Temnothorax curvispinosus

Temnothorax curvispinosus
Scientific classification
Kingdom:	Animalia
Phylum:	Arthropoda
Class:	Insecta
Order:	Hymenoptera
Family:	Formicidae
Subfamily:	Myrmicinae
Tribe:	Crematogastrini
Genus:	Temnothorax
Species group:	palearctic
Species:	T. curvispinosus
Binomial name
	Temnothorax curvispinosus; (Mayr, 1866) Specimen labels
Synonyms
	Stenamma gallarum Patton, 1879;

This is one of North America's better studied ant species. Many aspects of their basic biology has been examined and a variety of experiments have studied particulars about their nesting behavior, foraging and social dynamics.

At a Glance

• Polygynous • Cavity Nesting • Tandem running

Photo Gallery

Worker.
Queen from Groton, Massachusetts. Photo by Tom Murray.

Identification

Prebus (2017) - A member of the Palearctic clade.

Mackay (2000) - This is a small yellow ant with an 11-segmented antenna, which is very common in eastern USA. The very long, inwardly curved, closely spaced propodeal spines make this species immediately recognizable. The top and side of the mesosoma are covered with coarse rugae. The head is completely and coarsely punctate.

Keys including this Species

Distribution

USA. Throughout the eastern United States, from Maine to central Florida, and westward through eastern North Dakota down through the eastern half of Texas.

Latitudinal Distribution Pattern

Latitudinal Range: 44.34° to 30.05°.


North Temperate	North Subtropical	Tropical	South Subtropical	South Temperate

Source: AntMaps

Distribution based on Regional Taxon Lists

Nearctic Region: United States (type locality).

Distribution based on AntMaps

Distribution based on AntWeb specimens

Check data from AntWeb

Countries Occupied

Number of countries occupied by this species based on AntWiki Regional Taxon Lists. In general, fewer countries occupied indicates a narrower range, while more countries indicates a more widespread species.

Habitat

Forest.

Abundance

Common.

Biology

Worker from Groton, Massachusetts. Photo by Tom Murray

The following summary is from Mackay (2000): "The habits of this species are well known (Wheeler, 1916, 1917; Smith, 1924; Dennis, 1938; Cole, 1940; Wesson and Wesson, 1940; Headley. 1943; Buren, 1944; Gregg, 1944; Talbot. 1934, 1957, 1965; DuBoiS, 1985). Nests are found in plant cavities including hollow stems, under bark of living trees, in reeds, twigs, logs, acorns, nut shells, insect galls, puffballs, pine cones (Williams, 1989) and under rocks or in soil in forested areas (Wheeler, 1903a, 1905; Cole, 1940; DuBoiS. 1985). It is found at lower elevations in the southern Blue Ridge of Virginia (Van Pelt, 1963). Nest populations are about 80-100 workers with several queens (Wilson, 1974a), up to 113 workers in acorn nests (Talbot. 1957); average populations in Ohio were 235 (all castes), with the largest population of workers being 369 (total members 727), and nest densities of 0.6 nests per square meter (Headley, 1943). Alates are present from June-Aug. (Kannowski, 1959; Talbot, 1957; DuBoiS, 1985); flights occur in early July (Wesson and Wesson 1940). Workers reproduce in queenright colonies (Frumhoff and Ward. 1992). Larvae are found within nests throughout the year (Headley. 1943; Talbot. 1957). Nests are polygynous (Alloway et al.. 1982) and polydomous (Stuart. 1985, 1987a). The nest site may change after slight disturbances (Moeglich. 1978). The species is widely distributed in many different habitats, and is especially common in Tennessee (Dennis. 1938), Ohio (Wesson and Wesson. 1940), near Chicago (Gregg. 1944) and Mississippi (Smith. 1924). Herbers (1983) discussed the social organization and Wilson and Fagen (1974) estimated the total behavioral repertories. Stuart (1987a. 1987b) reported on transient nestmate recognition. Wilson (197 4b) studied the behavior of workers in laboratory colonies. It eats honeydew on leaves of trees and plants (Smith. 1924), but apparently does not tend aphids (Dennis,1938), and carries seeds (Heithaus. 1981) and presumably eats at least part of them. They also feed at the axillary nectaries of bracken fern (Douglas. 1983), and on dead insects (Fellers and Fellers. 1982). Foraging involves tandem running (Moeguch. 1979). Seasonally, foraging rate is highest in the spring and early summer, dropping off in the fall and being absent in the winter (Fellers. 1989). It is most active during the daylight hours (Fellers. 1989).

It is the host of Temnothorax duloticus (Talbot. 1957), Temnothorax americanus (Emery. 1895; Alloway et al.. 1982; Alloway and del Rio Pesado. 1979. 1983; pers. obs.), possibly T. minutissimus (Smith. 1942) and Limulodes parki (Coleoptera-Seevers and Dybas. 1943)."

Colony Attributes

Colonies typically contain less than one hundred workers although an occasional nest can be found that exceeds this number. Roughly half of the queens survive the winter, for workers roughly two thirds to slightly more than one half survive from fall to spring.

Nesting Habits

Nests are located in preformed cavities in structures found in the litter, e.g., in small sticks or nuts. Temnothorax curvispinosus is facultatively polydomous and their nesting arrangements vary with season. In the productive summer months, colonies can fragment and be arranged across numerous nest sites. These vary in queen number, from multiple queens to those that only have workers and brood.

