Lasius neoniger

In the northeastern United States, this commonly collected and widespread ant is found in open habitats, including farm fields, old fields and other early-successional habitats, beaches, and sand dunes. Mating swarms to occur on warm evenings following rains in late August or early September. It is likely to be ecologically very important because it is abundant in nearly every open habitat on the landscape. (Ellison et al., 2012)

Identification
Workers of this species are usually dark brown in color, and have relatively large eyes (at least 12 ommatidia in greatest diameter). The scape has several erect hairs, but the extensor surface of the anterior tibia has fewer than 6 erect hairs. The penultimate basal tooth is smaller than the 2 adjacent teeth, and may even be absent, leaving a gap between the other basal teeth. (Mackay and Mackay 2002)

Ellison et al. (2012) - The large-eyed Lasius americanus can be confused with either of the other three species in the niger group: Lasius pallitarsis, Lasius americanus or Lasius niger. The key features are that L. neoniger has an angular clypeal margin (best seen in full-face view with the mandibles open), equally-sized basal teeth, and has erect hairs on its scape and hind tibiae. The other three species have more gently-curved clypeal margins. Lasius americanus has no erect hairs on its scape or hind tibiae. Lasius pallitarsis and L. niger are hairy like L. neoniger, but L. pallitarsis and L. niger have curved clyepeal margins. The three basal teeth of Lasius neoniger are unequally spaced and the 2nd of these three teeth is smaller than the other two. In contrast, L. pallitarsis has a single offset, short, and upturned basal tooth on each mandible, and L. niger has equally-spaced and equally-sized basal teeth.

Distribution
Lasius neoniger is abundant in eastern North America and as far west as North Dakota, Iowa and Kansas. West of the Great Plains it apparently diminishes rapidly; it is rather uncommon in the southern Rockies and is thus far completely unknown from the Pacific Northwest. It is abundant and widespread in New England (Ellison et al., 2012).

Distribution based on Regional Taxon Lists
Nearctic Region: Canada, United States. Neotropical Region: Mexico.

Habitat
Open habitats.

Abundance
Common.

Biology


Wilson (1955) - This species nests almost exclusively in open areas, either under stones or in open soil in craters. In the eastern United States it is frequently the dominant ant in grassy road strips, lawns, cultivated fields, and other disturbed situations. By virtue of this ruderal adaptation it has become one of the most abundant and conspicuous insects within its range. The only natural habitats in which I have found it dominant are prairie (North Dakota, Montana) and old beach dunes (New England coast, Great Lakes). It never penetrates deep forests and is scarce in open woods and well shaded forest borders.

One of the most interesting features of its ecological distribution is its clearcut microgeographie replacement by the woodland species Lasius americanus. The two species overlap to a limited extent in open, sunny woodland, where both may be found nesting under stones. But the bulk of the neoniger population occurs in completely exposed situations, in craters and under stones, while most of the americanus are in rotting stumps and logs in well shaded woodland. This replacement pattern can be developed into an intricate mosaic where the habitats are suitable. Such a situation is found at Ipswich Beach, Massachusetts, for example, where the two species occur together in a small stretch of littoral woodland covering about fifty acres. The woods at this locality consist principally of small trees of Betula populifolia (grey birch), Prunus maritima (beach plum), Acer rubrum (red maple), Pinus rigida (pitch pine), and several species of oak. They are dense in low swampy areas where they provide leaf litter and humus of varying thickness to the loose, sandy soil. They are broken at intervals however, by foot paths, animal trails, and sand' 'blowouts". The latter are shallow, barren depressions in the sand in which only scattered grass and Hudsonia shrubs grow. neoniger is limited to these open areas, its craters often occurring in dense concentrations where the sand is firmly packed and moist several inches below the surface. L. americanus, on the other hand, is entirely limited to rotting stumps, logs, and branches on the shaded forest floor. Under natural conditions there is probably always an ample supply of such nesting sites, since Betula populifolia, the dominant tree, is very short-lived, and dead trees must always be in abundance. The two species are thus sharply segregated ecologically, with the neoniger population consisting of a series of enclaves within a more or less continuous woodland americanus population. No instances of overlap were found; the closest approach was a colony of americanus in a rotting log near the margin of a blowout and about seven feet from a cluster of neoniger craters.

