Camponotus pennsylvanicus

This species is the first North American ant to be described. It nests in living and dead trees, rotten logs or stumps in forested areas. This is an important, destructive pest that attacks fences, poles and buildings. This is probably the most destructive carpenter ant in North America (Wheeler and Wheeler 1963), although Creighton (1950) argued that its destructive capacities were somewhat exaggerated and that they only tunnel in decayed wood. It often forages inside homes, making it an important house pest. Reproductives were found in nests from April to October. This species is found in the same logs and stumps as members of the genera Lasius, Formica (i. e. Formica podzolica), and Leptothorax. Workers tend aphids, with the smaller workers collecting honeydew and transferring it to larger workers that carry it back to the nest. In addition, foragers feed on dead insects and plant juices.

Compare with Camponotus chromaiodes, Camponotus modoc, Camponotus herculeanus, Camponotus novaeboracensis, Camponotus schaefferi, Camponotus texanus.

The majors, minors, females and males of C. pennsylvanicus are black ants without erect setae on the cheeks or sides of the head, or posterior lateral corners, the scapes are without erect and suberect setae, except at the apex, but the mesosoma has numerous setae on the dorsal surface, as does the petiole. The gaster has erect and suberect setae scattered over the entire surface, the tibiae are without erect and suberect setae, except for 2 rows along the flexor surface, in which most setae are present along the distal half. Appressed pubescence is sparse on most surfaces, including the head and mesosoma, although the setae on the dorsum are more numerous, but non-overlapping or few overlap adjacent setae, the side of the propodeum has a few appressed setae.

The appressed or decumbent setae on the gaster are long (over 0.2 mm) and abundant, nearly all setae overlap adjacent setae and are nearly as long as the erect and suberect setae on the gaster. The appressed setae on the gaster of the minors and especially the females may be small and may not overlap adjacent setae.

Comparisons
Camponotus pennsylvanicus is a member of a small group of 4 similar species (together with Camponotus modoc, Camponotus herculeanus and Camponotus chromaiodes), which are difficult to separate. It has been shown to be genetically similar to C. modoc (Sämi Schär, et al., 2018). Majors of C. pennsylvanicus may be separated from those of Camponotus modoc (S Canada, mostly W US and N Mexico) as the appressed setae on the gaster of C. pennsylvanicus are nearly as long or as long as the erect and suberect setae (usually over 0.15 mm), whereas these setae on Camponotus modoc are generally shorter than ½ the length of the erect and suberect setae (usually less than 0.15 mm). The appressed setae on the gaster of C. pennsylvanicus overlap adjacent setae, whereas they rarely do in C. modoc. Unfortunately, the length of the appressed setae in these two species varies geographically, with the shortest setae being found at lower latitudes and higher longitude (west and north), the longer setae found at higher latitudes and lower longitudes (south and east, Fig. 256), further complicating the separation of the two species. The gradient suggests that they are both the same species, but as members of a single colony have appressed setae of similar lengths, and both species can be found at specific localities therefore they will be considered two species, until other evidence becomes available.

Camponotus pennsylvanicus can also be confused with Camponotus herculeanus (S Canada, US), but is nearly always concolorous black, whereas Camponotus herculeanus generally has a deep red mesosoma, or at least deep red legs. Along the western edge of the distribution (Wyoming south to northern New Mexico) specimens usually have red legs, which may be evidence of hybridization. The scape of the major of C. pennsylvanicus extends 1 - 2 funicular segments past the posterior lateral corner of the head, whereas the scape of majors of C. herculeanus rarely reaches the posterior lateral corner or extends less than 1 funicular segment past it. The appressed setae on the gaster are much denser in C. pennsylvanicus than in C. novaeboracensis. A single major from Ames, Iowa has short scapes (as in C. herculeanus) and long appressed pubescence, typical of C. pennsylvanicus. It could be a hybrid but is considered a member of the latter species.