During the winter nests coalesce and typically are found in a single structure. In the northeastern hardwoods forests this will typically be a nut (acorn, hickory) or small twig in the leaf litter. Nest mortality can be significant. From one third to one half of all nests are gone by the end of the winter. Some of these losses are colony deaths while others represent migration to a new nest site, which likely occurs during warmer winter days.

Nests of T. curvispinosus were found by Gibson et al. (2019) in bird nests.

MacGown (2006) examined ants nesting or occupying hickory nuts in the Tombigbee National Forest (Ackerman Unit), Mississippi. While several hickory species were present, ants were found almost exclusively in the nuts of Carya glabra. Nine colonies of Temnothorax curvispinosus were found, with 254 workers and 1 queen, 150 workers and 1 queen, 97 workers and 1 queen, 87 workers and 1 queen, 84 workers and 1 queen, 73 workers and 1 queen, 65 workers and 1 queen, 36 workers and 1 queen and 35 workers and 1 queen. These colonies fall within the size range of those observed by Headley (1943) in acorns. This is not surprising as Carya glabra hickory nuts are approximately the same size as many species of acorns, especially those of Quercus alba, which Headley mostly searched. It has been shown in laboratory tests that T. curvispinosus select larger cavities when given a choice and that they prefer a compact, high-ceilinged cavity over a thin, flat crevice, even if of equal volume (Pratt and Pierce, 2001). Combining this preference with the fact that the nuts of C. glabra have an extremely strong outer shell, it is clear that nuts and acorns provide a suitable and desirable shelter in all seasons for these ants. It also explains why T. curvispinosus colonies were found with relative ease and in large numbers in the hickory nuts while the species was otherwise only rarely encountered in the Tombigbee National Forest when using pitfall and other collecting methods (MacGown, 2006).

Reproduction

Queen number can vary by colony and season. New colonies are founded by pleoemetrosis and new queens are likely adopted into existing nests. The latter is evident from the presence within populations of both monogynous and polygonous nests. Reproductive queens contain 8 ovarioles.

Worker reproduction does occur with some male production possible from worker derived eggs. Reproductive workers contain 2 ovarioles.

New queens are produced in some queenless nests. These are presumed to be nests that are separated from a queenright nests or from a nest that had earlier lost its queen(s).

Kannowski (1959) noted the following concerning Temnothorax ambiguus reproduction in southeastern Michigan: "Alates were found in nests in bog communities from July 7 to August 6. In upland forest communities on the George Reserve, where curvispinosus is common, Talbot (1957, p. 451) found that alates were present in nests from late June to early August.

Flight Period

					X	X	X
Jan	Feb	Mar	Apr	May	Jun	Jul	Aug	Sep	Oct	Nov	Dec

Source: antkeeping.info.

Check details at Worldwide Ant Nuptial Flights Data, AntNupTracker and AntKeeping.
Explore: Show all Flight Month data or Search these data. See also a list of all data tables or learn how data is managed.

Association with Other Organisms

Explore: Show all Associate data or Search these data. See also a list of all data tables or learn how data is managed.

Other Ants

This species is parasitized by the slave-making ants Temnothorax americanus and Temnothorax duloticus and the workerless inquiline Temnothorax minutissimus (Beibl et al. 2005).

Life History Traits

Queen number: polygynous (Frumhoff & Ward, 1992)
Queen type: winged (Frumhoff & Ward, 1992) (queenless and queen-right worker reproduction)
Worker-produced males: present (Choe, 1988; Frumhoff & Ward, 1992)
Mean colony size: 50 (Headley, 1943; Talbot, 1965; Moglich, 1979; Beckers et al., 1989)
Foraging behaviour: tandem recruitment (Headley, 1943; Talbot, 1965; Moglich, 1979; Beckers et al., 1989)

Castes

Worker

Images from AntWeb


Worker. Specimen code casent0104032. Photographer April Nobile, uploaded by California Academy of Sciences.	Owned by ABS, Lake Placid, FL, USA.


Worker. Specimen code casent0104040. Photographer April Nobile, uploaded by California Academy of Sciences.	Owned by ABS, Lake Placid, FL, USA.

Queen

Images from AntWeb


Queen (alate/dealate). Specimen code casent0104039. Photographer April Nobile, uploaded by California Academy of Sciences.	Owned by ABS, Lake Placid, FL, USA.


.

Male

Images from AntWeb


Male (alate). Specimen code casent0104038. Photographer April Nobile, uploaded by California Academy of Sciences.	Owned by ABS, Lake Placid, FL, USA.

Nomenclature

The following information is derived from Barry Bolton's Online Catalogue of the Ants of the World.

curvispinosus. Leptothorax curvispinosus Mayr, 1866a: 508, pl., fig. 13 (w.) U.S.A. Mayr, 1886d: 451 (q.); Mackay, 2000: 338 (m.). Combination in L. (Myrafant): Smith, M.R. 1950: 30; in Temnothorax: Bolton, 2003: 271. Senior synonym of gallarum: Mayr, 1886d: 453. See also: Wheeler, W.M. 1903c: 239; Mackay, 2000: 337.
gallarum. Stenamma gallarum Patton, 1879: 126 (footnote) (w.q.) U.S.A. Junior synonym of curvispinosus: Mayr, 1886d: 453.

