Crematogaster laeviuscula

This species nests in wood, or under bark of dead logs (cottonwood), or in oak galls of the wasp Holcaspis cinerosus. Sexuals occur in nests in August. These ants are more aggressive than is typical for North American Crematogaster. (Mackay and Mackay 2002)

Identification
The workers of this species are predominantly shiny, specifically the head and side of the pronotum are shiny and glossy, the top of the pronotum is shiny, but finely sculptured. The mesopleuron and the sides of the propodeum are punctate or finely striate. The propodeal spines are well developed and long. It is generally light colored, with a pale reddish-yellow head and mesosoma and a darker gaster. (Mackay and Mackay 2002)

Morgan & Mackay (2017) - This common species is one of the most attractive, generally being bright red with a shiny black gaster. Key characteristics of C. laeviuscula are the entirely smooth and glossy head; the pronotum is lightly sculptured, and at least partially smooth and shining. Most the specimens are bicolored, mostly red with a black gaster.

There is considerable variation in C. laeviuscula in terms of color and sculpturing. Color ranges from the common bicolored individuals, completely reddish yellow, to completely dark brown. The head is mostly smooth and glossy, but the pronotum may be completely smooth and glossy (polished) to lightly sculptured (striated) and weakly shining.

Crematogaster atkinsoni
Morgan & Mackay (2017) - MacGown (pers. comm.) does not agree with our synonymizing C. atkinsoni, which makes unique and distinctive carton nests in gulf coastal grasses and is not the same thing as C. laeviuscula, that nests in stumps, cavities, etc., west of the Mississippi River. He has had considerable experience with southeastern ants, and we could be wrong.

Johnson 1988 - Crematogaster atkinsoni and Crematogaster pilosa may co-occur together in the same habitat and require care in identification.

The species typically occurs only about salt and freshwater marshes along the southern Coastal Plain and a distinct color variant having a rosy - red color over the whole body occurs in scattered localities along the salt marshes bordering the North Florida Gulf Coast. These ants are very distinct in color and the propodeal spines are slightly thicker in some specimens but no other morphological characters separate the two forms. The red variant constructs carton nests in and about the marsh, often above standing water, high enough to escape the tides. Other nests may occur within grass stems on dry land bordering the marsh, but even in these nests a carton component occurs at the base of the grass stems. Within the same marshes, the dark-colored, typical C. atkinsoni morph occurs nesting in a variety of circumstances but using a wide array of nest sites often with no carton involved.

Distribution
Central United States and south into Mexico.

Morgan & Mackay (2017) - This species is found throughout the United States, and has been collected as far south as Tabasco, México. Additional citations include Virginia (Guénard et al., 2012; Kjar, 2009), southeastern South Carolina (Davies, 2009, as C. atkinsoni), and Baja California (Johnson and Ward, 2002, as C. opaca cedrosensis, and C. hespera; Vázquez-Bolaños, 2011).

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

Habitat
Riparian Forest.

Biology
Morgan & Mackay (2017) - The habits of Crematogaster laeviuscula are diverse, mostly arboreal and relatively well known. Mackay and Mackay (unpublished) found they occasionally nest under stones, but generally live in living trees, in hollow twigs, under the bark and in branches (full of beetle burrows or termite tunnels), in completely rotten stumps with loose bark and in/under logs. They have often been taken in oak galls (Morgan, unpublished). They consistently nest in plants in Mississippi, including in dead dogwood, black willow, sumac, white ash, goldenrod, blackberry (Rubus laciniatus), common elder, mimosa, hickory, plume grass, rattan-vine, catalpa, bois d'arc, hawthorn, prickly ash, sweetgum, red oak, china-tree, pecan, black locust, and dead woody galls on red oak, often in preformed cavities (Tynes and Hutchins, 1964). They would often move the nest to logs and stumps when the colony outgrew its home (Tynes and Hutchins, 1964). They make carton nests in grass tussocks (as C. atkinsoni, Deyrup et al., 1988).

