Pheidole bicarinata

From Wilson (2003): P. bicarinata is highly adaptable across its range in both habitat and nest site. In Colorado, Gregg (1963) recorded bicarinata between 1000 and 2100 m, nesting in rotten logs as well as various types of soil beneath stones. The habitats occupied were very diverse, comprising pinyon-cedar and mixed deciduous woodland, canyon-bottom meadows, shortgrass prairie, and sagebrush desert. In Utah and Nevada, bicarinata occurs from sagebrush desert to montane woodland, nesting under stones and pieces of wood, as well as crater nests in open soil (Ingham 1959, 1963; Cole 1966b; Allred and Cole 1979). In western Texas, it has been found in xeric habitats in open soil and under stones, cow dung, a bale of hay, and in cracks in asphalt. Beck et al., studying scavenging ants of Utah, observed workers feeding on dead rodents. Males were found in one Colorado nest in early July. In the eastern United States the species favors sandy soils with some clay content. It is often abundant in open, disturbed grassy areas, such as lawns, golf courses, and abandoned fields (Stefan Cover, personal communication). It is omnivorous in diet, and regularly harvests seeds. Diana E. Wheeler and Frederik Nijhout used bicarinata in their pioneering studies to demonstrate the role of juvenile hormone in the determination of the major and minor subcastes; they traced development through four larval instars (see Wheeler 1991).

Identification
Mackay and Mackay (2002) - For co-occuring Pheidole species in New Mexico, Pheidole bicarinata is difficult to separate from Pheidole cerebrosior. It differs in that the vertex of the head of the minor worker is smooth and polished, or finely, transversely striolate, whereas the vertex of P. cerebrosior is densely punctate. There is a considerable amount of variation in the sculpturing of the vertex of the minor worker, and it is often difficult to identify a series, as different workers may have characteristics of both species. Pheidole bicarinata could be confused with Pheidole marcidula, but can be separated as the minor lacks the clavate hairs on the dorsum of the mesosoma.

See the description in the nomenclature section.

Distribution
New Jersey to northern Florida and west through Nebraska, Colorado, and Texas to Utah and Nevada. (Wilson 2003)

Distribution based on Regional Taxon Lists
Nearctic Region: United States.

Habitat
This species has a preference for open sites with sandy soils and habitats that are at the moister end of dry (e.g., grasslands and valley bottoms in arid areas), at least in the west. It also inhabitats sites that have some but not all of these characteristics. Gregg (1963) stated Pheidole bicarinata in Colorado was a Sonoran Zone species. In the eastern US (Carter, 1962; Wilson 2003) this ant is common in fields and open grassy locations, including in ruderal sites (sidewalks, lawns, and along the shoulder of roads).

Nests are found underneath ground-covering objects, within rotten downed wood, or in open ground with a small crater mound. Mounds are typically incomplete, resembling a wedge or crescent.

Colony Attributes
Colonies are small, often just a few hundred workers.

Foraging/Diet
Seeds are collected and may be stored in underground granaries that are part of the nest. Workers also visit extraflora nectories.

New Mexico
Mackay and Mackay (2002) - Occurs in grasslands (including arid sites with yuccas), scrub oak, pine and cedar, pinyon-juniper forests, oak forests, ponderosa pine forests. Pheidole bicarinata nests in rotten logs, as well as in the soil and under objects (especially stones) in grassy areas with fine sand. Brood was found in nests in March and August. They are attracted to baits on the soil surface (Vienna sausage). This species is the host of Solenopsis molesta.

Association with Other Organisms


The general biology of Eucharitid wasps are summarized nicely here (Lachaud, J., Cerdan, P. and Pérez-Lachaud, G. 2012 - see the original paper for the citations indicated by the numerals doi:10.1155/2012/393486):