Type Material

North America. As reported in Mackay (2000): One cotype worker in Museo Civico di Storia Naturale, Genoa [seen]. specimens could not be located in Roger's collection (Museum fur Naturkunde Zentralinstitut der Humboldt-Universitat zu Berlin). Unless otherwise noted the text for the remainder of this section is reported from the publication that includes the original description.

Description

Worker

Operada: Long. 2.4mm Flava, capite rubescente-flavo, abdominis segmento basali postice fascia fusescenti medio interrupta; caput, thorax, petiolus et abdomen pilis subclavatis erectis. scapus pilis simplicibus paulo abstantibus, pedes sine pilis abstantibus; mandibulae subtiliter striolatae; clypeus longitudinaliter striatus, medio non impressus et inermis; caput dense reticulato-punctulatus, fronte distincte, vertice antice indistincte subtiliter et longitudinaliter rugulosis; antennae 11articulatae; thorax longitrorsum convexus, sine impressione transversa,subtiliter et longitudinaliter rugulosus et partim reticulato-punctatus; metanotum spinis duabus longis, paulo curvatis, parte metanoti basali sublongioribus, inter spinas transverse rugosum; petioli nodi,reticulato-punctati; abdomen laevissimum et nitidum.

Queen

Mayr (1886d) Weibchen. Lange: 3-3·3mm. Rothlichgelb oder gelbroth, die Oberkiefer, Fiihler und Beille mehr gelb, del' Kopf mehr oder weniger gebraunt, ein Fleck in del' Mitte des Mesonotum, das Scutellum, a,uch theilweise das Metanotum,dann del' Petiolus ganz oder theilweise und oft auch die Seiten des Thorax lichtbraun, der Hinterleib schwarzbraun, die Vorderhalfte des ersten Abdominalsegmentes gelb. Die Behaarung wie beim Arbeiter. Die Oberkiefer langsgestreift und fiinfzahnig. Del' Kopf ist langsgerunzelt, die Fiihlergruben sind fein genetzt, del' Clypeus massig convex, die Fiihler eilfgliedl'ig. Das Pronotum ist streifig-gerunzelt, hie und da sind die Runzeln mitsammell netzartig verbnnden, zwischen den Runzeln fein genetzt; das Mesonotum ist langsgestreift,das Scutellum langsgerunzelt; das Metanotum mit zwei ziemlich starkell, kaum gekriimmten parallelen Dornen, welche, innen gemessen, etwas kiirzer sind, als ihre Spitzen von einander entfernt sind; zwischen den Dornen ist das Metanotum quergerunzelt oder genetzt, aussen ist es langsgerunzelt und fein genetzt. Del' Hinterleib ist glatt und glanzend.

Male

Mackay (2000): Mandible with apical and 1 or 2 subapical teeth well defined, others present as denticles; median anterior border of clypeus weakly convex. clypeus with well defined central, longitudinal carina and a few other, poorly defined lateral carinae; eyes very large, occupying most of side of head; ocelli well developed, diameters of all three greater than distance between them; scape short, extending about half distance to occipital corner; propodeal spines or angles absent, only indication is a poorly defined carina; petiole with low node, which is not much higher than peduncle.

Hairs erect and scattered on most surfaces, suberect on antennae and legs; decumbent pubescence sparse.

Sculpture consisting of rough, longitudinal striae on most of head, area posterior to eyes with concentric whorls, clypeus with central carina and poorly defined lateral carinae, sides of mesosoma mostly smooth and shining, but with a few rugae, propodeum mostly smooth, but finally striate, petiole striate.

Color: pale yellow with black eyes.

Male measurements: HL 0.52-0.53, HW 0.47-0.50, SL 0.17-0.20, EL 0.25-0.26, WL 0.97-1.02. PW 0.10-0. 14, PL 0.08-0.23, PPW 0.18-0.19, PPL 0.22. Indices: CI 90-94, SI 33-38, PI 61-125, PPI 82.

Karyotype

n = 23 (USA) (Fischer, 1987) (as Leptothorax curvispinosus).

Etymology

Morphological. A reference to the inwardly curving propodeal spines of the workers.

