbov, and S. D. Sheikin. 2012. Materials on the fauna and ecology of ants (formicidae) Southern part of the Tyumen region. Tyumen State University Herald 6: 86-91.
 * Glaser F. 2009. Die Ameisen des Fürstentums Liechtenstein. (Hymenoptera, Formicidae). Amtlicher Lehrmittelverlag, Vaduz, 2009 (Naturkundliche Forschung im Fürstentum Liechtenstein; Bd. 26).
 * Glaser F., A. Freitag, and H. Martz. 2012. Ants (Hymenoptera: Formicidae) in the Münstertal (Val Müstair)  a hot spot of regional species richness between Italy and Switzerland. Gredleriana 12: 273 - 284.
 * Glasier J. R. N., S. E. Nielsen, J. Acorn, and J. Pinzon. 2019. Boreal sand hills are areas of high diversity for Boreal ants (Hymenoptera: Formicidae). Diversity 11, 22; doi:10.3390/d11020022.
 * Glasier J. R. N., S. Nielsen, J. H. Acorn, L. H. Borysenko, and T. Radtke. 2016. A checklist of ants (Hymenoptera: Formicidae) of Saskatchewan. The Canadian Field-Naturalist 130(1): 40-48.
 * Grandi G. 1935. Contributi alla conoscenza degli Imenotteri Aculeati. XV. Boll. R. Ist. Entomol. Univ. Studi Bologna 8: 27-121.
 * Gratiashvili N., Barjadze S. 2008. Checklist of the ants (Formicidae Latreille, 1809) of Georgia. Proceedings of the Institute of Zoology (Tbilisi) 23: 130-146.
 * Gregg R. E. 1946. The ants of northeastern Minnesota. American Midland Naturalist 35: 747-755.
 * Gregg, R.T. 1963. The Ants of Colorado.
 * Guénard B., and R. R. Dunn. 2012. A checklist of the ants of China. Zootaxa 3558: 1-77.
 * Hayashida K. 1959. Ecological Distribution of Ants in Mt. Atusanupuri, An Active Volcano in Akan National Park, Hokkaido. Jour. Pac. Sci. Hokkaiao Univ. Ser. 4(14): 252-260.
 * Hayashida K. 1959. Ecological distribution of ants in Mt. Atusanupuri, an active volcano in Akan National Park, Hokkaido. Journal of the Faculty of Science, Hokkaido University. Series VI. Zoology 14:252-260.
 * Higgins R. J., and B. S. Lindgren. 2006. The fine scale physical attributes of coarse woody debris and effects of surrounding stand structure on its utilization by ants (Hymenoptera: Formicidae) in British Columbia, Canada. Gen. Tech. Rep. SRS-93. Asheville, NC: U.S. Department of Agriculture, Forest Service, Southern Research Station pp. 67-73.
 * Hoey-Chamberlain R. V., L. D. Hansen, J. H. Klotz and C. McNeeley. 2010. A survey of the ants of Washington and Surrounding areas in Idaho and Oregon focusing on disturbed sites (Hymenoptera: Formicidae). Sociobiology. 56: 195-207
 * Holgersen H. 1942. Ants of northern Norway (Hym., Form.). Tromso Mus. Årsh. 63(2): 1-34.
 * Holgersen H. 1943. Ant studies in Rogaland (south-western Norway). Avhandlingar utgitt av det Norske Videnskaps-Akademi i Oslo. I. Matematisk-Naturvidenskapelig Klasse 1943(7): 1-75.
 * Holgersen H. 1944. The ants of Norway (Hymenoptera, Formicidae). Nytt Magasin for Naturvidenskapene 84: 165-203.
 * Hua Li-zhong. 2006. List of Chinese insects Vol. IV. Pages 262-273. Sun Yat-sen university Press, Guangzhou. 539 pages.
 * Hågvar S. 2005. Altitudinal zonation of ants (Formicidae) in a steep fjord landscape in Sogndal, Western Norway. Norw. J. Entomol. 52: 3-12.
 * IZIKO South Africa Museum Collection
 * Ivanov, K. 2019. The ants of Ohio (Hymenoptera, Formicidae): an updated checklist. Journal of Hymenoptera Research 70: 65–87.
