Formica lugubris

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Formica lugubris
Conservation status
Scientific classification
Kingdom: Animalia
Phylum: Arthropoda
Class: Insecta
Order: Hymenoptera
Family: Formicidae
Subfamily: Formicinae
Tribe: Formicini
Genus: Formica
Species: F. lugubris
Binomial name
Formica lugubris
Zetterstedt, 1838

Formica lugubris casent0127733 profile 1.jpg

Formica lugubris casent0127733 dorsal 1.jpg

Specimen labels

Synonyms

This ant was found by Boulay et al. (2007) to be one of two important ant dispersers (also Camponotus cruentatus) of myrmecochorous seeds of the plant Helleborus feotidus. F. lugubris was responsible for 67% of the observed visits and 80% of the observed seed removals from a population from northwestern Spain.

At a Glance • Facultatively polygynous  • Temporary parasite  • Diploid male  

Identification

A sibling species of Formica paralugubris, these two species can be separated using queen pilosity characters and morphometrics (tables and discriminant functions to separate these two species and Formica aquilonia are provided by Seifert 1996).

Collingwood (1979) - Bicoloured with distinct but not well demarcated dark patch on promesonotum. Frontal groove distinctly shining. Large punctures coarse and deep, widely dispersed among close set microscopic puncturation. Occiput with a thick fringe of hairs extending forward over area between ocelli and sides of head and laterally round to the eyes. Eye hairs erect and prominent. Body pilosity including gula, tibiae and femora more or less densely pilose. Some populations have scape hairs. Head width of largest workers 2.1 mm. Length: 4.5-9.0 mm.

Keys including this Species

Distribution

Northern Eurosiberia and European mountains from Pyrenees to Kamchatka and Japan, Italy to North Norway (Collingwood 1979).

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: 69.687618° to 37.91362°.

 
North
Temperate 		North
Subtropical 		Tropical South
Subtropical 		South
Temperate

Distribution based on Regional Taxon Lists

Palaearctic Region: Albania, Andorra, Austria, Belarus, Bulgaria, China, Croatia, Czechia, Democratic Peoples Republic of Korea, Estonia, Finland, France, Germany, Greece, Iberian Peninsula, Ireland, Italy, Latvia, Mongolia, Montenegro, Norway (type locality), Poland, Republic of Korea, Romania, Russian Federation, Serbia, Slovakia, Slovenia, Spain, Sweden, Switzerland, Türkiye, Ukraine, United Kingdom of Great Britain and Northern Ireland.

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

Collingwood (1979) - This is a robust active species. Colonies are often in groups with inter-connecting nests. It has similar habits to Formica rufa but is able to forage at much lower temperatures and replaces F. rufa entirely from Central Fennoscandia to the far north. This species varies in the presence, abundance or absence of scape hairs in the female castes and some local populations in South Finland and in the Alps with such hairs have widely spaced micropunctures on the dorsum of the gaster as in F. rufa. Because of great variability among local populations in these areas it has not been possible to demarcate the extreme forms as a separate species but samples mainly from coastal areas and offshore islands in Nylandia include some extremely hairy specimens with queens consistently having wide spaced micropunctures which are well outside the range of F. lugubris as described by Yarrow (1955) and Betrem (1960). Bondroit (1917) briefly described a form, F. rufa var. nylanderi, as having long outstanding body and antennal hairs and F. nylanderi could be a suitable name for this form, if distinguished as a species.

F. lugubris spreads by colony fission but also by the adoption of fertile queens by Formica lemani. Such mixed incipient nests often under stones have frequently been seen in Norway and North Sweden (Collingwood, 1959).

Foraging/Diet

Formica lugubris collect large quantities of honeydew.

