Ponera pennsylvanica

A common member of most ant communities in deciduous forests of the eastern United States.

Identification
Taylor (1967) - A member of the coarctata species group, endemic to eastern North America. Distinguished from other species of the genus, including Ponera coarctata, by the characters listed on the latter species page. The larger size, presence of a distinct dorsal mesometanotal suture in the worker, and the undifferentiated antennal club in the female castes, distinguish pennsylvanica from Ponera exotica M. R. Smith, the only other known New World species of Ponera. Exotica is evidently not closely related to pennsylvanica, and is likely introduced from the Indo-Australian area, though it is not yet known from there.

Distribution
In Florida this species is known from north Florida south into Marion Co. The general distribution is from southern Nova Scotia south into northern Florida, west into eastern Minnesota and south into northeastern Texas (Taylor 1967). West of this area there are a few scattered records from North and South Dakota, Colorado and New Mexico (Taylor 1967), with an outlier in Michoacan in west-central Mexico (Mackay 1991). Like Ponera exotica, P. pennsylvanica presents a biogeographic riddle. It is unlikely to be a boreal lineage, even though its closest known relative occurs in Europe. A reasonable hypothesis is that a lineage moved north from the Indo-Australian region and expanded around the world in the band of Arctotertiaiy forest, which has vestiges in Europe and a better representation in eastern North America and the temperate Far East. Perhaps related species will turn up in China or Japan to support this hypothesis.

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

Biology
Taylor (1967) - P. pennsylvanica is characteristically found nesting in deciduous forests of various types, although it may penetrate more open habitats or coniferous forests. In the Chicago region Talbot (1934), and Gregg (1944) have reported it from stands of Black, Red, or White Oak, and mixed oak forest on sandy soil and from mixed oak forest on clay. The greatest number of colonies were found in such situations, but the species was often taken at woods margins, in fields, or along roadsides. Dennis (1938) found pennsylvanica widespread and common in Tennessee, in oak-pine and oak-chestnut forests over a wide elevational range (90 to 1515 m). It was found to be the most abundant of 15 species nesting in locust woods in Seneca County, Ohio, by Headley (1952). This author sampled the ants nesting in 15 square-meter plots during August 1958, and in 12 adjacent, similar sized plots in June, July and September 1949. In all 299 ant colonies were taken in the 27 square meters thus studied, with P. pennsylvanica by far the most common species, with 109 (36 %) of the colonies collected. The next most abundant species (Stenamma brevicorne Mayr) had only 35 nests (12% of the total). P. pennsylvanica has been listed as one of the most common ants in North Carolina by Carter (1962b). He found it especially abundant on the Piedmont Plateau in various forest associations, most notably in upland mature pine and oakhickory climax stands; the elevational range being 600 to 1030 m. P. pennsylvanica was stated by Carter to be "predictable" in coastal forest, but scarcer than on the Plateau. All of his records, except one, were from well shaded mesic forests. The extreme northwest record from Saganaga Lake, Minnesota, was made in coniferous forest (Gregg 1946).

Nests in forest areas are usually constructed in and under rotting logs, or in rotting stumps, at all but the earliest stages of decay (Talbot 1934); in small fragments of wood, acorns or other such objects; or in soil or leafmold. In drier habitats, especially non-forested areas, the colonies are usually found in soil under stones or other material, and at times the species may be common under stones in pastures (Gregg 1944). Workers forage in or on the surface of the ground, and may be readily collected using the Berlese funnel. Headley (1952) has given details concerning nest depth and proportions, and the size and proportion of colonies. He found 217 workers in 86 nests, undoubtedly a low count. During his August 1948, Ohio, survey he found no eggs and few larvae in 86 colonies. Pupae were, however, fairly abundant, with a mean of 11.2 per nest. The largest colony contained 36 pupae of all castes, one included 15 workers and 18 pupae, 10 of which were alates. The species apparently does not overwinter any brood (Talbot 1957).