Like other species of Lasius, neoniger is polyphagous, accepting both insect food and nectar. At Schroon Lake, N. Y. and Lake Constance, Ontario, I watched workers of this species foraging above ground in the early part of the night. The "trophophoric field" of the many nests observed did not seem to exceed two or three yards in radius, and the workers appeared to be engaged solely in gathering dead and crippled insects. A great variety of insects were accepted, including flies, leafhoppers, beetles, and spiders of various sizes. Using captive colonies maintained during a period of over a year, I have since learned that insects are readily accepted as food and are evidently necessary for successful colony growth.

The pastoral habits of this species have been dealt with in detail by S. A. Forbes' classic account (1894). He and later authors, including "W. M. Wheeler, have referred to the species studied as americanus (treated herein as a synonym of americanus), but there can be little doubt, judging from the descriptions of the ecology, nesting habits, etc., as well as from a drawing later published by Forbes, that it was really neoniger. The mutualistic relationship between the Lasius and the cornroot aphid (Aphis maidiradicus) as explored by Forbes is very similar to that later demonstrated by Eidmann (1926) between the European Lasius niger and an epigaeic species of aphid. The ants gather the eggs of the fall sexual brood of the aphid, store them in the nests over the winter, and in the spring excavate special galleries by means of which they are able to place the newly hatched nymphs directly on the corn roots. When strange aphids are freshly presented to the ants above ground they are promptly carried into the nest and placed on the roots also.

Talbot (1946) has studied diurnation in this species. In ordinary summer weather the workers are mainly nocturnal. Activity begins to rise in late afternoon, continues through the night, and declines through the following morning. The species prefers cool temperatures (50°-60° F.) and high humidity.

Forbes (1908) and Tanquary (1913) have studied the life history of this species in some detail. Nuptial flights are recorded by them for the period September 5-0ctober 18. Mr. B. D. Valentine (pers. commun.) has observed nuptial flights of this species on Long Island, New York, during the first week of September over a period of several years. It has been my observation that flights occur mainly in September in the Boston area also. The earliest record for a flight is one which I observed at Bishop, California, on July 30, 1952. In so far as I know the flights always take place during the second half of the afternoon; the one at Bishop continued to dusk. In the collections available to me, I have found winged forms associated in the nest with workers from July 25 (Salida, Colo.) to October 13 (Denver, Colo.). Forbes and Tanquary both showed that colony founding in this species resembles that later described for niger and Lasius brevicornis by Eidmann (1931). The fecundated queens overwinter without brood and start laying eggs in the following spring.

In New Mexico (Mackay and Mackay 2002) - Found along roadsides or in disturbed urban environments, grasslands, up to ponderosa pine-riparian. This species is usually found in mesic habitats. L. neoniger is uncommon in the state, and nests under stones or in the soil. Brood was found in nests in early July, reproductives in early July to October. It feeds on secretions of Homoptera.

Insects
This species is a host for the temporary parasites, , , and.

Nomenclature

 *  neoniger. Lasius niger var. neoniger Emery, 1893i: 639 (w.q.m.) U.S.A. (in text). Subspecies of niger: Wheeler, W.M. 1917a: 463; Buren, 1944a: 296; Creighton, 1950a: 420. Status as species: Gregg, E.V. 1945: 530; Wilson, 1955a: 97; Smith, D.R. 1979: 1436.

Description
Wilson (1955) - Worker and queen. Possessing the first two of the following characters in common with the other two members of the "neoniger complex" (Lasius crypticus and Lasius sitiens) and differing from these two species by the possession of the third.

(1) At least 20 per cent and often more than 90 per cent, of the nest series with the median of three basal teeth reduced in size relative to the two flanking teeth; in individuals where this tooth is absent, the space between the two remaining teeth is typically irregular in size and shape and usually larger in area than the basal-most tooth.

(2) The anterior border of the median clypeal lobe, when viewed in perfect full face (attaining maximum head length) and with the mandibles open, is obtusely angulate, the lateral faces straight and often meeting centrally to form a distinct angle; opposed to the broadly convex or parabolic condition of the "niger complex."