Novgorodova (2015b) investigated ant-aphid interactions of a dozen honeydew collecting ant species in Western Siberia pine and aspen-birch-pine forests (54°7´N, 83°06´E, 200 m, Novosibirsk) and mixed-grass-cereal steppes with aspen-birch groves (53°44´N, 78°02´E, 110 m, near Karasuk) in the Novosibirsk Region and coniferous forests in the northeastern Altai (north end of Lake Teletskoe, 51°48´N, 87°17´E, 434 m). All of the ants studied had workers that showed high fidelity to attending particular aphid colonies, i.e, individual ants tend to return to the same location, and group of aphids, every time they leave the nest. F. lugubris' honeydew collecting activities were highly coordinated during the summer months when the aphids and ants were most active. Individual foragers specialized on specific tasks and could be classified as shepherds (collect honeydew), guards (protect aphids from competitors), scouts (search for new aphid colonies) and transporters (transport honeydew to the nest). Individuals performed the same type of work day after day, with groups of the same workers, thereby forming teams. F. lugubris tended: Symydobius oblongus (Heyden) and Cinara laricis (Hartig).

Regional Notes

Borowiec and Salata (2022), for Greece - Few records come from coniferous forest. Nest in form of small mound to 110 cm in diameter. Mountain species, collected in area from an altitude 700 to 2200 m.

Nesting Habits

This species is polydomous and is considered to be a member of a Formica species group known as wood ants. Ellis and Robinson (2015) conducted a 3 year field-study of a population (2012-2014, Peak District, England) of Formica lugubris to ascertain the potential benefit of non-foraging nests. Abstract - A colony of red wood ants can inhabit more than one spatially separated nest, in a strategy called polydomy. Some nests within these polydomous colonies have no foraging trails to aphid colonies in the canopy. In this study we identify and investigate the possible roles of non-foraging nests in polydomous colonies of the wood ant Formica lugubris. To investigate the role of non-foraging nests we: (i) monitored colonies for three years; (ii) observed the resources being transported between non-foraging nests and the rest of the colony; (iii) measured the amount of extra-nest activity around non-foraging and foraging nests. We used these datasets to investigate the extent to which non-foraging nests within polydomous colonies are acting as: part of the colony expansion process; hunting and scavenging specialists; brood-development specialists; seasonal foragers; or a selfish strategy exploiting the foraging effort of the rest of the colony. We found that, rather than having a specialised role, non-foraging nests are part of the process of colony expansion. Polydomous colonies expand by founding new nests in the area surrounding the existing nests. Nests founded near food begin foraging and become part of the colony; other nests are not founded near food sources and do not initially forage. Some of these non-foraging nests eventually begin foraging; others do not and are abandoned. This is a method of colony growth not available to colonies inhabiting a single nest, and may be an important advantage of the polydomous nesting strategy, allowing the colony to expand into profitable areas.

This species is known to suffer from labial gland disease, a condition caused by an unknown agent, in Spain (Espadaler & Riasol, 1981; Elton, 1991).

Association with Other Organisms

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Formica lugubris is a temporary parasite of:

Other associations include:

  • This species is a xenobiont for the ant Formica fusca (a xenobiont) in Finland (Czechowski, 2004; Kanizsai et al., 2013) (Different successional series of rocky habitats. Mound of F. lugubris).
  • This species is a xenobiont for the ant Formicoxenus nitidulus (a xenobiont) (Holldobler & Wilson 1990; Busch 2001; Martin et al. 2007).
  • This species is a host for the beetle Monotoma angusticollis (Coleopotera: Monotomidae) (a myrmecophile) in Europe (Wagner et al., 2020).
  • This species is a host for the trematode Dicrocoelium dendriticum (a parasitoid) (Quevillon, 2018) (encounter mode primary; indirect transmission; transmission outside nest).
  • This species is a host for the fungus Aegeritella superficialis (a pathogen) (Espadaler & Santamaria, 2012).

The reported host/parasite relationship between Formica lugubris and Formica cinerea should be investigated in the field as Chernenko et al. (2013) found 100% F. lugubris queen mortality in lab introductions (de la Mora et al., 2021).

Hempitera

  • This species is a mutualist for the aphid Cinara pini (a trophobiont) (Sudd, 1983; Saddiqui et al., 2019).

Flight Period

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

Source: antkeeping.info.

