Genus Formica (Formicinae)
- Formica
Linnaeus, 1758: 579. Type species: Formica rufa Linnaeus, by subsequent designation of Curtis, 1829: 752. Replacement type by Yarrow, 1954c: 313-138.
- Formicina
Shuckard, in Swainson & Shuckard, 1840: 172. Type species: Formica rufa Linnaeus, by subsequent designation of Wheeler, W.M. 1911g: 164. Formica and Formicina share the same type species hence they are objective synonyms. Formicina unnecessarily and incorrectly used as replacement name for Lasius Fabricius: Emery, 1916a: 239; Bondroit, 1918: 19 (see Wheeler, 1916o: 172). Formicina incorrectly as junior synonym of Lasius: Emery, 1925d: 226. Synonymy:
Wheeler, W.M. 1911c: 860;
Wheeler, W.M. 1911g: 164
;
Donisthorpe, 1943g: 646;
Agosti, 1994b: 106
.
- Serviformica
Forel, 1913j: 361. Type species: Formica fusca Linnaeus, by original designation. Serviformica junior synonym of Formica : Wheeler, W.M. 1922i: 699 (footnote). Subgenus of Formica: Emery, 1925d: 245. Synonymy:
Francoeur, 1973: 34;
Smith, D.R. 1979: 1448;
Agosti, 1994b: 106
.
- Neoformica
Wheeler, W.M. 1913a: 82 (as a subgenus of Formica)
. Type species: Formica pallidefulva Latreille, by original designation. Also described as new by W.M. Wheeler, 1913i: 548. Synonymy:
Buren, 1968a: 39;
Agosti, 1994b: 107
.
- Raptiformica
Forel, 1913j: 361 (as a subgenus of Formica)
. Type species: Formica sanguinea Latreille, by original designation. Synonymy:
Wheeler, W.M. 1922i: 699 (footnote)
;
Smith, D.R. 1979: 1448;
Agosti, 1994b: 107
.
- Coptoformica
Müller, 1923b: 146 (as subgenus of Formica)
. Type species: Formica exsecta Nylander, by subsequent designation of Donisthorpe, 1941f: 37. Synonymy:
Smith, M.R. 1951c: 860;
Smith, D.R. 1979: 1448;
Agosti, 1994b: 107
.
- Adformica
Lomnicki, 1925a: 164 (as subgenus of Formica)
. Type species: Formica exsecta Nylander, by subsequent designation of Donisthorpe, 1927c: 316. Adformica and Coptoformica share the same type species, hence they are objective synonyms. Synonymy:
Stitz, 1939: 306 (Adformica junior synonym of Coptoformica)
;
Smith, M.R. 1951c: 860 (Coptoformica junior synonym of Formica)
.
- Iberoformica
Tinaut, 1990a: 282
(as subgenus of Formica)
. Type species: Formica subrufa Roger, by original designation. Synonymy:
Agosti, 1994b: 107
.
- OVERVIEW.
This is the largest genus in North America containing almost one-sixth of all Nearctic species. Its members exhibit an impressive variety of habits including slave making, temporary social parasitism of various types, and various methods of nest founding and nest construction. Formica is widely accepted as being a natural genus. Although the morphology based work of Agosti, 1994b failed to show a synapomorphy for the genus, more recent molecular evidence suggests that Formica is monophyletic (p. 133, Brady, Gadau & Ward, 2000. in Austin & Dowton (eds.) Hymenoptera, evolution, biodiversity and biological control. CSIRO Pub.). For most Formica species, little or nothing is known of their natural history. There are, however, a few species have been more closely studied. See the discussion and natural history accounts of F. exsectoides (exsecta species group), F. ulkei (exsecta species group), and F. obscuripes (rufa species group).
- DISTRIBUTION.
The distribution of Formica is exclusively Holarctic. It is rarely found in habitats with less than 250 mm of rainfall per year, and most of the species are found in the temperate zones with deciduous or coniferous trees. In North America the distribution extends further into xeric and desert areas than in the Palearctic region. Ants of genus Formica are more common, abundant, and diverse in the northern part of their range. In the far north it is not uncommon for them to be the dominant ants (Agosti, 1994b).
- ZOOGEOGRAPHICAL REGIONS. Neotropical, Nearctic, Palearctic, Oriental.
- NOMENCLATURE.
The approximately 160 species (Palearctic and Nearctic) have been divided according to several different plans (Creighton, 1950a; Wilson & Brown, 1955; Buren, 1968a; Smith, D.R. 1979; Tinaut, 1990). Here we follow the treatment of Smith, D.R. 1979 (and confirmed by Agosti, 1994b and Bolton, 1995b) using informal species groups that do not fall under the rules of zoological nomenclature. Further studies may clarify the species groupings and determine the desirability of establishing subgenera. The species groups are: exsecta, fusca, microgyna, neogagates, pallidefulva, rufa, and sanguinea. The microgyna and rufa groups are separated by properties of their queens so, for the purposes of keying workers to species, they are sometimes lumped together.
- RECOGNITION.
Size medium to large (usually 4.5-9 mm; occasionally 2.5-4.5 mm). Mandibles with 7 or more teeth or denticles. Lower rim of antennal socket nearly touching posterior border of clypeus. Frontal carinae short but distinct with moderately to sharply angulate summit (this is best observed by looking perpendicularly down on summit of ridge). Ocelli well developed. Profile of mesosoma clearly discontinuous and not evenly convex, metanotal impression moderate to strong. Propodeum often distinctly depressed below level of promesonotum; basal face of propodeum longer than declivitous face. Mesosoma in dorsal view not wedge-shaped, usually constricted at least slightly at middle.
- SIMILAR GENERA.
Many members of the Formicine genus Camponotus have similar size and general appearance. They can be separated from genus Formica by:
- Profile of mesosoma continuously and evenly convex; propodeum not depressed below level of promesonotum; metanotal region at most slightly impressed (usually not at all impressed).