Twenty-two sexual imagos were present in 4 of Headley's August colonies, but no flights were observed by him. Mating flight activity may extend from mid-August to early October, according to dates on the specimens I have seen, and those published by Headley (1943), Gregg (1944), and Kannowski (1959). Colony foundation is independent and non-claustral (Kannowski loco Cit.), and dealate females may be taken in rotting wood, soil, moss, or curled leaves. Some very incomplete mating flight observations are given by Haskins and Enzmann (1938).

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. A single colony of Ponera pennsylvanica, with 5 workers, was found. This is consistent with Headley (1943), who also reported finding nests of Ponera pennsylvanica in cavities, but in this case in Oak acorns.

Ecology
Taylor (1967) - The range of pennsylvanica corresponds remarkably well with the area carrying the Eastern Deciduous Forest Formation. The western and southern extensions of which are indicated on the map (see also Braun 1950); the western limit of the forest formation proper is shown as a dotted line; that of the forest-prairie transition belt as a solid line. Tongues of transition forest extend westwards into prairie grassland carrying with them forest pockets in which pennsylvanica occurs. The area of forest or transition country from which pennsylvanica has been recorded is shaded on the map; apparently this is the main range of the species. The few records known from outside the forest zones are indicated as dots.

The records appear to indicate a northwestern limit at about the latitude of northern Iowa. However, there are more northerly records from Saganaga Lake in extreme NE Minnesota (Gregg 1946), and from the Porcupine Mts in the upper peninsula of Michigan. These must be close to the limits of the range in this area.

Further east, in the southern Great Lakes area and New England, pennsylvanica seems fairly common at latitudes where it is reportedly rare in the west. The northernmost known records are from the vicinity of Quebec. The Digby, Nova Scotia, record may indicate present distribution at least as far as the "neck" of that province, however, the species is unknown from northern Maine and New Brunswick. P. pennsylvanica may be taken at low elevations in northern New England, but it is less common there than in the southern part of the area. Sturtevant (1931) noted that it was much less common on Cape Cod than near New York City.

The northern distributional limits are almost certainly set by deteriorating climatic conditions, and in fact correspond fairly well with the 40° Fahrenheit (4SC) mean annual temperature isotherm (see Vischer 1954: map 3). pennsylvanica apparently declines in abundance northwards, from a line running approximately through Milwaukee and Ottawa, on the map projection used in fig. 22. This corresponds almost exactly with the 45° Fahrenheit (7.4°C) mean annual isotherm as shown by Vischer. Ignoring the two extreme northwestern records, the main northern limit of pennsylvanica trends southwards as it moves inland towards the central plains. This trend is perhaps correlated with the increase in severity of climate at a given latitude moving westwards across the Great Lakes area.

The western limit of the main distributional range is clearly marked. All records available from South Dakota, Nebraska, Kansas, and Oklahoma, being towards their eastern areas; with the extreme western ones falling almost exactly in a north-south line. Details of the distribution in these States and in Iowa are interesting, for most records fall within the area of the Eastern Deciduous Forest Formation or its prairie-transition belt. The Iowa records of Smith and Buren are significant in this connection. This state is occupied mainly by country floristically part of the midwestern grassland-prairie association. Two areas are within the range of the forest-prairie transition belt: (1) the valleys of the Mississippi and Des Moines Rivers and (2) the drainage system of the Missouri River, which forms a long tongue extending northwards from Missouri, along the western Iowa border. All of the Iowa records listed above fall within the transitional forest zones, and not in the prairie zones. The single South Dakota record is also from transition country, and was made in the relatively small extension of the Missouri River valley "transition forest tongue in the extreme south east of the state. The Nebraska records, too, both lie within the limits of this transition forest extension, at the extreme east of the state. The only known Kansas records are from Riley and Dickinson Counties, which lie outside the broad limits of the forest transition belt; unfortunately no ecological details are available.

All Oklahoma records are from the east of the state, in transition country. Professor W. G. Carter, the collector, has provided more precise ecological details than are usually available. The Payne County specimens were taken in mesic forests of oaks and other hardwoods, growing in stream floodplains and deep ravines. The area in general is characterized by large tracts of tall or mixed grassland and there are also xeric scrub oak forests in certain highland areas. McCurtain County records were also from similar mesic ravine forest in primarily grassland areas. Professor Carter (in litt.) recently found pennsylvanica common in eastern Oklahoma.