Camponotus pennsylvanicus and Camponotus chromaiodes co-occur in Douglasville, Pennsylvania, suggesting that they are valid species. The majors appear to differ only in color, with the anterior edge of the gaster of C. pennsylvanicus being black instead of reddish as in C. chromaiodes.

The minors of the 4 species are very difficult to separate. The minors of Camponotus pennsylvanicus are usually concolorous black, whereas at least the legs of Camponotus modoc and Camponotus herculeanus are often dark red, and the minor of C. chromaiodes is bicolored, black with a red first tergum of the gaster, and the mesosoma is partially or even completely red. The appressed setae of C. modoc and C. herculeanus are usually shorter than they are in C. pennsylvanicus and C. chromaiodes.

The females of the 4 species often cannot be reliably separated as the appressed setae on the gaster of C. pennsylvanicus are often short and similar to those of C. modoc and C. herculeanus. The females of Camponotus pennsylvanicus are concolorous black, whereas the legs of the other 2 species are occasionally dark red. The female of C. chromaiodes is dark, but at least the anterior edge of the first gastral tergum is red, and the lower part of the mesosoma is often red. The scapes extend about the same distance past the posterior lateral corners of the head of the female in all 4 species (about 2 funicular segments past the posterior border of the head).

The males of the 3 species apparently cannot be distinguished and in general cannot be separated from most other species of Camponotus.

The clypeus of C. pennsylvanicus has weakly developed angles along the anterior border, similar to those of Camponotus schaefferi (AZ, NM) and Camponotus texanus (W TX). It can be easily separated as the side of the mesosoma is dull and punctate, not nearly smooth and glossy as in the latter two species.

Camponotus pennsylvanicus can be confused with Camponotus novaeboracensis (S Canada, US). They can be separated as the workers and often the females of C. novaeboracensis are bicolored, whereas they are nearly always concolorous black in C. pennsylvanicus. Questionable specimens can be separated as the setae on the gaster of workers and of the females of C. pennsylvanicus are longer and most overlap adjacent setae, whereas in C. novaeboracensis they are relatively short, with few or no appressed setae overlapping adjacent setae. Unfortunately, neither of these characteristics serve to separate the males, and there are no obvious characters which will separate them. Camponotus pennsylvanicus is sympatric with C. novaeboracensis in Iowa with no integration which suggests they are separate species (Buren, 1944).

Camponotus pennsylvanicus is founds in areas with clay or sandy brown soils. It is found in a variety of plant communities ranging from urban habitats, desert scrub with few creosote bushes, bocage farmland, grasslands, pine/spruce surrounded by grassland, oak dominated forest with palmetto understory, riparian weedy areas,  riparian hardwood forests, deciduous forest, bottomland forest, mixed hardwood forest, mixed hardwood/conifer forest, ponderosa pine with few pinyon pines, second growth pines/hardwoods, oak woodland, hardwood/oak woodlands,  oak evergreen forest wetlands, pine, elm cottonwood, larch forests and beach/conifer forest.

It is a habitat generalist that remains in urban forests in Canada (Thompson and McLachlan, 2007) and is found in urban habitat (Guénard et al., 2015). Wheeler and Wheeler (1963) and Oberg (2012) reported it from wooded areas. It is also found in open areas and deciduous and mixed forest (MacGown and Brown, 2006), including witch-hazel-red maple forests (Yitbarek et al., 2011), as well as grasslands and shrublands (Barber, 2015).

It is also found in marsh/emergent wetland, pocosin, swamps and bottomland hardwood, recently cleared land, upland pine, pine woodland/longleaf pine savanna, maritime forest, upland mixed forest, and grassland (Davis, 2009).

Distribution based on Regional Taxon Lists
Nearctic Region: Canada, United States. Neotropical Region: Bermuda.

Camponotus pennsylvanicus, the black carpenter ant, was the first North American ant to be described. Smith (1965) summarized what had been published and supplemented the summary with personal observations. He concluded it is one of our best known and most adaptable ants.