 * Ivanov K., L. Hightower, S. T. Dash, and J. B. Keiper. 2019. 150 years in the making: first comprehensive list of the ants (Hymenoptera: Formicidae) of Virginia, USA. Zootaxa 4554 (2): 532–560.
 * Jensen T. F., and Nielsen M. G. 1982. En status over udbredelsen af myreslaegten Camponotus i Danmark (Hymenoptera: Formicidae). Entomologiske Meddelelser 49: 113-116.
 * Karaman M. G. 2011. A catalogue of the ants (Hymenoptera, Formicidae) of Montenegro. Podgorica: Catalogues 3, Volume 2, Montenegrin Academy of Sciences and Arts, 140 pp.
 * Karavaiev V. 1912. Ameisen aus dem paläarktischen Faunengebiete. Rus. Entomol. Obozr. 12: 581-596.
 * Karavaiev V. 1926. Beiträge zur Ameisenfauna des Kaukasus, nebst einigen Bemerkungen über andere palaearktische Formen. (Schluss). Konowia 5: 187-199.
 * Karavaiev V. 1931. Beitrag zur Ameisenfauna Jakutiens. (Auf Grund der Sammelergebnisse der Expeditionen der Wissenschaften der UdSSR., ausgeführt in den Jahren 1925 und 1926.). Zool. Anz. 94: 104-117.
 * Kofler A. 1995. Nachtrag zur Ameisenfauna Osttirols (Tirol, Österreich) (Hymenoptera: Formicidae). Myrmecologische Nachrichten 1: 14-25.
 * Korlevic, A.. "Prilozi fauni hrvatskih opnokrilaca." Glasn. Hrv. Narav. Dr. 5 (1890): 189-250.
 * Krugova T. M. 2017. Ant's population in the chern taiga of Tigireksky mountain range (North-Western Altai). Proceedings of the Tigirek Reserve 9: 68-77.
 * Kvamme T. 1982. Atlas of the Formicidae of Norway (Hymenoptera: Aculeata). Insecta Norvegiae 2: 1-56.
 * Kvamme T., and A. Wetas. 2010. Revidert liste over norske maur  Inkludert dialektiske navn og forslag til nye norske navn og forslag til norske navn. Norsk institutt for skog og landskap, Ås. 127 pp
 * Lapeva-Gjonova, L., V. Antonova, A. G. Radchenko, and M. Atanasova. "Catalogue of the ants (Hymenoptera, Formicidae) of Bulgaria." ZooKeys 62 (2010): 1-124.
 * Lebas C., C. Galkowski, P. Wegnez, X. Espadaler, and R. Blatrix. 2015. The exceptional diversity of ants on mount Coronat (Pyrénées-Orientales), and Temnothorax gredosi(Hymenoptera, Formicidae) new to France. R.A.R.E., T. XXIV (1): 24  33
 * Legakis A. 1983. The Zoological Museum of the University of Athens 2. The collection of ants from Greece. Biologia Gallo-Hellenica 11(1): 85-87.
 * Legakis A. 2001. Ants (Hymenoptera, Formicidae) collected in the eastern Pyrenees and surrounding area, September 1999. Proceedings of the International Colloqium of the European Invertebrate Survey: Cartography and Conservation of Biodiversity Marcevol Priory, Arboussols, france 30.8-4.9.1999. OPIE.
 * Legakis Collection Database
 * Lelej A. S. 2012. Annotated catalogue of the Insects of Russian Far East. Volume 1. Hymenoptera. Dalnauka: Vladivostok. 635 p.
 * Li Z.h. 2006. List of Chinese Insects. Volume 4. Sun Yat-sen University Press
 * Lidgren, B.S. and A.M. MacIsaac. 2002. A Preliminary Study of Ant Diversity and of Ant Dependence on Dead Wood in Central Interior British Columbia. USDA Forest Service Gen. Tech. Rep. PSW-GTR-181.
 * Lindgren, B.S. and A.M. MacIsaac. 2002. Ant dependence on dead wood in Central Interior British Columbia. USDA Forest Service Gen. Tech. Rep.PSW-GTR-181
 * Longino, J.T. 2010. Personal Communication. Longino Collection Database
 * Lubertazi, D. Personal Communication. Specimen Data from Museum of Comparative Zoology at Harvard
 * Maavara V. 1953. Ants of Estonian SSR. ABIKS loodusevaatlejale 10: 1-44.