South of Oklahoma the known distributional limit has a strong easterly trend. Records from this area are scarce, but, it is notable that the 50% line of soil moisture saturation, as mapped by Vischer (1954: map 954, p. 359), follows about the same path. Considering that soil moisture values depend on interaction of numerous ecological parameters this may be significant, especially as pennsylvanica is soil inhabiting.

In Florida pennsylvanica is known only from Gainsville, in the north. It was not taken by Van Pelt (1956) in his careful survey of Welaka Reserve, slightly further south. The apparent southern limit indicated by these data almost exactly coincides with the limit of deciduous forest, and lies across an area where "one after another of the deciduous (forest) species reaches the southern limits of its range" (Braun 1950) exact limiting factors acting here on pennsylvanica are not clear, but considering its decline southward in Mississippi (M. R. Smith 1956) and possibly in other Gulf Coast states and Georgia (teste E. O. Wilson), climatic factors are probably involved. The possibility of competitive exclusion by other ant species should be considered.

In general the marginal limiting factors for P. pennsylvanica appear to be "density independent" ones-climatic deterioration in the north, decreasing humidity in the west and southwest, and high temperature (and humidity?) in the south and southeast. These factors seem to be effective even in areas where suitable forest occurs, such as the northwest and southwest of its range.

Although the main limits of P. pennsylvanica are approximately co-extensive with those of the Eastern Deciduous Forests, the ant ranges far to the west, where it is obviously very scarce. As Gregg (1963) has noted, it might be expected to occur sporadically in river valley forests throughout the central plains area.

Nomenclature

 *  pennsylvanica. Ponera pennsylvanica Buckley, 1866: 171 (w.) U.S.A. Emery, 1895c: 267 (q.m.); Wheeler, G.C. & Wheeler, J. 1952c: 631 (l.). Subspecies of coarctata: Emery, 1895c: 267; Dennis, 1938: 277; Creighton, 1950a: 48. Revived status as species: Taylor, 1967a: 29.

Type Material


No types are known to exist. This species is unlikely to be confused with any other, and usage of the name has been unambiguous; I have not, therefore, designated a neotype. (Taylor 1967)

Worker
Smith (1936) - Length: 3-3.75 mm. Head, excluding the mandibles, very much longer than broad, sides moderately convex, posterior border faintly but definitely excised in the middle. Eyes very small, not convex, composed of approximately 6 ommatidia, placed toward the front of the head about one-fourth the distance from the base of the mandibles. Ocelli absent. Clypeus convex medianally. No distinct frontal area. Frontal furrow extending approximately half the distance from the frontal carinae to the posterior border of the head. Mandibles with two or three prominent teeth anteriorly, and many small denticulae posteriorly. Scape robust, not attaining the posterior border of the head; funiculus clavate, the terminal segment almost, or about as long as the three preceding segments together. Thorax laterally compressed, moderately convex dorsally, wider anteriorly than posteriorly; pro-mesonotal and meso-epinotal sutures distinct. Epinotal declivity faintly concave with weakly marginate sides, meeting the base in a well rounded obtuse angle. Petiole thick antero-posteriorly, scarcely narrowed dorsally, convex anteriorly, concave posteriorly, beneath with a prominent rectangular tooth, the front of which bears a spiracular opening, the posterior of which terminates in a rather sharp tooth. Tibiae of legs each with a well developed pectinated spur.

Mandibles smooth and shining, with sparse, scattered punctures. Head and thorax subopaque, covered with dense and coarse, but not confluent, punctures; petiole and gaster more shining, finely and densely punctate or shagreened.

Entire body covered with rather dense, yellowish gray, appressed pubescence. Erect hairs on the head, thorax, petiole and gaster; most numerous on the latter. Hairs longest on the anterior and posterior ends of the body.

Typical specimens black; mandibles, clypeus, frontal carinae, antennae, legs, and tip of gaster reddish or yellowish. Occasionally individuals are reddish brown with varying amounts of infuscation.