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Life History Traits

  • Queen number: facultatively polygynous
  • Mean colony size: 34,600 (Breen, 1979) (5 nests examined by Breen (1979), with range of 10,000-70,000)
  • Maximum colony size: 70,000 (Breen, 1979)
  • Foraging behaviour: trunk trail (Rosengren, 1971; Breen, 1979; Beckers et al., 1989)

Castes

Worker

Images from AntWeb

Formica lugubris casent0173010 head 1.jpgFormica lugubris casent0173010 profile 1.jpgFormica lugubris casent0173010 dorsal 1.jpgFormica lugubris casent0173010 label 1.jpg
Worker. Specimen code casent0173010. Photographer April Nobile, uploaded by California Academy of Sciences. Owned by MCZ, Cambridge, MA, USA.
Formica lugubris casent0173155 head 1.jpgFormica lugubris casent0173155 profile 1.jpgFormica lugubris casent0173155 dorsal 1.jpgFormica lugubris casent0173155 label 1.jpg
Worker. Specimen code casent0173155. Photographer April Nobile, uploaded by California Academy of Sciences. Owned by CAS, San Francisco, CA, USA.

Queen

Images from AntWeb

Formica lugubris casent0173009 head 1.jpgFormica lugubris casent0173009 profile 1.jpgFormica lugubris casent0173009 dorsal 1.jpgFormica lugubris casent0173009 label 1.jpg
Queen (alate/dealate). Specimen code casent0173009. Photographer April Nobile, uploaded by California Academy of Sciences. Owned by MCZ, Cambridge, MA, USA.

Male

Images from AntWeb

Formica lugubris casent0173011 head 1.jpgFormica lugubris casent0173011 profile 1.jpgFormica lugubris casent0173011 profile 2.jpgFormica lugubris casent0173011 dorsal 1.jpgFormica lugubris casent0173011 dorsal 2.jpgFormica lugubris casent0173011 label 1.jpg
Worker. Specimen code casent0173011. Photographer April Nobile, uploaded by California Academy of Sciences. Owned by MCZ, Cambridge, MA, USA.

Diploid males are known to occur in this species (Pamilo et al., 1994; Cournault & Aron, 2009).

Nomenclature

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

  • lugubris. Formica lugubris Zetterstedt, 1838: 449 (m.) NORWAY. Junior synonym of rufa: Nylander, 1856b: 60; Emery & Forel, 1879: 450; Wheeler, W.M. 1913f: 425; Emery, 1925b: 253; Stitz, 1939: 328. Revived from synonymy and status as species: Yarrow, 1955a: 5; Betrem, 1960b: 77; Dlussky, 1967a: 91; Dlussky & Pisarski, 1971: 180; Baroni Urbani, 1971c: 218; Kutter, 1977c: 271; Gösswald, 1989: 19; Kupyanskaya, 1990: 198; Atanassov & Dlussky, 1992: 274. Senior synonym of congerens: Yarrow, 1955a: 5; Dlussky, 1967a: 91; Radchenko, 2007: 36; of nylanderi, santschii: Yarrow, 1955a: 5; of montana Sadil: Samsinak, 1964: 157; of unicolor: Dlussky, 1967a: 91; Dlussky & Pisarski, 1971: 180. Material of the unavailable name tir referred here by Yarrow, 1955a: 5.
  • congerens. Formica congerens Nylander, 1846a: 906 (w.) FINLAND. Nylander, 1849: 30 (m.); Foerster, 1850a: 17 (q.). Junior synonym of pratensis: Emery & Forel, 1879: 450; Nasonov, 1889: 17; Wheeler, W.M. 1913f: 428; Forel, 1915d: 57; Emery, 1916b: 256; Müller, 1923: 142. Revived from synonymy: Betrem, 1953: 324. Junior synonym of lugubris: Yarrow, 1955a: 5; Dlussky, 1967a: 91; Radchenko, 2007: 36.
  • alpina. Formica rufa var. alpina Santschi, 1911j: 349 (w.) ITALY. [Junior primary homonym of alpina Wheeler, above.] Replacement name: santschii Wheeler, 1913f: 428. Raised to species: Bondroit, 1918: 59.
  • santschii. Formica rufa var. santschii Wheeler, W.M. 1913f: 390 (in key). Replacement name for alpina Santschi, 1911j: 349. [Junior primary homonym of alpina Wheeler, W.M. 1909e: 85.] Junior synonym of lugubris: Yarrow, 1955a: 5.
  • nylanderi. Formica rufa var. nylanderi Bondroit, 1920a: 145 (q.) FRANCE. [Also described as new by Bondroit, 1920b: 300.] Junior synonym of lugubris: Yarrow, 1955a: 5; Seifert, 1996: 200.
  • unicolor. Formica pratensis subsp. unicolor Ruzsky, 1926: 110 (w.) RUSSIA. [First available use of Formica rufa subsp. pratensis var. unicolor Ruzsky, 1914b: 102; unavailable name.] Junior synonym of lugubris: Dlussky, 1967a: 91.
  • montana. Formica rufa var. montana Sadil, 1953b: 198, fig. 1 (q.) CZECHOSLOVAKIA. [Unresolved junior primary homonym of montana Wheeler, W.M., above.] Junior synonym of lugubris: Samsinak, 1964: 157.

