Camponotus herculeanus

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Camponotus herculeanus
Scientific classification
Kingdom: Animalia
Phylum: Arthropoda
Class: Insecta
Order: Hymenoptera
Family: Formicidae
Subfamily: Formicinae
Tribe: Camponotini
Genus: Camponotus
Subgenus: Camponotus
Species complex: herculeanus
Species: C. herculeanus
Binomial name
Camponotus herculeanus
(Linnaeus, 1758)

Camponotus herculeanus casent0103345 profile 1.jpg

Camponotus herculeanus casent0103345 dorsal 1.jpg

Specimen labels

Synonyms

A boreal species, C. herculeanus is common and has large colonies. They inhabit mountain conifer and mixed forests (larch, birch and pine), especially spruce, and arid mountain meadows. Nests of Camponotus herculeanusare located in rotten and live wood, stumps and tree hollows, and under partly buried logs in soil. Established colonies are most commonly found in conifers and dead wood of living poplars. In clay soils incipient nests can sometimes be found under stones. Males and winged queens have been observed in nests in July, and dealate queens collected outside the nest in late June to July. (Aibek & Yamane 2009; as C. sachalinensis, Mackay and Mackay 2002, Borowiec and Salata 2022, and Wheeler and Wheeler 1963)


Common Name
Karafuto-kuro-oo-ari
Language: Japanese
Notes: as Camponotus sachalinensis

Photo Gallery

  • Foraging major worker from Berlin, New Hampshire. Photo by Tom Murray.
  • Camponotus herculeanus queen. Photo by Michal Kukla.
  • Camponotus herculeanus queen, workers, brood. Photo by Michal Kukla.
  • A Camponotus herculeanus worker falls prey to a Aelurillus v-insignitus jumping spider. Photo by Michal Kukla.
  • Camponotus herculeanus and parasitic dipteran larvae. Photo by Michal Kukla.

Identification

Mackay (2019) - 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.

Keys including this Species

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.

Salata et al. (2023) - Camponotus herculeanus has not been detected in Cyprus following the records of Georghiou (1977). Nevertheless, it is known that Georghiou’s list excluded earlier publications and several species were misidentified (Preiss & Platia 2003; Austin et al. 2008). Camponotus herculeanus is a species which can be relatively easily recognized among ants of the subgenus Camponotus as the scape of the larger majors and females reaches or barely surpasses the posterior lateral corner of the head. No other species of the subgenus Camponotus was identified in our material, while the species is also absent from other Mediterranean islands such as Crete, Sicily, Sardinia, Corsica, and the Balearics despite of it being present in neighbouring mainland countries (Janicki et al. 2016; Guénard et al. 2017). Also, there is no habitat suitable for this species in Cyprus. Thus, we tentatively suggest the removal of C. herculeanus from the myrmecofauna of Cyprus.

Mackay (2019) - 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).

Latitudinal Distribution Pattern

Latitudinal Range: 70.377854° to 30.65861°.

   
North
Temperate 		North
Subtropical 		Tropical South
Subtropical 		South
Temperate

Distribution based on Regional Taxon Lists

Nearctic Region: Canada, United States.
Palaearctic Region: Andorra, Armenia, Austria, Azerbaijan, Belarus, Belgium, Bulgaria, China, Croatia, Czechia, Denmark, Estonia, Finland, Georgia, Germany, Greece, Hungary, Iberian Peninsula, Iceland, Italy, Kazakhstan, Kyrgyzstan, Latvia, Liechtenstein, Lithuania, Luxembourg, Netherlands, North Macedonia, Norway, Poland, Republic of Moldova, Romania, Russian Federation (type locality), Slovakia, Slovenia, Spain, Sweden, Switzerland, Türkiye, Ukraine.

Distribution based on AntMaps

AntMapLegend.png

Distribution based on AntWeb specimens

Check data from AntWeb

Countries Occupied

Number of countries occupied by this species based on AntWiki Regional Taxon Lists. In general, fewer countries occupied indicates a narrower range, while more countries indicates a more widespread species.
pChart


Biology

Mackay (2019) - 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.