Queen
Smith (1936) - Length: 4 mm. Resembling the worker in general appearance but larger and more robust. Head with large compound eyes which are placed slightly less than their greatest diameter from the base of the mandibles. Vertex with three ocelli in the form of a triangle. Petiole smaller, and more slender (anterio-posterially) than with the worker. Wings hyaline, ciliated, with yellowish veins and darker stigma; with one submarginal and a discoidal cell.

Taylor (1967) - 17 queens from all parts of the distributional range have the following measurements: HL 0.37-0.41 mm; HW 0.30-0.35 mm; SL 0.24-0.29 mm; CI 82-86; SI 80-84; PW 0.27-0.31 mm; PNL 0.12-0.15 mm; PH 0.28-0.31 mm; DPW 0.22-0.25 mm; PNI 76-80; maximum diameter of compound eye 0.09-0.11 mm; ocular index 27-31. Palpal formula (2 specimens dissected): Maxillary 2: Labial 2, as in the worker. Wing venation as in P. coarctata. Differing from the workers in the usual characters of full sexuality. Similar in general to P. coarctata, but larger in size. There is no overlap in the ranges for PW, PH, DPW, SI and PNI in queens of these 2 species. The petiolar node is distinctly broader in pennsylvanica (compare the PNI values), and thinner when viewed from above (cf. figs. 3 and 4). The sculpturation is much more heavily developed than in queens of coarctata; the diagnostically important sculptural characters being: mandibles, clypeus, frons, and scapes as in worker. Entire mesosoma opaque, with a close cover of punctures about 0.01 mm in diameter. Node and gaster feebly shining, with a finer more scattered puncturation than mesosoma-the punctae of the disc of the first gastric segment, for example, about 0.007 mm in diameter, separated by distances about equal to 1.5 to 2.5 X their maximum diameter. Color as in worker.

Male
Taylor (1967) - HL 0.60-0.63 mm; HW (across eyes) 0.64-0.70 mm; CI 107-113; WL 1.04-1.22 mm; PNL 0.20-0.23 mm; PH. 0.35-0.41 mm; DPW 0.28-0.31 mm; maximum diameter of eye 0.27-0.32 mm; ocular index 43-46; palpal formula (6 specimens dissected): Maxillary 5: Labial 3, as in P. coarctata. Wing venation as in queen. Color medium to rather dark brown; eyes black; clypeus, mandibles, legs and gastric apex lightly infuscated; wing veins yellowish.

Immature Forms
Taylor (1967) - The larva has 4 pairs of dorsal abdominal "mushroom" tubercles. A detailed description with figures has been published by G. C. & J. Wheeler (1952). The pupae of all castes are enclosed in cocoons.