Mackay and Mackay (unpublished) collected brood in nests in March, May, June and August, sexuals in August. A founding queen with one worker was found in a hollow twig in March. Workers tend to be sluggish and escape when the nest is disturbed, but are more active under warmer temperatures, and large nests can be aggressive, and the workers attack and bite, relatively painfully for Crematogaster. Colonies have hundreds of workers and reproductives, as well as brood (Tynes and Hutchins, 1964). Brood overwinter as larvae (Tynes and Hutchins, 1964).

They forage in the vegetation as well as loose on the ground. Morgan (unpublished) noted that they utilize hollow vegetation to camouflage their trails. She found them at Squaw Spring at the UTEP Indio Mountains Research Station of Hudspeth Co., Texas, a riparian habitat in the Chihuahuan Desert, where they were traveling almost unseen on cattail stalks. Many eggs and pupae (worker and queen) were found in the hollow stalks that were exposed to the sun. Workers were seen eating a cicada thorax, others were feeding on a discarded piece of meat (Mackay and Mackay, unpublished). They were attracted to meat baits in open areas (Morrison, 2004). They tend Hemiptera and build earthen tents to protect the animals (Tynes and Hutchins, 1964).

They are generally found in mesic sites, including urban areas in arid regions, but are also found in open desert with creosote bush scrub, mesquite and acacia, mixed shrubland (mesquite, cholla, big lobed cactus), sagebrush, grassland, juniper forest, pinyon pine riparian, oak forests, oak woodland with spiny vegetation, riparian forest, riparian cottonwoods, deciduous forest, mixed pine/hardwood trees and pine/spruce/fir forest (Mackay and Mackay, unpublished). Whitford et al. (1999, as C. hespera) found them in riparian woodlands in New Mexico, and Feener (1981) reported them from semi-deciduous woodlands in eastern Texas. They are found in Florida scrub and pine flatlands in Florida upland ecosystems (King, 2007). They occur in fresh and salt water marshes, grasslands, sunflowers and mesquite, they normally nest arboreally, but can be found in logs (Jusino-Atresino and Phillips, 1992). They even occur in caves (Cokendolpher et al., 2009), and adapt to urban ecosystems (Guénard et al., 2014).

They are found in areas with soils ranging from light brown, brown to dark brown clay, light to dark brown clay/loam, dark brown loam, light brown fine sand/clay, orange sand, light brown to brown rocky-loam, and light brown rocky sand to rocky red soil.

They are codominant with the red imported fire ant Solenopsis invicta, as well as with Solenopsis geminata, and are parasitized by phorid flies, Pseudacteon sp. (Morrison et al., 2000). They are also parasitized by the strepsipteran Caenocholax fenyesis (Cook, 2009; Kathirithamby et al., 2010). Crematogaster laeviuscula are predators on egg cases and spiderlings of the spider Peucetia viridans (Oxyopidae) in South Carolina (as C. clara, Willey and Adler, 1989). They eliminate relatively few nests of the wasp Polistes exclamans (Strassmann, 1981). Hung and Vinson (1975) studied the male reproductive system.

Deyrup (2017) includes details on the biology and distribution of this species as well as comparisons with other species occurring in Florida, as C. atkinsoni.