Association with Other Organisms

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  • This species is a xenobiont for the ant Formica fusca (a xenobiont) in Poland (Czechowski, 2005; Kanizsai et al., 2013) (Forest edge. Under wood).
  • This species is a xenobiont for the ant Lasius platythorax (a xenobiont) in Finland (Czechowski, 2004; Kanizsai et al., 2013) (In decaying wood.).

Diptera

  • This species is a host for the Microdon fly Microdon piperi (a predator) in Oregon, Montana, Washington (type) (Knab, 1917).
  • This species is a prey for the Microdon fly Microdon piperi (a predator) (Quevillon, 2018).
  • This species is a host for the phorid fly Menozziola schmitzi (a parasitoid) (Quevillon, 2018) (encounter mode primary; direct transmission; transmission outside nest).
  • This species is a host for the phorid fly Trucidophora camponoti (a parasitoid) (Quevillon, 2018) (encounter mode primary; direct transmission; transmission outside nest).

Hemiptera

This species is associated with the following aphids (Saddiqui et al., 2019 and included references): Aphis lugentis, Aphis salicariae, Aphis valerianae, Aphis varians, Chaitophorus populicola, Chaitophorus populifolii, Cinara apini, Cinara hottesi, Cinara pseudotsugae, Cinara splendens, Cinara vandykei, Obtusicauda frigidae, Pterocomma populeum and Schizaphis graminum.

Hymenoptera

Mites

  • This species is a host for the mite Holostaspis flexuosa (a parasite) in Western Siberia, Russia (Joharchi et al., 2020) (ectoparasite).

Orthoptera

  • This species is a host for the cricket Myrmecophilus pergandei (a myrmecophile) in United States.

Nematodes

  • This species is a host for the nematode Oscheius dolichura (a parasite) (Quevillon, 2018) (multiple encounter modes; indirect transmission; transmission outside nest).
  • This species is a host for the nematode Oscheius dolichura (a parasite) in Ontario (Nickle & Ayre, 1966).
  • This species is a host for the nematode Mermithidae (unspecified "Mermix") (a parasite) in Italy (Finzi, 1930; Laciny, 2021).

Trematode

  • This species is a host for the trematode Brachylecithum mosquensis (a parasitoid) (Quevillon, 2018) (encounter mode primary; indirect transmission; transmission outside nest).

Flight Period

X X X
Jan Feb Mar Apr May Jun Jul Aug Sep Oct Nov Dec

Source: Aibek & Yamane (2009); antkeeping.info.

Explore-icon.png Explore: Show all Flight Month data or Search these data. See also a list of all data tables or learn how data is managed.

Life History Traits

  • Queen number: monogynous (Rissing and Pollock, 1988; Frumhoff & Ward, 1992)
  • Queen type: winged (Rissing and Pollock, 1988; Frumhoff & Ward, 1992) (queenless worker reproduction)

Castes

Worker

Images from AntWeb

Camponotus herculeanus casent0173157 head 1.jpgCamponotus herculeanus casent0173157 profile 1.jpgCamponotus herculeanus casent0173157 dorsal 1.jpgCamponotus herculeanus casent0173157 label 1.jpg
Worker. Specimen code casent0173157. Photographer April Nobile, uploaded by California Academy of Sciences. Owned by CAS, San Francisco, CA, USA.