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

Description

Worker

Borowiec and Salata (2022) - Very large, HL: 1.270-2.160 (mean 1.705); HW: 1.048-1.960 (mean 1.449); SL: 1.048-1.740 (mean 1.406); EL: 0.321-0.587 (mean 0.444); ML: 1.72-2.97; MW: 0.78-1.41. Color. Head bicolours, clypeus, genae, sides behind eyes and ventral side yellowish red to red, rest of surface brown to black, yellowish red of anterior part of head gradually turn into the dark posterior part of head without sharp border between pale and dark parts, sometimes brown limited only to frons and frons; mesosoma rarely uniformly yellowish red to red, usually promesonotum at top with obscure spot with diffused borders but yellow or reddish always predominate, petiolar scale usually yellowish red to red, gaster brown to black with transparent white posterior margin of tergites, anterior slope of first tergite often yellowish brown, antennal scapi from yellowish brown to brown, funicle from yellowish brown to dark brown, legs from uniformly reddish to indistinctly infuscated, in the darkest form completely brown. Head. Broad, 1.1-1.2 times longer than wide, in front of eyes softly converging anterad, behind eyes softly rounded, occipital margin straight to slightly convex. Clypeus without or with obtuse median keel, on the whole surface distinctly microsculptured, slightly trapezoidal, its anterior margin convex, sides convergent posterad, posterior margin truncate, whole clypeal surface with very short and sparse appressed pubescence, without or with a row of 8-10 short setae close to the anterior margin and group of 2-3 long erected setae in lateral corners, rest of clypeal surface without or with 1-2 pairs of long erected setae, the longest anterior seta with length 0.238. Head distinctly microreticulate, appears mostly dull and opaque but at least partly with a mild silky shine, with very short and very sparse appressed pubescence not covering head surface, interocular and ocellar area usually with few moderately long, erected yellow setae, occasionally frons without setae, gena without setae, whole occipital area or only in lateral corners with erected setae, ventral side of head with several long, erected setae. Scape short, 0.9-1.0 times as long as width of head, thin, distinctly reaching beyond the occipital margin, distinctly, regularly widened from base to apex, its surface microreticulate, with short and dense appressed pubescence, erected setae absent. Funicular segments elongate, thin, first segment 1.5 times as long as second segment, the second segment 1.9-2.0 times as long as wide, not or only slightly shorter than third segment, the rest of funicular segments clearly longer than broad . Eyes big, elongate oval, approximately 0.26 length of head. Mesosoma. Elongate in dorsal view distinctly constricted in the middle, 2.1-2.4 times as long as wide, dorsally and laterally distinctly microreticulated, surface indistinctly dull and opaque. In lateral view promesonotum convex, mesonotal groove deep, propodeum strongly, obtusely convex. Whole mesosomal surface covered with moderately long and moderately dense appressed pubescence not covering the mesosomal surface, usually on dorsum with numerous short, erected setae, the longest with length 0.127 but setation varies in and between nests, sometimes is reduced to 1-3 setae on pronotum and 0-5 setae on mesonotum and propodeum and number of setae is not correlated with size of worker. Waist and gaster. Petiolar scale broad, moderately thick in lateral view, apex rounded or truncate but usually with very shallow median emargination, anterior sides usually with several short, erected setae. All tergites close to posterior margin without or with only few short setae, surface of tergites in Greek populations lacking erected setae or with only few setae, only first tergite on anterior slope always with numerous short to moderately long erected setae. Legs. Ventral surface of both fore and mid femora with row of 2-10 short erected setae.