Nomenclature

 * . Crematogaster laeviuscula Mayr, 1870b: 993 (w.) U.S.A. (Oklahoma).
 * Type-material: syntype workers (number not stated).
 * Type-locality: U.S.A.: Oklahoma, Fort Cobb, “on the border between Texas and the Indian Territories” (Norton).
 * Type-depository: NHMW.
 * [Misspelled as leviuscula by Dalla Torre, 1893: 83, and others.]
 * Wheeler, W.M. 1908e: 480 (q.); Wheeler, G.C. & Wheeler, J. 1952b: 255 (l.); Petralia & Vinson, 1980: 383 (l.).
 * Combination in C. (Acrocoelia): Emery, 1922e: 141;
 * combination in C. (Crematogaster): Buren, 1968b: 92.
 * Subspecies of lineolata: Emery, 1895c: 284; Wheeler, W.M. 1904e: 301; Wheeler, W.M. 1908e: 480; Wheeler, W.M. 1910g: 564; Santschi, 1911d: 5; Emery, 1922e: 141; Essig, 1926: 859; Wheeler, W.M. 1932a: 9.
 * Status as species: Mayr, 1886d: 463; Dalla Torre, 1893: 83; Wheeler, W.M. 1919g: 111; Smith, M.R. 1930a: 4; Dennis, 1938: 283; Enzmann, J. 1946c: 93; Creighton, 1950a: 210; Smith, M.R. 1951a: 809; Buren, in Smith, M.R. 1958c: 126; Buren, 1968b: 92 (in key); Smith, D.R. 1979: 1379; Johnson, C. 1988: 322 (redescription); Bolton, 1995b: 156; Mackay & Mackay, 2002: 93; Morgan & Mackay, 2017: 196 (redescription).
 * Senior synonym of arborea: Smith, M.R. 1951a: 809; Buren, in Smith, M.R. 1958c: 126; Smith, D.R. 1979: 1379; Bolton, 1995b: 156;; Morgan & Mackay, 2017: 196.
 * Senior synonym of clara: Mayr, 1886d: 463; Creighton, 1950a: 210; Johnson, C. 1988: 322; Bolton, 1995b: 156; Morgan & Mackay, 2017: 196.
 * Distribution: Mexico, U.S.A.
 * arborea. Oecodoma (Atta) arborea Buckley, 1867: 349 (w.q.) U.S.A. (Texas).
 * Type-material: syntype workers, syntype queens (number not stated).
 * Type-locality: U.S.A.: Texas (S.B. Buckley).
 * Type-depository: unknown (no material known to exist).
 * [Unresolved junior secondary homonym of Crematogaster arboreus Smith, F. 1858b: 138 (Bolton, 1995b: 147).]
 * Junior synonym of lineolata: McCook, 1880: 187; Mayr, 1886d: 462; Cresson, 1887: 259; Dalla Torre, 1893: 83; Pergande, 1893: 36.
 * Junior synonym of clara: Wheeler, W.M. 1902f: 30.
 * Junior synonym of laeviuscula: Smith, M.R. 1951a: 809; Buren, in Smith, M.R. 1958c: 126; Smith, D.R. 1979: 1379; Bolton, 1995b: 147; Morgan & Mackay, 2017: 196.
 * bicolor. Oecodoma (Atta) bicolor Buckley, 1867: 350 (w.) U.S.A. (Texas).
 * Type-material: syntype workers (number not stated0.
 * Type-locality: U.S.A.: Texas (S.B. Buckley).
 * Type-depository: unknown (no material known to exist).
 * [Junior secondary homonym of Crematogaster bicolor Smith, 1860b: 109.]
 * Junior synonym of laeviuscula: Cresson, 1887: 259.
 * Unidentifiable taxon: Wheeler, W.M. 1902f: 31.
 * Replacement name: Crematogaster clara Mayr, 1870b: 993.
 * [Note: clara junior synonym of bicolor Buckley (synonymy by McCook, 1880: 188); hence clara first available replacement name.]
 * biroi. Crematogaster biroi Emery, 1900c: 332, pl. 8, figs. 39-41 (w.) NEW GUINEA (Papua New Guinea).
 * Type-material: syntype workers (number not stated).
 * Type-locality: Papua New Guinea: (no further data) (L. Biró).
 * Type-depository: MSNG (perhaps also in HNHM).
 * [Junior primary homonym of Crematogaster biroi Mayr, 1897: 428.]
 * Replacement name: Crematogaster emeryi Forel, 1907a: 25.
 * clara. Crematogaster clara Mayr, 1870b: 993 (w.) U.S.A. (Oklahoma).
 * Type-material: syntype workers (number not stated).
 * Type-locality: U.S.A.: Oklahoma, Fort Cobb, “on the border between Texas and the Indian Territories” (Norton).
 * Type-depository: NHMW.
 * Replacement name for Oecodoma (Atta) bicolor Buckley, 1867: 350. [Junior secondary homonym of Crematogaster bicolor Smith, 1860b: 109.]
 * [Note: clara junior synonym of bicolor Buckley (synonymy by McCook, 1880: 188); hence clara first available replacement name.]
 * Wheeler, W.M. 1908e: 481 (q.m.).
 * Combination in C. (Acrocoelia): Emery, 1922e: 141;
 * combination in C. (Crematogaster): Buren, 1968b: 92.
 * As unavailable (infrasubspecific) name: Emery, 1895c: 285; Wheeler, W.M. 1908e: 481; Wheeler, W.M. 1910g: 564; Emery, 1922e: 141; Essig, 1926: 859; Cole, 1937a: 101.
 * Subspecies of laeviuscula: André, 1893b: 152; Emery, in Dalla Torre, 1893: 83; Wheeler, W.M. 1919g: 111; Dennis, 1938: 283; Enzmann, J. 1946c: 93; Smith, M.R. 1951a: 809.
 * Status as species: McCook, 1880: 188; Buren, in Smith, M.R. 1958c: 126; Carter, 1962a: 6 (in list); Buren, 1968b: 92 (in key); Smith, D.R. 1979: 1378.
 * Junior synonym of bicolor Buckley: McCook, 1880: 188; Emery, 1922e: 141.
 * Junior synonym of pilosa: Coovert, 2005: 62 (error).
 * Junior synonym of laeviuscula: Mayr, 1886d: 463; Creighton, 1950a: 210; Johnson, C. 1988: 322; Bolton, 1995b: 150; Morgan & Mackay, 2017: 196.