"Within Hymenoptera, the family Eucharitidae (subdivided in three subfamilies: Oraseminae, Eucharitinae, and the Indo-Pacific Gollumiellinae) is the most numerous and diverse group of ant parasitoids [1, 2]. All of the members of this family have a highly modified life cycle [3–6]. Unlike most parasitic wasp species, eucharitid females deposit their eggs away from hosts, in or on plant tissue [2, 7]. The active first instar larva, termed planidium, is responsible for gaining access to the host ant larvae by using various phoretic behaviors including either attachment to an intermediate host (most often a potential ant prey) or to foraging ant workers, with on occasion the presence of attractive substances in or on the eggs [2, 8]. Within the nest, planidia attach themselves to ant larvae, but development is only completed when the host pupates [6, 7, 9, 10]. In almost all of the cases, adults emerge among ant brood (but see [11]) and have to leave the host nest to reproduce. Ants show only moderate aggression to newly emerged eucharitids [7, 12–17] and transport them outside as if they were refuse [11, 15, 17], ultimately enhancing wasp dispersal. Parasitism is very variable and localized in time and space [13, 18, 19]. A very high local prevalence may lead to only a very low impact at the regional scale, suggesting that these parasitoids do not have a major influence on the dynamics of their host population [19]. However, they constitute a remarkable example of both host-parasitoid coevolution and host behavior manipulation."

Other Insects
The two wasps shown below were found by Stefan Cover with a colony of P. bicarinata  in Arizona (Specimen Labels).

Worker
Minor

Nomenclature

 *  bicarinata. Pheidole bicarinata Mayr, 1870b: 989 (s.) U.S.A. Senior synonym of hayesi: Creighton, 1950a: 171; of longula, vinelandica (and its junior synonyms buccalis, laeviuscula, nebrascensis): Wilson, 2003: 561.
 * vinelandica. Pheidole bicarinata r. vinelandica Forel, 1886b: xlv (s.w.q.m.) U.S.A. Combination in P. (Allopheidole): Forel, 1912f: 237. Raised to species: Mayr, 1886d: 458; Dalla Torre, 1893: 97; Naves, 1985: 66. Subspecies of bicarinata: Creighton, 1950a: 172; Gregg, 1959: 18. Senior synonym of laeviuscula: Creighton, 1950a: 172; of buccalis, nebrascensis: Gregg, 1959: 18. Junior synonym of bicarinata: Wilson, 2003: 561.
 * laeviuscula. Pheidole vinelandica subsp. laeviuscula Emery, 1895c: 292 (s.w.) U.S.A. Junior synonym of vinelandica: Creighton, 1950a: 172.
 * longula. Pheidole vinelandica var. longula Emery, 1895c: 292 (s.w.) U.S.A. Subspecies of bicarinata: Creighton, 1950a: 171; Smith, D.R. 1979: 1367. Raised to species: Cole, 1956c: 114. Junior synonym of bicarinata: Wilson, 2003: 561.
 * buccalis. Pheidole vinelandica subsp. buccalis Wheeler, W.M. 1908e: 454 (s.w.q.) U.S.A. Subspecies of bicarinata: Creighton, 1950a: 171. Material of the unavailable name castanea referred here by Creighton, 1950a: 171; of the unavailable name huachucana referred here by Smith, M.R. 1958c: 120. Junior synonym of vinelandica: Gregg, 1959: 18; of bicarinata: Cole, 1956c: 114; Wilson, 2003: 561.
 * hayesi. Pheidole hayesi Smith, M.R. 1924: 251 (s.w.) U.S.A. Junior synonym of bicarinata: Creighton, 1950a: 171.

Description
From Wilson (2003): DIAGNOSIS A member of the “bicarinata complex” belonging to the larger pilifera group and comprising Pheidole agricola, Pheidole aurea, Pheidole bajaensis, Pheidole barbata, Pheidole bicarinata, Pheidole centeotl, Pheidole cerebrosior, Pheidole ceres, Pheidole defecta, Pheidole macclendoni, Pheidole marcidula, Pheidole paiute, Pheidole pinealis, Pheidole psammophila, Pheidole xerophila, Pheidole yaqui and Pheidole yucatana, which complex is characterized by large to very large, forward-set eyes, especially in the minor; and in the major, the occipital lobes lacking any sculpturing (exception: aurea), the posterior half of the head capsule smooth and shiny, and the postpetiole from above oval, elliptical, or laterally angulate (conulate in cerebrosior). P. bicarinata is distinguished within the subgroup by the following combination of traits.