- Mesosoma in dorsal view wedge-shaped and tapering posteriorly.
- Lower rim of antennal socket not almost touching posterior border of clypeus.
Members of genus Polyergus take Formica workers as slaves and mixed colonies containing workers of both genera are not uncommon. Polyergus is easily distinguished from Formica by its falcate (sickle shaped) mandibles.
Workers of the genera Lasius, Paratrechina, and Prenolepis are usually much smaller than in Formica and are dark colored (never red or bicolored as some species of Formica). They are differentiated from Formica by: - Frontal carinae indistinct or absent (if present, each carina with a rounded summit (angulate summit in Formica)).
- Lower rim of antennal socket not almost touching posterior border of clypeus (set back at least one third maximum diameter of antennal socket).
Members of genus Acanthomyops are red, but they are also much smaller than Formica, and these subterranean ants are rarely seen above ground except during mating flights.
- TRIBE. Formicini.
- REVISIONS.
Emery, 1893k: 643654, 657665
(Nearctic species)
.
Wheeler, W.M. 1913i: 387399, 401536, 560565
(world species)
.
Creighton, 1940a: 110
(rufa species group in Nearctic region - out of date)
.
Creighton, 1950a: 450552
(Nearctic species)
.
Francoeur, 1973: 1316 (fusca species group in New World)
. - TAXONOMY.
Yarrow, 1950: 408.
Wheeler, G.C. & Wheeler, J. 1953c: 160 (larva)
.
Wilson & Brown, 1955: 108129
(notes on sanguinea and neogagates species groups, status of subgenera)
.
Yarrow, 1955: 313318 (type-species)
.
Cotti, 1963: 1413 (extensive bibliography)
.
Gregg, R.E. 1964: 5969 (distribution in mountains of CO)
.
Dlussky, 1965a: 1543 (Mongolia and northeastern Tibet)
.
Bernard, 1967a: 287 (France and Mediterranean basin)
.
Dlussky, 1967a: 1286 (Palearctic species (in Russian))
.
Buren, 1968a: 2540 (fundamental taxonomic problems in sanguinea species group (= subgenus Raptiformica), status of subgenera, key and descriptions of species in sanguinea species group)
.
Gregg, R.E. 1969a: 3849 (geographical distribution of genus in world)
.
Dlussky & Pisarski, 1971: 8590 (Poland)
.
Kutter, 1977c: 1298 (Palearctic species)
.
Arnol´di & Dlussky, 1978: 552 (former European USSR (in Russian))
.
Collingwood, 1978: 6595 (Palearctic - Iberian peninsula species)
.
Collingwood, 1979: 112154
(Fennoscandia and Denmark)
.
Kupyanskaya, 1980: 1120 (far eastern Russia)
.
Kupyanskaya, 1981: 3642 (far eastern Russia)
.
Agosti & Collingwood, 1987b: 261293 (Palearctic - Balkan species)
.
Agosti & Bolton, 1990b: 149156
(differentiating Formica from Lasius)
.
Tinaut, 1990a: 281291
(status of subgenera)
.
Wu, J. 1990: 18 (China (in Chinese))
.
Bolton, 1994: figs. 104, 105 (SEM (full face and lateral view) of w.)
.Bolton, 2003: 277.Ward, P.& Brady, S. 2003: 277.
exsecta species group (Genus Formica)
- OVERVIEW.
Ants of this group build large mounds in fields, woods, or at the edge of woods. Mounds are conspicuous not only because of their large size but also because they occur in clusters. They are more common in the eastern half of North America. The nest founding female may behave as a temporary social parasite, but colonies are also founded by budding. These fierce ants kill other ants by decapitation, a unique characteristic.
- NOMENCLATURE.
Subgenus Coptoformica was widely used for this group by earlier European authors.

Photo of Formica opaciventris Emery. Courtesy of Gary Alpert, Harvard University
- RECOGNITION.
Total length 3.5-7.5 mm. Bicolored. Ventral border of clypeus without a distinct notch. Larger workers with posterior margin of head distinctly concave. In profile, pronotum consisting of two distinct faces (not evenly convex). Viewed in profile, propodeum with basal and declivitous faces clearly differentiated and meeting at an angle.
- SIMILAR SPECIES GROUPS.
Species of the exsecta group resemble members of rufa species group in color and shape of propodeum. However, rufa group members have their posterior margin of the head fairly flat and, viewed in profile, the pronotum is a continuous curve and is not separated into two faces meeting at an angle.
- REVISIONS.
Seifert, 2000: 517568 (Palearctic, key to species)
.
- TAXONOMY.
Wheeler, W.M. 1913i: 481484
(description of species group and distribution)
.
Smith, M.R. 1947f: 620, 624 (description of speices group)
.
Creighton, 1950a: 510515
(Nearctic species)
.
Dlussky, 1964: 10261040 (exsecta group in former USSR (in Russian))
. - MISCELLANEOUS.
Hölldobler, B. & Wilson, 1990: 444, 450451 (colony founding by temporary social parasitism)
.
Formica exsectoides Forel. A large aggregation of mounds of this species may occur in a single locality. It sometimes damages bark and cambium of small trees and shrubs. Host: Formica fusca L. Locally called the Allegheny mound ant (Smith, D.R. 1979: 1456).
-
exsectoides. Formica exsectoides
Forel, 1886b: xxxviii (w.q.)
. [USA, NH; No types in USA.]
Emery, 1893k: 653
(m.)
.
-
davisi. Formica exsectoides var. davisi
Wheeler, W.M. 1913i: 396
(in key)
(w.q.) [USA, NJ, (Newfoundland)] First available use of the unavailable name Formica exsectoides subsp. exsectoides var. davisi Wheeler. Note that the type locality is not "Canada" as in Bolton, 1995b: 193. Synonymy:
Creighton, 1950a: 513
.
-
davisi. Formica exsectoides subsp. exsectoides var. davisi
Wheeler, W.M. 1913i: 484
(w.q.)