It nests almost exclusively in wood. Camponotus pennsylvanicus nests in many different species of trees, as well as in telephone poles; one nest was 75 feet up in a tree. Single females form their nests in preformed or self-formed cavities, usually under bark of a log or stump, and each nest has a single queen. There are approximately 2000 or more workers in a three to six-year-old nests. Development takes 60 days. Nuptial flights from overwintering sexuals occur from May to late July. Foragers are diurnal and nocturnal, feeding on living and dead insects. They collect honeydew from plant lice and treehoppers, as well as the juices of fruits and sap of certain plants. Forgers commonly feed on foods found in kitchens. It produces a lot of damage to wooden structures and destroys food in houses, especially when it is brought into houses with firewood. Camponotus pennsylvanicus is parasitized by at least two species of phorid flies, and three species of fungi.

Other researchers and our own experience add details to the biology of Camponotus pennsylvanicus. This is the common black carpenter ant, which is fond of nesting in logs, stumps, trees, etc. in South Carolina (Smith, 1934). It nests in living and dead trees, in /under bark of logs under rotten logs or in stumps, under bark of dead trees, in living cottonwood trees and rarely in soil under stones, as well as wooden structures ranging from telephone poles to buildings. It nests in rotten logs and in soil at the base of Quercus pagoda (MacGown and Brown, 2006).

It prefers mature hardwood (Quercus spp.) trees approximately 30 cm in diameter and pine logs (Pinus spp.) that were approximately 9 m in length. Defects in trees most often associated with nests included tree holes and crotches (Oswalt, 2007). One nest was in stack of newspapers in a box in a garage, two were from building walls (Wheeler and Wheeler, 1963).

Camponotus pennsylvanicus is unable to actively regulate its internal nest environment but can use the insulative properties of trees to dampen wide temperature fluctuations and provide a more stable nest microclimate (Oswalt, 2007). The internal nest temperature of occupied nests mirrored unoccupied nests throughout the year and differences were not statistically significant (Oswalt, 2007). In warm sites, C. pennsylvanicus tends to occupy relatively cool chambers (Diamond et al., 2012). They seal the nest during the winter, lose half of the fat reserves during the winter and fill their bodies with glycerol as antifreeze, freezing at temperatures as low as -22°C (Ellison et al., 2012).

Mature colonies can be large with 2000-2500 adults (Hölldobler and Wilson 1990) and may reach 10,000-15,000 workers (Hansen and Klotz 2005; Ellison et al., 2012) and are facultatively or weakly polydomous (Buczkowski, 2011).

Camponotus pennsylvanicus brood were found in nests from April to August; sexuals from April to October (Mackay and Mackay, 2002). Reproductives overwinter in nests (Buren, 1944).

Flights of C. pennsylvanicus occur in May, and sexuals can be captured in black light traps. Females start nests in or under logs, or under bark (Buren, 1944) in May and July; occasionally more than a single female may be found together.

Sanders (1972) studied foraging in C. pennsylvanicus and found that the start of seasonal activity was temperature dependent (Stuble et al., 2014), and activity peaked in mid-summer. Later in the season they became virtually nocturnal or are in general primarily nocturnal (Hansen and Klotz, 2005) when the weather becomes too hot (Klotz, 1984). Nuss et al. (2005) report that most foraging of C. pennsylvanicus is nocturnal and crepuscular.

The black carpenter ant tends honeydew producers and scavenges for dead insects and plant material and can be captured in surface baits. They are honeydew feeders (Oberg, 2012), tending aphids (Jones, 1929) and the calico scale Eulecanium cerasorum (Vanek and Potter, 2010). Cannon and Fell (2002) report on the macronutrients collected by foragers of Camponotus pennsylvanicus under natural conditions. Trophallaxis boosts disease resistance within the colony (Hamilton et al., 2010).