References

Alloway, T. M., A. Buschinger, M. Talbot, R. Stuart, and C. Thomas. 1983 (1982). Polygyny and polydomy in three North American species of the ant genus Leptothorax Mayr (Hymenoptera: Formicidae). Psyche (Cambridge). 89:249-274.
André, E. 1881c. Catalogue raisonné des Formicides provenant du voyage en Orient de M. Abeille de Perrin et description des espèces nouvelles. Ann. Soc. Entomol. Fr. (6)(1): 53-78 (page 72, worker described)
André, E. 1883a. Les fourmis. [part]. Pp. 281-344 in: André, Edm. 1881-1886. Species des Hyménoptères d'Europe et d'Algérie. Tome Deuxième. Beaune: Edmond André, 919 + 48 pp. (page 302, Replacement name: flavispinus)
Beckers R., Goss, S., Deneubourg, J.L., Pasteels, J.M. 1989. Colony size, communication and ant foraging Strategy. Psyche 96: 239-256 (doi:10.1155/1989/94279).
Beibl, J., R. J. Stuart, J. Heinze, and S. Foitzik. 2005. Six origins of slavery in formicoxenine ants. Insectes Sociaux. 52:291-297.
Beibl, J., Stuart, R.J., Heinze, J., Foitzik, S. 2005. Six origins of slavery in formicoxenine ants. Insectes Sociaux 52, 291–297 (doi:10.1007/s00040-005-0808-y).
Bolton, B. 2003. Synopsis and Classification of Formicidae. Mem. Am. Entomol. Inst. 71: 370pp (page 271, Combination in Temnothorax)
Booher, D., MacGown, J.A., Hubbell, S.P., Duffield, R.M. 2017. Density and dispersion of cavity dwelling ant species in nuts of eastern US forest floors. Transactions of the American Entomological Society 143: 79-93 (doi:10.3157/061.143.0105).
Buschinger, A.; Linksvayer, T. A. 2004. Novel blend of life history traits in an inquiline ant, Temnothorax minutissimus, with description of the male (Hymenoptera: Formicidae). Myrmecol. Nachr. 6: 67-76 (see also)
Carroll, T.M. 2011. The ants of Indiana (Hymenoptera: Formicidae). M.S. thesis, Purdue University.
Casacci, L.P., Barbero, F., Slipinski, P., Witek, M. 2021. The inquiline ant Myrmica karavajevi uses both chemical and vibroacoustic deception mechanisms to integrate into its host colonies. Biology 10, 654 (doi:10.3390/ biology10070654)..
Chernyshova, A.M. 2021. A genetic perspective on social insect castes: A synthetic review and empirical study. M.S. thesis, The University of Western Ontario. Electronic Thesis and Dissertation Repository 7771.
Chick, L.D., Lessard, J.-P., Dunn, R.R., Sanders, N.J. 2020. The coupled influence of thermal physiology and biotic interactions on the distribution and density of ant species along an elevational gradient. Diversity 12, 456 (doi:10.3390/d12120456).
Davis, T. 2009. The ants of South Carolina (thesis, Clemson University).
de la Mora, A., Sankovitz, M., Purcell, J. 2020. Ants (Hymenoptera: Formicidae) as host and intruder: recent advances and future directions in the study of exploitative strategies. Myrmecological News 30: 53-71 (doi:10.25849/MYRMECOL.NEWS_030:053).
Gibson, J. C., A. V. Suarez, D. Qazi, T. J. Benson, S. J. Chiavacci, and L. Merrill. 2019. Prevalence and consequences of ants and other arthropods in active nests of Midwestern birds. Canadian Journal of Zoology. 97:696-704. doi:10.1139/cjz-2018-0182
Gochnour, B.M., Suiter, D.R., Booher, D. 2019. Ant (Hymenoptera: Formicidae) fauna of the Marine Port of Savannah, Garden City, Georgia (USA). Journal of Entomological Science 54, 417-429 (doi:10.18474/jes18-132).
Headley, A. E. 1943. Population studies of two species of ants, Leptothorax longispinosus Roger and Leptothorax curvispinosus Mayr. Annals of the Entomological Society of America 36: 743-753 (doi:10.1093/aesa/36.4.743).
Herbers, J. M. and C. A. Johnson. 2007. Social structure and winter survival in acorn ants. Oikos (Copenhagen). 116:829-835.
Ipser, R.M., Brinkman, M.A., Gardner, W.A., Peeler, H.B. 2004. A survey of ground-dwelling ants (Hymenoptera: Formicidae) in Georgia. Florida Entomologist 87: 253-260.