. Unavailable name. [USA, NJ, (Newfoundland)] Synonymy:
Bolton, 1995b: 193.
-
hesperia. Formica exsectoides var. hesperia
Wheeler, W.M. 1913i: 396
(in key)
. [USA, CO, El Paso Co.] Variety of exsectoides: Wheeler, W.M. 1917a: 544. First available use of the unvailable name Formica exsectoides subsp. exsectoides var. hesperia Wheeler. Synonymy:
Creighton, 1950a: 513
.
-
hesperia. Formica exsectoides subsp. exsectoides var. hesperia
Wheeler, W.M. 1913i: 484
(w.)
. Unavailable name. [USA, CO, El Paso Co.] Synonymy:
Bolton, 1995b: 196.
- DISTRIBUTION. Nova Scotia south to GA, west to Ontario, WI, IA, south to CO and northern NM. At the latitude of NY it is more abundant in the hills than at sea level and from VA southward it is found at elevations between 3000 and 4000 feet (Creighton, 1950a: 513). Uncommon in the Rocky Mountains.
- NATURAL HISTORY. Formica exsectoides is found primarily in grassy or heath-like clearings. These habitats are relatively scarce and patchily distributed. They usually have a very sparse ant fauna which can often be fully occupied by dense population of exsectoides (Hölldobler, B. & Wilson, 1990: 214). The nests are conical mounds that can be 2.5 feet high and 9.5 feet in diameter (Wheeler, 1913i: 483). These mounds can be clumped together with up to several hundred mounds occurring at one location. Clumping of mounds is generally assumed to be evidence of new mound formation by budding although the budding process has never been observed in nature (Creighton, 1950a: 512). Large mounds have been found with a population of 237,000 workers and 1400 queens (Cory & Haviland, 1938: 50-56). Mean colony size is reported to be 140,000 workers (Kaspari & Vargo, 1995: 621). It is common to find several dozen fertile females in a single nest (Creighton, 1950a: 512) which is also a condition consistent with budding of new mounds. New colonies can also be formed by temporary social parasitism. This is usually inferred from finding nests with a mix of two species but only a queen of the socially parasitic species. Host species are F. fusca (Creighton, 1950a: 512) and F. subsericea (Wheeler, 1904i: 363). This fierce ant strongly defends its nest. It can kill other ants by decapitation, a method apparently restricted amongst the ants to the exsecta species group. Much of the diet comes from honeydew supplied by homopterans although this is supplemented by scavenging for dead arthropods, preying on spiders, flies, etc. They have also been observed feeding on the juices of dead nestmates. In warm weather they will forage around the clock, but in cooler weather, foraging is restricted to the warmer parts of the day (McCook, 1877: 273). This species sometimes employs trunk trails leading to the base of trees with a high concentration of homopterans (McCook, 1877: 273). There have been reports of possible division of labor in the foraging and food gathering. Workers will collect honeydew high up in the tree then descend to the base of the tree. The honeydew is transferred to other workers by trophylaxis at the base of the tree and under rocks near the base of the tree. Presumably, one worker gathers the honeydew and another for transporting it back to the nest. (This would be similar to observations on Camponotus (Camponotus) spp. in which the largest workers are the "tanker trucks" moving food between the base of the tree and the nest, and the smaller workers ascend the tree to gather the honeydew). Additionally, sentinels are placed along the trunk of the tree apparently in defense of the foraging area (McCook, 1877: 277). Some workers becomes repletes with greatly distended gasters storing large quantities of liquid food.
- TAXONOMY.
Emery, 1893k: 643, 653
(description w.m.)
.
Wheeler, W.M. 1913i: 396, 481483
(description w.q.m., distribution, nest structure)
.
Creighton, 1950a: 511514
.
Cole, A.C. 1954c: 166
.
- MISCELLANEOUS.
McCook, 1877c: 253295 (extensive natural history observations, foraging, nest structure, diet, as F. rufa)
.
Wheeler, W.M. 1904i: 363364 (temporary social parasite of F. subsericea Say)
.
Wheeler, W.M. 1906e: 7174
.
Andrews, 1925: 7587 (growth of mounds)
.
Andrews, 1926: 127150 (long term observations on colony movement and growth of mounds)
.
Andrews, 1927: 120.
Andrews, 1929a: 369391 (association with treehoppers)
.
Andrews, 1929b: 248257 (census of 1 mound)
.
Cory & Haviland, 1938: 5056 (census of 2 mounds, large - 237,000 w. 1400 q., medium - 41,300 w. 40q.)
.
Dreyer, 1938: 3849 (body weight and water content of workers, no variation in water content when hibernating)
.
Andrews, 1941: 530533.
Buren, 1944a: 300307
(brief natural history, distribution in IA)
.
Haviland, 1947: 413418 (biology and control)
.
Wheeler, G.C. & Wheeler, J. 1953c: 164 (life cycle of larva)
.
Cole, A.C. 1954c: 166
(brief natural history, first occurrence in NM)
.
Gregg, R.E. 1963: 597599
(discussion of junior synonyms, occurrence in CO)
.
Christensen & Quick, 1970: 207209.
Hölldobler, B. & Wilson, 1990: 214, 216, 373 (number of queens, nest structure)
.
Bishop & Bristow, 2001: 3340.
- ALL REFERENCES
- PHOTOS.
Formica opaciventris Emery. This ant may construct either earthen mounds or thatched nests (Smith, D.R. 1979: 1456).
-
opaciventris. Formica exsectoides var. opaciventris
Emery, 1893k: 653
(w.m.)
. [USA, CO, Summit Co.; AMNH, MCZ]
Gregg, R.E. 1952b: 14
(q.)
. Subspecies of exsectoides: Wheeler, W.M. 1913i: 484. Raised to species: Creighton, 1950a: 514.
- DISTRIBUTION. The Rocky Mountain states and ND.
- TAXONOMY.
Wheeler, W.M. 1913i: 396, 484485
(description w.m., natural history)
.
