Formica sanguinea

Formica sanguinea is a slave-maker of many ants from the subgenus Serviformica. It uses two strategies to exploit its Serviformica hosts. The newly mated F. sanguinea queens usurp Serviformica nests and kill the resident Serviformica queens, thereby avoiding the costs of independent nest founding. Formica sanguinea workers from established colonies are also capable of raiding worker pupae from nearby Serviformica nests. This raiding can provide food and new Serviformica workers. After eclosion these slaves behave as regular members of the society, performing normal worker tasks and taking care of the slave-maker brood. Since F. sanguinea is a facultative social parasite, the workers are capable of nursing broods also independently and slaves are not found in all colonies. Queens may also start new colonies by fission. In Russia it is most common in dry pine forests and forest margins where large nests are constructed of plant material or soil (Haapaniemi and Pamilo 2015, Zryanin and Zryanina 2007) while in Greece it prefers mountain coniferous forests and in Achaia it has been observed inside humid fir forest and in border of mountain meadows, pastures and fir forest (Borowiec & Salata, 2021).

Identification
Mesosoma and front of head bright red; gaster and ocellar region of head brownish black. Proportion of dark to red varies from north to south with some arctic samples having proponderantly dark heads and dusky red mesosoma contrasting with southern samples which may have whole body excluding gaster bright red. Frontal triangle dull; eyes without hairs. Clypeus with median anterior notch variable in size and shape but always present. Head and mandibles broad relative to alitrunk and gaster. Length: 6.0-9.0 mm (Collingwood 1979).

Distribution
Throughout Eurasia from Portugal to Japan and Iran to Arctic Norway (Collingwood 1979). The southernmost known Japanese locality in Honshu is Mt. Fuji (Japanese Ant Image Database).

Distribution based on Regional Taxon Lists
Oriental Region: India, Pakistan. Palaearctic Region: Afghanistan, Albania, Andorra, Armenia, Austria, Balearic Islands, Belarus, Belgium, Bulgaria, Canary Islands, China, Croatia, Czech Republic, Democratic Peoples Republic of Korea, Denmark, Estonia, Finland, France, Georgia, Germany, Gibraltar, Greece, Hungary, Iberian Peninsula, Iran, Japan, Kazakhstan, Kyrgyzstan, Latvia, Lithuania, Luxembourg, Mongolia, Netherlands, Norway, Poland, Portugal, Republic of Korea, Republic of Macedonia, Republic of Moldova, Romania, Russian Federation, Slovakia, Slovenia, Spain, Sweden, Switzerland, Turkey, Ukraine, United Kingdom of Great Britain and Northern Ireland.

Biology
Collingwood (1979) - This is the well known aggressive red slave-making ant raiding nests of any species in its neighbourhood during the summer and removing pupae of the Formica fusca group of species for rearing in the parent nest as auxiliaries and also as food. All Fennoscandian species of the F. fusca group have been found in mixed colonies with F. sanguinea including Formica rufibarbis, Formica cinerea, Formica gagatoides and Formica transkaucasica as well as the more frequent Formica fusca or Formica lemani. F. sanguinea tends to avoid in-fighting but overwhelms other species by abrupt aggressive movements. Nests are situated under stones or in tree stumps with a small accumulation of leaf litter. It is often a dominant species in cleared woodland and in some localities all other Formica species have been eliminated and in such cases only pure F. sanguinea colonies are to be found, usually with a high proportion of small workers to act as nurses. Colonies spread by nest splitting and also by individual queens entering nests of the auxiliary species and appropriating a part of the brood, the host queen or queens being subsequently destroyed. Alatae are developed in July pairing often occurring in the vicinity of the nest.

Haapaniemi and Pamilo (2015) found workers of Formica fusca, Formica lemani, Formica gagatoides and Formica picea in the nests of Formica sanguinea they studied in Finland.

This species is known to suffer from labial gland disease, a condition caused by an (Elton, 1991).

Ant Hosts


The record of F. sanguinea being enslaved by Formica pratensis was based on the observation of a mixed colony and is likely an ephemeral association rather than true enslavement (Czechowski, 2001; de la Mora et al., 2021; Ruano et al., 2018; Seifert, 2018).

The record of Formica truncorum enslaving this species (Ruano et al., 2018) is unlikely based on biology (Seifert, pers. comm., in de la Mora et al., 2021).

Other Insects

 * This ant has been associated with the butterfly (Obregon et al. 2015).
 * This ant has been associated with the butterfly (Obregon et al. 2015).
 * This ant has been associated with the butterfly (Obregon et al. 2015).
 * This ant has been associated with the butterfly (Obregon et al. 2015).
 * This ant has been associated with the butterfly (Obregon et al. 2015).
 * This ant has been associated with the butterfly (Obregon et al. 2015).
 * This ant has been associated with the butterfly (Obregon et al. 2015).
 * This ant has been associated with the butterfly (Obregon et al. 2015).
 * This ant has been associated with the butterfly (Obregon et al. 2015).
 * This ant has been associated with the butterfly (Obregon et al. 2015).
 * This ant has been associated with the butterfly (Obregon et al. 2015).
 * This ant has been associated with the butterfly (Obregon et al. 2015).

Fungi

 * This taxon is a host for the fungi (Shrestha et al., 2017) and  (Espadaler & Santamaria, 2012).