Major: humerus in dorsal-oblique view dentate to subangulate; propodeal spine moderately long and stout; petiolar node in side view tapered; lateral margins of postpetiolar node seen from about acute-angular; anterior fringe of pronotal dorsum transversely carinulate, remainder mostly smooth and shiny; lateral margins of pronotal dorsum carinulate or not; mesonotal dorsum foveolate and opaque to mostly smooth and shiny; dorsum of propodeum foveolate and opaque and carinulate or not.

Minor: humerus in dorsal-oblique view obtusely subangulate; petiole variable in size and shape, from small and equilaterally triangular to moderately elongate. P. bicarinata is a taxonomically difficult species (or, possibly, complex of closely similar species). It is highly variable, especially geographically, in propodeal and pronotal sculpturing, and as suggested above, in development of the propodeal spine, and in color. The variation among different characters is discordant to some degree, and intermediates within the respective characters occur. I believe it prudent for the present to leave the described infraspecific forms as a single species (I am indebted to Philip S. Ward for valuable information and advice on the variation and status of bicarinata).

MEASUREMENTS (mm) Major (Brown Co. State Park, Indiana): HW 1.04, HL 1.14, SL 0.54, EL 0.14, PW 0.52. Minor (Brown Co. State Park): HW 0.54, HL 0.58, SL 0.50, EL 0.12, PW 0.34.

COLOR Major: body varying geographically, from clear yellow in the western parts of the range to brownish yellow to dark brown in the east.

Minor: clear yellow to medium brown.



'''Figure. Upper: “typical” bicarinata major. Lower: “typical” bicarinata minor. INDIANA: Brown County State Park, col. E. O. Wilson. (Lower propodeum is of “vinelandica” variant major from Calvert Cliffs State Park, Calvert Co., Maryland; see description of other variation in Diagnosis and Color.) Scale bars = 1 mm.'''

Type Material
USA. Illinois, - as reported in Wilson (2003)

Etymology
L bicarinata, double-ridged, two-carinal, possibly referring to the carinae of the clypeus. (Wilson 2003)