(3) Scapes and fore tibiae with abundant hairs, many of which are standing. The density declines allometrically, and nanitic workers (with PW about 0.40 mm.) may have seta counts of 0.

Worker
Wilson (1955) - PW 0.39 mm. (incipient colony) to 0.64 mm.; the range and dispersion are apparently about the same as for Lasius americanus (q. v.). SI of series of average-sized to large workers falling in the niger-americanus regression zone, although two nanitics measured (Sudbury, Mass. incipient colony; Wilson leg.; MCZ) fell lower, just above the extrapolated brunneus zone — SL/HW ratios were 0.59/0.57 mm. and 0.59/0.58 mm. respectively. A similar dip in the regression zone has been observed in the sibling species Lasius crypticus, while the third member of the neoniger complex, Lasius sitiens, falls along the extrapolated Lasius brunneus zone. This indicates that the neoniger complex differs generally from the niger complex by having steeper regression zones. In body form, head shape, petiolar variation, etc., neoniger closely resembles niger and americanus. Appendage pilosity sparser and averaging shorter than in niger, with more hairs inclining to decumbent-subdecumbent, which condition occurs in niger in the Asiatic population only. Maximum color variation has been encountered within collections from the western U. S.: series from the White Sands National Monument, New Mexico, range to pale yellowish brown, with the head slightly darker than the remainder of the body; a series from Bishop, California, is a shade of dark brown about average for niger and americanus. The usual neoniger coloration is a light brown with feebly contrasting darker head.

Queen
Wilson (1955) - Size range and dispersion apparently about the same as for americanus. The dentition and clypeal characters of the worker are repeated in this caste with exaggerated effect. The clypeus is typically very angular, contrasting with the evenly convex condition of the niger complex and the flat-to-emarginate condition of Lasius pallitarsis. A peculiarity of the dentition in this caste is the frequent occurrence of an offset basal tooth similar to that characterizing pallitarsis; but when this occurs, it is often present on one mandible only, it is usually larger and more acute than in pallitarsis, and it is rarely if ever accompanied by secondary teeth on the basal border. The wings are more opaque than in the niger complex but lack the pattern of basal infuscation characterizing pallitarsis and brunneus.

Male
Wilson (1955) - Very similar to small individuals of niger except that the anterior border of the median clypeal lobe tends to be angulate as in the worker and queen. In the male this character is highly variable, however, and not all isolated individuals can be determined to species with certainty.

HW 0.74-0.86 mm., dispersion apparently about the same as for the North American populations of niger and americanus. Genitalia size and configuration, including the posterior lobing of the subgenital plate, apparently the same as for these two species.

Type Material
Wilson (1955) - LECTOTYPE. By present selection, a worker in the Emery Collection labelled "Va.".