 * Mackay W. P., and E. E. Mackay. 2002. The ants of New Mexico (Hymenoptera: Formicidae). Lewiston, New York: Edwin Mellen Press, 400 pp.
 * Mackay, W., D. Lowrie, A. Fisher, E. Mackay, F. Barnes and D. Lowrie. 1988. The ants of Los Alamos County, New Mexico (Hymenoptera: Formicidae). pages 79-131 in J.C. Trager, editor, Advances in Myrmecololgy.
 * Majzlan O., and P. Devan. 2009. Selected insect groups (Hymenoptera, Neuroptera, Mecoptera, Raphidioptera) of the Rokoš Massif (Strážovské vrchy Mts.). Rosalia (Nitra), 20, p. 63–70.
 * Malozemova L. A. 1972. Ants of steppe forests, their distribution by habitats, and perspectives of their utilization for protection of forests (north Kazakhstan). [In Russian.]. Zoologicheskii Zhurnal 51: 57-68.
 * Marko B., K. Kiss, and L. Galle. 2004. Mosaic structure of ant communities (Hymenoptera: Formicidae) in Eastern Carpathian marshes: regional versus local scales. Acta Zoologica Academiae Scientiarum Hungaricae 50(2): 7795.
 * Markó B., A. Ionescu-Hirsch, and A. Szász-Len. 2009. Genus Camponotus Mayr, 1861 (Hymenoptera: Formicidae) in Romania: distribution and identification key to the worker caste. Entomologica Romanica 14: 29-41.
 * Markó B., B. Sipos, S. Csősz, K. Kiss, I. Boros, and L. Gallé. 2006. A comprehensive list of the ants of Romania (Hymenoptera: Formicidae). Myrmecologische Nachrichten 9: 65-76.
 * Martorell, M.. "Catálogos sinonímicos de insectos encontrados en Cataluña." Barcelona (1879).
 * Menozzi C. 1922. Nota complementare per la distinzione specifica dei Camponotus herculeanus L. e ligniperda Latr. (Hym.-Formic.). Bollettino della Società Entomologica Italiana. 54: 141-145.
 * Merle W. W. 1939. An Annotated List of the Ants of Maine (Hymenoptera: Formicidae). Entomological News. 50: 161-165
 * Moscaliuc L. 2008. Notes on the ant fauna (Hymenoptera: Formicidae) of Rarau mountain. Analele Facult??ii de Biologie, Univ A.I. Cuza Ia?i 54: 53-55.
 * Nadig A. 1918. Alcune note sulla fauna dell'alta Valsesia. Formicidae. Atti Soc. Ital. Sci. Nat. Mus. Civ. Stor. Nat. Milano 56: 331-341.
 * Neumeyer R., and B. Seifert. 2005. Commented check list of free living ant (Hymenoptera: Formicidae) species of Switzerland. Bulletin de la Societe Entomologique Suisse 78: 1-17.
 * Nielsen M. G. 1987. The ant fauna (Hymenoptera: Formicidae) in northern and interior Alaska. A survey along the trans-Alaskan pipeline and a few highways. Entomological News 98:74-88
 * Nielsen M. G. 2011. A check list of Danish ants and proposed common names. Ent. Meddr. 79: 13-18.
 * Nielsen, M.G. 1986. Respiratory rates of ants from different climatic areas. Journal of Insect Physiology 32(2): 125-131
 * Nielsen, M.G. 1987. The ant fauna (Hymenoptera:Formicidae) in northern and Interior Alaska. Entomological News 98(2):74-88
 * Novgorodova T. A., A. S. Ryabinin. 2015. Trophobiotic associations between ants (Hymenoptera, Formicidae) and aphids (Hemiptera, Aphidomorpha) in South Zauralye. News of Saratov University. Chemistry Series, Biology, Ecology 2(15): 98-107.