Ivanov, K. 2019. The ants of Ohio (Hymenoptera, Formicidae): an updated checklist. Journal of Hymenoptera Research 70: 65–87 (doi:10.3897@jhr.70.35207).
Kannowski, P. B. 1959. The flight activities and colony-founding behavior of bog ants in southeastern Michigan. Insectes Sociaux. 6:115-162.
Kaur, R., Joseph, J., Anoop, K., Sumana, A. 2017. Characterization of recruitment through tandem running in an Indian queenless ant Diacamma indicum. Royal Society Open Science 41, 160476 (doi:10.1098/rsos.160476).
Khimoun, A., Doums, C., Molet, M., Kaufmann, B., Peronnet, R., Eyer, P.A., Mona, S. 2020. Urbanization without isolation: the absence of genetic structure among cities and forests in the tiny acorn ant Temnothorax nylanderi. Biology Letters 16, 20190741 (doi:10.1098/rsbl.2019.0741).
MacGown, J.A. 2006. Hickory nuts used as nesting sites by ants (Hymenoptera: Formicidae). Marginalia Insecta 1(1): 1-3.
MacKay, W. P. 2000. A review of the New World ants of the subgenus Myrafant, (genus Leptothorax) (Hymenoptera: Formicidae). Sociobiology 36: 265-444 (page 337, see also)
Mackay, W. P. and E. Mackay. 2002. The ants of New Mexico (Hymenoptera: Formicidae). Edwin Mellen Press, Lewiston, NY.
Mayr, G. 1866a. Myrmecologische Beiträge. Sitzungsber. Kais. Akad. Wiss. Wien Math.-Naturwiss. Cl. Abt. I 53: 484-517 (page 508, pl., fig. 13 worker described)
Mayr, G. 1886d. Die Formiciden der Vereinigten Staaten von Nordamerika. Verh. K-K. Zool.-Bot. Ges. Wien 36: 419-464 (page 451, queen described, page 453, Senior synonym of gallarum)
Pratt, S.C. 2010. Collective intelligence. Encyclopedia of Animal Behavior (Second Edition), pp. 303-309.
Pratt, S.C., Pierce, N.E. 2001. The cavity-dwelling ant Leptothorax curvispinosus uses nest geometry to discriminate between potential homes. Animal Behaviour 62: 281-287 (doi:10.1006/anbe.2001.1777).
Prebus, M.M. 2020. Phylogenomic species delimitation in the ants of the Temnothorax salvini group (Hymenoptera: Formicidae): an integrative approach. Systematic Entomology. 46:307-326. doi:https://doi.org/10.1111/syen.12463
Prebus, M.M. 2021. Taxonomic revision of the Temnothorax salvini clade (Hymenoptera: Formicidae), with a key to the clades of New World Temnothorax. PeerJ 9, e11514 (doi:10.7717/peerj.11514).
Rericha, L. 2007. Ants of Indiana. Indiana Department of Natural Resources, 51pp.
Schultner, E., Pulliainen, U. 2020. Brood recognition and discrimination in ants. Insectes Sociaux 67, 11–34 (doi:10.1007/s00040-019-00747-3).
Smith, M. R. 1950a. On the status of Leptothorax Mayr and some of its subgenera. Psyche (Camb.) 57: 29-30 (page 30, Combination in L. (Myrafant))
Stukalyuk, S., Radchenko, Y., Gonchar, O., Akhmedov, A., Stelia, V., Reshetov, A., Shymanskyi, A. 2021. Mixed colonies of Lasius umbratus and Lasius fuliginosus (Hymenoptera, Formicidae): when superparasitism may potentially develop into coexistence: a case study in Ukraine and Moldova. Halteres 12, 25–48 (doi:10.5281/zenodo.5753121).
Talbot, M. 1957. Population studies of the slave-making ant Leptothorax duloticus and its slave, Leptothorax curvispinosus. Ecology. 38:449-456.
Warren II, R.J., Bayba, S., Krupp, K.T. 2018. Interacting effects of urbanization and coastal gradients on ant thermal responses. Journal of Urban Ecology 4: 1-11 (doi:10.1093/jue/juy026).
Waters, J.S., Keough, N.W., Burt, J., Eckel, J.D., Hutchinson, T., Ewanchuk, J., Rock, M., Markert, J.A., Axen, H.J., Gregg, D. 2022. Survey of ants (Hymenoptera, Formicidae) in the city of Providence (Rhode Island, United States) and a new northern-most record for Brachyponera chinensis (Emery, 1895). Check List 18(6), 1347–1368 (doi:10.15560/18.6.1347).
Wheeler, W. M. 1903d. A revision of the North American ants of the genus Leptothorax Mayr. Proc. Acad. Nat. Sci. Phila. 55: 215-260 (page 239, see also)