References based on Global Ant Biodiversity Informatics

 * 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.
 * Beck D. E., D. M. Allred, W. J. Despain. 1967. Predaceous-scavenger ants in Utah. Great Basin Naturalist 27: 67-78
 * Beckmann R. L., and J. M. Stucky. 1981. Extrafloral Nectaries and Plant Guarding in Ipomoea pandurata (L.) G. F. W. Mey. (Convolvulaceae). American Journal of Botany 68(1): 72-79.
 * 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.
 * Buczkowski G., and D. S. Richmond. 2012. The Effect of Urbanization on Ant Abundance and Diversity: A Temporal Examination of Factors Affecting Biodiversity. PLoS ONE 7(8): e41729. doi:10.1371/journal.pone.0041729
 * 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.
 * Carroll T. M. 2011. The ants of Indiana (Hymenoptera: Formicidae). Master's Thesis Purdue university, 385 pages.
 * Cole A. C. 1940. A Guide to the Ants of the Great Smoky Mountains National Park, Tennessee. American Midland Naturalist 24(1): 1-88.
 * Cole A. C., Jr. 1956. Observations of some members of the genus Pheidole in the southwestern United States with synonymy (Hymenoptera: Formicidae). Journal of the Tennessee Academy of Science 31: 112-118.
 * 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
 * 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
 * Davis W. T., and J. Bequaert. 1922. An annoted list of the ants of Staten Island and Long Island, N. Y. Bulletin of the Brooklyn Entomological Society 17(1): 1-25.
 * Del Toro, I. 2010. PERSONAL COMMUNICATION. MUSEUM RECORDS COLLATED BY ISRAEL DEL TORO
 * Dennis C. A. 1938. The distribution of ant species in Tennessee with reference to ecological factors. Annals of the Entomological Society of America 31: 267-308.
 * Deyrup M. 2016. Ants of Florida: identification and natural history. CRC Press, 423 pages.
 * Deyrup, M. 2003. An updated list of Florida ants (Hymenoptera: Formicidae). Florida Entomologist 86(1):43-48.
 * DuBois M. B. 1985. Distribution of ants in Kansas: subfamilies Ponerinae, Ecitoninae, and Myrmicinae (Hymenoptera: Formicidae). Sociobiology 11: 153-1000
 * DuBois M. B. 1985. Distribution of ants in Kansas: subfamilies Ponerinae, Ecitoninae, and Myrmicinae (Hymenoptera: Formicidae). Sociobiology 11: 153-1001
 * DuBois M. B. 1985. Distribution of ants in Kansas: subfamilies Ponerinae, Ecitoninae, and Myrmicinae (Hymenoptera: Formicidae). Sociobiology 11: 153-1002
 * DuBois M. B. 1985. Distribution of ants in Kansas: subfamilies Ponerinae, Ecitoninae, and Myrmicinae (Hymenoptera: Formicidae). Sociobiology 11: 153-1003
 * DuBois M. B. 1985. Distribution of ants in Kansas: subfamilies Ponerinae, Ecitoninae, and Myrmicinae (Hymenoptera: Formicidae). Sociobiology 11: 153-1004
 * DuBois M. B. 1985. Distribution of ants in Kansas: subfamilies Ponerinae, Ecitoninae, and Myrmicinae (Hymenoptera: Formicidae). Sociobiology 11: 153-1005
 * DuBois M. B. 1985. Distribution of ants in Kansas: subfamilies Ponerinae, Ecitoninae, and Myrmicinae (Hymenoptera: Formicidae). Sociobiology 11: 153-1006
 * DuBois M. B. 1985. Distribution of ants in Kansas: subfamilies Ponerinae, Ecitoninae, and Myrmicinae (Hymenoptera: Formicidae). Sociobiology 11: 153-1007
 * DuBois M. B. 1985. Distribution of ants in Kansas: subfamilies Ponerinae, Ecitoninae, and Myrmicinae (Hymenoptera: Formicidae). Sociobiology 11: 153-1008
 * DuBois M. B. 1985. Distribution of ants in Kansas: subfamilies Ponerinae, Ecitoninae, and Myrmicinae (Hymenoptera: Formicidae). Sociobiology 11: 153-1009
 * DuBois M. B. 1985. Distribution of ants in Kansas: subfamilies Ponerinae, Ecitoninae, and Myrmicinae (Hymenoptera: Formicidae). Sociobiology 11: 153-1010
 * DuBois M. B. 1985. Distribution of ants in Kansas: subfamilies Ponerinae, Ecitoninae, and Myrmicinae (Hymenoptera: Formicidae). Sociobiology 11: 153-1011
 * DuBois M. B. 1985. Distribution of ants in Kansas: subfamilies Ponerinae, Ecitoninae, and Myrmicinae (Hymenoptera: Formicidae). Sociobiology 11: 153-1012
 * DuBois M. B. 1985. Distribution of ants in Kansas: subfamilies Ponerinae, Ecitoninae, and Myrmicinae (Hymenoptera: Formicidae). Sociobiology 11: 153-1013