Oswald (2007) completed a series of feeding experiments. Camponotus pennsylvanicus selected a higher concentration of sucrose as the distance from a food patch to the nest increased to 15 m. Foragers imbibed sucrose from both concentrations at 1 m with no preference noted between the two solutions. Mean feeding durations indicated that foraging black carpenter ants fed deferentially on casein solutions, depending on concentration or distance. Overall casein mean feeding time was significant, suggesting C. pennsylvanicus feeds longer on 30% casein solution regardless of the distance involved. Additional analysis indicated that the effect of distance on feeding duration was only significant at a 5% solution concentration. Overall mean feeding time was significant, suggesting feeding intervals were greater on 30% casein over both distances (Oswalt, 2007).

Camponotus pennsylvanicus is important in seed removal (Stuble et al., 2014) and preys on the red oak borer Enaphalodes rufulus (Coleoptera: Cerambycidae) (Muilenburg et al., 2008; Verble and Stephen, 2009).

Foragers are aggressive (Stuble et al., 2014), and patrol bracken fern (Pteridium aquilinum) during the sensitive crozier growth stage (Oberg, 2012) and remove herbivorous insects from rapidly expanding fronds (Oldenkamp and Douglas, 2011). They removed seeds from bait stations (Warren et al., 2010). Scout ants recruit nestmates to a new food source using alerting motor displays (Traniello, 1977). The nest mates follow a recruitment trail that is apparently composed of materials from the hindgut and poison gland (formic acid). The pheromone is produced by the hindgut and the trails consist of streaks, that are not reinforced by gorged workers passing over them (Hartwick et al., 1977). They show topochemical learning in which the foragers navigate using preexisting chemicals of plant substrates (Helmy and Jander, 2003). When a strong direct light is available, the workers use light orientation and stop making trails (Hartwick et al., 1977).

Camponotus pennsylvanicus is a behaviorally dominant ant (Oberg, 2012), and the most common carpenter ant in New England (Ellison et al., 2012). There was a strong negative association between the congeners Camponotus novaeboracensis and C. pennsylvanicus (Thompson and McLachlan, 2007). The proportion of trees colonized by C. pennsylvanicus was significantly higher in urban plots (with fewer trees), suggesting that intraspecific competition for nesting sites may be especially high in areas with lower tree density (Buczkowski, 2011). Del Toro et al. (2013) found it to be a neutrally interacting species found mainly in forested plots.

Camponotus pennsylvanicus is the host of the endosymbiotic proteobacteria Candidatus Blochmannia (Sauer et al., 2000; Degnan et al., 2004), which has close relatives that occur in aphids and the tsetse fly (Sauer et al., 2000). The bacteria apparently lack abilities to replicate outside of host cells, and the insect hosts depend on endosymbiont functions that are often nutritional in nature (Wernegreen, 2015).

They carry phoretic mites (Campbell et al., 2013), are an alternate host for Nemadus triangulum (Coleoptera: Leiodidae) and a minor host for N. parasitus (Peck and Cook, 2007). Reemer (2012) discusses the parasitism by the syrphid fly parasite Microdon sp. and C. pennsylvanicus is specifically the host of Microdon cothurnatus (Duffield, 1981). They are not infected by the fungus Ophiocordyceps unilateralis sensu lato, whereas Camponotus americanus and Camponotus castaneus were (de Bakker et al., 2014).

Camponotus pennsylvanicus is one of the major ant pests in the United States (Santos, 2016) and one of the four most common house infesting ants in Illinois (Walker, 2013). Hansen and Klotz (2005) conclude that it is the principal structural pest in eastern North America. It causes damage to buildings and forages in homes, and also attacks fences, poles and buildings (Wheeler, 1905a). Buren (1944) found it in beams of frame houses. The nest is often outside 200-300 meters away from a home so it is not easily exterminated (Ellison et al., 2012). This species may bite but generally avoids confrontation (Forester, 2003).

Additional Notes
In southern New England this species is one of the most abundant and adaptable species across a wide range of forest habitats.

