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).

Distribution based on Regional Taxon Lists
Nearctic Region: Canada, United States. Palaearctic Region: Andorra, Armenia, Austria, 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, Republic of 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 dolichurus (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.

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.

Nomenclature

 *  herculeanus. Formica herculeana Linnaeus, 1758: 579 (q.) EUROPE. Lepeletier de Saint-Fargeau, 1835: 209 (w.q.m.); Mayr, 1855: 308 (w.q.m.); Wheeler, G.C. & Wheeler, J. 1953e: 185 (l.). Combination in Camponotus: Mayr, 1861: 36; in C. (Camponotus): Forel, 1914a: 259. Senior synonym of atra, intermedia: Nylander, 1846a: 894; of castanea: Mayr, 1863: 399; of whymperi: Creighton, 1950a: 367; of nadigi: Yasumatsu & Brown, 1951: 30; of montanus, shitkowi: Emery, 1925b: 72; Karavaiev, 1936: 178; Arnol'di, 1967: 1819; of caucasicus: Arakelian, 1994: 85; of eudokiae: Radchenko, 1997a: 555. See also: Tarbinsky, 1976: 148; Kutter, 1977c: 204; Collingwood, 1979: 90; Smith, D.R. 1979: 1426; Atanassov & Dlussky, 1992: 210.
 * castanea. Formica castanea Lepeletier de Saint-Fargeau, 1835: 215 (w.q.m.) U.S.A. [Unresolved junior primary homonym of castanea Latreille, above.] Junior synonym of herculeanus: Mayr, 1863: 399.
 * atra. Formica atra Zetterstedt, 1838: 450 (m.) SWEDEN. Junior synonym of herculeanus: Nylander, 1846a: 894.
 * intermedia. Formica intermedia Zetterstedt, 1838: 448 (w.) SWEDEN. Junior synonym of herculeanus: Nylander, 1846a: 894.
 * whymperi. Camponotus herculeanus var. whymperi Forel, 1902i: 699 (w.q.) CANADA. Wheeler, W.M. 1910d: 330 (s.m.). Raised to species: Ruzsky, 1926: 108. Subspecies of herculeanus: Ruzsky, 1936: 90. Junior synonym of herculeanus: Creighton, 1950a: 367.
 * montanus. Camponotus herculeanus var. montanus Ruzsky, 1904a: 293 (w.) RUSSIA. Karavaiev, 1926e: 192 (s.w.q.). [Unresolved junior primary homonym of montanus Emery, above.] Subspecies of herculeanus: Ruzsky, 1926: 108. Junior synonym of herculeanus: Emery, 1925b: 72; Karavaiev, 1936: 178.
 * shitkowi. Camponotus herculeanus var. shitkowi Ruzsky, 1904a: 292 (w.q.) RUSSIA. Junior synonym of herculeanus: Emery, 1925b: 72; Karavaiev, 1936: 178; Arnol'di, 1967: 1819.
 * nadigi. Camponotus herculeanus var. nadigi Menozzi, 1922c: 142 (s.w.q.m.) ITALY. Material of the nomen nudum vagusherculeanus referred here by Emery, 1925b: 72; Kutter, 1977c: 205. Junior synonym of herculeanus: Yasumatsu & Brown, 1951: 30; Baroni Urbani, 1971c: 176.
 * eudokiae. Camponotus herculeanus var. eudokiae Ruzsky, 1926: 108 (w.) RUSSIA. Junior synonym of herculeanus: Radchenko, 1997a: 555.
 * caucasicus. Camponotus herculeanus subsp. caucasicus Arnol'di, 1967: 1822 (s.w.q.m.) CAUCASUS. Junior synonym of herculeanus: Arakelian, 1994: 85.
 * 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: Chapman & Capco, 1951: 221.
 * Junior synonym of saxatilis: Kuznetsov-Ugamsky, 1928b: 4; Kuznetsov-Ugamsky, 1929a: 18.
 * Status as species: Collingwood, 1981: 29; Bolton, 1995b: 121; Collingwood & Heatwole, 2000: 12; Imai, et al. 2003: 39.
 * Subspecies of herculeanus: Ruzsky, 1905b: 222; Wheeler, W.M. 1906c: 325; Yano, 1910: 422; Karavaiev, 1912b: 592; Ruzsky, 1926: 108; Arnol'di, 1967: 1821; Pisarski, 1969a: 230; Pisarski, 1969b: 304; Dlussky & Pisarski, 1970: 86; Collingwood, 1976: 306; 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.
 * Senior synonym of altaica: Arnol'di, 1967: 1821.
 * Senior synonym of jacuticus: Arnol'di, 1967: 1821.
 * Junior synonym of herculeanus: Yasumatsu & Brown, 1957: 49; Schar et al., 2018: 6.
 * altaica. Camponotus herculeanus subsp. altaica Ruzsky, 1926: 108 (w.) RUSSIA.
 * [First available use of Camponotus herculeanus saxatilis var. altaica Ruzsky, 1915b: 6; unavailable name.]
 * Subspecies of saxatilis: Ruzsky, 1936: 90.
 * Junior synonym of sachalinensis: Arnol'di, 1967: 1821.
 * Junior synonym of herculeanus: Schar et al., 2018: 6.
 * jacuticus. Camponotus (Camponotus) herculeanus var. jacuticus Karavaiev, 1929b: 210 (s.w.q.m.) RUSSIA.
 * Junior synonym of sachalinensis: Arnol'di, 1967: 1821.
 * Junior synonym of herculeanus: Yasumatsu & Brown, 1951: 30; Schar et al., 2018: 6.

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.

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