Creighton, 1950a: 512, 514515
.
- MISCELLANEOUS.
Cole, A.C. 1954c: 166
(brief natural history, distribution in NM)
.
Scherba, 1961: 7187 (reproduction, nest structure)
.
Gregg, R.E. 1963: 599601
(valid species, nest structure)
.
Scherba, 1963: 219231.
Wheeler, G.C. & Wheeler, J. 1963: 219231.
Scherba, 1964: 231.
Allred, 1982: 473 (distribution in UT, habitat, elevation)
.
Hölldobler, B. & Wilson, 1990: 216, 216 (number of queens)
.
- ALL REFERENCES
- PHOTOS.
Formica ulkei Emery. This is a water and sun loving ant. Mounds are found on forest edges or clearings but never within forests. The mounds are situated to receive direct sunlight much of the day (Dreyer & Park, 1932). The large mounds occasionally have a thin layer of plant debris on their surfaces (Smith, D.R. 1979: 1456). It is most commonly found near water, and a mound must have close proximity to shrubs and trees occupied by homopterans (Talbot, 1961. Ecology 42: 202). Colony founding is primarily by budding (Scherba, 1958. Insectes Soc. 5: 201-213). Rarely, mixed colonies of F. fusca and ulkei are found (Creighton, 1934: 199; Talbot, 1961. Ecology 24: 203) providing evidence that temporary social parasitism is an alternate method of colony formation. As with exsectoides, trunk trails and mass recruitment are employed for the tending of homopterans (Deslippe & Savolainen, 1995). For over 20 years a group of 400 mounds in Palos Park, Illinois was studied by University of Chicago researchers (Holmquist, 1928. Ecology 9; Dreyer & Park, 1932; Dreyer, 1942; Scherba, 1958. Insectes Soc. 5). This is one of the few long term studies of an ant population, and its changes over time.
-
ulkei. Formica ulkei
Emery, 1893k: 653, pl. 22, fig. 7
(w.)
. [worker: USA, SD, Pennington Co., queen: Canada, Nova Scotia; MCZ (queen)]
Wheeler, W.M. 1913i: 486
(q.m.)
. Note that the correct page number is "Hung, 1969: 456 (k.)." not "Hung, 1969: 455 (k.)." as in Bolton, 1995b: 205.
-
hebescens. Formica ulkei var. hebescens
Wheeler, W.M. 1913i: 487
(w.)
. [USA, IN, Stark Co.] Synonymy:
Creighton, 1950a: 515
.
- DISTRIBUTION. Northern Midwest. In Canada, Nova Scotia west to Manitoba.
- TAXONOMY.
Wheeler, W.M. 1913i: 396, 485487
(description w.q.m., distribution, natural history)
.
Gregg, R.E. 1945a: 472
.
Creighton, 1950a: 512, 515
.
- MISCELLANEOUS. Holmquist, 1928: 7087.Holmquist, 1928: 325327.
Park, 1929: 195215.
Dreyer & Park, 1932: 127133 (local distribution of mound in correlation with light intensity and soil moisture)
.
Creighton, 1934: 199200 (nest founding by temporary social parasitism)
.
Dreyer, 1942: 486490 (follow up on survey of mound locations in Dreyer & Park, 1932)
.
Buren, 1944a: 300, 307
(brief natural history, distribution in IA)
.
Scherba, 1958: 201213 (reproduction, budding of new mounds)
.
Kannowski, 1959b: 142143 (incipient colony)
.
Scherba, 1959: 499508 (moisture regulation in mounds)
.
Talbot, 1959: 128132 (flight activities)
.
Talbot, 1961: 202205 (mounds)
.
Scherba, 1962: 373385 (temperature regulation in mounds)
.
Wheeler, G.C. & Wheeler, J. 1963: 256258 (natural history)
.
Hölldobler, B. & Wilson, 1990: 373, 374 (nest structure and humidity regulation)
.
- ALL REFERENCES
- PHOTOS.
fusca species group (Genus Formica)
- OVERVIEW.
Members of this group nest in the soil, and the nest is commonly started under objects or at the base of tufts of grass. Excavated soil may be spread out in the form of an irregular crater or low mound about the nest openings. Because of their ubiquity and usual docility, they are common hosts for many of the slave-making species of Formica. In his 1973 revision, Francoeur add 14 new species to this group. When consulting literature prior to 1973, one must take into account the fact that older identifications may not reflect the current taxonomy of the group. Francoeur further divided this group into 11 complexes as follows (Francoeur, 1973: 34-35):
- cinerea complex = {altipetens, canadensis, francoeuri, montana, pulla }
- fusca complex = {accreta, fusca, microphthalma }
- lepida complex = {aerata, lepida, longipilosa, pauchucana, pacifica, transmontanis }
- neoclara complex = {neoclara, occulta, propatula }
- neorufibarbis complex = {hewitti, neorufibarbis }
- occidua complex = {moki, xerophila}
- rufibarbis complex = {browni, foreliana, gnava}
- sibylla complex = { sibylla, subelongata}
- subcyanea complex = {subcyanea}
- subpolita complex = {subpolita}
- subsericea complex = {argentea, glacialis, podzolica, subsericea}
- NOMENCLATURE.
Species group fusca was formerly considered to be subgenus Serviformica.

Photo of Formica fusca Linnaeus from CA, Santa Barbara Co.. Courtesy of Ant Web, U. of Calif. Davis
- RECOGNITION.
Length 2.5-7.5 mm. Ventral border of clypeus without a distinct notch. Posterior margin of head border not distinctly concave. Pronotum, in profile, evenly convex and not separated into two distinct faces. Propodeum usually angulate in profile (basal and declivitous faces clearly differentiated). Usually concolorous black or brown; if bicolored, mesosoma lighter than gaster or upper portion of head.
- SIMILAR SPECIES GROUPS.
