Hypoponera opacior

Hypoponera opacior is a wide ranging and highly variable species that may actually be a species complex. It nests in rotten stumps in partial shade, grass sod, or beneath stones (Coovert 2005, Nemec et al. 2012).

Identification
Smith (1936) - P. opacior is highly variable in color, sculpture and the shape of the petiole. From other species the worker can be distinguished by its slender, graceful appearance, finely punctate body, and distinctly triangular shaped petiole (viewed in lateral profile).

This species may be part of a species complex, which is consistent with its variable morphology.

Workers and queens have a high petiole that is strongly narrowed above in lateral view. Small, reddish brown individuals can be confused with Hypoponera punctatissima, but the latter species has a smooth, shining mesopleuron, and a less elevated petiole than H. opacior. This last character state is very useful for sorting specimens, but is not easily quantified and requires a little practice in its use. Workers of H. opacior also have conspicuous punctures on the central portion of the mandible, while this area in H. punctatissima is smooth and shining. Male H. opacior have light brown wing veins (unlike H. opaciceps), and are most similar to males of Hypoponera inexorata. The latter species is considerably larger (3mm or more in length). We have occasionally found ergatoid queens (or workers with large eyes).

Distribution
Virginia south through Florida and the West Indies, west into Oregon, south through Central and South America into Chile and Argentina.

In some areas the distribution of H. opacior may be discontinuous and the populations small: in a survey of the ants of Nevada, Wheeler and Wheeler (1986) found this species only once, while in Florida it is found wherever there is cursory sampling.

Distribution based on Regional Taxon Lists
Nearctic Region: United States. Neotropical Region: Argentina, Costa Rica, Cuba, Dominican Republic, Ecuador, Greater Antilles, Guadeloupe, Haiti, Lesser Antilles, Mexico, Netherlands Antilles, Puerto Rico, Saint Vincent and the Grenadines, Trinidad and Tobago.

Reproduction
Fiotzik and colleagues have studied aspects of the reproductive biology of this species.

From the abstract of Fiotzik et al. (2002): Wingless (apterous) males of Hypoponera opacior sat on top of queen cocoons, inserted their genitalia into the cocoons and remained in copula with cocooned queens for up to 40 h. These males were tolerant of each other; fighting was never recorded. Our observations therefore suggest that wingless males of H. opacior ensure reproduction by copulatory mate guarding. This strategy, although time consuming, presumably reduces the likelihood of subsequent inseminations by other males. Apterous H. opacior males have only a limited amount of sperm available: histological preparations showed that, in contrast to Cardiocondyla fighter males, the testes degenerate in early adult life. Males of H. opacior have relatively few mating opportunities. Although some wingless males were reproductively active for more than 3 weeks, we observed a maximum of only six matings per male, with a mean slightly above one. Some H. opacior males used an alternative reproductive tactic of dispersal and outbreeding. We found colonies headed by single, dealate queens, which did not rear wingless sexuals but presumably reproduced through winged reproductives that mate in nuptial flights. The social structure of those colonies contrasted with nests containing wingless reproductives, which were highly polygynous and polydomous.

From the abstract of Fiotzik et al. (2010): The ant Hypoponera opacior exhibits alternative reproductive morphs of males and females associated with distinct sexual behaviours. Our long-term study reports strong seasonality in sexual production with a mating season in early and one in late summer. Winged (alate) reproductives emerge in June, swarm during the monsoon season and establish new colonies independently. In contrast, wingless worker-like (ergatoid) reproductives that appear in late August mate within their natal or adjacent nests and either do not disperse or establish new nests close by. These divergent dispersal patterns allowed us to analyse the impact of local factors on investment strategies by comparing sex allocation between and within the two reproductive events. The optimal sex ratio for ergatoid reproductives should be influenced both by competition for matings between brothers (local mate competition) and rivalry among young locally dispersing queens for workers, nest sites or food (local resource competition). The greater importance of local resource competition was demonstrated both by a male-biased sex ratio for wingless sexuals and a stronger increase in the number of males with total sexual production than for the number of queens. Microsatellite analysis revealed that inter-nest variation in relatedness asymmetry cannot explain split sex ratios in the August generation. Instead, nests with related ergatoid males raised a male-biased sex ratio contrary to the expectations under local mate competition. In conclusion, male bias in wingless H. opacior indicates that local mate competition is less strong than local resource competition among ergatoid queens over the help of workers during nest foundation.

From the abstract of Fiotzik et al. (2011): The ant Hypoponera opacior exhibits alternative reproductive morphs associated with distinct sexual behaviours. Wingless, worker-like gynes mate in fall in their mother nest. Subsequently, these polygynous nests divide and wingless queens disperse on foot accompanied by workers, which help during nest foundation. Local resource competition amongst wingless queens was indicated by male-biased sex ratios. Queens compete over access to workers, and this competition manifests in dominance interactions. We colour-marked queens and workers to study this behaviour. Marked queens were treated aggressively and frequently killed by nestmates, whilst similarly labelled workers were readopted. We hypothesise that mated wingless queens—although they resemble workers in external morphology—differ chemically and that fertile queens with divergent odours are not tolerated. Thereby, sterile H. opacior workers avoid the risk to raise offsprings of unrelated queens. However, when nests split, workers should base their decision which queen to follow not only on relatedness, but also on queen fertility. Cuticular hydrocarbon profiles of queens could serve as an honest signal for workers. Indeed, gas chromatography showed that hydrocarbon composition differs between workers and mated queens and is linked to ovary development in queens. Workers and unmated queens exhibit complex profiles of short-chained hydrocarbons, whilst fertile queens carry few, long-chained alkenes and branched alkanes. Furthermore, ovary length was correlated to the relative amounts of characteristic hydrocarbons of queens. The chemical profile was also associated with eye size—an unalterable trait in adult insects—suggesting that larval nutrition and consequently body size affects adult ovary development.