Queen

Images from AntWeb

Camponotus herculeanus casent0103346 head 1.jpgCamponotus herculeanus casent0103346 profile 1.jpgCamponotus herculeanus casent0103346 dorsal 1.jpgCamponotus herculeanus casent0103346 label 1.jpg
Queen (alate/dealate). Specimen code casent0103346. Photographer April Nobile, uploaded by California Academy of Sciences. Owned by USNM, Washington, DC, USA.
Camponotus herculeanus casent0178766 head 1.jpgCamponotus herculeanus casent0178766 profile 1.jpgCamponotus herculeanus casent0178766 dorsal 1.jpgCamponotus herculeanus casent0178766 label 1.jpg
Queen (alate/dealate). Specimen code casent0178766. Photographer April Nobile, uploaded by California Academy of Sciences. Owned by CAS, San Francisco, CA, USA.
Camponotus sachalinensis casent0909955 h 1 high.jpgCamponotus sachalinensis casent0909955 p 1 high.jpgCamponotus sachalinensis casent0909955 p 2 high.jpgCamponotus sachalinensis casent0909955 p 3 high.jpgCamponotus sachalinensis casent0909955 d 1 high.jpgCamponotus sachalinensis casent0909955 l 1 high.jpg
Syntype of Camponotus herculeanus sachalinensis. Queen (alate/dealate). Specimen code casent0909955. Photographer Will Ericson, uploaded by California Academy of Sciences. Owned by MHNG, Geneva, Switzerland.

Nomenclature

The following information is derived from Barry Bolton's Online Catalogue of the Ants of the World.