Crematogaster lineolata cedrosensis
Eight specimens from Cedros Island (VI.5.'95).

Crematogaster hespera
Phoenix, Arizona

Types: Holotype worker and eleven para type workers from Phoenix, Arizona, July 20, 1918- A. W. Morrill, with the note “attacking unripe peaches, very destructive.” I have another series from Phoenix of ten paratype workers and three females collected Dec. 25, 1895. The latter series was seen by Emery and marked by him as transitional from laeviuscula to cerasi. I have marked as paratypes also numerous specimens from these localities: Big Dalton Canyon, Glendora, Calif.- A. Mallis and J. Schwartz; Col. Desert, Calif. -A. L. Pickens; Glenville, Calif.- A. Wetmore; Pacific Grove, Calif.- W. M. Mann; Sacaton, Ariz.- F. S. Strickney; Nogales, Ruby, and Patagonia, Ariz.- L. F. Byars; Las Cruces, N. Mex.- A. C. Cole; Nogales and Patagonia, Ariz., and Ysleta and El Paso, Tex.- W. F. Buren; Sacramento, Calif.- P. H. Arnaud; Santa Ysabel and Mt. Laguna, San Diego Co., Calif.; Ft. Huachuca, Wickenberg, and Patagonia, Ariz.; and Imuris, Sonora, Mex.- Vv. S. Creighton; and Tempe, Ariz., W. M. Wheeler. A few other paratype specimens are known from "Los Angeles Riv., Calif.," Ventura and Pasadena, Calif., Tucson, Ariz., and Zion Canyon, Utah.

Holotype and many of the above paratype specimens are to be returned to the National Museum. Other para types will be returned to the California Academy of Natural Sciences and to the private collections of Dr. A. C. Cole and Dr. Wm. S. Creighton. I will retain a few paratypes from some of the series and deposit some of the paratypes in the Museum of Comparative Zoology.