 * Ouellette G. D., F. A. Drummond, B. Choate and E. Groden. 2010. Ant diversity and distribution in Acadia National Park, Maine. Environmental Entomology 39: 1447-1556
 * Paik W.H. 1984. A checklist of Formicidae (Hymenoptera) of Korea. Korean J. Plant Prot. 23(3): 193-195.
 * Paraschivescu D. 1972. Fauna mirmecologica din zonele saline ale Romaniei. Studii si Cercetari de Biologie. Seria Zoologie 24: 489-495.
 * Paraschivescu D. 1978. Elemente balcanice in mirmecofauna R. S. Romania. Nymphaea 6: 463- 474.
 * Paukkunen J., and M. V. Kozlov. 2015. Stinging wasps, ants and bees (Hy menoptera: Aculeata) of the Murmansk region, Northwest Russia. — Entomol. Fennica. 26: 53–73.
 * Petal J. M. 1963. Faune des fourmis de la reserve de tourbiere en projet a Rakowskie Bagno pres de Frampol (voivodie de Lublin). Annales Universitatis Mariae Curie-Sk?odowska 58(7): 143-174.
 * Petrov I. Z. 1986. Contribution to myrmecofauna in some oak-tree communities on the mountain Jastrebac. Prirodnjackog Muzeja i Beogradu Seriya B Bioloske Nauke Supplement: No. 41: 109-114.
 * Petrov I. Z. 2002. Contribution to the myrmecofauna (Formicidae, Hymenoptera) of the Banat Province (Serbia). Archives of Biological Sciences, Belgrade, 54(12): 57-64.
 * Petrov I. Z. 2005. Contribution to the myrmecofauna (Formicidae, Hymenoptera) of Mt. Kopaonik (Serbia). Archives of Biological Sciences 57: 17-18.
 * Petrov I. Z., and C. A. Collingwood. 1992. Survey of the myrmecofauna (Formicidae, Hymenoptera) of Yugoslavia. Archives of Biological Sciences (Belgrade) 44: 79-91.
 * Pisarski B. 1969. Fourmis (Hymenoptera: Formicidae) de la Mongolie. Fragmenta Faunistica (Warsaw). 15: 221-236.
 * Powell, J.M. 1971. The arthropod fauna collected from the comandra blister rust, Cronartium comandrae, on lodgepole pine in Alberta. Canadian Entomologist 103:908-918
 * Prest W. H., and H. Piers. 1922. List of a Small Collection of Ants (Formicidae) obtained in Queen's County, Nova Scotia. Nova Scotian Institute of Science 15(4): 169-173.
 * Procter W. 1938. Biological survey of the Mount Desert Region. Part VI. The insect fauna. Philadelphia: Wistar Institute of Anatomy and Biology, 496 pp.
 * Punttila P., Y. Haila, J. Niemela, and T. Pajunen. 1994. Ant communities in fragments of old-growth taiga and managed surroundings. Ann. Zool. Fennici 31: 131-144.
 * Punttila P., and Y. Haila. 1996. Colonisation of a burned forest by ants in the southern Finnish Boreal forest. Silva Fennica 30(4): 421-435.
 * Pusvaskyte O. 1979. Myrmecofauna of the Lituanian SSR. Acta Entomologica Lituanica 4: 99-105.
 * Radchenko, A. 2005. Monographic revision of the ants (Hymenoptera: Formicidae) of North Korea. Annales Zoologici (Warsaw) 55: 127-221.
 * Ran H., and S. Y. Zhou. 2011. Checklist of Chinese Ants: the Formicomorph Subfamilies (Hymenoptera: Formicidae) (I). Journal of Guangxi Normal University: Natural Science Edition. 29(3): 65-73.
 * Rees D. M., and A. W. Grundmann. 1940. A preliminary list of the ants of Utah. Bulletin of the University of Utah, 31(5): 1-12.
 * Rigato F., and R. Sciaky. 1989. Contributo alla conoscenza della mirmecofauna della Val Gesso (alpi Marittime) (Hymenoptera, Formicidae). Boll. Mus. Reg. Sci. Nat. Torino 7(2): 427-442.
 * Rigato F., and R. Sciaky. 1991. The myrmecofauna of the Gesso Valley (Maritime Alps) (Hymenoptera Formicidae). Ethology Ecology and Evolution Special Issue 1: 87-89.