Karyotype

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  • n = 26, 2n = 52 (Finland; Switzerland) (Hauschteck-Jungen & Jungen, 1976; Rosengren et al., 1980).

References

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
  • AntArea. Accessed on February 5th 2014 at http://antarea.fr/fourmi/
  • Antarea (Personal Communication - Rumsais Blatrix- 27 April 2018)
  • Antarea (at www.antarea.fr on June 11th 2017)
  • Antonov I. A. 2012. Ant complexes of Baikalsk town. The Bulletin of Irkutsk State University 4: 143-146.
  • Antonov I. A. 2013. Ant Assemblages (Hymenoptera: Formicidae) of Cities of the Temperate Zone of Eurasia. Russian Journal of Ecology 44(6): 523–526.
  • 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)
  • Asociacion Iberica de Mirmecologia. 2011. List of species collected during the Taxomara Lisboa 2011. Iberomyrmex 3: 30-31.
  • Astric A., and H. Callot. 2006. Presence d'Anergates atratulus (Schenck, 1852) et Strongylognathus testaceus (Schenck, 1852) en Alsace. Bull. Soc. Ent. Mulhouse 62(2): 19-21.
  • Banert P, and B. Pisarski. 1972. Mrówki (Formicidae) Sudetów. Fragmenta Faunistica (Warsaw) 18: 345-359.
  • Baroni Urbani C., and C. A. Collingwood. 1976. A Numerical Analysis of the Distribution of British Formicidae (Hymenoptera, Aculeata). Verhandlungen der Naturforschenden Gesellschaft in Basel 85: 51-91.
  • Baroni Urbani C., and C. A. Collingwood. 1977. The zoogeography of ants (Hymenoptera, Formicidae) in Northern Europe. Acta Zoologica Fennica 152: 1-34.
  • Barrett K. E. J. 1970. Ants in France, 1968-69. Entomologist 103: 270-274.
  • 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., 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. 1967. Faune de l'Europe et du Bassin Méditerranéen. 3. Les fourmis (Hymenoptera Formicidae) d'Europe occidentale et septentrionale. Paris: Masson, 411 pp.
  • Bernasconi C., A. Maeder, A. Freitag and D. Cherix. 2006. Formica paralugubris (Hymenoptera: Formicidae) in the Italian Alps from new data and old data revisited. Myrmecologische Nachrichten. 8: 251-256
  • Bezdecka P., and K. Bezdeckova. 2012. Updated list of the ants of the Czech Republic. Pp 7-12. Bezd??ka P. & Bezd??ková K. (eds) 2012: Blanok?ídlí v ?eských zemích a na Slovensku 8, Chaloupky, 1.-3. ?ervna 2012, sborník abstrakt? z konference. – MVJ Jihlava, 37 pp.
  • 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.
  • Blinov V. V. 1984. New for the fauna of Byelorussia ant species. Vestsi Akademii Navuk BSSR. Seryia Biialahichnykh Navuk 1984(5): 113-115.
  • 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.
  • 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.
  • Carniel A. 1988. Contributo alla conoscenza della mirmecofauna del Cansiglio (Prealpi Carniche). Boll. Soc. Ent. Ital., Genova 119(3): 179-190.
  • Carniel A. 1998. Ricerche sulla mirmecofauna delle Prealpi Orobiche (Lombardia) (Insecta, Hymenoptera, Formicidae). Atti. Mus. Civ. Stor. Nat. Morbegno 9: 29-39.
  • Casevitz-Weulersse J., and C. Galkowski. 2009. Liste actualisee des Fourmis de France (Hymenoptera, Formicidae). Bull. Soc. Entomol. Fr. 114: 475-510.
  • Ceballos, P., and G. Ronchetti. "Le formiche del gruppo Formica rufa sui Pirenei orientali spagnoli, nelle province di Lerida e Gerona." Memorie della Società Entomologica Italiana 45 (1966): 153-168.