References based on Global Ant Biodiversity Informatics

Amstutz M. E. 1943. The ants of the Kildeer plain area of Ohio (Hymenoptera, Formicidae). The Ohio Journal of Science 43(4): 165-173.
Beibl, J., R.J. Stuart, J. Heinze and S. Foitzik. 2005. Six origins of slavery in formicoxenine ants. Insectes Sociaux 52:291-297
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.
Booher D., J. A. MacGown, R. M. Duffield, and S. P. Hubbell. 2012. Density and Dispersion of Cavity Dwelling Ant Species in Nuts of Eastern US Forest Floors. Entomological Society of America annual meeting Knoxville, 2012.
Buren W. F. 1941. A preliminary list of Iowa ants. Iowa State College Journal of Science 15: 111-117
Buren W. F. 1944. A list of Iowa ants. Iowa State College Journal of Science 18:277-312
Carroll T. M. 2011. The ants of Indiana (Hymenoptera: Formicidae). Master's Thesis Purdue university, 385 pages.
Clark A. T., J. J. Rykken, and B. D. Farrell. 2011. The Effects of Biogeography on Ant Diversity and Activity on the Boston Harbor Islands, Massachusetts, U.S.A. PloS One 6(11): 1-13.
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. 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
Creighton W. S. 1927. The slave-raids of Harpagoxenus americanus. Psyche (Cambridge) 34: 11-29.
Dash S. T. and L. M. Hooper-Bui. 2008. Species diversity of ants (Hymenoptera: Formicidae) in Louisiana. Conservation Biology and Biodiversity. 101: 1056-1066
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., K. Towle, D. N. Morrison, and S. L. Pelini. 2013. Community Structure, Ecological and Behavioral Traits of Ants (Hymenoptera: Formicidae) in Massachusetts Open and Forested Habitats. Northeastern Naturalist 20: 1-12.
Del Toro, I. 2010. PERSONAL COMMUNICATION. MUSEUM RECORDS COLLATED BY ISRAEL DEL TORO
Deyrup M., C. Johnson, G. C. Wheeler, J. Wheeler. 1989. A preliminary list of the ants of Florida. Florida Entomologist 72: 91-101
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: 153-725
DuBois M. B. 1985. Distribution of ants in Kansas: subfamilies Ponerinae, Ecitoninae, and Myrmicinae (Hymenoptera: Formicidae). Sociobiology 11: 153-726
DuBois M. B. 1985. Distribution of ants in Kansas: subfamilies Ponerinae, Ecitoninae, and Myrmicinae (Hymenoptera: Formicidae). Sociobiology 11: 153-727
DuBois M. B. 1985. Distribution of ants in Kansas: subfamilies Ponerinae, Ecitoninae, and Myrmicinae (Hymenoptera: Formicidae). Sociobiology 11: 153-728
DuBois M. B. 1985. Distribution of ants in Kansas: subfamilies Ponerinae, Ecitoninae, and Myrmicinae (Hymenoptera: Formicidae). Sociobiology 11: 153-729
DuBois M. B. 1985. Distribution of ants in Kansas: subfamilies Ponerinae, Ecitoninae, and Myrmicinae (Hymenoptera: Formicidae). Sociobiology 11: 153-730
DuBois M. B. 1985. Distribution of ants in Kansas: subfamilies Ponerinae, Ecitoninae, and Myrmicinae (Hymenoptera: Formicidae). Sociobiology 11: 153-731
DuBois M. B. 1985. Distribution of ants in Kansas: subfamilies Ponerinae, Ecitoninae, and Myrmicinae (Hymenoptera: Formicidae). Sociobiology 11: 153-732
DuBois M. B. 1985. Distribution of ants in Kansas: subfamilies Ponerinae, Ecitoninae, and Myrmicinae (Hymenoptera: Formicidae). Sociobiology 11: 153-733
DuBois M. B. 1985. Distribution of ants in Kansas: subfamilies Ponerinae, Ecitoninae, and Myrmicinae (Hymenoptera: Formicidae). Sociobiology 11: 153-734
DuBois M. B. 1985. Distribution of ants in Kansas: subfamilies Ponerinae, Ecitoninae, and Myrmicinae (Hymenoptera: Formicidae). Sociobiology 11: 153-735
DuBois M. B. 1985. Distribution of ants in Kansas: subfamilies Ponerinae, Ecitoninae, and Myrmicinae (Hymenoptera: Formicidae). Sociobiology 11: 153-736
DuBois M. B. 1985. Distribution of ants in Kansas: subfamilies Ponerinae, Ecitoninae, and Myrmicinae (Hymenoptera: Formicidae). Sociobiology 11: 153-737
DuBois M. B. 1985. Distribution of ants in Kansas: subfamilies Ponerinae, Ecitoninae, and Myrmicinae (Hymenoptera: Formicidae). Sociobiology 11: 153-738
DuBois M. B. 1985. Distribution of ants in Kansas: subfamilies Ponerinae, Ecitoninae, and Myrmicinae (Hymenoptera: Formicidae). Sociobiology 11: 153-739
DuBois M. B. 1985. Distribution of ants in Kansas: subfamilies Ponerinae, Ecitoninae, and Myrmicinae (Hymenoptera: Formicidae). Sociobiology 11: 153-740
DuBois M. B. 1985. Distribution of ants in Kansas: subfamilies Ponerinae, Ecitoninae, and Myrmicinae (Hymenoptera: Formicidae). Sociobiology 11: 153-741
DuBois M. B. 1985. Distribution of ants in Kansas: subfamilies Ponerinae, Ecitoninae, and Myrmicinae (Hymenoptera: Formicidae). Sociobiology 11: 153-742
DuBois M. B. 1985. Distribution of ants in Kansas: subfamilies Ponerinae, Ecitoninae, and Myrmicinae (Hymenoptera: Formicidae). Sociobiology 11: 153-743
DuBois M. B. 1985. Distribution of ants in Kansas: subfamilies Ponerinae, Ecitoninae, and Myrmicinae (Hymenoptera: Formicidae). Sociobiology 11: 153-744
DuBois M. B. 1985. Distribution of ants in Kansas: subfamilies Ponerinae, Ecitoninae, and Myrmicinae (Hymenoptera: Formicidae). Sociobiology 11: 153-745
DuBois M. B. 1985. Distribution of ants in Kansas: subfamilies Ponerinae, Ecitoninae, and Myrmicinae (Hymenoptera: Formicidae). Sociobiology 11: 153-746