  • herculeanus. Formica herculeana Linnaeus, 1758: 579 (w.) “Europe and North America”.
    • Type-material: syntype workers (number not stated).
    • Type-localities: Europe & America: (“Habitat in Europae”…. “in America septentrionali. Kalm.”) (no further data).
    • [Note: Linnaeus, 1761: 426, gives the type-locality as Sweden.]
    • Type-depositories: LSLC, ZMLS.
    • [Note: Day & Fitton, 1978: 196, cite 3 specimens LSLC.]
    • [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.]
    • Olivier, 1792: 491 (q.); Lepeletier de Saint-Fargeau, 1835: 209 (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 (redescription); Zetterstedt, 1838: 448; Schilling, 1839: 52; Nylander, 1846a: 894 (redescription); 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; Borowiec, M.L. et al. 2009: 378; 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; Steiner, et al. 2017: 7; Salata & Borowiec, 2018c: 43; Schär, Talavera, et al. 2018: 6; Seifert, 2018: 256; Werner, et al. 2018: 6; Mackay, 2019: 213 (redescription); Borowiec, L. & Salata, 2022: 59.
    • [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.
    • Distribution
    • Nearctic: Canada, U.S.A.
    • Palaearctic: Andorra, Armenia, Austria, Belarus, Belgium, Bosnia & Herzegovina, Bulgaria, China, Croatia, Czech Republic, Denmark, Estonia, Finland, France, Georgia, Germany, Greece, Hungary, Italy, Japan, Korean Peninsula, Kyrgyzstan, Latvia, Liechtenstein, Lithuania, Luxembourg, Moldova, Mongolia, Montenegro, Norway, Poland, Romania, Russia, Serbia, Slovakia, Slovenia, Spain, Sweden, Switzerland, Turkey, Ukraine.
    • [Note: Palaearctic distribution based on Borowiec, L. 2014: 32.]
  • 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.]
    • Type-material: syntype workers (number not stated).
    • Type-locality: Russia: Tomsk gub., Cherga, Muyuta, road to Anosu, valley slopes of river Katun, 1914-15 (M. Ruzsky).
    • Type-depository: unknown (syntypes lost (Radchenko, 1997a: 556)).
    • 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.
    • Type-material: holotype (?) male.
    • [Note: no indication of number of specimens is given.]
    • Type-locality: Sweden: Lapponia Tornensi, Junosuando, River Tornense, 9.vi. (J.W. Zetterstedt).
    • Type-depository: unknown (perhaps ZMLS).
    • 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.
    • Type-material: holotype major worker, 25 paratype major and minor workers, 2 paratype queens 3 paratype males.
    • Type-locality: holotype Georgia (“Western Caucasus”): Avalkhara, 6.vi.1958 (T.I. Zhizhilashvili); paratypes: with same data, Georgia: Svenetiya, Zeshko (Anguladze).
    • Type-depository: ZMUM.
    • 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.
    • Type-material: syntype workers (number not stated).
    • Type-locality: Russia: Siberia, Verkhneudinsk (= Ulan-Ude), 19.vii.1916 (M. Ruzsky).
    • Type-depository: unknown (syntypes lost (Radchenko, 1997a: 555)).
    • 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.
    • Type-material: syntype workers (number not stated).
    • Type-locality: Sweden: Lapponia, Juckasjervi (J.W. Zetterstedt).
    • Type-depository: unknown (perhaps ZMLS).
    • [Note: Horn & Kahle, 1937: 310, refer only to Diptera specimens in ZMLS.]
    • 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.
  • jacutica. Camponotus (Camponotus) herculeanus var. jacutica Karavaiev, 1929b: 210 (s.w.q.) RUSSIA.
    • Type-material: syntype major and minor workers, syntype queens (numbers not stated).
    • Type-localities: Russia: Kurort Kuldur, 32 km. N Stat. Birakan Ussuri-Bahn (vorm. Amursche Bahn), 26.vi.1928, no. 3902 (G. Kotshubey), with same data (P. Verestshak), Russia: Sakhalin I., Poronav river, 5-7.viii.1901, no. 4022 (P. Schmidt), Sakhalin I., Posten Tichmeniovsk, 23-24.viii.1901, no. 3592 (P. Schmidt), Russia: Irkutsk, Kaban Dist., vic. Dvoretz village, vi.1926 (P. Verestshak).
    • Type-depository: SIZK.
    • 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 (footnote) (w.) RUSSIA.
    • Type-material: lectotype worker (by designation of Radchenko, 1997a: 556).
    • Type-localities: Caucasus, and Tian Shan (M. Ruzsky?).
    • Type-depository: ZISP.
    • [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.
    • Type-material: syntype major and minor workers, syntype queen(s), syntype male(s) (numbers not stated).
    • Type-locality: Italy: Alagna, Riva e Val d’Artogna (Alta Valsesia) (A. Nadig).
    • Type-depository: IEUB.
    • 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.
    • Type-material: 7 syntype queens.
    • Type-localities: 6 queens Russia: Sakhalin I. (Suprunenko), 1 queen Mongolia: N slope Bogdo-Ula, S Urga, 15.vi.1894 (Kashkarov).
    • Type-depository: ZISP.
    • 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; Pfeiffer, et al. 2007: 4.
    • Status as species: Aibek & Yamane, 2009: 101 (redescription); 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.
    • Type-material: syntype workers, syntype queens (numbers not stated).
    • Type-localities: Russia: Archangel Prov., White Sea, Cape Keretsky, coast Bilago morya, 19.vi.1902 (B.M. Zhitkov), Archangel Prov., banks of Mezen, 30.vi.1902 (B.M. Zhitkov), Archangel Prov., Kuloi River, nr Dolgaya Shchel village, 2.vii.1902 (B.M. Zhitkov), Archangel Prov., River Kameni mouth, right bank river Kuloya, 23.vii.1902 (B.M. Zhitkov), Archangel Prov., Yazhmy River mouth, 23.vii.1902) (B.M. Zhitkov).
    • Type-depository: unknown (syntypes lost (Radchenko, 1997a: 555)).
    • [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).
    • Type-material: syntype workers (number not stated), 1 syntype queen.
    • Type-localities: workers Canada : Alberta, Ice River Valley, 4800-5000 ft (E. Whymper), queen Canada: Alberta, Banff, 4500 ft (E. Whymper).
    • [Note: Wheeler, W.M. 1910d: 331, incorrectly refers to Alberta specimens from Field, Vermilion Pass, and Lake Louise as “cotypes”; these localities are not mentioned in the original description but instead derive from Forel, 1904a: 152.]
    • Type-depository: MHNG.
    • 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.

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. Unless otherwise noted the text for the remainder of this section is reported from the publication that includes the original description.

Description

Mackay (2019):

Worker

Major worker 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 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.

Queen

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

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.

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