References based on Global Ant Biodiversity Informatics

 * Armbrecht M., and N. J. Gotelli. 2001. Spatial and temporal niche partitioning in grassland ants. Oecologia 126: 134-141.
 * Banschbach V. S., and E. Ogilvy. 2014. Long-term Impacts of Controlled Burns on the Ant Community (Hymenoptera: Formicidae) of a Sandplain Forest in Vermont. Northeastern Naturalist 21(1): 1-12.
 * Bare O. S. 1929. A taxonomic study of Nebraska ants, or Formicidae (Hymenoptera). Thesis, University of Nebraska, Lincoln, USA.
 * Belcher A. K., M. R. Berenbaum, and A. V. Suarez. 2016. Urbana House Ants 2.0.: revisiting M. R. Smith's 1926 survey of house-infesting ants in central Illinois after 87 years. American Entomologist 62(3): 182-193.
 * Bestelmeyer B. T., and J. A. Wiens. 2001. Local and regional-scale responses of ant diversity to a semiarid biome transition. Ecography 24: 381-392.
 * Borchert, H.F. and N.L. Anderson. 1973. The Ants of the Bearpaw Mountains of Montana (Hymenoptera: Formicidae). Journal of the Kansas Entomological Society 46(2):200-224
 * Braman C. A., and B. T. Forschler. 2018. Survey of Formicidae attracted to protein baits on Georgia’s Barrier Island dunes. Southeastern Naturalist 17(4): 645-653.
 * Buren W. F. 1944. A list of Iowa ants. Iowa State College Journal of Science 18:277-312
 * Callcott A. M. A., D. H. oi, H. L. Collins, D. F. Williams, and T. C. Lockley. 2000. Seasonal Studies of an Isolated Red Imported Fire Ant (Hymenoptera: Formicidae) Population in Eastern Tennessee. Environmental Entomology, 29(4): 788-794.
 * Campbell K. U., and T. O. Crist. 2017. Ant species assembly in constructed grasslands isstructured at patch and landscape levels. Insect Conservation and Diversity doi: 10.1111/icad.12215
 * Carroll T. M. 2011. The ants of Indiana (Hymenoptera: Formicidae). Master's Thesis Purdue university, 385 pages.
 * Choate B., and F. A. Drummond. 2012. Ant Diversity and Distribution (Hymenoptera: Formicidae) Throughout Maine Lowbush Blueberry Fields in Hancock and Washington Counties. Environ. Entomol. 41(2): 222-232.
 * Choate B., and F. A. Drummond. 2013. The influence of insecticides and vegetation in structuring Formica Mound ant communities (Hymenoptera: Formicidae) in Maine lowbush blueberry. Environ. Entomol. 41(2): 222-232.
 * Clark A. T., J. J. Rykken, and B. D. Farrell. 2011. The Effects of Biogeography on Ant Diversity and Activity on the Boston Harbor Islands, Massachusetts, U.S.A. PloS One 6(11): 1-13.
 * Clark Adam. Personal communication on November 25th 2013.
 * Cokendolpher J. C., and O. F. Francke. 1990. The ants (Hymenoptera, Formicidae) of western Texas. Part II. Subfamilies Ecitoninae, Ponerinae, Pseudomyrmecinae, Dolichoderinae, and Formicinae. Special Publications, the Museum. Texas Tech University 30:1-76.
 * Cole A. C., Jr. 1942. The ants of Utah. American Midland Naturalist 28: 358-388.
 * Cole, A.C. 1936. An annotated list of the ants of Idaho (Hymenoptera; Formicidae). Canadian Entomologist 68(2):34-39
 * Coovert G. A. 2005. The Ants of Ohio (Hymenoptera: Formicidae). Ohio Biological Survey, Inc. 15(2): 1-207.
 * Coovert, G.A. 2005. The Ants of Ohio (Hymenoptera: Formicidae) Ohio Biological Survey Bulletin New Series Volume 15(2):1-196
 * Del Toro I., K. Towle, D. N. Morrison, and S. L. Pelini. 2013. Community Structure, Ecological and Behavioral Traits of Ants (Hymenoptera: Formicidae) in Massachusetts Open and Forested Habitats. Northeastern Naturalist 20: 1-12.
 * Del Toro, I. 2010. PERSONAL COMMUNICATION. MUSEUM RECORDS COLLATED BY ISRAEL DEL TORO