Kureck et al. (2011) investigated whether wingless sexuals of the ant Hypoponera opacior adjust mate-guarding behaviour to the level of competition in the nest. Males mate with young nestmate females shortly before these emerge from the cocoon. Aggressive interactions among adult males have never been observed, but males embrace and guard the cocoons of their mating partners for up to 2 days. In laboratory experiments, the duration of pupal guarding increased with the number of adult males in the nest, but decreased with an increasing number of mating partners per male. These findings demonstrate that males are aware of the competitive situation in the nest and adjust their mating behaviour in an adaptive manner. Males also guarded and attempted to copulate with sterile worker and male pupae. These misdirected behaviours might be the result of identification errors, as we found that the cuticular hydrocarbon profiles of young individuals of the different castes were very similar. Copulatory behaviour towards sterile workers is certainly maladaptive, whereas interactions with young males may provide a fitness benefit: We found a high mortality rate of young males that were embraced and guarded by adult males. Adult maleemale pupae interactions predominantly occurred when only a single male was present in the nest. In addition, single-male nests were found at unusually high frequencies. These findings suggest that wingless males try to kill their pupal rivals through embracing when there are few adult competitors in the nest, but switch to mate-guarding behaviour when intramale competition is high.

Brazil
DaRocha et al. (2015) studied the diversity of ants found in bromeliads of a single large tree of Erythrina, a common cocoa shade tree, at an agricultural research center in Ilhéus, Brazil. Forty-seven species of ants were found in 36 of 52 the bromeliads examined. Bromeliads with suspended soil and those that were larger had higher ant diversity. Hypoponera opacior was found in 4 different bromeliads and was associated with the suspended soil and litter of the plants.

Florida
Deyrup, Davis and Cover (unpublished): This species forms small colonies in soil, rotten logs, and in accumulations of leaf litter. Is is the most abundant species of Hypoponera in Florida, and one of the most common ants in leaf litter samples.

Van Pelt (1958), reporting on this species within the Weleka Reserve in Florida: "In all respects except total length, the Welaka specimens agree with Smith's description in which he cites the length of workers of opacior as 2-2.3 mm. Workers from the Reserve measure 2.4-2.7 mm in total length.

P. t. opacior tends to prefer the higher, drier plant associations. It was taken commonly in xeric hammock; occasionally to commonly in turkey oak; occasionally in bayhead and Plummer slash pine flatwoods, in all welldrained areas except Pomello scrubby flatwoods, and in the hammocks; and rarely in the other flatwoods stations. No collections have been made from Pomello scrubby flatwoods or the seasonally flooded areas of the river swamp and marsh. Cole (1940) points out that in the Great Smoky Mountains the ant does not nest in dense wet woods, but prefers rather open areas where the soil is able to contain an appreciable amount of moisture.

The majority of the nests occupied the surface stratum. The several nests recorded from sand, moreover, did not extend more than a few inches into the sand, but were mostly under litter. The nests in the surface stratum were usually associated with debris, although some nests were found with little or no debris. Several of the nests taken in fallen logs were found under the bark against fairly hard wood. The order of preference of nests in the surface stratum is: 1) fallen logs, 2) dead stumps, 3) bases of living trees" 4) litter, 5) palmetto roots on ground, 6) under mat of palmetto roots, and 7) under mat of palmetto trunks.

Of four nests, taken from wood, the number of workers varied from 7 to 21, with an average of 13. None of these nests contained queens. Immature forms have been found in all months. Males have been observed in flight in December and February. No information has been obtained concerning females.

P. t. opacior is relatively fast moving, and characteristically evasive. Individuals, are difficult to see because they are very nearly the color of the wood or litter surrounding their nests. They seek immediately the first available crevice in which to hide.

Nests are occasionally found in the same logs and stumps as nests of Odontomachus baematoda insularis."

New Mexico
Mackay and Mackay (2002) - Habitat. Ranges from Chihuahuan Desert mixed basin bajada to short grass prairie, sagebrush into areas of pinyon-juniper, alligator bark juniper, Chihuahua pines, ponderosa pine and emery oak. Biology. These ants nest under stones, primarily in shady areas with abundant stones.

Costa Rica
Longino (1999): The form I call opacior s.s. seems to have a preference for synanthropic and seasonally dry habitats. I have scattered collections from Costa Rica, as follows:

Santa Rosa National Park: in Winkler sample collected by P. S. Ward.

Finca La Pacifica, a dry-forest riparian habitat: in Winkler sample of sifted litter from the forest floor.

Carara Biological Reserve: in Winkler sample collected by P. S. Ward.

Casa Plastico, a 600m elevation wet site on the Atlantic slope, in young second growth at the edge of a pasture: under a mossmat at the base of a tree.

A coffee farm near Heredia in the Central Valley: collected in studies of coffee farm fauna by Ivette Perfecto.

Sirena in Corcovado National Park, a lowland wet forest site: in Winkler sample of sifted litter from the forest floor.

A roadside in Monteverde, where an isolated and epiphyte-laden tree had recently fallen: a nest was under epiphytes near the ground.

La Selva Biological Station, a mature lowland wet forest site: a nest was under epiphytes on a dead branch recently fallen from the canopy.

Fila Cruces near San Vito: a nest was under a stone at the edge of a gravel road through pastures and scrubby forest.

Parque Nacional, a small landscaped park in the middle of San Jose: among a collection of stray foragers.

Genetics
From the abstract of Ruger et al. (2005): In the species Hypoponera opacior, winged and wingless males and queens co-occur and molecular markers are essential to study the reproductive success of these sexual tactics. Primers were developed for five polymorphic microsatellite loci isolated from H. opacior. Their variability was tested on 34 colonies from a population in the United States. Nine to 21 alleles per locus were found with observed heterozygosities between 0.1 and 0.7. A significantly positive Fis value suggests inbreeding in this ant with predominant intracolonial matings.