Nomenclature

 *  sanguinea. Formica sanguinea Latreille, 1798: 37 (w.) FRANCE. Jurine, 1807: 272 (q.m.); Wheeler, G.C. & Wheeler, J. 1953c: 169 (l.); Imai, 1966: 119 (k.). Combination in F. (Raptiformica): Forel, 1913i: 361. Senior synonym of dominula: Smith, F. 1851: 115; Schenck, 1852: 36; Mayr, 1855: 337; Radchenko, 2007: 37; of arenicola, borea, clarior, fusciceps, griseopubescens, leninei, mollesonae, monticola, rotundata, strennua, tristis: Dlussky, 1965a: 16; of flavorubra: Dlussky, 1967a: 97; Collingwood, 1978: 74. Material of the unavailable name minuta referred here by Dlussky, 1965a: 16. See also: Wheeler, W.M. 1913f: 401; Donisthorpe, 1915d: 280; Stitz, 1939: 316; Dlussky, 1967a: 97; Tarbinsky, 1976: 186; Kutter, 1977c: 289; Collingwood, 1979: 137; Kupyanskaya, 1990: 191; Atanassov & Dlussky, 1992: 280.
 * dominula. Formica dominula Nylander, 1846a: 905, pl. 18, fig. 15 (w.q.m.) FINLAND. Junior synonym of sanguinea: Smith, F. 1851: 115; Schenck, 1852: 36; Mayr, 1855: 337; Radchenko, 2007: 37.
 * fusciceps. Formica sanguinea var. fusciceps Emery, 1895c: 335 (footnote) (w.) JAPAN. Junior synonym of sanguinea: Dlussky, 1965a: 16.
 * mollesonae. Formica sanguinea var. mollesonae Ruzsky, 1903c: 206 (w.) RUSSIA. Junior synonym of sanguinea: Dlussky, 1965a: 16.
 * clarior. Formica (Raptiformica) sanguinea var. clarior Ruzsky, 1905b: 420 (w.) CAUCASUS. Junior synonym of sanguinea: Dlussky, 1965a: 16.
 * flavorubra. Formica sanguinea var. flavorubra Forel, 1909c: 105 (w.) SPAIN. Junior synonym of sanguinea: Dlussky, 1967a: 97; Collingwood, 1978: 74.
 * borea. Formica (Raptiformica) sanguinea var. borea Santschi, 1925g: 351 (w.m.) FINLAND. Junior synonym of sanguinea: Dlussky, 1965a: 16.
 * strennua. Formica (Raptiformica) sanguinea var. strennua Santschi, 1925g: 352 (w.) SPAIN. [Unresolved junior primary homonym of strenua Haliday, above.] Junior synonym of sanguinea: Dlussky, 1965a: 16.
 * griseopubescens. Formica sanguinea var. griseopubescens Kuznetsov-Ugamsky, 1926b: 95 (w.) KAZAKHSTAN. Junior synonym of sanguinea: Dlussky, 1965a: 16.
 * monticola. Formica sanguinea subsp. monticola Kuznetsov-Ugamsky, 1926b: 95, figs. 3, 4 (w.) KAZAKHSTAN. [Unresolved junior primary homonym of monticola Buckley, 1866: 157, above.] Junior synonym of sanguinea: Dlussky, 1965a: 16.
 * rotundata. Formica sanguinea var. rotundata Kuznetsov-Ugamsky, 1926b: 95, fig. 2 (w.) KAZAKHSTAN. Raised to species: Dlussky, 1962: 182. Junior synonym of sanguinea: Dlussky, 1965a: 16.
 * clara. Formica (Raptiformica) sanguinea var. clara Karavaiev, 1927d: 345 (w.) UKRAINE. [Junior primary homonym of clara Forel, 1886f: 206.] Replacement name: leninei Santschi, 1928b: 46.
 * arenicola. Formica sanguinea subsp. arenicola Kuznetsov-Ugamsky, 1928b: 15 (w.) RUSSIA. [Unresolved junior primary homonym of arenicola Buckley, above.] Junior synonym of sanguinea: Dlussky, 1965a: 16.
 * leninei. Formica (Raptiformica) sanguinea var. leninei Santschi, 1928b: 46. Replacement name for clara Karavaiev, 1927d: 345. [Junior primary homonym of clara Forel, 1886f: 206.] Junior synonym of sanguinea: Dlussky, 1965a: 16.
 * tristis. Formica (Raptiformica) sanguinea var. tristis Karavaiev, 1929b: 217 (w.) RUSSIA. [Unresolved junior primary homonym of tristis Christ, above.] Junior synonym of sanguinea: Dlussky, 1965a: 16.

Description
Worker

Bingham (1903): Head, thorax and legs a clear light red, slightly fuscous on the head; abdomen fuscous brown or black. Head, thorax and abdomen dull and semiopaque, with a few scattered erect red hairs on the front of the head, on the underside and at the apex of the abdomen; pubescence extremely minute and thin, giving the insect a silky appearance. Head without the mandibles nearly square; mandibles strong, broad and very clearly dentate; clypeus convex, medially vertically carinate and incised or emarginate at the apex of the carina; antenna more massive than in F. truncicola, F. fusca, &c. ; head posteriorly distinctly widely emarginate. Thorax narrower than the head, the emargination at the meso-metanotal suture deep and broad. Node of the pedicel sub-triangular, slightly biconvex, very broad and transverse above, the lateral angles rounded; abdomen depressed, broad, as broad as long.

Length: 6 - 7 mm

Queen

Bingham (1903): Very similar to the worker but larger, the head, thorax and abdomen a lighter brighter yellow, the abdomen slightly darker, in greater contrast to the colour of the head and thorax.

Length: 9 - 9.5 mm

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