 * Deyrup M., C. Johnson, G. C. Wheeler, J. Wheeler. 1989. A preliminary list of the ants of Florida. Florida Entomologist 72: 91-101
 * Deyrup, M. 2003. An updated list of Florida ants (Hymenoptera: Formicidae). Florida Entomologist 86(1):43-48.
 * Drummond F. A., A. M. llison, E. Groden, and G. D. Ouellette. 2012. The ants (Formicidae). In Biodiversity of the Schoodic Peninsula: Results of the Insect and Arachnid Bioblitzes at the Schoodic District of Acadia National Park, Maine. Maine Agricultural and forest experiment station, The University of Maine, Technical Bulletin 206. 217 pages
 * DuBois M. B. 1981. New records of ants in Kansas, III. State Biological Survey of Kansas. Technical Publications 10: 32-44
 * Dubois, M.B. and W.E. Laberge. 1988. An Annotated list of the ants of Illionois. pages 133-156 in Advances in Myrmecology, J. Trager
 * Ellison A. M., and E. J. Farnsworth. 2014. Targeted sampling increases knowledge and improves estimates of ant species richness in Rhode Island. Northeastern Naturalist 21(1): NENHC-13NENHC-24.
 * Forster J.A. 2005. The Ants (hymenoptera: Formicidae) of Alabama. Master of Science, Auburn University. 242 pages.
 * Francoeur A. 1997. Ants (Hymenoptera: Formicidae) of the Yukon. Biological Survey of Canada Monograph Series 2: 901-910.
 * Francoeur A., and R. Béique. 1966. Les Formicides (Hyménoptères) de Provancher. Can. Entomol. 98: 140-145.
 * Francoeur, A. 1997. Ants (Hymenoptera: Formicidae) of the Yukon. Pages 901 910 in H.V. Danks and J.A. Downes (Eds.), Insects of the Yukon. Biological Survey of Canada (Terrestrial Arthropods), Ottawa.
 * Frye J. A., T. Frye, and T. W. Suman. 2014. The ant fauna of inland sand dune communities in Worcester County, Maryland. Northeastern Naturalist, 21(3): 446-471.
 * General D. M., and L. C. Thompson. 2011. New Distributional Records of Ants in Arkansas for 2009 and 2010 with Comments on Previous Records. Journal of the Arkansas Academy of Science 65: 166-168.
 * Glasier J. R. N., S. E. Nielsen, J. Acorn, and J. Pinzon. 2019. Boreal sand hills are areas of high diversity for Boreal ants (Hymenoptera: Formicidae). Diversity 11, 22; doi:10.3390/d11020022.
 * Gotelli, N.J. and A.M. Ellison. 2002. Biogeography at a Regional Scale: Determinants of Ant Species Density in New England Bogs and Forests. Ecology 83(6):1604-1609
 * Graham, J.H., A.J. Krzysik, D.A. Kovacic, J.J. Duda, D.C. Freeman, J.M. Emlen, J.C. Zak, W.R. Long, M.P. Wallace, C. Chamberlin-Graham, J.P. Nutter and H.E. Balbach. 2008. Ant Community Composition across a Gradient of Disturbed Military Landscapes at Fort Benning, Georgia. Southeastern Naturalist 7(3):429-448
 * Gregg E. V. 1945. A statistical study of taxonomic categories in ants (Formicidae: Lasius neoniger and Lasius americanus). Annals of the Entomological Society of America 38: 529-548.
 * Gregg R. E. 1945 (1944). The ants of the Chicago region. Annals of the Entomological Society of America 37: 447-480
 * Gregg, R.T. 1963. The Ants of Colorado.
 * Guénard B., K. A. Mccaffrey, A. Lucky, and R. R. Dunn. 2012. Ants of North Carolina: an updated list (Hymenoptera: Formicidae). Zootaxa 3552: 1-36.
 * Headley A. E. 1943. The ants of Ashtabula County, Ohio (Hymenoptera, Formicidae). The Ohio Journal of Science 43(1): 22-31.
 * Heithaus R. E., and M. Humes. 2003. Variation in Communities of Seed-Dispersing Ants in Habitats with Different Disturbance in Knox County, Ohio. OHIO J. SCI. 103 (4): 89-97.
 * Hill J.G. & Brown R. L. 2010. The Ant (Hymenoptera: Formicidae) Fauna of Black Belt Prairie Remnants in Alabama and Mississippi. Southeastern Naturalist. 9: 73-84