Nomenclature

 *  opacior. Ponera trigona var. opacior Forel, 1893g: 363 (w.q.) WEST INDIES. Emery, 1895c: 268 (m.); Wheeler, G.C. & Wheeler, J. 1964b: 454 (l.). Combination in Hypoponera: Taylor, 1968a: 65. Subspecies of trigona: Emery, 1906c: 116. Raised to species: Kempf, 1962b: 10.

Taxonomic Notes
There is some variation in size and color, as in many other ponerines. This variation does not seem to be correlated with any structural variation, and it is easy to set up series of specimens showing transitions between light and dark, or large and small. There is also some regional variation. Arizona specimens that we have seen are relatively large and reddish; specimens from coastal Oregon are dark brown with a yellowish tip to the gaster. It is possible that such populations represent distinct species, but it would require genetic work, or some major difference in males, to show that this is the case. It is normal for widely disjunct populations in different habitats to look a little different.

Longino (1999): Hypoponera opacior is probably a species complex; a brief perusal of collections from throughout its range reveals abundant geographic variation in details of color, pilosity, and eye size. For example, Californian material that has traditionally been identified as opacior can be separated into two or three distinct species (P. S. Ward, pers. comm.). This pattern suggests an older lineage whose current distribution is the result of vicariance rather than recent dispersal, at least over parts of its range. It is also possible that some parts of the range are the result of recent dispersal. For example, is its current distribution on many Caribbean islands a result of old vicariance or recent introductions?

Worker
Smith (1936) - Length: 2-2.3 mm. Head, excluding the mandibles, longer than broad, posterior border faintly emarginate, posterior angles rounded, sides convex. Eyes small, rounded, apparently composed of about 6 facets, placed about the anterior fourth of the sides of the head. Mandibles rather irregularly toothed. Scapes somewhat robust, not attaining the posterior border of the head; funicular segments 3-6 much broader than long, the terminal segment scarcely as long as the three preceding segments taken together. Thorax short, fairly robust, with distinct pro-mesonotal and meso-epinotal sutures; viewed in lateral profile the base of the epinotum appears to be of the same length as the declivity and meets the latter in a very distinct obtuse angle, the declivity faintly concave. Petiole when viewed in lateral profile not as robust as in the other species, very distinctly narrower (antero-posteriorly) dorsally than ventrally, strongly convex anteriorly, faintly convex posteriorly; viewed posteriorly the rounded superior border meets the lateral borders in such a way as to give an almost circular effect.

Body, due to the faint sculpturing and the nature of the pubescence rather glabrous, in some lights however somewhat subopaque. Head covered with extremely fine, dense punctures; thorax much less so; pleurae of the meso-and metathorax, especially the lower parts, finely and longitudinally striated.

Pubescence extremely fine on all parts of the body but probably most apparent on the gaster and head, pubescence scarcely hiding the general surface of the body, but in some lights giving it a pruinose effect. Pilosity poorly developed; hairs grayish, most noticeable on the dorsal surfaces of the head and gaster, on the former strikingly long, on the latter short, except toward the apex of the gaster.

Color variable; ranging from light brown to practically black, appendages lighter.

Queen
Length: 2.75-3 mm. Very similar to the worker but larger and more robust. As with the worker funicular segments 3-6 are much broader than long, and the terminal segment shorter than the preceding segments taken together. The most distinctive character is the shape of the petiole which is very narrow antero-posteriorly and clearly thinner above than at the base. The wing veins and stigma very light brown.

Type Material
Island of St. Vincent, Lesser Antilles.