Taxonomic Notes
Morgan & Mackay (2017) - Mayr described two species in 1870 from Ft. Cobb, Oklahoma: C. laeviuscula and C. clara. These two species differed by size and intensity of sculpturing on the face and mesosoma. Crematogaster laeviuscula is smaller with less intense sculpture and C. clara is larger with more intense sculpture. Dalla Torre (1893) and Wheeler (1919) made C. clara a variety of C. laeviuscula. Wheeler (1908b) found 2 nests with series of mixed sizes and sculpturing and contained ants that ranged from the largest C. clara to the smallest C. laeviuscula giving credence to synonymy. The smaller C. laeviuscula have been found in smaller nests, possibly because they are newer colonies that have been recently established. Wheeler concluded that C. laeviuscula were merely minims of C. clara, with which we agree, based on several series. Since C. laeviuscula had page preference, the species was assigned that name (Johnson, 1988).

Crematogaster lineolata var. cedrosensis has smaller spines, but the shape of the spines, pilosity and sculpture are consistent with C. laeviuscula and it is proposed as a synonym.

Crematogaster atkinsoni keys out to C. laeviuscula in both Creighton (1950) and Buren (1968) when using type material borrowed from the. We have also compared type material of C. atkinsoni and type material of C. laeviuscula. The types two species appear to be identical leading us to consider Crematogaster atkinsoni a junior synonym of C. laeviuscula.

We have compared paratypes of Crematogaster hespera Buren to syntypes of C. clara (now C. laeviuscula) and find no apparent differences. Buren’s description of C. hespera could also be applied to C. clara. In Buren’s key to eastern species, C. hespera keys out to C. laeviuscula. We did not find much variation in the Buren paratypes; however, we have seen considerable variation within nest series from CWEM leading us to consider Crematogaster hespera Buren a junior synonym of C. laeviuscula.