 * Ruzsky M. 1920. Ants of Kamchatka. Izv. Inst. Issled. Sib. 2: 76-80
 * Salata S. 2014. Ants (Hymenoptera: Formicidae) of the National Park of the Stołowe Mts. Przyroda Sudetow 17: 161-172.
 * Schar S., G Talavera, X. Espadaler, J. D. Rana, A. A. Andersen, S. P. Cover, and R. Vila. 2018. Do Holarctic ant species exist? Trans-Beringian dispersal and homoplasy in the Formicidae. Journal of Biogeography 00: 1-12.
 * Schlick-Steiner B. C., and F. M. Steiner. 1999. Faunistisch-ökologische Untersuchungen an den freilebenden Ameisen (Hymenoptera: Formicidae) Wiens. Myrmecologische Nachrichten 3: 9-53.
 * Schultz, R., A. G. Radchenko, and B. Seifert. "A critical checklist of the ants of Kyrgyzstan (Hymenoptera: Formicidae)." Myrmecologische Nachrichten 8 (2006): 201-207.
 * Seifert B. 2019. Hybridization in the European carpenter ants Camponotus herculeanus and C. ligniperda (Hymenoptera: Formicidae). Insectes Sociaux https://doi.org/10.1007/s00040-019-00693-0
 * Sharplin, J. 1966. An annotated list of the Formicidae (Hymenoptera) of Central and Southern Alberta. Quaetiones Entomoligcae 2:243-253
 * Shlyakhtenok A. S. 2007. Hymenoptera Aculeata of Raised Bogs in Belarus. Entomological Review 87(2): 136147.
 * Siberian Zoological Museum. Website available at http://szmn.sbras.ru/old/Hymenop/Formicid.htm. Accessed on January 27th 2014.
 * Smith F. 1941. A list of the ants of Washington State. The Pan-Pacific Entomologist 17(1): 23-28.
 * Smith M. R. 1952. On the collection of ants made by Titus Ulke in the Black Hills of South Dakota in the early nineties. Journal of the New York Entomological Society 60: 55-63.
 * Sonnenburg H. 2005. Die Ameisenfauna (Hymenoptera: Formicidae) Niedersachsens und Bremens. Braunschweiger Naturkundliche Schriften 7: 377-441.
 * Sonobe R. 1971. Ant survey of the Mt. Daisetsu area. Faunal Survey of the Mt. Daisetsu area. JIBP main area XIV. Annual Report of JIBP/CT-S: 199-210.
 * Soulie J. 1962. Fourmis des Hautes-Pyrenees. Bull. Soc. Hist. Nat. Toulouse 97: 35-37.
 * Steiner F. M., S. Schödl, and B. C. Schlick-Steiner. 2002. Liste der Ameisen Österreichs (Hymenoptera: Formicidae), Stand Oktober 2002. Beiträge zur Entomofaunistik 3: 17-25.
 * Tausan I. 2010. Notes on the ant fauna (Hymenoptera, Formicidae) of the Rodna Mountains National Park and it's surroundings (Transylvania-Maramures, Romania). Transylv. Rev. Syst. Ecol. Res. 9: 159-166.
 * Tausan I. 2016. First record of Camponotus lateralis (Olivier, 1792)(Hymenoptera: Formicidae) from Dobrogea (Romania). Brukenthal. Acta Musei, 11: 443-446.
 * Tausan I., M. M. Jerpel, I. R. Puscasu, C. Sadeanu, R. E. Brutatu, L. A. Radutiu, and V. Giurescu. 2012. Ant fauna (Hymenoptera: Formicidae) of Sibiu County (Transylvania, Romania). Brukenthal. Acta Musei 7(3): 499-520.
 * Tausan I., and I C. Negrila. 2015. The ant collection (Hymenoptera: Formicidae) of the natural history museum of Sibiu (Romania). Brukenthal Acta Musei 3: 409-418.
 * Terayama M. 1992. Structure of ant communities in East Asia. A. Regional differences and species richness. Bulletin of the Bio-geographical Society of Japan 47: 1-31.
 * Terayama M., Choi, B.M., Kim, C.H. 1992. A check list of ants from Korea, with taxonomic notes. Bulletin of the Toho Gakuen 7:19-54.