References based on Global Ant Biodiversity Informatics

 * Achury R., and A.V. Suarez. 2017. Richness and composition of ground-dwelling ants in tropical rainforest and surrounding landscapes in the Colombian Inter-Andean valley. Neotropical Entomology https://doi.org/10.1007/s13744-017-0565-4
 * Adams T. A., W. J. Staubus, and W. M. Meyer. 2018. Fire impacts on ant assemblages in California sage scrub. Southwestern Entomologist 43(2): 323-334.
 * Ahuatzin D. A., E. J. Corro, A. Aguirre Jaimes, J. E. Valenzuela Gonzalez, R. Machado Feitosa, M. Cezar Ribeiro, J. Carlos Lopez Acosta, R. Coates, W. Dattilo. 2019. Forest cover drives leaf litter ant diversity in primary rainforest remnants within human-modified tropical landscapes. Biodiversity and Conservation 28(5): 1091-1107.
 * Alayo D. P. 1974. Introduccion al estudio de los Himenopteros de Cuba. Superfamilia Formicoidea. Academia de Ciencias de Cuba. Instituto de Zoologia. Serie Biologica no.53: 58 pp. La Habana.
 * Albrecht M. 1995. New Species Distributions of Ants in Oklahoma, including a South American Invader. Proc. Okla. Acad. Sci. 75: 21-24.
 * Allred D. M. 1982. Ants of Utah. The Great Basin Naturalist 42: 415-511.
 * Allred, D.M. 1982. The ants of Utah. Great Basin Naturalist 42:415-511.
 * Annotated Ant Species List Ordway-Swisher Biological Station. Downloaded at http://ordway-swisher.ufl.edu/species/os-hymenoptera.htm on 5th Oct 2010.
 * Atchison R. A., J. Hulcr, and A. Lucky. 2018. Managed fire frequency significantly influences the litter arthropod community in longleaf pine flatwoods. Environmental Entomology 20(10): 1-11.
 * 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.
 * Boer P. 2019. Ants of Curacao, species list. Accessed on January 22 2019 at http://www.nlmieren.nl/websitepages/SPECIES%20LIST%20CURACAO.html
 * Boer P. 2019. Ants of Saba, species list. Accessed on January 22 2019 at http://www.nlmieren.nl/websitepages/SPECIES%20LIST%20SABA.html
 * Brandao, C.R.F. 1991. Adendos ao catalogo abreviado das formigas da regiao neotropical (Hymenoptera: Formicidae). Rev. Bras. Entomol. 35: 319-412.
 * Bruch C. 1914. Catálogo sistemático de los formícidos argentinos. Revista del Museo de La Plata 19: 211-234.
 * Carroll T. M. 2011. The ants of Indiana (Hymenoptera: Formicidae). Master's Thesis Purdue university, 385 pages.
 * Castano-Meneses, G., M. Vasquez-Bolanos, J. L. Navarrete-Heredia, G. A. Quiroz-Rocha, and I. Alcala-Martinez. 2015. Avances de Formicidae de Mexico. Universidad Nacional Autonoma de Mexico.
 * Clouse R. 1999. Leaf-Litter Inhabitants of a Brazilian Pepper Stand in Everglades National Park. The Florida Entomologist. 82: 388-403
 * 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.
 * Cokendolpher J.C., Reddell J.R., Taylor S.J, Krejca J.K., Suarez A.V. and Pekins C.E. 2009. Further ants (Hymenoptera: Formicidae) from caves of Texas [Hormigas (Hymenoptera: Formicdae) adicionales de cuevas de Texas]. Texas Memorial Museum Speleological Monographs, 7. Studies on the cave and endogean fauna of North America, V. Pp. 151-168
 * Colby, D. and D. Prowell. 2006. Ants (Hymenoptera: Formicidae) in Wet Longleaf Pine Savannas in Louisiana. Florida Entomologist 89(2):266-269
 * Cole A. C., Jr. 1949. The ants of Mountain Lake, Virginia. Journal of the Tennessee Academy of Science 24: 155-156.
 * 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
 * Corassa J. N., I. C. Magistrali, J. C. Moreno, E. B. Cantarelli, and A. Corassa. Effect of formicid granulated baits on non-target ants biodiversity in eucalyptus plantations litter. Comunicata Scientiae 4(1): 35-42.
 * Cover S. P., and R. A. Johnson. 20011. Checklist of Arizona Ants. Downloaded on January 7th at http://www.asu.edu/clas/sirgtools/AZants-2011%20updatev2.pdf
 * Cuezzo, F. 1998. Formicidae. Chapter 42 in Morrone J.J., and S. Coscaron (dirs) Biodiversidad de artropodos argentinos: una perspectiva biotaxonomica Ediciones Sur, La Plata. Pages 452-462.
 * Dash S. T. 2011. A taxonomic revision of the New World Hypoponera Santschi, 1938 (Hymenoptera: Formicidae). PhD Thesis University of Texas at El Paso, 296 pages.
 * Dash S. T. and L. M. Hooper-Bui. 2008. Species diversity of ants (Hymenoptera: Formicidae) in Louisiana. Conservation Biology and Biodiversity. 101: 1056-1066
 * Dattilo W. et al. 2019. MEXICO ANTS: incidence and abundance along the Nearctic-Neotropical interface. Ecology https://doi.org/10.1002/ecy.2944
 * De La Mora, A., and S. M. Philpott. 2010. Wood-nesting ants and their parasites in forests and coffee agroecosystems. Environmental Entomology 39: 1473-1481.
 * Dean D. A., and S. R. Dean. 2018. A survey of the ant fauna and seasonal alate nuptial flights at two locations in South-Central Texas. Southwestern Entomologist 43(3): 639-647.
 * Del Toro, I. 2010. PERSONAL COMMUNICATION. MUSEUM RECORDS COLLATED BY ISRAEL DEL TORO
 * Del Toro, I., M. Vázquez, W.P. Mackay, P. Rojas and R. Zapata-Mata. Hormigas (Hymenoptera: Formicidae) de Tabasco: explorando la diversidad de la mirmecofauna en las selvas tropicales de baja altitud. Dugesiana 16(1):1-14.
 * Des Lauriers J., and D. Ikeda. 2017. The ants (Hymenoptera: Formicidae) of the San Gabriel Mountains of Southern California, USA with an annotated list. In: Reynolds R. E. (Ed.) Desert Studies Symposium. California State University Desert Studies Consortium, 342 pp. Pages 264-277.