References based on Global Ant Biodiversity Informatics

 * Addison D. S., I. Bartoszek, V. Booher, M. A. Deyrup, M. Schuman, J. Schmid, and K. Worley. 2016. Baseline surveys for ants (Hymenoptera: Formicidae) of the western Everglades, Collier County, Florida. Florida Entomologist 99(3): 389-394.
 * 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.
 * 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.
 * 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. 1968. A Review of the species of Crematogaster, Sensu Stricto, in North America (Hymenoptera, Formicidae) Part 2. Descriptions of New Species. J. Georgia Entomol. Soc. 3: 91-121
 * 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.
 * 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
 * Cole A. C. 1940. A Guide to the Ants of the Great Smoky Mountains National Park, Tennessee. American Midland Naturalist 24(1): 1-88.
 * Dattilo W. et al. 2019. MEXICO ANTS: incidence and abundance along the Nearctic-Neotropical interface. Ecology https://doi.org/10.1002/ecy.2944
 * 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., C. Johnson, G. C. Wheeler, J. Wheeler. 1989. A preliminary list of the ants of Florida. Florida Entomologist 72: 91-101
 * 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. 1985. Distribution of ants in Kansas: subfamilies Ponerinae, Ecitoninae, and Myrmicinae (Hymenoptera: Formicidae). Sociobiology 11: 153-505
 * DuBois M. B. 1985. Distribution of ants in Kansas: subfamilies Ponerinae, Ecitoninae, and Myrmicinae (Hymenoptera: Formicidae). Sociobiology 11: 153-506
 * DuBois M. B. 1985. Distribution of ants in Kansas: subfamilies Ponerinae, Ecitoninae, and Myrmicinae (Hymenoptera: Formicidae). Sociobiology 11: 153-507
 * DuBois M. B. 1985. Distribution of ants in Kansas: subfamilies Ponerinae, Ecitoninae, and Myrmicinae (Hymenoptera: Formicidae). Sociobiology 11: 153-508
 * DuBois M. B. 1985. Distribution of ants in Kansas: subfamilies Ponerinae, Ecitoninae, and Myrmicinae (Hymenoptera: Formicidae). Sociobiology 11: 153-509
 * DuBois M. B. 1985. Distribution of ants in Kansas: subfamilies Ponerinae, Ecitoninae, and Myrmicinae (Hymenoptera: Formicidae). Sociobiology 11: 153-510
 * DuBois M. B. 1985. Distribution of ants in Kansas: subfamilies Ponerinae, Ecitoninae, and Myrmicinae (Hymenoptera: Formicidae). Sociobiology 11: 153-511
 * DuBois M. B. 1985. Distribution of ants in Kansas: subfamilies Ponerinae, Ecitoninae, and Myrmicinae (Hymenoptera: Formicidae). Sociobiology 11: 153-514
 * DuBois M. B. 1985. Distribution of ants in Kansas: subfamilies Ponerinae, Ecitoninae, and Myrmicinae (Hymenoptera: Formicidae). Sociobiology 11: 153-515
 * DuBois M. B. 1985. Distribution of ants in Kansas: subfamilies Ponerinae, Ecitoninae, and Myrmicinae (Hymenoptera: Formicidae). Sociobiology 11: 153-516
 * DuBois M. B. 1985. Distribution of ants in Kansas: subfamilies Ponerinae, Ecitoninae, and Myrmicinae (Hymenoptera: Formicidae). Sociobiology 11: 153-517
 * DuBois M. B. 1985. Distribution of ants in Kansas: subfamilies Ponerinae, Ecitoninae, and Myrmicinae (Hymenoptera: Formicidae). Sociobiology 11: 153-518
 * DuBois M. B. 1985. Distribution of ants in Kansas: subfamilies Ponerinae, Ecitoninae, and Myrmicinae (Hymenoptera: Formicidae). Sociobiology 11: 153-519
 * DuBois M. B. 1985. Distribution of ants in Kansas: subfamilies Ponerinae, Ecitoninae, and Myrmicinae (Hymenoptera: Formicidae). Sociobiology 11: 153-520
 * DuBois M. B. 1985. Distribution of ants in Kansas: subfamilies Ponerinae, Ecitoninae, and Myrmicinae (Hymenoptera: Formicidae). Sociobiology 11: 153-521
 * DuBois M. B. 1985. Distribution of ants in Kansas: subfamilies Ponerinae, Ecitoninae, and Myrmicinae (Hymenoptera: Formicidae). Sociobiology 11: 153-522
 * DuBois M. B. 1985. Distribution of ants in Kansas: subfamilies Ponerinae, Ecitoninae, and Myrmicinae (Hymenoptera: Formicidae). Sociobiology 11: 153-523
 * DuBois M. B. 1985. Distribution of ants in Kansas: subfamilies Ponerinae, Ecitoninae, and Myrmicinae (Hymenoptera: Formicidae). Sociobiology 11: 153-524