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.
 * Banschbach V. S., and E. Ogilvy. 2014. Long-term Impacts of Controlled Burns on the Ant Community (Hymenoptera: Formicidae) of a Sandplain Forest in Vermont. Northeastern Naturalist 21(1): 1-12.
 * 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.
 * Buren W. F. 1944. A list of Iowa ants. Iowa State College Journal of Science 18:277-312
 * Campbell J. W., S. M. Grodsky, D. A. Halbritter, P. A. Vigueira, C. C. Vigueira, O. Keller, and C. H. Greenberg. 2019. Asian needle ant (Brachyponera chinensis) and woodland ant responses to repeated applications of fuel reduction methods. Ecosphere 10(1): e02547.
 * Campbell K. U., and T. O. Crist. 2017. Ant species assembly in constructed grasslands isstructured at patch and landscape levels. Insect Conservation and Diversity doi: 10.1111/icad.12215
 * Canadensys Database. Dowloaded on 5th February 2014 at http://www.canadensys.net/
 * Carrias E. S. 2009. A comparison of leaf litter ants in (Hymenoptera: Formicidae) the pine forests of North and Central America. Master's Thesis, Faculty of the Department Biological Sciences, Sam Houston State University, 73 pages.
 * Carroll T. M. 2011. The ants of Indiana (Hymenoptera: Formicidae). Master's Thesis Purdue university, 385 pages.
 * Choate B., and F. A. Drummond. 2012. Ant Diversity and Distribution (Hymenoptera: Formicidae) Throughout Maine Lowbush Blueberry Fields in Hancock and Washington Counties. Environ. Entomol. 41(2): 222-232.
 * Choate B., and F. A. Drummond. 2013. The influence of insecticides and vegetation in structuring Formica Mound ant communities (Hymenoptera: Formicidae) in Maine lowbush blueberry. Environ. Entomol. 41(2): 222-232.
 * Clark A. T., J. J. Rykken, and B. D. Farrell. 2011. The Effects of Biogeography on Ant Diversity and Activity on the Boston Harbor Islands, Massachusetts, U.S.A. PloS One 6(11): 1-13.
 * Clark Adam. Personal communication on November 25th 2013.
 * Cokendolpher J. C., and O. F. Francke. 1990. The ants (Hymenoptera, Formicidae) of western Texas. Part II. Subfamilies Ecitoninae, Ponerinae, Pseudomyrmecinae, Dolichoderinae, and Formicinae. Special Publications, the Museum. Texas Tech University 30:1-76.
 * Colby, D. and D. Prowell. 2006. Ants (Hymenoptera: Formicidae) in Wet Longleaf Pine Savannas in Louisiana. Florida Entomologist 89(2):266-269
 * Cole A. C., Jr. 1949. The ants of Mountain Lake, Virginia. Journal of the Tennessee Academy of Science 24: 155-156.
 * Coovert G. A. 2005. The Ants of Ohio (Hymenoptera: Formicidae). Ohio Biological Survey, Inc. 15(2): 1-207.
 * Coovert, G.A. 2005. The Ants of Ohio (Hymenoptera: Formicidae) Ohio Biological Survey Bulletin New Series Volume 15(2):1-196
 * Dattilo W. et al. 2019. MEXICO ANTS: incidence and abundance along the Nearctic-Neotropical interface. Ecology https://doi.org/10.1002/ecy.2944
 * 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. 1998. Smithistruma memorialis (Hymenoptera: Formicidae), a new species of ant from the Kentucky Cumberland Plateau. Entomological News 109: 81-87.
 * Deyrup, M. 2003. An updated list of Florida ants (Hymenoptera: Formicidae). Florida Entomologist 86(1):43-48.
 * Drummond F. A., A. M. llison, E. Groden, and G. D. Ouellette. 2012. The ants (Formicidae). In Biodiversity of the Schoodic Peninsula: Results of the Insect and Arachnid Bioblitzes at the Schoodic District of Acadia National Park, Maine. Maine Agricultural and forest experiment station, The University of Maine, Technical Bulletin 206. 217 pages
 * DuBois M. B. 1981. New records of ants in Kansas, III. State Biological Survey of Kansas. Technical Publications 10: 32-44