Species group fusca is most similar to the rufa group. Most fusca group species are considerably smaller than members of the rufa group, and the major workers in the rufa group are considerably larger than in the fusca group. Most fusca species (but not all) are concolorous black or brown whereas rufas are bicolored. When they are bicolored, fusca group workers have a different color pattern than rufa group workers. Bicolored fusca group workers will have the mesosoma lighter than both the head and gaster whereas rufa group workers have both head and mesosoma lighter than the dark gaster. Members of the fusca group are approximately the same size as member of the neogagates species group. Members of neogagates group are shiny, have longer scapes and legs, and propodeum rounded in profile (basal and declivitous faces not clearly differentiated).
- REVISIONS.
Wheeler, W.M. 1913i: 494533
.
Creighton, 1950a: 515543
.
Yarrow, 1954a: 229244 (British species)
.
Francoeur, 1973: 1316.
Wheeler, G.C. & Wheeler, J. 1977b: 12 (figures of an ant with mid and hing coxae removed - fig. 1 - lateral view, figs. 2-11 dorsal view of mesosoma)
.
Wheeler, G.C. & Wheeler, J. 1986g: 8792
(figures useful for species identification)
.
Formica accreta Francoeur.
-
accreta. Formica accreta
Francoeur, 1973: 182, figs. 308323 (w.q.m.)
. [Canada, BC; CDA, DBUL, LACM, MCZ] Junior synonym of fusca: Wheeler, G.C. & Wheeler, J. 1986g: 16. Revived from synonymy: Bolton, 1995b: 190.
- DISTRIBUTION. Far western states and British Columbia.
- TAXONOMY.
Francoeur, 1973: 182189, figs. 308323 (figure (full face view and lateral view of mesosoma and petiole) w.q.; full face view of m.)
.
- MISCELLANEOUS.
Allred, 1982: 461462 (distribution in UT)
.
- ALL REFERENCES
- PHOTOS.
Formica aerata Francoeur. This ant is found in sandy soil where it nests under rocks.
-
aerata. Formica aerata
Francoeur, 1973: 116, figs. 183189 (w.q.)
. [USA, CA, Humboldt Co.; USNM, LACM, DBUL]
Formica altipetens Wheeler. Nests are in forested and open areas in the soil; low mounds are sometimes built, but nests are also under objects (Smith, D.R. 1979: 1452).
-
altipetens. Formica cinerea var. altipetens
Wheeler, W.M. 1913i: 399
(diagnosis in key)
(w.q.m.) [USA, CO, Teller Co.; AMNH, MCZ, DBUL, USNM]
Wheeler, G.C. & Wheeler, J. 1953c: 160, pl. IV, figs. 15 (l.)
. Variety / subspecies of cinerea: Wheeler, W.M. 1917a: 550; Wheeler, G.C. & Wheeler, E.W. 1944: 259. Raised to species: Creighton, 1950a: 531. First available use of the unavailable name Formica cinerea subsp. cinerea var. altipetens Wheeler, W.M.1913.
-
altipetens. Formica cinerea subsp. cinerea var. altipetens
Wheeler, W.M. 1913i: 523
(w.q.m.)
. Unavailable name.
- DISTRIBUTION. MT south through the Rockies to Mexico and west to the Pacific Coast.
- TAXONOMY.
Creighton, 1950a: 531
.
Francoeur, 1973: 5260, figs. 3955 (figure (full face view and lateral view of mesosoma and petiole) w.; full face view of m.)
.
- MISCELLANEOUS.
Cole, A.C. 1954c: 166
(brief natural history; distribution in NM)
.
Gregg, R.E. 1963: 507509
.
Wheeler, G.C. & Wheeler, J. 1963: 260261 (diagnosis of w., habitat, distribution in ND)
.
Funk, 1975: 261262 (association with Lycaenidae)
.
Bernstein, R.A. 1979: table 2 (temperature range for foraging)
.
Allred, 1982: 462 (distribution in UT, habitat, elevation)
.
Cushman, Martinsen & Mazeroll, 1988: 522525 (evidence for intraspecific competition based on nest spacing)
.
- PHOTOS.
Formica argentea Wheeler. This ant is found in open or semiopen situations usually in sandy soil under rocks or with a low mound (Smith, D.R. 1979: 1452).
-
argentea. Formica fusca var. argentea
Wheeler, W.M. 1912c: 90 (w.)
. [USA, IL, Winnebago Co.; MCZ, AMNH, CE ,DBUL, USNM]
Wheeler, W.M. 1913i: 502
(q.m.)
. Combination in F. (Serviformica): Emery, 1925d: 248. Subspecies of fusca: Buren, 1944a: 301. Junior synonym of fusca: Creighton, 1950a: 532. Revived from synonymy and raised to species: Francoeur, 1973: 141. Replacement name for the junior homonym F. argentata Wheeler.
-
argentata. Formica fusca var. argentata
Wheeler, W.M. 1902f: 952 (footnote)
(w.) Junior primary homonym of F. argentata Fabricius, 1804. Replacement name is F. fusca argentea Wheeler. Synonymy:
Bolton, 1995b: 191.
-
blanda. Formica fusca var. blanda
Wheeler, W.M. 1913i: 398
(in key)
(w.) [USA, WA, Thurston Co.] Variety of fusca: Wheeler, 1917a: 548. Unresolved junior primary homonym of F. blanda F. Smith, 1858a: 43. First available use of the unavailable name Formica fusca fusca var. blanda Wheeler. Synonymy:
Francoeur, 1973: 141.
-
blanda. Formica fusca subsp. fusca var. blanda
Wheeler, W.M. 1913i: 510
(w.)
. Unavailable name. [USA, WA, Thurston Co.]
- DISTRIBUTION. Quebec west to British Columbia south to SC, OH, IL, IA, SD, NM, AZ, CA.
- TAXONOMY.
Wheeler, W.M. 1913i: 501503
(description w.q.m.; distrubution; natural history)
.
Francoeur, 1973: 141152, figs. 233248 (also natural history)
.