 * DuBois M. B. 1985. Distribution of ants in Kansas: subfamilies Ponerinae, Ecitoninae, and Myrmicinae (Hymenoptera: Formicidae). Sociobiology 11: 153-525
 * DuBois M. B. 1985. Distribution of ants in Kansas: subfamilies Ponerinae, Ecitoninae, and Myrmicinae (Hymenoptera: Formicidae). Sociobiology 11: 153-526
 * DuBois M. B. 1985. Distribution of ants in Kansas: subfamilies Ponerinae, Ecitoninae, and Myrmicinae (Hymenoptera: Formicidae). Sociobiology 11: 153-527
 * DuBois M. B. 1985. Distribution of ants in Kansas: subfamilies Ponerinae, Ecitoninae, and Myrmicinae (Hymenoptera: Formicidae). Sociobiology 11: 153-528
 * DuBois M. B. 1985. Distribution of ants in Kansas: subfamilies Ponerinae, Ecitoninae, and Myrmicinae (Hymenoptera: Formicidae). Sociobiology 11: 153-529
 * DuBois M. B. 1985. Distribution of ants in Kansas: subfamilies Ponerinae, Ecitoninae, and Myrmicinae (Hymenoptera: Formicidae). Sociobiology 11: 153-530
 * DuBois M. B. 1985. Distribution of ants in Kansas: subfamilies Ponerinae, Ecitoninae, and Myrmicinae (Hymenoptera: Formicidae). Sociobiology 11: 153-531
 * DuBois M. B. 1985. Distribution of ants in Kansas: subfamilies Ponerinae, Ecitoninae, and Myrmicinae (Hymenoptera: Formicidae). Sociobiology 11: 153-532
 * DuBois M. B. 1985. Distribution of ants in Kansas: subfamilies Ponerinae, Ecitoninae, and Myrmicinae (Hymenoptera: Formicidae). Sociobiology 11: 153-533
 * DuBois M. B. 1985. Distribution of ants in Kansas: subfamilies Ponerinae, Ecitoninae, and Myrmicinae (Hymenoptera: Formicidae). Sociobiology 11: 153-534
 * DuBois M. B. 1985. Distribution of ants in Kansas: subfamilies Ponerinae, Ecitoninae, and Myrmicinae (Hymenoptera: Formicidae). Sociobiology 11: 153-535
 * DuBois M. B. 1985. Distribution of ants in Kansas: subfamilies Ponerinae, Ecitoninae, and Myrmicinae (Hymenoptera: Formicidae). Sociobiology 11: 153-536
 * DuBois M. B. 1985. Distribution of ants in Kansas: subfamilies Ponerinae, Ecitoninae, and Myrmicinae (Hymenoptera: Formicidae). Sociobiology 11: 153-537
 * DuBois M. B. 1985. Distribution of ants in Kansas: subfamilies Ponerinae, Ecitoninae, and Myrmicinae (Hymenoptera: Formicidae). Sociobiology 11: 153-538
 * DuBois M. B. 1985. Distribution of ants in Kansas: subfamilies Ponerinae, Ecitoninae, and Myrmicinae (Hymenoptera: Formicidae). Sociobiology 11: 153-539
 * DuBois M. B. 1985. Distribution of ants in Kansas: subfamilies Ponerinae, Ecitoninae, and Myrmicinae (Hymenoptera: Formicidae). Sociobiology 11: 153-540
 * DuBois M. B. 1985. Distribution of ants in Kansas: subfamilies Ponerinae, Ecitoninae, and Myrmicinae (Hymenoptera: Formicidae). Sociobiology 11: 153-541
 * DuBois M. B. 1985. Distribution of ants in Kansas: subfamilies Ponerinae, Ecitoninae, and Myrmicinae (Hymenoptera: Formicidae). Sociobiology 11: 153-542
 * DuBois M. B. 1985. Distribution of ants in Kansas: subfamilies Ponerinae, Ecitoninae, and Myrmicinae (Hymenoptera: Formicidae). Sociobiology 11: 153-543
 * DuBois M. B. 1985. Distribution of ants in Kansas: subfamilies Ponerinae, Ecitoninae, and Myrmicinae (Hymenoptera: Formicidae). Sociobiology 11: 153-544
 * DuBois M. B. 1985. Distribution of ants in Kansas: subfamilies Ponerinae, Ecitoninae, and Myrmicinae (Hymenoptera: Formicidae). Sociobiology 11: 153-545
 * DuBois M. B. 1985. Distribution of ants in Kansas: subfamilies Ponerinae, Ecitoninae, and Myrmicinae (Hymenoptera: Formicidae). Sociobiology 11: 153-548
 * DuBois M. B. 1985. Distribution of ants in Kansas: subfamilies Ponerinae, Ecitoninae, and Myrmicinae (Hymenoptera: Formicidae). Sociobiology 11: 153-550
 * Enzmann J. 1946. Crematogaster lineolata cerasi, the cherry ant of Asa Fitsch; (with a survey of the American forms of Crematogaster, subgenus Acrocoelia). J. N. Y. Entomol. Soc. 54: 89-97.
 * Epperson, D.M. and C.R. Allen. 2010. Red Imported Fire Ant Impacts on Upland Arthropods in Southern Mississippi. American Midland Naturalist, 163(1):54-63.
 * Gans M. J., J. R. Arnold, A. Cohuo, L. Castro, D. Lam, and C. Wiley. 2016. Survey of ant species in Rockwall County, Texas. Southwestern Entomologist 41(2): 373-378.