  • 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.
  • Collingwood C. A. 1951. The distribution of ants in north-west Scotland. Scottish Naturalist 63: 45-49
  • Collingwood C. A. 1951. The distribution of ants in north-west Scotland. Scottish Naturalist 63: 45-49.
  • Collingwood C. A. 1956. Ant hunting in France. Entomologist 89: 106-108.
  • Collingwood C. A. 1961. Ants in the Scottish Highlands. Scotish Naturalist 70: 12-21.
  • 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. A., and J. Hughes. 1987. Ant species in Yorkshire, England. Naturalist (Leeds) 112: 95-101.
  • Collingwood C.A. 1959. Ants in the Scottish Highlands. The Scottish Naturalist. 70: 12-21
  • Collingwood C.A. 1959. Scandinavian Ants. Entomol. Rec. 71: 78-83
  • Collingwood C.A. 1961. Ants in Finland. Entomol. Rec. 73: 190-195
  • Collingwood C.A. 1961. New Vice-County Records for British Ants. Entomologist. 73: 90-93
  • Collingwood C.A. and Satchell J.E. 1956. The Ants of the South Lake District. Journal of the Society for British Entomology. 5: 159-164
  • Collingwood, C. A. 1958b. A key to the species of ants (Hymenoptera, Formicidae) found in Britain. Trans. Soc. Br. Entomol. 13: 69-96
  • Collingwood, C. A. 1964. The Identification of British Ants (Hym. Formicidae). Transactions of the Society for British Entomology. 16:93-121.
  • Collingwood, C. A. 1974. A revised list of Norwegian ants (Hymenoptera: Formicidae). Norsk Entomologisk Tidsskrift 21: 31-35.
  • Collingwood, C. A., and I. H. H. Yarrow. "A survey of Iberian Formicidae." EOS (Revista española de entomología) 44 (1969): 53-101.
  • Collingwood, C. A.. "The Formicidae (Hymenoptera) of Fennoscandia and Denmark." Fauna Entomologica Scandinavica 8 (1979): 1-174.
  • Collingwood, C.A. 1958. A survey of Irish Formicidae. Proceedings of the Royal Irish Academy 59B:213-219
  • 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
  • Donisthorpe H. 1914. Myrmecophilous notes for 1913. Entomologist's Record and Journal of Variation 26: 37-45.
  • 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.
  • Else G., B. Bolton, and G. Broad. 2016. Checklist of British and Irish Hymenoptera - aculeates (Apoidea, Chrysidoidea and Vespoidea). Biodiversity Data Journal 4: e8050. doi: 10.3897/BDJ.4.e8050
  • Entomological Society of Latvia. 2003. http://leb.daba.lv/Formicidae.htm (Accessed on December 1st 2013).
  • 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.. "Contribución al conocimiento de los formícidos (Hymenoptera, Formicidae) del Pirineo catalán." Tesis Universida (1979): 285 pp.
  • Field Museum Collection, Chicago, Illinois (C. Moreau)
  • Fowles, A.P. 1996. A provisional checklist of the invertebrates recorded from Wales. 2. Aculeate wasps, bees and ants (Hymenoptera: Aculeata). Countryside Council for Wales
  • Gaspar C. 1968. Les fourmis de la Drome et des Basses-Alpes, en France (Hymenoptera, Formicidae). Naturaliste can. 95: 747-766.
  • 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.
  • Gjershaug J. O., F. Odegaard, A. Staverlokk, and K. M. Olsen. 2016. Records of bilateral gynandromorphism in three species of ants (Hymenoptera, Formicidae) in Norway. Norwegian. Journal of Entomology 63: 65–70.
  • Glaser F. 2009. Die Ameisen des Fürstentums Liechtenstein. (Hymenoptera, Formicidae). Amtlicher Lehrmittelverlag, Vaduz, 2009 (Naturkundliche Forschung im Fürstentum Liechtenstein; Bd. 26).