DuBois M. B. 1985. Distribution of ants in Kansas: subfamilies Ponerinae, Ecitoninae, and Myrmicinae (Hymenoptera: Formicidae). Sociobiology 11: 153-747
DuBois M. B. 1985. Distribution of ants in Kansas: subfamilies Ponerinae, Ecitoninae, and Myrmicinae (Hymenoptera: Formicidae). Sociobiology 11: 153-748
DuBois M. B. 1985. Distribution of ants in Kansas: subfamilies Ponerinae, Ecitoninae, and Myrmicinae (Hymenoptera: Formicidae). Sociobiology 11: 153-749
DuBois M. B. 1985. Distribution of ants in Kansas: subfamilies Ponerinae, Ecitoninae, and Myrmicinae (Hymenoptera: Formicidae). Sociobiology 11: 153-750
DuBois M. B. 1985. Distribution of ants in Kansas: subfamilies Ponerinae, Ecitoninae, and Myrmicinae (Hymenoptera: Formicidae). Sociobiology 11: 153-751
DuBois M. B. 1985. Distribution of ants in Kansas: subfamilies Ponerinae, Ecitoninae, and Myrmicinae (Hymenoptera: Formicidae). Sociobiology 11: 153-752
DuBois M. B. 1985. Distribution of ants in Kansas: subfamilies Ponerinae, Ecitoninae, and Myrmicinae (Hymenoptera: Formicidae). Sociobiology 11: 153-753
DuBois M. B. 1985. Distribution of ants in Kansas: subfamilies Ponerinae, Ecitoninae, and Myrmicinae (Hymenoptera: Formicidae). Sociobiology 11: 153-754
DuBois M. B. 1985. Distribution of ants in Kansas: subfamilies Ponerinae, Ecitoninae, and Myrmicinae (Hymenoptera: Formicidae). Sociobiology 11: 153-755
DuBois M. B. 1985. Distribution of ants in Kansas: subfamilies Ponerinae, Ecitoninae, and Myrmicinae (Hymenoptera: Formicidae). Sociobiology 11: 153-756
DuBois M. B. 1985. Distribution of ants in Kansas: subfamilies Ponerinae, Ecitoninae, and Myrmicinae (Hymenoptera: Formicidae). Sociobiology 11: 153-757
DuBois M. B. 1985. Distribution of ants in Kansas: subfamilies Ponerinae, Ecitoninae, and Myrmicinae (Hymenoptera: Formicidae). Sociobiology 11: 153-758
DuBois M. B. 1985. Distribution of ants in Kansas: subfamilies Ponerinae, Ecitoninae, and Myrmicinae (Hymenoptera: Formicidae). Sociobiology 11: 153-759
DuBois M. B. 1985. Distribution of ants in Kansas: subfamilies Ponerinae, Ecitoninae, and Myrmicinae (Hymenoptera: Formicidae). Sociobiology 11: 153-760
DuBois M. B. 1985. Distribution of ants in Kansas: subfamilies Ponerinae, Ecitoninae, and Myrmicinae (Hymenoptera: Formicidae). Sociobiology 11: 153-761
DuBois M. B. 1985. Distribution of ants in Kansas: subfamilies Ponerinae, Ecitoninae, and Myrmicinae (Hymenoptera: Formicidae). Sociobiology 11: 153-762
DuBois M. B. 1985. Distribution of ants in Kansas: subfamilies Ponerinae, Ecitoninae, and Myrmicinae (Hymenoptera: Formicidae). Sociobiology 11: 153-763
Dubois, M.B. and W.E. Laberge. 1988. An Annotated list of the ants of Illionois. pages 133-156 in Advances in Myrmecology, J. Trager
Ellison A. M., E. J. Farnsworth, and N. J. Gotelli. 2002. Ant diversity in pitcher-plant bogs of Massachussetts. Northeastern Naturalist 9(3): 267-284.
Ellison A. M., S. Record, A. Arguello, and N. J. Gotelli. 2007. Rapid Inventory of the Ant Assemblage in a Temperate Hardwood Forest: Species Composition and Assessment of Sampling Methods. Environ. Entomol. 36(4): 766-775.
Ellison A. M., and E. J. Farnsworth. 2014. Targeted sampling increases knowledge and improves estimates of ant species richness in Rhode Island. Northeastern Naturalist 21(1): NENHC-13NENHC-24.
Emery C. 1895. Beiträge zur Kenntniss der nordamerikanischen Ameisenfauna. (Schluss). Zoologische Jahrbücher. Abteilung für Systematik, Geographie und Biologie der Tiere 8: 257-360.
Evans, J.D. and N.E. Pierce. 1995. Effects of Diet Quality and Queen Number on Growth in Leptothoracine Ant Colonies (Hymenoptera: Formicidae). Journal of the New York Entomological Society 103(1):91-99
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., and L. Thompson. 2008. New distributional records of ants in Arkansas. Journal of the Arkansas Academy of Science 62: 148-150.
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
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
Guénard B., K. A. Mccaffrey, A. Lucky, and R. R. Dunn. 2012. Ants of North Carolina: an updated list (Hymenoptera: Formicidae). Zootaxa 3552: 1-36.
Headley A. E. 1943. The ants of Ashtabula County, Ohio (Hymenoptera, Formicidae). The Ohio Journal of Science 43(1): 22-31.
Heithaus R. E., and M. Humes. 2003. Variation in Communities of Seed-Dispersing Ants in Habitats with Different Disturbance in Knox County, Ohio. OHIO J. SCI. 103 (4): 89-97.
Herbers J. M. 2011. Nineteen years of field data on ant communities (Hymenoptera: Formicidae): what can we learn. Myrmecological News 15: 43-52.
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
Hill, J.G. 2006. Ants collected at Okatibbee Lake, Lauderdale County, Mississippi
Ipser R. M. 2004. Native and exotic ants (Hymenoptera: Formicidae) of Georgia: Ecological Relationships with implications for development of biologically-based management strategies. Doctor of Philosophy thesis, University of Georgia. 165 pages.
Ipser, R.M., M.A. Brinkman, W.A. Gardner and H.B. Peeler. 2004. A Survey of Ground-Dwelling Ants (Hymenoptera: Formicidae) in Georgia. The Florida Entomologist 87(3) 253-260.
Ivanov, K. 2019. The ants of Ohio (Hymenoptera, Formicidae): an updated checklist. Journal of Hymenoptera Research 70: 65–87.