 * Deyrup M., L. Davis, and S. Buckner. 1998. Composition of the ant fauna of three Bahamian islands. Proceedings of the seventh symposium on the natural history of the Bahamas. 23-32. Bahamian Field Station, San Salvador, Bahamas
 * Deyrup, M. and J. Trager. 1986. Ants of the Archbold Biological Station, Highlands County, Florida (Hymenoptera: Formicidae). Florida Entomologist 69(1):206-228
 * DuBois M. B. 1981. New records of ants in Kansas, III. State Biological Survey of Kansas. Technical Publications 10: 32-44
 * DuBois M. B. 1985. Distribution of ants in Kansas: subfamilies Ponerinae, Ecitoninae, and Myrmicinae (Hymenoptera: Formicidae). Sociobiology 11: 153-200
 * DuBois M. B. 1985. Distribution of ants in Kansas: subfamilies Ponerinae, Ecitoninae, and Myrmicinae (Hymenoptera: Formicidae). Sociobiology 11: 153-201
 * DuBois M. B. 1985. Distribution of ants in Kansas: subfamilies Ponerinae, Ecitoninae, and Myrmicinae (Hymenoptera: Formicidae). Sociobiology 11: 153-202
 * DuBois M. B. 1985. Distribution of ants in Kansas: subfamilies Ponerinae, Ecitoninae, and Myrmicinae (Hymenoptera: Formicidae). Sociobiology 11: 153-203
 * DuBois M. B. 1985. Distribution of ants in Kansas: subfamilies Ponerinae, Ecitoninae, and Myrmicinae (Hymenoptera: Formicidae). Sociobiology 11: 153-204
 * DuBois M. B. 1985. Distribution of ants in Kansas: subfamilies Ponerinae, Ecitoninae, and Myrmicinae (Hymenoptera: Formicidae). Sociobiology 11: 153-205
 * DuBois M. B. 1985. Distribution of ants in Kansas: subfamilies Ponerinae, Ecitoninae, and Myrmicinae (Hymenoptera: Formicidae). Sociobiology 11: 153-206
 * DuBois M. B. 1985. Distribution of ants in Kansas: subfamilies Ponerinae, Ecitoninae, and Myrmicinae (Hymenoptera: Formicidae). Sociobiology 11: 153-207
 * DuBois M. B. 1985. Distribution of ants in Kansas: subfamilies Ponerinae, Ecitoninae, and Myrmicinae (Hymenoptera: Formicidae). Sociobiology 11: 153-208
 * DuBois M. B. 1985. Distribution of ants in Kansas: subfamilies Ponerinae, Ecitoninae, and Myrmicinae (Hymenoptera: Formicidae). Sociobiology 11: 153-209
 * DuBois M. B. 1985. Distribution of ants in Kansas: subfamilies Ponerinae, Ecitoninae, and Myrmicinae (Hymenoptera: Formicidae). Sociobiology 11: 153-210
 * DuBois M. B. 1985. Distribution of ants in Kansas: subfamilies Ponerinae, Ecitoninae, and Myrmicinae (Hymenoptera: Formicidae). Sociobiology 11: 153-211
 * DuBois M. B. 1985. Distribution of ants in Kansas: subfamilies Ponerinae, Ecitoninae, and Myrmicinae (Hymenoptera: Formicidae). Sociobiology 11: 153-212
 * DuBois M. B. 1985. Distribution of ants in Kansas: subfamilies Ponerinae, Ecitoninae, and Myrmicinae (Hymenoptera: Formicidae). Sociobiology 11: 153-213
 * DuBois M. B. 1985. Distribution of ants in Kansas: subfamilies Ponerinae, Ecitoninae, and Myrmicinae (Hymenoptera: Formicidae). Sociobiology 11: 153-214
 * DuBois M. B. 1985. Distribution of ants in Kansas: subfamilies Ponerinae, Ecitoninae, and Myrmicinae (Hymenoptera: Formicidae). Sociobiology 11: 153-215
 * DuBois M. B. 1985. Distribution of ants in Kansas: subfamilies Ponerinae, Ecitoninae, and Myrmicinae (Hymenoptera: Formicidae). Sociobiology 11: 153-216
 * DuBois M. B. 1985. Distribution of ants in Kansas: subfamilies Ponerinae, Ecitoninae, and Myrmicinae (Hymenoptera: Formicidae). Sociobiology 11: 153-217
 * DuBois M. B. 1985. Distribution of ants in Kansas: subfamilies Ponerinae, Ecitoninae, and Myrmicinae (Hymenoptera: Formicidae). Sociobiology 11: 153-218
 * DuBois M. B. 1985. Distribution of ants in Kansas: subfamilies Ponerinae, Ecitoninae, and Myrmicinae (Hymenoptera: Formicidae). Sociobiology 11: 153-219
 * 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
 * Emery C. 1906. Studi sulle formiche della fauna neotropica. XXVI. Bullettino della Società Entomologica Italiana 37: 107-194.
 * Emery C. 1911. Hymenoptera. Fam. Formicidae. Subfam. Ponerinae. Genera Insectorum 118: 1-125.
 * Emery, C.. "Sopra alcune formiche della fauna mediterranea." Memorie della Reale Accademia delle Scienze dell'Istituto di Bologna (5)5 (1895): 59-75 [pagination of separate: 291-307].
 * Favretto M. A., E. Bortolon dos Santos, and C. J. Geuster. 2013. Entomofauna from West of Santa Catarina State, South of Brazil. EntomoBrasilis 6 (1): 42-63.
 * Fernandes, P.R. XXXX. Los hormigas del suelo en Mexico: Diversidad, distribucion e importancia (Hymenoptera: Formicidae).
 * Fernández F., and T. M. Arias-Penna. 2008. Las hormigas cazadoras en la región Neotropical. Pp. 3-39 in: Jiménez, E.; Fernández, F.; Arias, T.M.; Lozano-Zambrano, F. H. (eds.) 2008. Sistemática, biogeografía y conservación de las hormigas cazadoras de Colombia. Bogotá: Instituto de Investigación de Recursos Biológicos Alexander von Humboldt, xiv + 609 pp.
 * Fernández, F. and S. Sendoya. 2004. Lista de las hormigas neotropicales. Biota Colombiana Volume 5, Number 1.
 * Fichaux M., B. Bechade, J. Donald, A. Weyna, J. H. C. Delabie, J. Murienne, C. Baraloto, and J. Orivel. 2019. Habitats shape taxonomic and functional composition of Neotropical ant assemblages. Oecologia 189(2): 501-513.
 * Foitzik, S., J. Froba, M.H. Ruger and V.Witte. 2011. Competition over workers: fertility signalling in wingless queens of Hypoponera opacior. Insectes Sociaux 58:271-278
 * Fontanla Rizo J.L. 1997. Lista preliminar de las hormigas de Cuba. Cocuyo 6: 18-21.