 * DuBois M. B. 1985. Distribution of ants in Kansas: subfamilies Ponerinae, Ecitoninae, and Myrmicinae (Hymenoptera: Formicidae). Sociobiology 11: 153-227
 * DuBois M. B. 1985. Distribution of ants in Kansas: subfamilies Ponerinae, Ecitoninae, and Myrmicinae (Hymenoptera: Formicidae). Sociobiology 11: 153-228
 * DuBois M. B. 1985. Distribution of ants in Kansas: subfamilies Ponerinae, Ecitoninae, and Myrmicinae (Hymenoptera: Formicidae). Sociobiology 11: 153-229
 * DuBois M. B. 1985. Distribution of ants in Kansas: subfamilies Ponerinae, Ecitoninae, and Myrmicinae (Hymenoptera: Formicidae). Sociobiology 11: 153-230
 * DuBois M. B. 1985. Distribution of ants in Kansas: subfamilies Ponerinae, Ecitoninae, and Myrmicinae (Hymenoptera: Formicidae). Sociobiology 11: 153-232
 * DuBois M. B. 1985. Distribution of ants in Kansas: subfamilies Ponerinae, Ecitoninae, and Myrmicinae (Hymenoptera: Formicidae). Sociobiology 11: 153-233
 * DuBois M. B. 1985. Distribution of ants in Kansas: subfamilies Ponerinae, Ecitoninae, and Myrmicinae (Hymenoptera: Formicidae). Sociobiology 11: 153-234
 * DuBois M. B. 1985. Distribution of ants in Kansas: subfamilies Ponerinae, Ecitoninae, and Myrmicinae (Hymenoptera: Formicidae). Sociobiology 11: 153-235
 * DuBois M. B. 1985. Distribution of ants in Kansas: subfamilies Ponerinae, Ecitoninae, and Myrmicinae (Hymenoptera: Formicidae). Sociobiology 11: 153-236
 * DuBois M. B. 1985. Distribution of ants in Kansas: subfamilies Ponerinae, Ecitoninae, and Myrmicinae (Hymenoptera: Formicidae). Sociobiology 11: 153-237
 * DuBois M. B. 1985. Distribution of ants in Kansas: subfamilies Ponerinae, Ecitoninae, and Myrmicinae (Hymenoptera: Formicidae). Sociobiology 11: 153-238
 * DuBois M. B. 1985. Distribution of ants in Kansas: subfamilies Ponerinae, Ecitoninae, and Myrmicinae (Hymenoptera: Formicidae). Sociobiology 11: 153-239
 * DuBois M. B. 1985. Distribution of ants in Kansas: subfamilies Ponerinae, Ecitoninae, and Myrmicinae (Hymenoptera: Formicidae). Sociobiology 11: 153-240
 * DuBois M. B. 1985. Distribution of ants in Kansas: subfamilies Ponerinae, Ecitoninae, and Myrmicinae (Hymenoptera: Formicidae). Sociobiology 11: 153-241
 * DuBois M. B. 1985. Distribution of ants in Kansas: subfamilies Ponerinae, Ecitoninae, and Myrmicinae (Hymenoptera: Formicidae). Sociobiology 11: 153-242
 * DuBois M. B. 1985. Distribution of ants in Kansas: subfamilies Ponerinae, Ecitoninae, and Myrmicinae (Hymenoptera: Formicidae). Sociobiology 11: 153-243
 * DuBois M. B. 1985. Distribution of ants in Kansas: subfamilies Ponerinae, Ecitoninae, and Myrmicinae (Hymenoptera: Formicidae). Sociobiology 11: 153-244
 * DuBois M. B. 1985. Distribution of ants in Kansas: subfamilies Ponerinae, Ecitoninae, and Myrmicinae (Hymenoptera: Formicidae). Sociobiology 11: 153-245
 * DuBois M. B. 1985. Distribution of ants in Kansas: subfamilies Ponerinae, Ecitoninae, and Myrmicinae (Hymenoptera: Formicidae). Sociobiology 11: 153-246
 * DuBois M. B. 1985. Distribution of ants in Kansas: subfamilies Ponerinae, Ecitoninae, and Myrmicinae (Hymenoptera: Formicidae). Sociobiology 11: 153-247
 * DuBois M. B. 1985. Distribution of ants in Kansas: subfamilies Ponerinae, Ecitoninae, and Myrmicinae (Hymenoptera: Formicidae). Sociobiology 11: 153-248
 * DuBois M. B. 1985. Distribution of ants in Kansas: subfamilies Ponerinae, Ecitoninae, and Myrmicinae (Hymenoptera: Formicidae). Sociobiology 11: 153-249
 * DuBois M. B. 1985. Distribution of ants in Kansas: subfamilies Ponerinae, Ecitoninae, and Myrmicinae (Hymenoptera: Formicidae). Sociobiology 11: 153-250
 * DuBois M. B. 1985. Distribution of ants in Kansas: subfamilies Ponerinae, Ecitoninae, and Myrmicinae (Hymenoptera: Formicidae). Sociobiology 11: 153-251