- MISCELLANEOUS.
Mallis, 1941: 83 (distribution in CA)
.
Buren, 1944a: 301
(distribution in IA)
.
Gregg, R.E. 1963: 521525
.
Wheeler, G.C. & Wheeler, J. 1977b: 1718 (natural history)
.
Allred, 1982: 462 (distribution in UT, habitat, elevation)
.
MacKay, Lowrie, et. al. 1988: 113
(habitat)
.
- ALL REFERENCES
- PHOTOS.
Formica canadensis Santschi. Nests are in open or wooded areas in the soil, sometimes with a low mound. This species has been confused with lepida in much of the literature (Smith, D.R. 1979: 1452).
-
canadensis. Formica cinerea var. canadensis
Santschi, 1914a: 435 (w.q.)
. [Canada, SK; Santschi Collection - Mus. of Natural History, Bale, Switzerland.]
Wheeler, W.M. 1917a: 550
(q.)
. Combination in F. (Serviformica): Emery, 1925d: 246. Junior synonym of lepida: Smith, M.R. 1961a: 252. Revivied from synonymy and raised to species: Francoeur, 1973: 60.
- DISTRIBUTION. MB, SK, ND, SD, KS, AB, WY, CO, NM, ID, UT, AZ, CA.
- TAXONOMY.
Wheeler, W.M. 1917a: 550
(description of w.q.)
.
Creighton, 1950a: 531
.
Francoeur, 1973: 6067, figs. 5772 (figure (full face view and lateral view of mesosoma and petiole) w.; full face view of m.)
.
Wheeler, G.C. & Wheeler, J. 1977b: 20 (considered a junior synonym of F. montana)
.
- MISCELLANEOUS.
Allred, 1982: 463 (distribution in UT, habitat, elevation)
.
- ALL REFERENCES
- PHOTOS.
Formica foreliana Wheeler.
-
foreliana. Formica foreliana
Wheeler, W.M. 1913i: 451
(w.)
. [USA, AZ, Cochise Co.; MCZ, USNM]
Francoeur, 1973: 235 (q.m.)
. Junior synonym of gnava: Creighton, 1950a: 539. Revived from synonymy: Francoeur, 1973: 235.
Formica francoeuri Bolton. This ant is most common at low elevations in the Coastal Range and San Bernadino Mountains. It makes crater or mound nests in sandy soil, sometimes under rocks (Smith, D.R. 1979: 1455).
-
francoeuri. Formica francoeuri
Bolton, 1995b: 195 (w.q.m.)
. [USA, CA, San Benito Co.; Emery Collection, CM, DBUL.] Replacement name for junior primary homonym Formica pilicornis Emery, 1893.
-
pilicornis. Formica pilicornis
Emery, 1893k: 664
(w.q.m.)
. [USA, CA, San Benito Co.; Emery Collection, CM, DBUL.] Subspecies of cinerea: Wheeler, W.M. 1913i: 527. Revived status as species: Creighton, 1950a: 538. Junior primary homonym of Formica pilicornis Roger, 1859. Replacement name is Formica francoeuri Bolton.
- DISTRIBUTION. CA ranging from San Francisco south to Mexico.
- NOMENCLATURE. Prior to 1995 this species was called Formica pilicornis Emery.
- TAXONOMY.
Wheeler, W.M. 1913i: 527, 528
(description w.q.m., distribution (as F. cinearea pilicornis Emery))
.
Creighton, 1950a: 538
.
Francoeur, 1973: 7480, figs. 90106 (description w.q.m., figure (full face & lateral view of mesosoma and petiole) of w.m., geographic variation, natural history)
.
- MISCELLANEOUS.
Cole, A.C. 1934b: 401.
Eckert & Mallis, 1937: 7, 29.
Mallis, 1941: 85 (natural history)
.
- ALL REFERENCES
- PHOTOS.
Formica fuliginothorax Blacker. This ant was found in short turf or crumbling banks.
-
fuliginothorax. Formica fuliginothorax
Blacker, 1992: 11 (w.)
. [Canada, BC, (Victoria Island). No types known to exist.]
Formica fusca Linnaeus. Nests are found in a wide variety of situations, in forests or open areas, under rocks, logs, in soil, or in rotting wood. Two forms are recognized by Francoeur (1973), form subaenescem which is mostly eastern and form marcida which is mostly western in distribution (Smith, D.R. 1979: 1452).
-
fusca. Formica fusca
Linnaeus, 1758: 580 (w.)
. [Europe; No types known to exist.]
Latreille, 1802a: 159 (q.m.)
. Combination in F. (Serviformica): Forel, 1913j: 361. Current subspecies nominal plus alpicola, fuscolemani, hyrcana, maura, tombeuri.
-
libera. Formica libera
Scopoli, 1763: 313 (w.)
. [Austria] Synonymy:
Latreille, 1802a: 159 (citation from Bolton, 1995b: 197. Which reference of Latreille's in 1802 is uncertain.)
.
-
flavipes. Formica flavipes
Geoffroy, in Fourcroy, 1785: 452 (m.)
. [France] Synonymy:
Latreille, 1802a: 161.
-
barbata. Formica barbata
Razoumowski, 1789: 225, fig. 12 (w.)
. [Switzerland] Synonymy:
Emery & Forel, 1879: 451.
-
tristis. Formica tristis
Christ, 1791: 513, pl. 60, fig. 13 (m.)
. [No type locality given.] Synonymy:
Emery, 1892c: 162.
-
chevrolatti. Formica chevrolatti
Romand, 1846: xxxii, pl. 2, fig. 5 (w.)
. [France] Synonymy:
Dalla Torre, 1893: 193.
-
marcida. Formica fusca var. marcida
Wheeler, W.M. 1913i: 398
(in key)
(w.q.) [Canada, BC, (Prairie Hills)] Variety of fusca: Wheeler, W.M. 1917a: 546. Raised to species: Creighton, 1950a: 534. First available use of the unavailable name Formica fusca subsp. fusca var. marcida Wheeler, W.M. 1913. Synonymy:
Francoeur, 1973: 189.