Wheeler (1910) - C. pennsylvanicus ranges over southern Canada and the states as far west as Victoria, Texas, and the Pine Ridge Indian Reservation in South Dakota. Unlike whymperi and modoc it is a lowland form. It is far and away the most abundant Camponotus in the North Atlantic states and Middle West, showing a range of adaptability to differences in temperature and humidity second only to that of Lasius americanus and Formica subsericea. On this account, it is the only one of our Camponoti that has attracted general attention. It is commonly found nesting in old logs and stumps or in the dead wood of standing trees, but occasionally it nests in old houses. In such places it may do considerable damage by tunneling in beams and rafters and may become a domestic nuisance by visiting the sweet food-stuffs in kitchens and pantries. Its habits were first studied by McCook. Recently Pricer and Miss Edith Buckingham have made many interesting observations on its polymorphism and behavior.

Gripshover et al. (2018) examined the mechanics of how workers of this species are able to move through water.

Morphology
Draft et al. (2018) - A study of antenna olfactory behavior. Abstract:Directed and meaningful animal behavior depends on the ability to sense key features in the environment. Among the different environmental signals, olfactory cues are critically important for foraging, navigation and social communication in many species, including ants. Ants use their two antennae to explore the olfactory world, but how they do so remains largely unknown. In this study, we used high-resolution videography to characterize the antennae dynamics of carpenter ants (Camponotus pennsylvanicus). Antennae are highly active during both odor tracking and exploratory behavior. When tracking, ants used several distinct behavioral strategies with stereotyped antennae sampling patterns (which we call 'sinusoidal', 'probing' and 'trail following'). In all behaviors, left and right antennae movements were anti-correlated, and tracking ants exhibited biases in the use of left versus right antenna to sample the odor trail. These results suggest non-redundant roles for the two antennae. In one of the behavioral modules (trail following), ants used both antennae to detect trail edges and direct subsequent turns, suggesting a specialized form of tropotaxis. Lastly, removal of an antenna resulted not only in less accurate tracking but also in changes in the sampling pattern of the remaining antenna. Our quantitative characterization of odor trail tracking lays a foundation to build better models of olfactory sensory processing and sensorimotor behavior in terrestrial insects.