 * Fontenla J. L., and J. Alfonso-Simonetti. 2018. Classification of Cuban ants (Hymenoptera: Formicidae) into functional groups. Poeyana Revista Cubana de Zoologia 506: 21-30.
 * Fontenla Rizo J. L. 1997. Lista preliminar de las hormigas de Cuba (Hymenoptera: Formicidae). Cocuyo 6: 18-21.
 * Forster J.A. 2005. The Ants (hymenoptera: Formicidae) of Alabama. Master of Science, Auburn University. 242 pages.
 * Franco W., N. Ladino, J. H. C. Delabie, A. Dejean, J. Orivel, M. Fichaux, S. Groc, M. Leponce, and R. M. Feitosa. 2019. First checklist of the ants (Hymenoptera: Formicidae) of French Guiana. Zootaxa 4674(5): 509-543.
 * 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.
 * Galkowski C. 2016. New data on the ants from the Guadeloupe (Hymenoptera, Formicidae). Bull. Soc. Linn. Bordeaux 151, 44(1): 25-36.
 * Gallardo A. 1918. Las hormigas de la República Argentina. Subfamilia Ponerinas. Anales del Museo Nacional de Historia Natural de Buenos Aires 30: 1-112.
 * 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.
 * General D., and L. Thompson. 2008. Ants of Arkansas Post National Memorial: How and Where Collected. Journal of the Arkansas Academy of Science 62: 52-60.
 * General D.M. & Thompson L.C. 2007. Ants (Hymenoptera: Formicidae) of Arkansas Post National Memorial. Journal of the Arkansas Acaedemy of Science. 61: 59-64
 * General D.M. & Thompson L.C. 2008. New Distributional Records of Ants in Arkansas for 2008. Journal of the Arkansas Academy of Science. 63: 182-184
 * Goetsch, W., and C. Menozzi. "Die Ameisen Chiles." Konowia 14 (1935): 94-102.
 * Gregg, R.T. 1963. The Ants of Colorado.
 * Groc S., J. H. C. Delabie, F. Fernandez, F. Petitclerc, B. Corbara, M. Leponce, R. Cereghino, and A. Dejean. 2017. Litter-dwelling ants as bioindicators to gauge the sustainability of small arboreal monocultures embedded in the Amazonian rainforest. Ecological Indicators 82: 43-49.
 * Groc S., J. H. C. Delabie, F. Fernandez, M. Leponce, J. Orivel, R. Silvestre, Heraldo L. Vasconcelos, and A. Dejean. 2013. Leaf-litter ant communities (Hymenoptera: Formicidae) in a pristine Guianese rainforest: stable functional structure versus high species turnover. Myrmecological News 19: 43-51.
 * Grodsky S. M., J. W. Campbell, S. R. Fritts, T. B. Wigley, and C. E. Moorman. 2018. Variable responses of non-native and native ants to coarse woody debris removal following forest bioenergy harvests. Forest Ecology and Management doi.org/10.1016/j.foreco.2018.02.010
 * Hess C. G. 1958. The ants of Dallas County, Texas, and their nesting sites; with particular reference to soil texture as an ecological factor. Field and Laboratory 26: 3-72.
 * 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
 * Holway D.A. 1998. Effect of Argentine ant invasions on ground-dwelling arthropods in northern California riparian woodlands. Oecologia. 116: 252-258
 * INBio Collection (via Gbif)
 * Ilha C., J. A. Lutinski, D. Von Muller Pereira, F. R. Mello Garcia. 2009. Riqueza de formigas (Hymenoptera: Formicidae) de Bacia da Sanga Caramuru, municipio de Chapeco-SC. Biotemas 22(4): 95-105.
 * 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.
 * Johnson C. 1986. A north Florida ant fauna (Hymenoptera: Formicidae). Insecta Mundi 1: 243-246
 * Johnson R. Personnal Database. Accessed on February 5th 2014 at http://www.asu.edu/clas/sirgtools/resources.htm
 * Kempf W. W. 1970. Catálogo das formigas do Chile. Papeis Avulsos de Zoologia (São Paulo) 23: 17-43.
 * Kempf W. W., and K. Lenko. 1976. Levantamento da formicifauna no litoral norte e ilhas adjacentes do Estado de São Paulo, Brasil. I. Subfamilias Dorylinae, Ponerinae e Pseudomyrmecinae (Hym., Formicidae). Studia Entomologica 19: 45-66.
 * Kempf, W.W. 1972. Catalago abreviado das formigas da regiao Neotropical (Hym. Formicidae) Studia Entomologica 15(1-4).
 * LeBrun E. G., R. M. Plowes, and L. E. Gilbert. 2015. Imported fire ants near the edge of their range: disturbance and moisture determine prevalence and impact of an invasive social insect. Journal of Animal Ecology,81: 884–895.
 * Longino J. T., and R. K. Colwell. 2011. Density compensation, species composition, and richness of ants on a neotropical elevational gradient. Ecosphere 2(3): 16pp.
 * Longino J. et al. ADMAC project. Accessed on March 24th 2017 at https://sites.google.com/site/admacsite/
 * Longino, J.T. 2010. Personal Communication. Longino Collection Database
 * Lubertazi, D. Personal Communication. Specimen Data from Museum of Comparative Zoology at Harvard
 * Lutinski J. A., F. R. Mello Garcia, C. J. Lutinska, and S. Iop. 2008. Ants diversity in Floresta Nacional de Chapecó in Santa Catarina State, Brazil. Ciência Rural, Santa Maria 38(7): 1810-1816.
 * 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, R. L. Brown, T. L. Schiefer, J. G. Lewis. 2012. Ant diversity at Noxubee National Wildlife Refuge in Oktibbeha, Noxubee, and Winston Counties, Mississippi. Mississippi Agricultural and Forestry Experiment Station Bulletin 1197: 1-30
 * MacGown J. A., J. G. Hill, and M. Deyrup. 2009. Ants (Hymenoptera: Formicidae) of the Little Ohoopee River Dunes, Emanuel County, Georgia. J. Entomol. Sci. 44(3): 193-197.
 * 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 R. L. Brown. 2006. Survey of the ants (Hymenoptera: Formicidae) of the Tombigbee National Forest in Mississippi. Journal of the Kansas Entomological Society 79(4):325-340.