-
marcida. Formica fusca subsp. fusca var. marcida
Wheeler, W.M. 1913i: 503
(w.q.)
. Unavailable name. [Canada, BC, (Prairie Hills)] Synonymy:
Bolton, 1995b: 198.
-
pallipes. Formica fusca var. pallipes
Kuznetsov-Ugamsky, 1926c: 97 (w.)
. [Turkestan] Unresolved junior primary homonym of F. pallipes Latreille, 1787. Synonymy:
Dlussky, 1967a: 58;
Tarbinsky, 1976: 180.
-
rufipes. Formica fusca var. rufipes
Stitz, 1930a: 238 (w.q.)
. [Russia] Unresolved junior primary homonym of Formica rufipes Fabricius, 1775. Synonymy:
Dlussky, 1967a: 58.
- DISTRIBUTION. Newfoundland west to AK and the entire northern half of the USA. The southern limits of its range lies near the 38° parallel except in the larger mountain ranges where it extends further to the south (Creighton, 1950a: 532). Holarctic.
- TAXONOMY.
Wheeler, W.M. 1913i: 494497
(description w.q.m., distribution, comparison with fusca in Europe)
.
Donisthorpe, 1915f: 304.
Stitz, 1939: 348.
Creighton, 1950a: 532, 534
(as marcida)
.
Yarrow, 1954a: 229244 (redescription)
.
Dlussky, 1967a: 58.
Warren & Rouse, 1969: 58, 60 (diagnosis of w., brief natural history, distribution in AR)
.
Dlussky & Pisarski, 1971: 148.
Francoeur, 1973: 189201, figs. 326341.
Tarbinsky, 1976: 180.
Collingwood, 1979: 114, 115, 120, 121, figs. 176179
(diagnosis of w.q.m., figure (lateral view) w.q.m., full face q.m.)
.
Kupyanskaya, 1990a: 183.
Atanassov & Dlussky, 1992: 260.
- MISCELLANEOUS.
Mallis, 1941: 83, 84 (distribution in CA, natural history (as F. (F.) fusca L. and F. (F.) fusca var. marcida Wheeler))
.
Buren, 1944a: 300, 301
(brief natural history, distribution in IA (as Formica (Formica) fusca subaenescens Emery and Formica (Formica) fusca neoclara Emery)
.
King, R.L. 1949: 367370 (in mixed colony with Formica reflexa Buren)
.
King, R.L. 1951: 487489.
Cole, A.C. 1954c: 166
(brief natural history, distribution in NM, very common between 6000 and 10,000 ft.)
.
Kannowski, 1959b: 141142 (mating flights)
.
Wallis, 1960: 187190 (spinning movements of larvae)
.
Carter, 1962b: 201 (habitat, distribution in NC)
.
Gregg, R.E. 1963: 514525
.
Wheeler, G.C. & Wheeler, J. 1963: 264268 (diagnosis of w., habitat, distribution in ND, nesting sites)
.
Cole, A.C. 1966b: 23
(Nev. Test Site)
.
Wheeler, G.C. & Wheeler, J. 1977b: 1819 (nautral history)
.
Allred, 1982: 464465 (distribution in UT, habitat, elevation)
.
Nielsen, 1987: 77, map 9 (distribution in AK)
.
MacKay, Lowrie, et. al. 1988: 114
(habitat, nest, occurrence in NM)
.
Hölldobler, B. & Wilson, 1990: 229, 267, 276, 295, 341, 399, 450 (Dufour's gland, recruitment, division of labor, preyed upon as food by F. polyctena)
.
Nannonen & Sundström, 2002: 961973 (determinants of reproductive skew)
.
- ALL REFERENCES
- PHOTOS.
Formica glacialis Wheeler. This ant is found in woods or open fields where it builds its nests in the soil and with a low mound commonly covered with grass. This species has been confused with F. fusca L. in much of the literature (Smith, D.R. 1979: 1453).
-
glacialis. Formica fusca var. glacialis
Wheeler, W.M. 1908i: 624
(w.q.m.)
. [USA, ME, Cumberland Co.; MCZ, AMNH, DBUL] Combination in F. (Serviformica): Emery, 1925d: 247. Junior synonym of fusca: Wheeler, W.M. 1913i: 494; Creighton, 1950a: 532. Revived from synonymy and raised to species: Francoeur, 1973: 152. Junior synonym of subsericea: Wheeler, G.C. & Wheeler, J. 1986g: 92. Revived from synonymy: Bolton, 1995b: 196.
-
fairchildi. Formica fusca var. fairchildi
Wheeler, W.M. 1927a: 40 (w.)
. [Sumatra (locality in error)] Synonymy:
Francoeur, 1977a: 11.
Formica gnava Buckley. Formica gnava is one of the few members of the genus that are found in desert and semi-desert areas. It can also be found in open woods in the mountains. It nests in the soil, normally under rocks (Smith, D.R. 1979: 1453). This species is enslaved by Polyergus breviceps at the study site of Howard Topoff in southwestern Arizona. As a result, it has been a subject of long term studies especially in its relationship to P. breviceps.
-
gnava. Formica gnava
Buckley, S.B. 1866: 156 (w.q.m.)
. [USA, TX, Travis Co.] Combination in F. (Serviformica): Emery, 1925d: 250. Variety / subspecies of fusca: Wheeler, W.M. 1902g: 20; of rufibarbis: Wheeler, W.M. 1913i: 518; Creighton, 1950a: 539. Revived status as species: Francoeur, 1973: 238.
-
subsericeoneorufibarbis. Formica subsericeoneorufibarbis
Wheeler, W.M. 1901c: 519
. Nomen nudum. Attributed to Emery. Synonymy:
Wheeler, W.M. 1913i: 518
.