 * Terayama M., K. Ogata, and B.M. Choi. 1994. Distribution records of ants in 47 prefectures of Japan. Ari (report of the Myrmecologists Society of Japan) 18: 5-17.
 * Terayama. M. 2004. Geological and ecological distribution of Japanese ants communities. (translated from Japanese) Reports of the Saitama Prefecture Animal Research Association. 48:24
 * Tie Ru, and Xu Shengquan. 2004. Variety and distribution of ants in Northwest China. Journal of Ningxia Agricultural College 25(3): 4-9.
 * Vagalinski B., and A. Lapeva-Gjonova. 2012. The ants (Hymenoptera, Formicidae) of Plana Mountain (Bulgaria). Historia naturalis bulgarica 20: 87-101.
 * Varady-Szabo, H. 2004. Spiders and Ants Associated with Fallen Logs in Forillon National Park of Canada, Quebec. Masters Thesis, MgGill University, Montreal, Canada
 * Vele A., J. Holusa, J. Frouz, and O. Konvicka. 2011. Local and landscape drivers of ant and carabid beetle communities during spruce forest succession. European Journal of Soil Biology 47: 349-356.
 * Vele A., J. Holusa, and J. Frouz. 2009. Sampling for ants in different-aged spruce forests: A comparison of methods. European Journal of Soil Biology 45(4): 1-6.
 * Vepsalainen K., H. Ikonene, and M. J. Koivula. 2008. The structure of ant assembalges in an urban area of Helsinki, southern Finland. Ann. Zool. Fennici 45: 109-127.
 * Vepsalainen K., R. Savolainen, J. Tiainen, and J. Vilen. 2000. Successional changes of ant assemblages: from virgin and ditched bogs to forests. Ann. Zool. Fennici 37: 135-149.
 * Vespalainen K., B. Pisarski, R. Kantorek, and K. J. Laine. 1984. Formicidae (Hymenoptera) of Inari Lapland. Kevo Notes 7: 115-116.
 * Vogrin, V.. "Prilog fauni Hymenoptera - Aculeata Jugoslavije." Zast. Bilja 31(suppl.) (1955): 1-74.
 * Wang Ji-Fei et al. 2009. Study on the Species and Ecological Distribution of Ants on Ningxia Helan Mountain. Journal of Anhui Agri. Sci. 37(23): 11032-11034.
 * Wang 2008. Ant species diversity on Shangqiu forest park, Henan province. Sichuan Journal of Zoology 27(6): 1041-1044.
 * Wang C., Xiao G. and Wu J.. 1989. Taxonomic studies on the genus Camponotus in China (Hymenoptera: Formicidae) (conclusion). Forest Research 2: 221-228
 * Wang Lu-ling. 2008. Ant species diversity on Shangqiu forest Park, Henan province. Sichuan Journal of Zoology 27(6): 1041-1044.
 * Wang S. Z. et al. 2011. Investigation on ant species for garden tree in Chengdu. Journal of Anhui Agri. Sci. 39(9): 5045-5046.
 * Weber N. A. 1943. The ants of the Imatong Mountains, Anglo-Egyptian Sudan. Bulletin of the Museum of Comparative Zoology 93: 263-389.
 * Wegnez P., and A. Ronk. 2017. Découverte de Camponotus herculeanus (Linnaeus, 1758) et signalement de quelques autres espèces rares de fourmis au Luxembourg (Hymenoptera : Formicidae). Bulletin de la Société des naturalistes luxembourgeois 119 : 153–159.
 * Wegnez P., and F. Mourey. 2016. Formica uralensis Ruzsky, 1895 une espèce encore présente en France mais pour combien de temps ? (Hymenoptera: Formicidae). Bulletin de la Société royale belge d’Entomologie 152: 72-80.
 * Wheeler G. C., J. N. Wheeler, and P. B. Kannowski. 1994. Checklist of the ants of Michigan (Hymenoptera: Formicidae). The Great Lakes Entomologist 26(4): 297-310
 * Wheeler G. C., and E. W. Wheeler. 1944. Ants of North Dakota. North Dakota Historical Quarterly 11:231-271.