 * Hoey-Chamberlain R. V., L. D. Hansen, J. H. Klotz and C. McNeeley. 2010. A survey of the ants of Washington and Surrounding areas in Idaho and Oregon focusing on disturbed sites (Hymenoptera: Formicidae). Sociobiology. 56: 195-207
 * Ipser R. M. 2004. Native and exotic ants (Hymenoptera: Formicidae) of Georgia: Ecological Relationships with implications for development of biologically-based management strategies. Doctor of Philosophy thesis, University of Georgia. 165 pages.
 * Ipser, R.M., M.A. Brinkman, W.A. Gardner and H.B. Peeler. 2004. A Survey of Ground-Dwelling Ants (Hymenoptera: Formicidae) in Georgia. The Florida Entomologist 87(3) 253-260.
 * Ivanov K. 2019. The ants of Ohio (Hymenoptera, Formicidae): an updated checklist. Journal of Hymenoptera Research 70: 65–87.
 * Ivanov K., L. Hightower, S. T. Dash, and J. B. Keiper. 2019. 150 years in the making: first comprehensive list of the ants (Hymenoptera: Formicidae) of Virginia, USA. Zootaxa 4554 (2): 532–560.
 * Jeanne R. J. 1979. A latitudinal gradient in rates of ant predation. Ecology 60(6): 1211-1224.
 * Johnson C. 1986. A north Florida ant fauna (Hymenoptera: Formicidae). Insecta Mundi 1: 243-246
 * Kannowski P. B. 1956. The ants of Ramsey County, North Dakota. American Midland Naturalist 56(1): 168-185.
 * Kittelson P. M., M. P. Priebe, and P. J. Graeve. 2008. Ant Diversity in Two Southern Minnesota Tallgrass Prairie Restoration Sites. Jour. Iowa Acad. Sci. 115(14): 2832.
 * Kupianskaya A. N., Lelej, A.S., and Urbain, B. K. 2000. The Ants (Hymenoptera: Formicidae) of the Kuril Islands. Far Eastern Entomologist. 92:1-21.
 * Kupianskaya, A. N., Lelej, A.S., and Urbain, B. K. 2000. The Ants (Hymenoptera: Formicidae) of the Kuril Islands. Far Eastern Entomologist. 92:1-21.
 * La Rivers I. 1968. A first listing of the ants of Nevada. Biological Society of Nevada, Occasional Papers 17: 1-12.
 * Longino, J.T. 2010. Personal Communication. Longino Collection Database
 * Lopez R., and D. A. Potter. 2003. Biodiversity of ants (Hymenoptera: Formicidae) in golf course and lawn turf habitats in Kentucky. Sociobiology 42(3): 701-713.
 * Lubertazi, D. Personal Communication. Specimen Data from Museum of Comparative Zoology at Harvard
 * Lynch J. F. 1981. Seasonal, successional, and vertical segregation in a Maryland ant community. Oikos 37: 183-198.
 * Lynch J. F. 1988. An annotated checklist and key to the species of ants (Hymenoptera: Formicidae) of the Chesapeake Bay region. The Maryland Naturalist 31: 61-106
 * MacGown J. A., J. G. Hill, and R. L. Brown. 2010.  Native and exotic ant in Mississippi state parks.  Proceedings:  Imported Fire Ant Conference, Charleston, South Carolina, March 24-26, 2008: 74-80.
 * MacGown, J.A and J.A. Forster. 2005. A preliminary list of the ants (Hymenoptera: Formicidae) of Alabama, U.S.A. Entomological News 116(2):61-74
 * MacGown, J.A. and JV.G. Hill. Ants of the Great Smoky Mountains National Park (Tennessee and North Carolina).
 * MacGown. J. 2011. Ants collected during the 25th Annual Cross Expedition at Tims Ford State Park, Franklin County, Tennessee
 * Mackay W. P., and E. E. Mackay. 2002. The ants of New Mexico (Hymenoptera: Formicidae). Lewiston, New York: Edwin Mellen Press, 400 pp.
 * Mackay, W., D. Lowrie, A. Fisher, E. Mackay, F. Barnes and D. Lowrie. 1988. The ants of Los Alamos County, New Mexico (Hymenoptera: Formicidae). pages 79-131 in J.C. Trager, editor, Advances in Myrmecololgy.
 * Mackay, W.P. and E. Mackay. XXXX. The Ants of New Mexico