-
subsericeorufibarbis. Formica subsericeorufibarbis
Emery, 1925d: 250
. Incorrect subsequent spelling of Formica subsericeoneorufibarbis Wheeler, W.M. 1901. Synonymy:
Bolton, 1995b: 196.
- DISTRIBUTION. Southwestern USA and Mexico. .
- TAXONOMY.
Wheeler, W.M. 1913i: 390, 518521
(description w.q.m., distribution, natural history)
.
Cole, A.C. 1942: 377, 380, 383
.
Creighton, 1950a: 539541
.
Francoeur, 1973: 238245, figs. 425440 (description w.q.m., figure (full face and lateral view of mesosoma) of w.m., goegraphic vairation, distribution, natural history)
.
- MISCELLANEOUS.
Wheeler, W.M. 1902g: 20.
Wheeler, W.M. 1917a: 541, 550
.
Cole, A.C. 1934b: 388 (common in eastern part of Mojave Desert especially near Needles, CA)
.
Mallis, 1941: 86 (habitat)
.
Gregg, R.E. 1963: 539541
.
Allred, 1982: 465466 (nest sites, habitat, distribution in Utah)
.
Topoff, Inez-Pagani, Mack & Goldstein, 1985: 259267.
Topoff, Pagani, et.al. 1985: 10411046.
Goodloe & Topoff, 1987: 293302 (acceptance of pupae of P. breviceps)
.
Topoff, Bodoni, et.al. 1987: 261270.
Topoff, Cover, et.al. 1988: 209218.
Topoff, Cover & Jacobs, 1989: 545556.
Cokendolpher & Francke, 1990: 41 (habitat, nest)
.
Topoff & Mendez, 1990: 819821.
Topoff & Zimmerli, 1993: 479486.
- ALL REFERENCES
- PHOTOS.
Formica hewitti Wheeler. This ant is found in open or semiopen woods and forests where it nests in the soil under rocks or in rotting wood (Smith, D.R. 1979: 1453).
-
hewitti. Formica hewitti
Wheeler, W.M. 1917a: 552
(w.q.m.)
. [Canada, BC, Kootenay-East Co.; AMNH, MCZ]
- DISTRIBUTION. Quebec, ME west to British Columbia south to MN, ND, NM, UT, CA.
- TAXONOMY.
Creighton, 1950a: 533534
.
Francoeur, 1973: 208215, figs. 363378.
- MISCELLANEOUS.
Gregg, R.E. 1963: 526527
.
Wheeler, G.C. & Wheeler, J. 1977b: 19.
Allred, 1982: 464 (distribution in UT)
.
MacKay, Lowrie, et. al. 1988: 116
(habitat, nest, a common species in northern NM)
.
- ALL REFERENCES
- PHOTOS.
Formica lepida Wheeler.
-
lepida. Formica cinerea var. lepida
Wheeler, W.M. 1913i: 399
(diagnosis in key)
(w.) [USA, CA, Humboldt Co.; MCZ, USNM, LACM, DBUL]
Wheeler, W.M. 1917a: 551
(q.)
. Variety / subspecies of cinerea: Wheeler, W.M. 1917a: 551; Creighton, 1950a: 531. Raised to species: Francoeur, 1973: 122. First available us of the unavailable name Formica cinerea subsp. cinerea var. lepida Wheeler, W.M. 1913.) [ Type: USA, CA, Humboldt Co.; MCZ, USNM, LACM, DBUL
-
lepida. Formica cinerea subsp. cinerea var. lepida
Wheeler, W.M. 1913i: 526
(w.)
. Unavailable name. [USA, CA, Humboldt Co.; MCZ, USNM, LACM, DBUL] Synonymy:
Bolton, 1995b: 197.
Formica longipilosa Francoeur.
Formica microphthalma Francoeur. Found in the mountains, this ant may construct small crater nests in the soil (Smith, D.R. 1979: 1453).
-
microphthalma. Formica microphthalma
Francoeur, 1973: 201, figs. 344359 (w.m.)
. [USA, CA, Tuolumne Co.; MCZ, DBUL, CAUC]
Formica moki Wheeler. Nests are in soil under rocks. The nest is unmarked and the entrance is usually a hole near a bush or tree. Francoeur (1973) chose to use the name occidua for this species because occidua has been used more in the literature; however, the case has not been referred to the Commission (Smith, D.R. 1979: 1453). Formerly placed in species group pallidefulva.
-
moki. Formica moki
Wheeler, W.M. 1906i: 343
(w.)
. [USA, AZ, Coconini Co.; AMNH, MCZ]
Cole, A.C. 1966b: 25
(m.)
. Combination in F. (Neoformica): Wheeler, W.M. 1913i: 558; in fusca species group: Francoeur, 1973: 254; Smith, D.R. 1979: 1455. Junior synonym of occidua: Francoeur, 1973: 254. Note that moki has priority over occidua and hence is the valid name: Smith, D.R. 1979: 1453; Bolton, 1995b: 199.
-
occidentalis. Formica fusca var. occidentalis
Wheeler, W.M. 1908k: 409
(w.)
. Nomen nudum. [USA] Synonymy:
Smith, D.R. 1979: 1453.
-
occidentalis. Formica rufibarbis var. occidentalis
Wheeler, W.M. 1910b: 570. Nomen nudum. [USA] Synonymy:
Bolton, 1995b: 200.
-
occidua. Formica rufibarbis var. occidua
Wheeler, W.M. 1912c: 90. Unnecesary replacement name for F. rufibarbis occidentalis Wheeler. Synonymy:
Smith, D.R. 1979: 1453.
-
occidua. Formica rufibarbis var. occidua
Wheeler, W.M. 1913i: 517
(w.q.)
. [USA, CA, Santa Clara Co.] Subspecies of rufibarbis: Creighton, 1950a: 539. Raised to species: Francoeur, 1973: 254. Incorrectly stated as the senior synonym of moki: Francoeur, 1973: 254. Synonymy:
Smith, D.R. 1979: 1453.