 * 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
 * Glancey B. M., Wojcik D. P., Craig C. H. and Mitchell J. A. 1976. Ants of Mobile County, AL, as monitored by bait transects. Journal of the Georgia Entomological Society 11: 191-197
 * Glancey, B.M., Wojcik, D.P., Craig, C.H. and Mitchell, J.A. 1976. Ants of Mobile County, AL, as monitored by bait transects. Journal of the Georgia Entomological Society 11(3):191-197
 * 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.
 * Holway D.A. 1998. Effect of Argentine ant invasions on ground-dwelling arthropods in northern California riparian woodlands. Oecologia. 116: 252-258
 * 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
 * Johnson C. 1988. Species identification in the eastern Crematogaster (Hymenoptera: Formicidae). Journal of Entomological Science 23: 314-332.
 * Johnson R. Personnal Database. Accessed on February 5th 2014 at http://www.asu.edu/clas/sirgtools/resources.htm
 * Johnson, R.A. and P.S. Ward. 2002. Biogeography and endemism of ants (Hymenoptera: Formicidae) in Baja California, Mexico: a first overview. Journal of Biogeography 29:10091026/
 * Jusino-Atresino R., and S. A. Phillips, Jr. 1992. New ant records for Taylor Co., Texas. The Southern Naturalist 34(4): 430-433.
 * La Rivers I. 1968. A first listing of the ants of Nevada. Biological Society of Nevada, Occasional Papers 17: 1-12.
 * 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. 2010. Personal Communication. Longino Collection Database
 * 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
 * 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
 * Matsuda T., G. Turschak, C. Brehme, C. Rochester, M. Mitrovich, and R. Fisher. 2011. Effects of Large-Scale Wildfires on Ground Foraging Ants (Hymenoptera: Formicidae) in Southern California. Environmental Entomology 40(2): 204-216.
 * 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.
 * 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.
 * 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
 * Morgan C., and W. P. Mackay. 2017. The North America acrobat ants of the hyperdiverse genus Crematogaster. Mauritius: LAP LAMBERT Academic Publishing, 540 pp.
 * Morgan, C.E. 2009. Revision of the ant genus Crematogaster (Hymenoptera: Formicidae) in North America. Ph.D. Dissertation, University of Texas at El Paso, 268 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
 * 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.
 * Rajagopal T., S. P. Sevarkodiyone, and A. Manimozhi. 2005. Ant diversity in some selected localities of Sattur Taluk, Virudhunagar district, Tamil Nadu. Zoos' Print Journal 20(6): 1887-1888.
 * 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.
 * 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.
 * 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. 1962. A new species of exotic Ponera from North Carolina (Hymenoptera, Formicidae). Acta Hymenopterologica 1: 377-382.
 * Turner C. R., and J. L. Cook. 1998. The ants (Hymenoptera: Formicidae) of the Caddo Lake region of northeast Texas. Texas Journal of Science 50: 171-173.
 * Tynes J. S., and R. E. Hutchins. 1964. Studies of plant-nesting ants in east central Mississippi. American Midland Naturalist 72(1): 152-156.
 * 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. 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.
 * Vasquez-Bolanos M. 2011. Checklist of the ants (Hymenoptera: Formicidae) from Mexico. Dugesiana 18(1): 95-133.
 * 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. and J.A. Moore. 2005. Red Imported Fire Ants (Hymenoptera: Formicidae) at Gopher Tortoise (Testudines: Testudinidae) Burrows. The Florida Entomologist 88(4):349-354
 * 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. 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. 1919. A new paper-making Crematogaster from the southeastern United States. Psyche (Cambridge) 26: 107-112.
 * Wheeler W. M. 1932. A list of the ants of Florida with descriptions of new forms. J. N. Y. Entomol. Soc. 40: 1-17.
 * 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