Nomenclature

 * . Formica pensylvanica De Geer, 1773: 603, pl. 31, figs. 9, 10 (s.w.q.m.) U.S.A. (Pennsylvania).
 * [Justified emendation of spelling to pennsylvanica: Buckley, 1866: 155.]
 * Wheeler, G.C. & Wheeler, J. 1953e: 187 (l.).
 * Combination in Camponotus: Mayr, 1862: 666;
 * combination in C. (Camponotus): Forel, 1914a: 266.
 * Subspecies of herculeanus: Forel, 1879a: 57; Provancher, 1881b: 355; Mayr, 1886d: 420; Cresson, 1887: 256; Emery, 1893i: 675; Emery, 1895k: 477; Emery, 1896d: 372 (in list); Wheeler, W.M. 1900c: 47; Viereck, 1903: 72; Wheeler, W.M. 1905f: 402; Wheeler, W.M. 1906b: 22; Wheeler, W.M. 1910d: 335 (redescription); Wheeler, W.M. 1910g: 571; Santschi, 1911d: 7; Wheeler, W.M. 1913c: 117; Forel, 1914a: 266; Donisthorpe, 1915d: 348; Wheeler, W.M. 1916m: 600; Wheeler, W.M. 1917i: 465; Emery, 1925b: 73; Donisthorpe, 1927b: 401; Karavaiev, 1929b: 211; Wheeler, W.M. 1932a: 13; Dennis, 1938: 301; Wing, 1939: 163; Wesson, L.G. & Wesson, R.G. 1940: 103; Buren, 1944a: 293; Wheeler, G.C. & Wheeler, E.W. 1944: 251; Smith, M.R. 1951a: 840.
 * Status as species: Retzius, 1783: 75; Olivier, 1792: 501; Latreille, 1802c: 99; Lepeletier de Saint-Fargeau, 1835: 213; Smith, F. 1858b: 53; Mayr, 1862: 666 (redescription); Mayr, 1863: 400; Roger, 1863b: 6; Buckley, 1866: 155; Provancher, 1881b: 355; André, 1882a: 141 (in key); Provancher, 1883: 598; Provancher, 1887: 229 (in key); Nasonov, 1889: 13; Dalla Torre, 1893: 246; Ruzsky, 1896: 67; Emery, 1898a: 225; Forel, 1901h: 70; Ruzsky, 1903c: 205; Ruzsky, 1905b: 227; Forel, 1907e: 10; Wheeler, W.M. 1910d: 335; Emery, 1920b: 255; Creighton, 1950a: 367; Smith, M.R. 1958c: 143; Carter, 1962a: 7 (in list); Smith, M.R. 1967: 366; Francoeur, 1975: 264; Francoeur, 1977b: 207; Smith, D.R. 1979: 1427; DuBois & LaBerge, 1988: 146; Deyrup, et al. 1989: 100; Wheeler, G.C., et al. 1994: 305; Bolton, 1995b: 116; Mackay & Mackay, 2002: 301; Deyrup, 2003: 44; Wetterer & Wetterer, 2004: 215; Coovert, 2005: 167; Hansen & Klotz, 2005: 86; MacGown & Forster, 2005: 66; MacGown, et al. 2007: 11; Ellison, et al. 2012: 124; Deyrup, 2017: 196; Mackay, 2019: 292 (redescription).
 * Senior synonym of herculeanopennsylvanicus: Creighton, 1950a: 367; Smith, D.R. 1979: 1427; Bolton, 1995b: 117; Coovert, 2005: 167.
 * Senior synonym of mohican: Creighton, 1950a: 367; Smith, D.R. 1979: 1427; Bolton, 1995b: 117.
 * herculeanopennsylvanicus. Camponotus herculeanus var. herculeanopennsylvanicus Dalla Torre, 1893: 235.
 * [First available use of Camponotus herculeanus r. herculeanus var. herculeanopennsylvanicus Forel, 1879a: 57 (w.) U.S.A. (New York, Illinois, South Carolina); unavailable (infrasubspecific) name.]
 * As unavailable (infrasubspecific) name: André, 1882a: 143; Emery, 1896d: 372 (in list); Smith, M.R. 1951a: 842; Smith, M.R. 1958c: 161.
 * Subspecies of herculeanus: Ruzsky, 1896: 68.
 * Junior synonym of pennsylvanicus: Creighton, 1950a: 367; Smith, D.R. 1979: 1427; Bolton, 1995b: 103; Coovert, 2005: 167; Mackay, 2019: 293.
 * mohican. Camponotus herculeanus var. mohican Emery, 1925b: 72.
 * [First available use of Camponotus herculeanus subsp. pennsylvanicus var. mahican Wheeler, W.M. 1910d: 338 (s.w.) U.S.A. (Massachusetts, New Jersey); unavailable (infrasubspecific) name.]
 * Unnecessary replacement name for herculeanopennsylvanicus; hence junior synonym of the latter: Emery, 1925b: 72; Smith, M.R. 1951a: 842; Smith, D.R. 1979: 1427; Bolton, 1995b: 110; Mackay, 2019: 293.
 * Subspecies of herculeanus: Wheeler, G.C. & Wheeler, E.W. 1944: 250.
 * Junior synonym of pennsylvanicus: Creighton, 1950a: 367; Smith, D.R. 1979: 1427.

Major worker measurements (mm): HL 2.62 - 3.16, HW 2.68 - 3.34, SL 2.58 - 2.84, EL 0.59 - 0.73, CL 0.89 - 1.04, CW 1.13 - 1.34, WL 3.70 - 4.20, FFL 2.36 - 2.66, FFW 0.70 - 0.89. Indices: CI 102 - 106, SI 90 - 98, CLI 127 - 129, FFI 30 - 33.