 * MacGown J. A., and R. Whitehouse. 2015. A preliminary report of the ants of West Ship Island. A report submitted to the Gulf Islands National Seashore. Mississippi Entomological Museum Report #2015-02. 9 pp.
 * MacGown, J. and J.G. Hill. Ants collected at Palestinean Gardens, George County Mississippi.
 * 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.A. and R.L. Brown. 2006. Observations on the High Diversity of Native Ant Species Coexisting with Imported Fire Ants at a Microspatial Scale in Mississippi. Southeastern Naturalist 5(4):573-586
 * MacGown, J.A. and R.L. Brown. 2006. Survey of the ants (Hymenoptera: Formicidae) of the Tombigbee National Forest in Mississippi. Journal of the Kansas Entomological Society 79(4):325-340.
 * MacGown, J.A. and T. Lockley. Ants of Horn Island, Jackson County, Mississippi
 * MacGown, J.A., J.G. Hill, R.L. Brown and T.L. 2009. Ant Diversity at Noxubee National Wildlife Refuge in Oktibbeha, Noxubee, and Winston Counties, Mississippi Report #2009-01. Schiefer. 2009.
 * MacGown. J. 2011. Ants collected during the 25th Annual Cross Expedition at Tims Ford State Park, Franklin County, Tennessee
 * Macgown J. A., S. Y. Wang, J. G. Hill, and R. J. Whitehouse. 2017. A List of Ants (Hymenoptera: Formicidae) Collected During the 2017 William H. Cross Expedition to the Ouachita Mountains of Arkansas with New State Records. Transactions of the American Entomological Society, 143(4): 735-740.
 * Mackay W. P., and E. E. Mackay. 2002. The ants of New Mexico (Hymenoptera: Formicidae). Lewiston, New York: Edwin Mellen Press, 400 pp.
 * 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.
 * Menozzi C, Russo G. 1930. Contributo alla conoscenza della mirmecofauna della Repubblica Dominicana (Antille). Bollettino del Laboratorio di Zoologia Generale e Agraria della Reale Scuola Superiore d'Agricoltura. Portici. 24: 148-173.
 * Menozzi C. 1935. Fauna Chilensis. II. (Nach Sammlungen von W. Goetsch). Le formiche del Cile. Zoologische Jahrbücher. Abteilung für Systematik, Ökologie und Geographie der Tiere. 67: 319-336.
 * Moreau C. S., M. A. Deyrup, and L. R. David Jr. 2014. Ants of the Florida Keys: Species Accounts, Biogeography, and Conservation (Hymenoptera: Formicidae). J. Insect Sci. 14(295): DOI: 10.1093/jisesa/ieu157
 * Morrison L. W. 1998. A review of Bahamian ant (Hymenoptera: Formicidae) biogeography. Journal of Biogeography 25: 561-571.
 * Munsee J. R. 1968. Nine species of ants (Formicidae) recently recorded from Indiana. Proc. Indiana Acad. Sci. 77: 222-227.
 * Nemec K. T., J. C. Trager, E. Manley, and C. R. Allen. Five new records of ants (Hymenoptera: Formicidae0 from Nebraska. The Prairie Naturalist 44(10: 63-65.
 * 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.
 * Parys K. A., M. L. Gimmel, and S. J. Johnson. 2013. Checklist of Insects Associated with Salvinia minima Baker in Louisiana, USA. Check List 9(6): 14881495.
 * Perez-Gelabert D. E. 2008. Arthropods of Hispaniola (Dominican Republic and Haiti): A checklist and bibliography. Zootaxa 1831:1-530.
 * Pires de Prado L., R. M. Feitosa, S. Pinzon Triana, J. A. Munoz Gutierrez, G. X. Rousseau, R. Alves Silva, G. M. Siqueira, C. L. Caldas dos Santos, F. Veras Silva, T. Sanches Ranzani da Silva, A. Casadei-Ferreira, R. Rosa da Silva, and J. Andrade-Silva. 2019. An overview of the ant fauna (Hymenoptera: Formicidae) of the state of Maranhao, Brazil. Pap. Avulsos Zool. 59: e20195938.
 * Portuondo E. F., and J. L. Reyes. 2002. Mirmecofauna de los macizos montañosos de Sierra Maestra y Nipe-Sagua-Baracoa. Cocuyo 12: 10-13
 * Portuondo Ferrer E., and J. L. Fernández Triana. 2005. Species of hymenopterans (bees, wasps, and ants) recorded in Alejandro de Humboldt National Park, from literature records, revision of the collection at BIOECO, and collections before and during the rapid inventory, 12-22 February 2004. In Fong G., A., D. Maceira F., W. S. Alverson, y/and T. Wachter, eds. 2005. Cuba: Parque Nacional Alejandro de Humboldt. Rapid Biological Inventories Report 14. The Field Museum, Chicago.
 * Portuondo Ferrer, E. and J. Fernandez Triana. Biodiversidad del orden Hymenoptera en Los Macizos Montanosos de Cuba Oriental. Boletin S.E.A. 35:121-136.
 * Reddell J. R., and J. C. Cokendolpher. 2001. Ants (Hymenoptera: Formicidae) from caves of Belize, Mexico, and California and Texas (U.S.A.) Texas. Texas Memorial Museum Speleological Monographs 5: 129-154.
 * Reyes, J. L.. "Inventario de la colección de hormigas (Hymenoptera: Formicidae) del Centro Oriental de Ecosistemas y Biodiversidad, Santiago de Cuba, Cuba." Boletín de la Sociedad Aragonesa 36 (2005): 279-283.
 * 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.
 * Rosa da Silva R. 1999. Formigas (Hymenoptera: Formicidae) do oeste de Santa Catarina: historico das coletas e lista atualizada das especies do Estado de Santa Catarina. Biotemas 12(2): 75-100.
 * Rüger, M.H., J. Fröba and S. Foitzik. 2008. Larval cannibalism and worker-induced separation of larvae in Hypoponera ants: a case of conflict over caste determination? Insectes Sociaux 55(1): 12-21.
 * Salinas P. J. 2010. Catalogue of the ants of the Táchira State, Venezuela, with notes on their biodiversity, biogeography and ecology (Hymenoptera: Formicidae: Amblyioponinae, Ponerinae, Proceratiinae, Myrmicinae, Ecitoninae, Formicinae, Pseudomyrmecinae, Dolichoderinae). Boletín de la SEA 47: 315-328.