Ivanov K., L. Hightower, S. T. Dash, and J. B. Keiper. 2019. 150 years in the making: first comprehensive list of the ants (Hymenoptera: Formicidae) of Virginia, USA. Zootaxa 4554 (2): 532–560.
Jeanne R. J. 1979. A latitudinal gradient in rates of ant predation. Ecology 60(6): 1211-1224.
Kjar D. 2009. The ant community of a riparian forest in the Dyke Marsh Preserve, Fairfax County, Virginiam and a checklist of Mid-Atlantic Formicidae. Banisteria 33: 3-17.
Kjar D., and E. M. Barrows. 2004. Arthropod community heterogeneity in a mid-Atlantic forest highly invaded by alien organisms. Banisteria 23: 26-37.
Kjar D., and Z. Park. 2016. Increased ant (Hymenoptera: Formicidae) incidence and richness are associated with alien plant cover in a small mid-Atlantic riparian forest. Myrmecological News 22: 109-117.
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.
Lessard, J.-P., R. R. Dunn and N. J. Sanders. 2009. Temperature-mediated coexistence in temperate forest ant communities. Insectes Sociaux 56(2):149-456.
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
Lynch J. F., and A. K. Johnson. 1988. Spatial and temporal variation in the abundance and diversity of ants (Hymenoptera: Formicidae) in the soild and litter layers of a Maryland forest. American Midland Naturalist 119(1): 31-44.
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 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
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. 2000. A review of the New World ants of the subgenus Myrafant, (genus Leptothorax) (Hymenoptera: Formicidae). Sociobiology 36: 265-444.
Mahon M. B., K. U. Campbell, and T. O. Crist. 2017. Effectiveness of Winkler litter extraction and pitfall traps in sampling ant communities and functional groups in a temperate forest. Environmental Entomology 46(3): 470–479.
Mann H. R., E. Rowe, J. Selfridge, and D. L. Price. 2018. Leaf litter and arboreal ants (Hymenoptera: Formicidae) in a Mid-Atlantic Forest. Northeastern Naturalist 25(2): 341-354.
Martelli, M.G., M.M. Ward and Ann M. Fraser. 2004. Ant Diversity Sampling on the Southern Cumberland Plateau: A Comparison of Litter Sifting and Pitfall Trapping. Southeastern Naturalist 3(1): 113-126
Menke S. B., E. Gaulke, A. Hamel, and N. Vachter. 2015. The effects of restoration age and prescribed burns on grassland ant community structure. Environmental Entomology http://dx.doi.org/10.1093/ee/nvv110
Menke S. B., and N. Vachter. 2014. A comparison of the effectiveness of pitfall traps and winkler litter samples for characterization of terrestrial ant (Formicidae) communities in temperate savannas. The Great Lakes Entomologist 47(3-4): 149-165.
Menzel T. O., and T. E. Nebeker. 2008. Distribution of Hybrid Imported Fire Ants (Hymenoptera: Formicidae) and Some Native Ant Species in Relation to Local Environmental Conditions and Interspecific Competition in Mississippi Forests. Ann. Entomol. Soc. Am. 101(1): 119-127.
Merle W. W. 1939. An Annotated List of the Ants of Maine (Hymenoptera: Formicidae). Entomological News. 50: 161-165
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.
O'Neill J.C. and Dowling A.P.G. 2011. A Survey of the Ants (hymenoptera: Formicidae) of Arkansas and the Ozark Mountains. An Undergraduate Honors, University of Arkansas. 18pages.
Resasco J., S. L. Pelini, K. L. Stuble, N. J. Sanders, R. R. Dunn, S. E. Diamond, A. M. Ellison, N. J. Gotelli, and D. J. Levey. 2014. Using Historical and Experimental Data to Reveal Warming Effects on Ant Assemblages. PLoS ONE 9(2): e88029. doi:10.1371/journal.pone.0088029
Rowles, A.D. and J. Silverman. 2009. Carbohydrate supply limits invasion of natural communities by Argentine ants. Oecologia 161(1):161-171
Smith M. R. 1962. A new species of exotic Ponera from North Carolina (Hymenoptera, Formicidae). Acta Hymenopterologica 1: 377-382.
Sturtevant A. H. 1931. Ants collected on Cape Cod, Massachusetts. Psyche (Cambridge) 38: 73-79
Talbot M. 1957. Populations of ants in a Missouri woodland. Insectes Sociaux 4(4): 375-384.
Talbot M. 1976. A list of the ants (Hymenoptera: Formicidae) of the Edwin S. George Reserve, Livingston County, Michigan. Great Lakes Entomologist 8: 245-246.
Toennisson T. A., N. J. Sanders, W. E. Klingeman, and K. M. Vail. 2011. Influences on the Structure of Suburban Ant (Hymenoptera: Formicidae) Communities and the Abundance of Tapinoma sessile. Environ. Entomol. 40(6): 1397-1404.
Van Pelt A., 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.
Wang C., J. Strazanac and L. Butler. 2000. Abundance, diversity and activity of ants (Hymenoptera: Formicidae) in oak-dominated mixed Appalachian forests treated with microbial pesticides. Environmental Entomology. 29: 579-586
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. 1987. A Checklist of the Ants of South Dakota. Prairie Nat. 19(3): 199-208.
Wheeler W. M. 1903. A revision of the North American ants of the genus Leptothorax Mayr. Proceedings of the Academy of Natural Sciences of Philadelphia. 55: 215-260.
Wheeler W. M. 1917. A list of Indiana ants. Proceedings of the Indiana Academy of Science 26: 460-466.
Wheeler, G.C. and J. Wheeler. 1985. A checklist of Texas ants. Prairie Naturalist 17:49-64.
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.
Wing M. W. 1939. An annotated list of the ants of Maine (Hymenoptera: Formicidae). Entomological News 50:161-165.