 * DuBois M. B. 1985. Distribution of ants in Kansas: subfamilies Ponerinae, Ecitoninae, and Myrmicinae (Hymenoptera: Formicidae). Sociobiology 11: 153-252
 * DuBois M. B. 1985. Distribution of ants in Kansas: subfamilies Ponerinae, Ecitoninae, and Myrmicinae (Hymenoptera: Formicidae). Sociobiology 11: 153-253
 * DuBois M. B. 1985. Distribution of ants in Kansas: subfamilies Ponerinae, Ecitoninae, and Myrmicinae (Hymenoptera: Formicidae). Sociobiology 11: 153-254
 * DuBois M. B. 1985. Distribution of ants in Kansas: subfamilies Ponerinae, Ecitoninae, and Myrmicinae (Hymenoptera: Formicidae). Sociobiology 11: 153-255
 * DuBois M. B. 1985. Distribution of ants in Kansas: subfamilies Ponerinae, Ecitoninae, and Myrmicinae (Hymenoptera: Formicidae). Sociobiology 11: 153-256
 * DuBois M. B. 1985. Distribution of ants in Kansas: subfamilies Ponerinae, Ecitoninae, and Myrmicinae (Hymenoptera: Formicidae). Sociobiology 11: 153-257
 * DuBois M. B. 1985. Distribution of ants in Kansas: subfamilies Ponerinae, Ecitoninae, and Myrmicinae (Hymenoptera: Formicidae). Sociobiology 11: 153-258
 * DuBois M. B. 1985. Distribution of ants in Kansas: subfamilies Ponerinae, Ecitoninae, and Myrmicinae (Hymenoptera: Formicidae). Sociobiology 11: 153-259
 * 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., and E. J. Farnsworth. 2014. Targeted sampling increases knowledge and improves estimates of ant species richness in Rhode Island. Northeastern Naturalist 21(1): NENHC-13NENHC-24.
 * Emery C. 1895. Beiträge zur Kenntniss der nordamerikanischen Ameisenfauna. (Schluss). Zoologische Jahrbücher. Abteilung für Systematik, Geographie und Biologie der Tiere 8: 257-360.
 * Fernandes, P.R. XXXX. Los hormigas del suelo en Mexico: Diversidad, distribucion e importancia (Hymenoptera: Formicidae).
 * Forster J.A. 2005. The Ants (hymenoptera: Formicidae) of Alabama. Master of Science, Auburn University. 242 pages.
 * Francoeur, A. 1979. Le fourmis du Quebec. Annales de la Societe Entomologique du Quebec 24(1):48-64.
 * 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., 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. 2008. New Distributional Records of Ants in Arkansas for 2008. Journal of the Arkansas Academy of Science. 63: 182-184
 * Gibbs M. M., P. L. Lambdin, J. F. Grant, and A. M. Saxton. 2003. Ground-inhabiting ants collected in a mixed hardwood southern Appalachian forest in Eastern Tennessee. Journal of the Tennessee Academy of Science 78(2): 45-49.
 * 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.
 * Headley A. E. 1943. The ants of Ashtabula County, Ohio (Hymenoptera, Formicidae). The Ohio Journal of Science 43(1): 22-31.
 * 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.
 * 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.
 * Ivanov K., and J. Keiper. 2010. Ant (Hymenoptera: Formicidae) diversity and community composition along sharp urban forest edges. Biodivers Conserv 19: 39173933.
 * Johnson C. 1986. A north Florida ant fauna (Hymenoptera: Formicidae). Insecta Mundi 1: 243-246
 * Kittelson P. M., M. P. Priebe, and P. J. Graeve. 2008. Ant Diversity in Two Southern Minnesota Tallgrass Prairie Restoration Sites. Jour. Iowa Acad. Sci. 115(14): 2832.
 * 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.
 * Longino J. T., and D. B. Booher. 2019. Expect the unexpected: a new ant from a backyard in Utah. Western North American Naturalist 79(4): 496–499.
 * Lopez R., and D. A. Potter. 2003. Biodiversity of ants (Hymenoptera: Formicidae) in golf course and lawn turf habitats in Kentucky. Sociobiology 42(3): 701-713.
 * 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., Hill J.G. and Skvarla M. 2011. New Records of Ants (Hymenoptera: Formicidae) for Arkansas with a Synopsis of Previous Records. Midsouth Entomologist. 4: 29-38