 * DuBois M. B. 1985. Distribution of ants in Kansas: subfamilies Ponerinae, Ecitoninae, and Myrmicinae (Hymenoptera: Formicidae). Sociobiology 11: 153-1014
 * DuBois M. B. 1985. Distribution of ants in Kansas: subfamilies Ponerinae, Ecitoninae, and Myrmicinae (Hymenoptera: Formicidae). Sociobiology 11: 153-1015
 * DuBois M. B. 1985. Distribution of ants in Kansas: subfamilies Ponerinae, Ecitoninae, and Myrmicinae (Hymenoptera: Formicidae). Sociobiology 11: 153-1016
 * DuBois M. B. 1985. Distribution of ants in Kansas: subfamilies Ponerinae, Ecitoninae, and Myrmicinae (Hymenoptera: Formicidae). Sociobiology 11: 153-1017
 * DuBois M. B. 1985. Distribution of ants in Kansas: subfamilies Ponerinae, Ecitoninae, and Myrmicinae (Hymenoptera: Formicidae). Sociobiology 11: 153-959
 * DuBois M. B. 1985. Distribution of ants in Kansas: subfamilies Ponerinae, Ecitoninae, and Myrmicinae (Hymenoptera: Formicidae). Sociobiology 11: 153-960
 * DuBois M. B. 1985. Distribution of ants in Kansas: subfamilies Ponerinae, Ecitoninae, and Myrmicinae (Hymenoptera: Formicidae). Sociobiology 11: 153-961
 * DuBois M. B. 1985. Distribution of ants in Kansas: subfamilies Ponerinae, Ecitoninae, and Myrmicinae (Hymenoptera: Formicidae). Sociobiology 11: 153-962
 * DuBois M. B. 1985. Distribution of ants in Kansas: subfamilies Ponerinae, Ecitoninae, and Myrmicinae (Hymenoptera: Formicidae). Sociobiology 11: 153-963
 * DuBois M. B. 1985. Distribution of ants in Kansas: subfamilies Ponerinae, Ecitoninae, and Myrmicinae (Hymenoptera: Formicidae). Sociobiology 11: 153-964
 * DuBois M. B. 1985. Distribution of ants in Kansas: subfamilies Ponerinae, Ecitoninae, and Myrmicinae (Hymenoptera: Formicidae). Sociobiology 11: 153-965
 * DuBois M. B. 1985. Distribution of ants in Kansas: subfamilies Ponerinae, Ecitoninae, and Myrmicinae (Hymenoptera: Formicidae). Sociobiology 11: 153-966
 * DuBois M. B. 1985. Distribution of ants in Kansas: subfamilies Ponerinae, Ecitoninae, and Myrmicinae (Hymenoptera: Formicidae). Sociobiology 11: 153-967
 * DuBois M. B. 1985. Distribution of ants in Kansas: subfamilies Ponerinae, Ecitoninae, and Myrmicinae (Hymenoptera: Formicidae). Sociobiology 11: 153-968
 * DuBois M. B. 1985. Distribution of ants in Kansas: subfamilies Ponerinae, Ecitoninae, and Myrmicinae (Hymenoptera: Formicidae). Sociobiology 11: 153-969
 * DuBois M. B. 1985. Distribution of ants in Kansas: subfamilies Ponerinae, Ecitoninae, and Myrmicinae (Hymenoptera: Formicidae). Sociobiology 11: 153-970
 * DuBois M. B. 1985. Distribution of ants in Kansas: subfamilies Ponerinae, Ecitoninae, and Myrmicinae (Hymenoptera: Formicidae). Sociobiology 11: 153-971
 * DuBois M. B. 1985. Distribution of ants in Kansas: subfamilies Ponerinae, Ecitoninae, and Myrmicinae (Hymenoptera: Formicidae). Sociobiology 11: 153-972
 * DuBois M. B. 1985. Distribution of ants in Kansas: subfamilies Ponerinae, Ecitoninae, and Myrmicinae (Hymenoptera: Formicidae). Sociobiology 11: 153-973
 * DuBois M. B. 1985. Distribution of ants in Kansas: subfamilies Ponerinae, Ecitoninae, and Myrmicinae (Hymenoptera: Formicidae). Sociobiology 11: 153-974
 * DuBois M. B. 1985. Distribution of ants in Kansas: subfamilies Ponerinae, Ecitoninae, and Myrmicinae (Hymenoptera: Formicidae). Sociobiology 11: 153-975
 * DuBois M. B. 1985. Distribution of ants in Kansas: subfamilies Ponerinae, Ecitoninae, and Myrmicinae (Hymenoptera: Formicidae). Sociobiology 11: 153-976
 * DuBois M. B. 1985. Distribution of ants in Kansas: subfamilies Ponerinae, Ecitoninae, and Myrmicinae (Hymenoptera: Formicidae). Sociobiology 11: 153-977
 * DuBois M. B. 1985. Distribution of ants in Kansas: subfamilies Ponerinae, Ecitoninae, and Myrmicinae (Hymenoptera: Formicidae). Sociobiology 11: 153-978
 * DuBois M. B. 1985. Distribution of ants in Kansas: subfamilies Ponerinae, Ecitoninae, and Myrmicinae (Hymenoptera: Formicidae). Sociobiology 11: 153-979
 * DuBois M. B. 1985. Distribution of ants in Kansas: subfamilies Ponerinae, Ecitoninae, and Myrmicinae (Hymenoptera: Formicidae). Sociobiology 11: 153-980