 * Mallis A. 1941. A list of the ants of California with notes on their habits and distribution. Bulletin of the Southern California Academy of Sciences 40: 61-100.
 * Menke S. B., E. Gaulke, A. Hamel, and N. Vachter. 2015. The effects of restoration age and prescribed burns on grassland ant community structure. Environmental Entomology http://dx.doi.org/10.1093/ee/nvv110
 * Menke S. B., and N. Vachter. 2014. A comparison of the effectiveness of pitfall traps and winkler litter samples for characterization of terrestrial ant (Formicidae) communities in temperate savannas. The Great Lakes Entomologist 47(3-4): 149-165.
 * Merle W. W. 1939. An Annotated List of the Ants of Maine (Hymenoptera: Formicidae). Entomological News. 50: 161-165
 * Michigan State University, The Albert J. Cook Arthropod Research Collection. Accessed on January 7th 2014 at http://www.arc.ent.msu.edu:8080/collection/index.jsp
 * Nuhn, T.P. and C.G. Wright. 1979. An Ecological Survey of Ants (Hymenoptera: Formicidae) in a Landscaped Suburban Habitat. American Midland Naturalist 102(2):353-362
 * O'Keefe S. T., J. L. Cook, T. Dudek, D. F. Wunneburger, M. D. Guzman, R. N. Coulson, and S. B. Vinson. 2000. The Distribution of Texas Ants. The Southwestern Entomologist 22: 1-92.
 * Ouellette G. D. and A. Francoeur. 2012. Formicidae [Hymenoptera] diversity from the Lower Kennebec Valley Region of Maine. Journal of the Acadian Entomological Society 8: 48-51
 * Ouellette G. D., F. A. Drummond, B. Choate and E. Groden. 2010. Ant diversity and distribution in Acadia National Park, Maine. Environmental Entomology 39: 1447-1556
 * Pecarevic M., J. Danoff-Burg, and R. R. Dunn. 2010. Biodiversity on Broadway - Enigmatic Diversity of the Societies of Ants (Formicidae) on the Streets of New York City. PLoS ONE 5(10): e13222. doi:10.1371/journal.pone.0013222
 * Procter W. 1938. Biological survey of the Mount Desert Region. Part VI. The insect fauna. Philadelphia: Wistar Institute of Anatomy and Biology, 496 pp.
 * Rees D. M., and A. W. Grundmann. 1940. A preliminary list of the ants of Utah. Bulletin of the University of Utah, 31(5): 1-12.
 * Roeder K. A., and D. V. Roeder. 2016. A checklist and assemblage comparison of ants (Hymenoptera: Formicidae) from the Wichita Mountains Wildlife Refuge in Oklahoma. Check List 12(4): 1935.
 * Sharplin, J. 1966. An annotated list of the Formicidae (Hymenoptera) of Central and Southern Alberta. Quaetiones Entomoligcae 2:243-253
 * Shik, J., A. Francoeur and C. Buddle. 2005. The effect of human activity on ant species (Hymenoptera: Formicidae) richness at the Mont St. Hilaire Biosphere Reserve, Quebec. Canadian Field-Naturalist 119(1): 38-42.
 * Smith F. 1941. A list of the ants of Washington State. The Pan-Pacific Entomologist 17(1): 23-28.
 * Smith M. R. 1934. A list of the ants of South Carolina. Journal of the New York Entomological Society 42: 353-361.
 * Smith M. R. 1935. A list of the ants of Oklahoma (Hymen.: Formicidae) (continued from page 241). Entomological News 46: 261-264.
 * Sturtevant A. H. 1931. Ants collected on Cape Cod, Massachusetts. Psyche (Cambridge) 38: 73-79
 * Talbot M. 1976. A list of the ants (Hymenoptera: Formicidae) of the Edwin S. George Reserve, Livingston County, Michigan. Great Lakes Entomologist 8: 245-246.
 * Toennisson T. A., N. J. Sanders, W. E. Klingeman, and K. M. Vail. 2011. Influences on the Structure of Suburban Ant (Hymenoptera: Formicidae) Communities and the Abundance of Tapinoma sessile. Environ. Entomol. 40(6): 1397-1404.
 * Van Pelt A. F. 1966. Activity and density of old-field ants of the Savannah River Plant, South Carolina. Journal of the Elisha Mitchell Scientific Society 82: 35-43.