Mandibles with 5 teeth; anterior border of clypeus weakly concave; sides of head narrowed anteriorly, convex, posterior margin concave; eyes failing to reach sides of head by about ½ - 1 minimum diameter; scape extending 1 - 2 funicular segments past posterior lateral corner of head; propodeum weakly angulate between 2 faces, which are about equal in length, propodeal spiracle elongate; petiole narrow in profile, apex rounded and convex as seen from behind. Erect and suberect setae sparse on most surfaces, present on clypeus, mostly along margins, present along frontal carinae extending back to posterior margin, absent on cheeks, sides of head, posterior lateral corners, scapes (except apex), few setae scattered on ventral surface of head, numerous on mesosoma, petiole and gaster; appressed setae sparse on head, present on dorsum of mesosoma down along upper edge of propodeum, very abundant on gaster, touching adjacent setae.

Most surfaces dull, head densely and evenly punctate, with scattered larger punctures, mesosoma punctate, merging to coriaceous on dorsum of mesosoma, gaster very finely punctate, with scattered larger punctures. Concolorous black.

Minor worker measurements (mm): HL 1.58 - 2.56, HW 1.24 - 2.34, SL 1.86 - 2.56, EL 0.41 - 0.59, CL 0.48 - 0.81, CW 0.73 - 1.06, WL 2.28 - 3.48, FFL 1.56 - 2.36, FFW 0.48 - 0.70. Indices: CI 78 - 91, SI 100 - 118, CLI 131 - 153, FFI 30.

Similar to major worker, except eyes come within ½ minimum diameter of the side of the head, sides of head weakly narrowed anteriorly, posterior margin nearly straight to rounded, scapes extending about ½ length past posterior lateral corner of head.

Pilosity sculpture and color as in major worker.

Female measurements (mm): HL 3.10 - 3.52, HW 3.38 - 3.80, SL 2.82 - 2.90, EL 0.80 - 0.89, CL 0.95 - 1.14, CW 1.31 - 1.40, WL 5.66 - 5.70, FFL 2.76 - 2.86, FFW 0.76 - 0.91. Indices: CI 108 - 109, SI 82 - 91, CLI 123 - 138, FFI 28 - 32.

Mandible with 5 teeth; clypeus weakly concave anteriorly, lateral angles poorly developed; head narrowed anteriorly, posterior margin nearly straight to convex; eyes failing to reach sides of head by less than 1 minimum diameter; scape extending 1 or 2 funicular segments past posterior lateral corner of head; propodeal spiracle elongate, narrow; petiole narrowed as seen in profile, apex moderately sharp.

Erect and suberect setae sparse, present along margins of clypeus, setae along frontal carinae extend to posterior margin, absent on posterior lateral corners, sides of head, scapes (except apex), few setae on dorsum of mesosoma, on petiole and dorsal surface of gaster; appressed pubescence sparse, few tiny setae on head and dorsal surface of mesosoma, those on gaster longer (0.01 - 0.02 mm), but shorter than setae of major.

Sculpture and color as in major worker.

Male measurements (mm): HL 1.58 - 1.70, HW 1.42 - 1.54, SL 1.74 - 2.04, EL 0.54 - 0.60, CL 0.36 - 0.45, CW 0.71 - 0.79, WL 3.64 - 3.82, FFL 2.26 - 2.46, FFW 0.46 - 0.51. Indices: CI 88 - 91, SI 110 - 126, CLI 175 - 197, FFI 20 - 21.

Black specimen, which apparently possesses no characteristics that would separate it from those of other similar species.