  • Glaser, F., M. J. Lush, and B. Seifert. "Rediscovered after 140 years at two localities: Myrmica myrmicoxena Forel, 1895 (Hymenoptera: Formicidae)." Myrmecological News 14 (2010): 107-111.
  • Gulzar A. 2014. Classification and Distribution of ants in Kashgar, Xinjiang. Master's Thesis Shaanxi Normal University, 75 pages.
  • Guénard B., and R. R. Dunn. 2012. A checklist of the ants of China. Zootaxa 3558: 1-77.
  • Heatwole H. 1989. Changes in ant assemblages across an artic treeline. Revue d'Entomologie du Quebec 34(1-2): 10-22.
  • Huang Ren-Xing, Ouyang Tong, Wu Wei, and Fan Zhao-tian. 2004. Forty two new record species of Family Formicidae (Hymenoptera: Formicoidea) from Xinjiang, China. Entomotaxonomia 26 (2): 156-160.
  • Hågvar S. 2005. Altitudinal zonation of ants (Formicidae) in a steep fjord landscape in Sogndal, Western Norway. Norw. J. Entomol. 52: 3-12.
  • Kim B., S. Park, J. Kim, K. Kim. 1997. Ants from Mount Moak (Hymenoptera: Formicidae). Korean Journal of Soil Zoology 2(1): 41-58.
  • Kim B.J. 1996. Synonymic list and distribution of Formicidae (Hymenoptera) in Korea. Entomological Research Bulletin Supplement 169-196.
  • Kim B.J., S.J. Park, and J.H. Kim. 1996. Ants from Naejangsan national park (Hymenoptera: Formicidae). Korean J. Soil. Zoology &(2): 120-133.
  • Kofler A. 1995. Nachtrag zur Ameisenfauna Osttirols (Tirol, Österreich) (Hymenoptera: Formicidae). Myrmecologische Nachrichten 1: 14-25.
  • Kupianskaia A.N. 1990. Murav'I (Hymenoptera, Formicidae) Dal'nego Vostoka SSSR (1989). Vladivostok. 258 pages.
  • Kvamme T. 1982. Atlas of the Formicidae of Norway (Hymenoptera: Aculeata). Insecta Norvegiae 2: 1-56.
  • Lapeva-Gjonova, L., V. Antonova, A. G. Radchenko, and M. Atanasova. "Catalogue of the ants (Hymenoptera, Formicidae) of Bulgaria." ZooKeys 62 (2010): 1-124.
  • Le Moli F., and A. Zaccone. 1995. Ricerche sulla mirmecofauna del Cansiglio (Prealpi Carniche). Soc. Ven. Sc. Nat. 20: 33-52.
  • 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 Collection Database
  • 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.
  • Martin, S.J., E.A. Jenner and F.P. Drijfhout. 2007. Chemical Deterrent Enables a Socially Parasitic Ant to Invade Multiple Hosts. Proceedings: Biological Sciences 274(1626):2717-2721
  • 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. 2011. A check list of Danish ants and proposed common names. Ent. Meddr. 79: 13-18.
  • Nylander, W.. "Synopsis des Formicides de France et d'Algérie." Annales des Sciences Naturelles, Zoologie (4)5 (1856): 51-109.
  • Paik W.H. 1984. A checklist of Formicidae (Hymenoptera) of Korea. Korean J. Plant Prot. 23(3): 193-195.
  • 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.
  • Petrov I. Z., and C. A. Collingwood. 1992. Survey of the myrmecofauna (Formicidae, Hymenoptera) of Yugoslavia. Archives of Biological Sciences (Belgrade) 44: 79-91.
  • Poldi B., M. Mei, and F. Rigato. 1995. Hymenoptera, Formicidae. Vol 102. Checklist delle specie della fauna Italiana: 1-10.
  • Punttila P., Y. Haila, and H. Tukia. 1996. Ant communities in taiga clearcuts: habitat effects and species interactions. Ecography 19: 16-28.
  • 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.