 * Wheeler G. C., and J. Wheeler. 1986. The ants of Nevada. Los Angeles: Natural History Museum of Los Angeles County, vii + 138 pp.
 * Wheeler G. C., and J. Wheeler. 1987. A Checklist of the Ants of South Dakota. Prairie Nat. 19(3): 199-208.
 * Wheeler J. N., G. C. Wheeler, R. J. Lavigne, T. A. Christiansen, and D. E. Wheeler. 2014. The ants of Yellowstone National Park. Lexington, Ky. : CreateSpace Independent Publishing Platform, 2013. 112 pages.
 * Wheeler W. M. 1906. The ants of Japan. Bulletin of the American Museum of Natural History 22: 301-328.
 * Wheeler W. M. 1908. The ants of Casco Bay, Maine, with observations on two races of Formica sanguinea Latreille. Bulletin of the American Museum of Natural History 24: 619-645.
 * Wheeler W. M. 1910. The North American ants of the genus Camponotus Mayr. Annals of the New York Academy of Sciences 20: 295-354.
 * Wheeler W. M. 1917. The mountain ants of western North America. Proceedings of the American Academy of Arts and Sciences 52: 457-569.
 * Wheeler, G.C. and J. Wheeler. 1988. A checklist of the ants of Montana. Psyche 95:101-114
 * Wheeler, G.C. and J. Wheeler. 1988. A checklist of the ants of Wyoming. Insecta Mundi 2(3&4):230-239
 * Wheeler, G.C., J. Wheeler and P.B. Kannowski. 1994. CHECKLIST OF THE ANTS OF MICHIGAN (HYMENOPTERA: FORMICIDAE). Great Lakes Entomologist 26:1:297-310
 * Wheeler, G.C., J. Wheeler, T.D. Galloway and G.L. Ayre. 1989. A list of the ants of Manitoba. Proceedings of the Entomological Society of Manitoba 45:34-49
 * Wiezik M. 2007. Ants (Hymenoptera: Formicidae) of mountain and alpine ecosystems at Southern part of Krá?ovoho?ské Tatry Mts. Naturae Tutela 11: 85-90.
 * Wiezik M. 2008. Vertical distribution of ants from alpine habitats at Nízke Tatry mts. Naturae Tutela 12: 61-67.
 * Wiezik M., A. Wiezikova, and J. Murin. 2008. Ant assemblages at horná orava peatbogs. Naturae Tutela 12: 55-60.
 * Wing M. W. 1939. An annotated list of the ants of Maine (Hymenoptera: Formicidae). Entomological News 50:161-165.
 * Xin M., Y. Ma, and D. He. 2011. Fauna composition of Formicidae in Ningxia. Journal of Ningxia University (Natural Science Edition) 32(4): 403-412.
 * Yang Xiuzhi, and Wang Junsen. 1994. Two kinds of ant chromosomes from Heilongjiang. Zoological Research 15(2): 93-96.
 * Yasumatsu K., and W. L. Brown, Jr. 1951. Revisional notes on Camponotus herculeanus Linné and close relatives in Palearctic regions (Hymenoptera: Formicidae). J. Fac. Agric. Kyushu Univ. 10: 29-44.
 * Yusupov Z. M. 2014. On the fauna of ants (Hymenoptera, Formicidae) in Teberda State Natural Biospheric Reserve. Bulletin of the Adyghe State University. Series 4: Natural and Mathematical and Technical Sciences 147: 121-124.
 * Zhang W. and Zheng Z.. 2002. Studies of ant (Hymenoptera: Formicidae) fauna in Sichuan Province. Entomotaxonomia 24(3): 216-222.
 * Zhigulskaya Z. A. 2009. The ants of the Chuya Depression and the Yustyd river basin in Southeastern Altai. Contemporary Problems of Ecology 2009 2(3): 210-215.
 * Zhuytszyuan D. 2016. The ants (Hymenoptera, Formicidae) Nizhne-Svirsky reserve and their environmental features. Master's thesis Saint Petersburg State University.
 * Zryanin V. A., and T. A. Zryanina. 2007. New data on the ant fauna Hymenoptera, Formicidae in the middle Volga River Basin. Uspekhi Sovremennoi Biologii 127(2): 226-240.