 * DuBois M. B. 1985. Distribution of ants in Kansas: subfamilies Ponerinae, Ecitoninae, and Myrmicinae (Hymenoptera: Formicidae). Sociobiology 11: 153-981
 * DuBois M. B. 1985. Distribution of ants in Kansas: subfamilies Ponerinae, Ecitoninae, and Myrmicinae (Hymenoptera: Formicidae). Sociobiology 11: 153-982
 * DuBois M. B. 1985. Distribution of ants in Kansas: subfamilies Ponerinae, Ecitoninae, and Myrmicinae (Hymenoptera: Formicidae). Sociobiology 11: 153-983
 * DuBois M. B. 1985. Distribution of ants in Kansas: subfamilies Ponerinae, Ecitoninae, and Myrmicinae (Hymenoptera: Formicidae). Sociobiology 11: 153-984
 * DuBois M. B. 1985. Distribution of ants in Kansas: subfamilies Ponerinae, Ecitoninae, and Myrmicinae (Hymenoptera: Formicidae). Sociobiology 11: 153-985
 * DuBois M. B. 1985. Distribution of ants in Kansas: subfamilies Ponerinae, Ecitoninae, and Myrmicinae (Hymenoptera: Formicidae). Sociobiology 11: 153-986
 * DuBois M. B. 1985. Distribution of ants in Kansas: subfamilies Ponerinae, Ecitoninae, and Myrmicinae (Hymenoptera: Formicidae). Sociobiology 11: 153-987
 * DuBois M. B. 1985. Distribution of ants in Kansas: subfamilies Ponerinae, Ecitoninae, and Myrmicinae (Hymenoptera: Formicidae). Sociobiology 11: 153-988
 * DuBois M. B. 1985. Distribution of ants in Kansas: subfamilies Ponerinae, Ecitoninae, and Myrmicinae (Hymenoptera: Formicidae). Sociobiology 11: 153-989
 * DuBois M. B. 1985. Distribution of ants in Kansas: subfamilies Ponerinae, Ecitoninae, and Myrmicinae (Hymenoptera: Formicidae). Sociobiology 11: 153-990
 * DuBois M. B. 1985. Distribution of ants in Kansas: subfamilies Ponerinae, Ecitoninae, and Myrmicinae (Hymenoptera: Formicidae). Sociobiology 11: 153-991
 * DuBois M. B. 1985. Distribution of ants in Kansas: subfamilies Ponerinae, Ecitoninae, and Myrmicinae (Hymenoptera: Formicidae). Sociobiology 11: 153-992
 * DuBois M. B. 1985. Distribution of ants in Kansas: subfamilies Ponerinae, Ecitoninae, and Myrmicinae (Hymenoptera: Formicidae). Sociobiology 11: 153-993
 * DuBois M. B. 1985. Distribution of ants in Kansas: subfamilies Ponerinae, Ecitoninae, and Myrmicinae (Hymenoptera: Formicidae). Sociobiology 11: 153-994
 * DuBois M. B. 1985. Distribution of ants in Kansas: subfamilies Ponerinae, Ecitoninae, and Myrmicinae (Hymenoptera: Formicidae). Sociobiology 11: 153-995
 * DuBois M. B. 1985. Distribution of ants in Kansas: subfamilies Ponerinae, Ecitoninae, and Myrmicinae (Hymenoptera: Formicidae). Sociobiology 11: 153-996
 * DuBois M. B. 1985. Distribution of ants in Kansas: subfamilies Ponerinae, Ecitoninae, and Myrmicinae (Hymenoptera: Formicidae). Sociobiology 11: 153-997
 * DuBois M. B. 1985. Distribution of ants in Kansas: subfamilies Ponerinae, Ecitoninae, and Myrmicinae (Hymenoptera: Formicidae). Sociobiology 11: 153-998
 * DuBois M. B. 1985. Distribution of ants in Kansas: subfamilies Ponerinae, Ecitoninae, and Myrmicinae (Hymenoptera: Formicidae). Sociobiology 11: 153-999
 * 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
 * Eastlake Chew A. and Chew R. M. 1980. Body size as a determinant of small-scale distributions of ants in evergreen woodland southeastern Arizona. Insectes Sociaux 27: 189-202
 * Emery C. 1895. Beiträge zur Kenntniss der nordamerikanischen Ameisenfauna. (Schluss). Zoologische Jahrbücher. Abteilung für Systematik, Geographie und Biologie der Tiere 8: 257-360.
 * Forel A. 1901. Variétés myrmécologiques. Annales de la Société Entomologique de Belgique 45: 334-382.
 * Forster J.A. 2005. The Ants (hymenoptera: Formicidae) of Alabama. Master of Science, Auburn University. 242 pages.
 * 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.
 * 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
 * Graham J.H., H.H. Hughie, S. Jones, K. Wrinn, A.J. Krzysik, J.J. Duda, D.C. Freeman, J.M. Emlen, J.C. Zak, D.A. Kovacic, C. Chamberlin-Graham, H. Balbach. 2004. Habitat disturbance and the diversity and abundance of ants (Formicidae) in the Southeastern Fall-Line Sandhills. 15pp. Journal of Insect Science. 4: 30