 * Van Pelt A., and J. B. Gentry. 1985. The ants (Hymenoptera: Formicidae) of the Savannah River Plant, South Carolina. Dept. Energy, Savannah River Ecology Lab., Aiken, SC., Report SRO-NERP-14, 56 p.
 * Warren, L.O. and E.P. Rouse. 1969. The Ants of Arkansas. Bulletin of the Agricultural Experiment Station 742:1-67
 * Wenner A. M. 1959. The ants of Bidwell Park, Chico, California. American Midland Naturalist 62: 174-183
 * Wheeler G. C., J. N. Wheeler, and P. B. Kannowski. 1994. Checklist of the ants of Michigan (Hymenoptera: Formicidae). The Great Lakes Entomologist 26(4): 297-310
 * Wheeler G. C., and E. W. Wheeler. 1944. Ants of North Dakota. North Dakota Historical Quarterly 11:231-271.
 * Wheeler G. C., and J. Wheeler J. 1989. A checklist of the ants of Oklahoma. Prairie Naturalist 21: 203-210.
 * Wheeler G. C., and J. Wheeler. 1986. The ants of Nevada. Los Angeles: Natural History Museum of Los Angeles County, vii + 138 pp.
 * Wheeler G. C., and J. Wheeler. 1987. A Checklist of the Ants of South Dakota. Prairie Nat. 19(3): 199-208.
 * Wheeler W. M. 1904. The ants of North Carolina. Bulletin of the American Museum of Natural History. 20: 299-306.
 * Wheeler W. M. 1906. Fauna of New England. 7. List of the Formicidae. Occasional Papers of the Boston Society of Natural History 7: 1-24
 * Wheeler W. M. 1906. Fauna of New England. 7. List of the Formicidae. Occasional Papers of the Boston Society of Natural History 7: 1-24.
 * Wheeler W. M. 1908. The ants of Casco Bay, Maine, with observations on two races of Formica sanguinea Latreille. Bulletin of the American Museum of Natural History 24: 619-645.
 * Wheeler W. M. 1916. Formicoidea. Formicidae. Pp. 577-601 in: Viereck, H. L. 1916. Guide to the insects of Connecticut. Part III. The Hymenoptera, or wasp-like insects, of Connecticut. Connecticut State Geological and Natural History Survey. Bulletin 22: 1-824.
 * Wheeler W. M. 1917. The mountain ants of western North America. Proceedings of the American Academy of Arts and Sciences 52: 457-569.
 * Wheeler, G.C. and J. Wheeler. 1978. Mountain ants of Nevada. Great Basin Naturalist 35(4):379-396
 * Wheeler, G.C. and J. Wheeler. 1985. A checklist of Texas ants. Prairie Naturalist 17:49-64.
 * Wheeler, G.C. and J. Wheeler. 1988. A checklist of the ants of Montana. Psyche 95:101-114
 * Wheeler, G.C. and J. Wheeler. 1988. A checklist of the ants of Wyoming. Insecta Mundi 2(3&4):230-239
 * Wheeler, G.C., J. Wheeler and P.B. Kannowski. 1994. CHECKLIST OF THE ANTS OF MICHIGAN (HYMENOPTERA: FORMICIDAE). Great Lakes Entomologist 26:1:297-310
 * Wheeler, G.C., J. Wheeler, T.D. Galloway and G.L. Ayre. 1989. A list of the ants of Manitoba. Proceedings of the Entomological Society of Manitoba 45:34-49
 * Whitcomb W. H., H. A. Denmark, A. P. Bhatkar, and G. L. Greene. 1972. Preliminary studies on the ants of Florida soybean fields. Florida Entomologist 55: 129-142.
 * Wilson E. O. 1955. A monographic revision of the ant genus Lasius. Bulletin of the Museum of Comparative Zoology 113: 1-201
 * Wing M. W. 1939. An annotated list of the ants of Maine (Hymenoptera: Formicidae). Entomological News 50:161-165.
 * Wing M. W. 1968. Taxonomic revision of the Nearctic genus Acanthomyops (Hymenoptera: Formicidae). Memoirs of the Cornell University Agricultural Experiment Station 405: 1-173.
 * Young J., and D. E. Howell. 1964. Ants of Oklahoma. Miscellaneous Publication. Oklahoma Agricultural Experimental Station 71: 1-42.
 * Young, J. and D.E. Howell. 1964. Ants of Oklahoma. Miscellaneous Publications of Oklahoma State University MP-71