 * 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 Anderson R.S. 1991.New Distributional Records for the Ant Genus Ponera (Hymenoptera: Formicidae)
 * Macgown J. A., S. Y. Wang, J. G. Hill, and R. J. Whitehouse. 2017. A List of Ants (Hymenoptera: Formicidae) Collected During the 2017 William H. Cross Expedition to the Ouachita Mountains of Arkansas with New State Records. Transactions of the American Entomological Society, 143(4): 735-740.
 * Mackay W. P., and E. E. Mackay. 2002. The ants of New Mexico (Hymenoptera: Formicidae). Lewiston, New York: Edwin Mellen Press, 400 pp.
 * Mackay, W.P. and E. Mackay. XXXX. The Ants of New Mexico
 * 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.
 * 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.
 * Ouellette G. D. and A. Francoeur. 2012. Formicidae [Hymenoptera] diversity from the Lower Kennebec Valley Region of Maine. Journal of the Acadian Entomological Society 8: 48-51
 * Ouellette G. D., F. A. Drummond, B. Choate and E. Groden. 2010. Ant diversity and distribution in Acadia National Park, Maine. Environmental Entomology 39: 1447-1556
 * Procter W. 1938. Biological survey of the Mount Desert Region. Part VI. The insect fauna. Philadelphia: Wistar Institute of Anatomy and Biology, 496 pp.
 * Santschi F. 1911. Formicides récoltés par Mr. le Prof. F. Silvestri aux Etats Unis en 1908. Bullettino della Società Entomologica Italiana 41: 3-7.
 * Shik, J., A. Francoeur and C. Buddle. 2005. The effect of human activity on ant species (Hymenoptera: Formicidae) richness at the Mont St. Hilaire Biosphere Reserve, Quebec. Canadian Field-Naturalist 119(1): 38-42.
 * Smith M. R. 1924. An annotated list of the ants of Mississippi (Hym.). Entomological News 35: 47-54.
 * Smith M. R. 1936. Ants of the genus Ponera in America, north of Mexico. Annals of the Entomological Society of America 29: 420-430.
 * Smith M. R. 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.
 * Taylor R. W. 1967. A monographic revision of the ant genus Ponera Latreille (Hymenoptera: Formicidae). Pacific Insects Monograph 13: 1-112.
 * 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. 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.
 * Van Pelt, A. 1983. Ants of the Chisos Mountains, Texas (Hymenoptera: Formicidae) . Southwestern Naturalist 28:137-142.
 * Vásquez-Bolaños M. 2011. Lista de especies de hormigas (Hymenoptera: Formicidae) para México. Dugesiana 18: 95-133
 * 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
 * Wesson L. G., and R. G. Wesson. 1939. Notes on Strumigenys from southern Ohio, with descriptions of six new species. Psyche (Cambridge) 46: 91-112.
 * 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. 1900. The habits of Ponera and Stigmatomma. Biological Bulletin (Woods Hole). 2: 43-69.
 * 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. 1906. Fauna of New England. 7. List of the Formicidae. Occasional Papers of the Boston Society of Natural History 7: 1-24
 * Wheeler W. M. 1908. The ants of Casco Bay, Maine, with observations on two races of Formica sanguinea Latreille. Bulletin of the American Museum of Natural History 24: 619-645.
 * Wheeler W. M. 1913. Ants collected in Georgia by Dr. J. C. Bradley and Mr. W. T. Davis. Psyche (Cambridge) 20: 112-117.
 * Wheeler W. M. 1928. Ants of Nantucket Island, Mass. Psyche (Cambridge) 35: 10-11.
 * Wheeler W. M. 1931. Concerning some ant gynandromorphs. Psyche (Cambridge) 38: 80-85.
 * 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
 * Wing M. W. 1939. An annotated list of the ants of Maine (Hymenoptera: Formicidae). Entomological News 50:161-165.
 * 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