 * 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 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.
 * 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
 * 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.
 * 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 R. Personnal Database. Accessed on February 5th 2014 at http://www.asu.edu/clas/sirgtools/resources.htm
 * Kansas State Entomology Collection. Dowloaded the 30th of May 2011 at http://biodis.k-state.edu/collections/entomology/
 * La Rivers I. 1968. A first listing of the ants of Nevada. Biological Society of Nevada, Occasional Papers 17: 1-12.
 * LaBrun, E.G. and D.H. Feener Jr. 2007. When trade-offs interact: balance of terror enforces dominance discovery trade-off in a local ant assemblage. Journal of Animal Ecology 76:58-64
 * LeBrun, E.G. 2005. Who Is the Top Dog in Ant Communities? Resources, Parasitoids, and Multiple Competitive Hierarchies. Oecologia 142(4):643-652
 * Lessard J. P., R. R. Dunn, C. R. Parker, and N. J. Sanders. 2007. Rarity and Diversity in Forest Ant Assemblages of Great Smoky Mountains National Park. Southeastern Naturalist 1: 215-228.
 * 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. 2015. Report on the ants collected on Spring Island, Beaufort County, South Carolina. A report submitted to Spring Island Nature Preserve, May 2015. Mississippi Entomological Museum Report #2015-01. 8 pp
 * 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 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 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., 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
 * 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.P. and E. Mackay. XXXX. The Ants of New Mexico
 * McDonald D. L., D. R. Hoffpauir, and J. L. Cook. 2016. Survey yields seven new Texas county records and documents further spread of Red Imported Fire Ant, Solenopsis invicta Buren. Southwestern Entomologist, 41(4): 913-920.
 * Miguelena J. G., and P. B. Baker. 2019. Effects of urbanization on the diversity, abundance, and composition of ant assemblages in an arid city. Environmental Entomology doi: 10.1093/ee/nvz069.
 * Moody J. V., and O. F. Francke. 1982. The Ants (Hymenoptera, Formicidae) of Western Texas Part 1: Subfamily Myrmicinae. Graduate Studies Texas Tech University 27: 80 pp.
 * Morrison, L.W. 2002. Long-Term Impacts of an Arthropod-Community Invasion by the Imported Fire Ant, Solenopsis invicta. Ecology 83(8):2337-2345
 * 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.
 * 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.
 * Roeder K. A., and D. V. Roeder. 2017. The Pheidole (Hymenoptera: Formicidae) of Oklahoma: new species records and distributional notes. Check List 13(2): 2071.
 * Smith M. R. 1924. A new species of ant from Kansas (Hym.: Formicidae). Entomological News 35: 250-253.
 * Smith M. R. 1934. Dates on which the immature or mature sexual phases of ants have been observed (Hymen.: Formicoidea). Entomological News 45: 247-251.
 * Smith M. R. 1936. A list of the ants of Texas. Journal of the New York Entomological Society 44: 155-170.
 * 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
 * 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 J. Wheeler J. 1989. A checklist of the ants of Oklahoma. Prairie Naturalist 21: 203-210.
 * 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. 1905. An annotated list of the ants of New Jersey. Bulletin of the American Museum of Natural History. 21: 371-403.
 * 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, 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 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
 * 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.
 * Wilkinson, E.B. and D.H. Feener Jr. 2007. Habitat Complexity Modifies Ant-Parasitoid Interactions: Implications for Community Dynamics and the Role of Disturbance. Oecologia 152(1):151-161
 * Wilson, E.O. 2003. Pheidole in the New World: A Dominant, Hyperdiverse Genus. Harvard University Press
 * 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