  • Radchenko A. G. 2007. The ants (Hymenoptera, Formicidae) in the collection of William Nylander. Fragmenta Faunistica (Warsaw) 50: 27-41.
  • Radchenko, A. 2005. Monographic revision of the ants (Hymenoptera: Formicidae) of North Korea. Annales Zoologici (Warsaw) 55: 127-221.
  • Radchenko, A. 2005. Monographic revision of the ants (Hymenoptera, Formicidae) of North Korea. Annales Zoologici 55(2): 127-221.
  • Ran H., and S. Y. Zhou. 2012. Checklist of chinese ants: formicomorph subfamilies (Hymenoptera: Formicidae) II. Journal of Guangxi Normal University: Natural Science Edition 30(4): 81-91.
  • 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.
  • Santschi, F.. "Une nouvelle variété de Formica rufa L. (Hym. Formicidae)." Bulletin de la Société Entomologique de France 1911 (1911): 349-350.
  • Scupola A. 2016. First records of the ant genus Formica Linnaeus 1758 (Hymenoptera Formicidae) from the coastal pine woods of Salento Peninsula (Apulia, southern Italy). Thalassia Salentina 38: 137-141.
  • Seifert B. 1996. Formica paralugubris nov. spec. - a sympatric sibling species of Formica lugubris from the western Alps (Insecta: Hymenoptera: Formicoidea: Formicidae). Reichenbachia 31: 193-201.
  • Seifert B. 1998. Rote Liste der Ameisen. - in: M. Binot, R. Bless, P. Boye, H. Gruttke und P. Pretscher: Rote Liste gefährdeter Tiere Deutschlands. Bonn-Bad Godesberg 1998: 130-133.
  • Seifert B. 2007. Die Ameisen Mittel- und Nordeuropas. Tauer: lutra Verlags- und Vertriebsgesellschaft, 368 pp.
  • Siberian Zoological Museum. Website available at http://szmn.sbras.ru/old/Hymenop/Formicid.htm. Accessed on January 27th 2014.
  • 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. 2004. Geological and ecological distribution of Japanese ants communities. (translated from Japanese) Reports of the Saitama Prefecture Animal Research Association. 48:24
  • Vagalinski B., and A. Lapeva-Gjonova. 2012. The ants (Hymenoptera, Formicidae) of Plana Mountain (Bulgaria). Historia naturalis bulgarica 20: 87-101.
  • Vesni? A. 2011: Revidirani sistematski prijegled mrava Bosne i Hercegovine. Unutar : S. Lelo (urednik), Fauna Bosne i Hercegovine – Biosistematski pregledi. 7. izmijenjeno i popravljeno interno izdanje Udruženja za inventarizaciju i zaštitu životinja, Ilijaš, Kanton Sarajevo, pp: 205-207.
  • Vespalainen K., B. Pisarski, R. Kantorek, and K. J. Laine. 1984. Formicidae (Hymenoptera) of Inari Lapland. Kevo Notes 7: 115-116.
  • Wegnez P., D. Ignace, E. Lommelen, M. Hardy, J. Bogaert, and C. Nilsson. 2015. Redécouverte de Teleutomyrmex schneideriKutter, 1950 dans les Alpes françaises (Hymenoptera: Formicidae). Bulletin de la Société royale belge d’Entomologie 151: 52-57.
  • Wegnez P., S. De Greef, C. Degache, D. Ignace, and W. Dekoninck. 2011. Observations recentes de la fourmi Formicoxenus nitidulus (NYLANDER, 1846) en Belgique et en France (Hymenoptera Formicidae). Bulletin de la Societe Royale Belge d'Entomologie 20-26.
  • 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 W. M. 1913. A revision of the ants of the genus Formica (Linné) Mayr. Bulletin of the Museum of Comparative Zoology 53: 379-565.
  • Yarrow I. H. H. 1955. The British ants allied to Formica rufa L. (Hym., Formicidae). Trans. Soc. Br. Entomol. 12: 1-48.
  • 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.
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