 * Sanders, N.J. 2004.Immediate Effects of Fire on the Invasive Argentine Ant, Linepithema humile. The Southwestern Naturalist 49(2) :246-250
 * Santschi F. 1930. Quelques fourmis de Cuba et du Brésil. Bulletin. Société Entomologique d'Egypte. 14: 75-83.
 * Schonberg, L.A., J.T. Longino, N.M. Nadkarni and S.P. Yanoviak. 2004. Arboreal Ant Species Richness in Primary Forest, Secondary Forest, and Pasture Habitats of a Tropical Montane Landscape. Biotropica 36(3):402-409.
 * 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). Entomological News 46: 235-241.
 * Smith M. R. 1936. A list of the ants of Texas. Journal of the New York Entomological Society 44: 155-170.
 * Smith M. R. 1936. Ants of the genus Ponera in America, north of Mexico. Annals of the Entomological Society of America 29: 420-430.
 * Smith M. R. 1937. The ants of Puerto Rico. Journal of Agriculture of the University of Puerto Rico 20: 819-875.
 * Smith, Marion R. 1954. Ants of the Bimini Island Group, Bahamas, British West Indies. American Museum of Natural History. 1671. 1-16.
 * Snelling R. R., and J. H. Hunt. 1975. The ants of Chile (Hymenoptera: Formicidae) Revista Chilena de Entomología 9: 63-129.
 * Solomon, S.E. and A.S. Mikheyev. 2005. The ant (Hymenoptera: Formicidae) fauna of Cocos Island, Costa Rica. Florida Entomologist 88(4):415-423
 * Staubus W. J., E. S. Boyd, T. A. Adams, D. M. Spear, M. M. Dipman, W. M. Meyer III. 2015. Ant communities in native sage scrub, non-native grassland, and suburban habitats in Los Angeles County, USA: conservation implications. Journal of Insect Conservervation 19:669–680
 * Theunis, L., M. Gilbert, Y. Roisin and M. Leponce. 2005. Spatial structure of litter-dwelling ant distribution in a subtropical dry forest. Insectes Sociaux. 52:366377.
 * 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.
 * Torres J.A. 1984. Niches and Coexistence of Ant Communities in Puerto Rico: Repeated Patterns. Biotropica 16(4): 284-295.
 * Torres, Juan A. and Roy R. Snelling. 1997. Biogeography of Puerto Rican ants: a non-equilibrium case?. Biodiversity and Conservation 6:1103-1121.
 * Valenzuela-Gonzalez J., A. V. Parra-Cabanas, L. Quiroz-Robledo, D. L. Martinez-Tlapa, and E. D. Montes-de-Oca-Torres. 2013. Variación de la mirmecofauna en un gradiente altitudinal en la región central de Veracruz, México. In Formicidae de Mexico (eds. M. Vasquez-Bolanos, G. Castano-Meneses, A. Cisneros-Caballero, G. A. Quiroz-Rocha, and J. L. Navarrete-Heredia) p 75-82.
 * Van Pelt A. F. 1948. A Preliminary Key to the Worker Ants of Alachua County, Florida. The Florida Entomologist 30(4): 57-67
 * Van Pelt A. F. 1956. The ecology of the ants of the Welaka Reserve, Florida (Hymenoptera: Formicidae). American Midland Naturalist 56: 358-387
 * 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.
 * Van Pelt, A. 1983. Ants of the Chisos Mountains, Texas (Hymenoptera: Formicidae) . Southwestern Naturalist 28:137-142.
 * Vittar, F. 2008. Hormigas (Hymenoptera: Formicidae) de la Mesopotamia Argentina. INSUGEO Miscelania 17(2):447-466
 * Vásquez-Bolaños M. 2011. Lista de especies de hormigas (Hymenoptera: Formicidae) para México. Dugesiana 18: 95-133
 * Ward P. S. 1987. Distribution of the introduced Argentine ant (Iridomyrmex humilis) in natural habitats of the lower Sacramento Valley and its effects on the indigenous ant fauna. Hilgardia 55: 1-16
 * Warren, L.O. and E.P. Rouse. 1969. The Ants of Arkansas. Bulletin of the Agricultural Experiment Station 742:1-67
 * Wetterer, J. K.; Ward, P. S.; Wetterer, A. L.; Longino, J. T.; Trager, J. C.; Miller, S. E. 2000. Ants (Hymenoptera: Formicidae) of Santa Cruz Island, California. Bulletin of the Southern California Academy of Sciences 99:25-31.
 * Wetterer, J.K., P.S. Ward, A.L. Wetterer, J.T. Longino, J.C. Trager and S.E. Miller. 2000. Ants (Hymenoptera:Formicidae) of Santa Cruz Island, California. Bulletin of the Southern California Academy of Science 99(1):25-31.
 * 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 W. M. 1905. The ants of the Bahamas, with a list of the known West Indian species. Bulletin of the American Museum of Natural History 21: 79-135.
 * Wheeler W. M. 1908. The ants of Texas, New Mexico and Arizona. (Part I.). Bulletin of the American Museum of Natural History 24: 399-485.
 * Wheeler W. M. 1913. Ants collected in Georgia by Dr. J. C. Bradley and Mr. W. T. Davis. Psyche (Cambridge) 20: 112-117.
 * Wheeler W. M. 1913. The ants of Cuba. Bulletin of the Museum of Comparative Zoology 54: 477-505.
 * Wheeler W. M. 1917. Jamaican ants collected by Prof. C. T. Brues. Bulletin of the Museum of Comparative Zoology 61: 457-471.
 * Wheeler W. M. 1922. The ants of Trinidad. American Museum Novitates 45: 1-16.
 * Wheeler W. M. 1937. Ants mostly from the mountains of Cuba. Bulletin of the Museum of Comparative Zoology. 81: 439-465.
 * Wheeler, G.C. and J. Wheeler. 1985. A checklist of Texas ants. Prairie Naturalist 17:49-64.
 * 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.
 * 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
 * Zettler J. A., M. D. Taylor, C. R. Allen, and T. P. Spira. 2004. Consequences of Forest Clear-Cuts for Native and Nonindigenous Ants (Hymenoptera: Formicidae). Ann. Entomol. Soc. Am. 97(3): 513-518.