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.
 * Bare O. S. 1929. A taxonomic study of Nebraska ants, or Formicidae (Hymenoptera). Thesis, University of Nebraska, Lincoln, USA.
 * 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.
 * Bowers D. Q. 2018. An investigation of the distribution and behavior of the Dark Rover Ant, Brachymyrmex patagonicus Mayr, in South Carolina. Clemson University Master's Thesis 99pp.
 * Buren W. F. 1941. A preliminary list of Iowa ants. Iowa State College Journal of Science 15: 111-117
 * Canadensys Database. Dowloaded on 5th February 2014 at http://www.canadensys.net/
 * Carroll T. M. 2011. The ants of Indiana (Hymenoptera: Formicidae). Master's Thesis Purdue university, 385 pages.
 * Chen X., B. Adams, A. Sabo, T. Crupi, L. Hooper-Bui. 2016. Ant assemblages and co-occurrence patterns in Cypress-Tupelo swamp. Wetlands 36: 849–861.
 * 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. 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
 * 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
 * Dennis C. A. 1938. The distribution of ant species in Tennessee with reference to ecological factors. Annals of the Entomological Society of America 31: 267-308.
 * Deyrup M. 2016. Ants of Florida: identification and natural history. CRC Press, 423 pages.
 * Deyrup M., C. Johnson, G. C. Wheeler, J. Wheeler. 1989. A preliminary list of the ants of Florida. Florida Entomologist 72: 91-101
 * Downing H., and J. Clark. 2018. Ant biodiversity in the Northern Black Hills, South Dakota (Hymenoptera, Formicidae). Journal of the Kansas Entomological Society 91(2): 119-132.
 * 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. 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., J. Chen, D. Díaz, C. Kammerer-Burnham, and M. Lau. 2005. Changes in ant community structure and composition associated with hemlock decline in New England. Pages 280-289 in B. Onken and R. Reardon, editors. Proceedings of the 3rd Symposium on Hemlock Woolly Adelgid in the Eastern United States. US Department of Agriculture - US Forest Service - Forest Health Technology Enterprise Team, Morgantown, West Virginia.
 * 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-13NENHC-24.
 * 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. & Thompson L. 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. 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
 * 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.
 * 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
 * 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.
 * Hayes W. P. 1925. A preliminary list of the ants of Kansas (Hymenoptera, Formicidae). [concl.]. Entomological News 36: 69-73
 * 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.
 * Hess C. G. 1958. The ants of Dallas County, Texas, and their nesting sites; with particular reference to soil texture as an ecological factor. Field and Laboratory 26: 3-72.
 * Hill, J.G. 2006. Ants collected at Okatibbee Lake, Lauderdale County, Mississippi
 * Ionescu-Hirsch A. 2009. An annotated list of Camponotus of Israel (Hymenoptera: Formicidae), with a key and descriptions of new species. ISRAEL JOURNAL OF ENTOMOLOGY 39: 5798.
 * 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.
 * Johnson C. 1986. A north Florida ant fauna (Hymenoptera: Formicidae). Insecta Mundi 1: 243-246
 * Kannowski P. B. 1956. The ants of Ramsey County, North Dakota. American Midland Naturalist 56(1): 168-185.
 * 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.
 * Longino, J.T. 2010. Personal Communication. Longino Collection Database
 * Lubertazi, D. Personal Communication. Specimen Data from Museum of Comparative Zoology at Harvard
 * Lynch J. F. 1988. An annotated checklist and key to the species of ants (Hymenoptera: Formicidae) of the Chesapeake Bay region. The Maryland Naturalist 31: 61-106
 * MacGown J. A., J. G. Hill, R. L. Brown, T. L. Schiefer, J. G. Lewis. 2012. Ant diversity at Noxubee National Wildlife Refuge in Oktibbeha, Noxubee, and Winston Counties, Mississippi. Mississippi Agricultural and Forestry Experiment Station Bulletin 1197: 1-30
 * MacGown J. A., J. G. Hill, and 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., R. L. Brown, J. G. Hill, and B. Layton. 2007. Carpenter ants of Mississippi. Mississippi Agricultural and Forestry Experiment Station Bulletin 1158: 1-35.
 * 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., 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, W. P. and E. Mackay. 2002. The ants of New Mexico (Hymenoptera: Formicidae). Edwin Mellen Press, Lewiston, NY.
 * 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.
 * Mallis A. 1941. A list of the ants of California with notes on their habits and distribution. Bulletin of the Southern California Academy of Sciences 40: 61-100.
 * 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.
 * Mann W. M. 1911. On some Northwestern ants and their guests. Psyche (Cambridge) 18: 102-109.
 * 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
 * McClelland L. A. 1978. The Nebraska distribution of the ant genus Camponotus Mayr (Hymenoptera: Formicidae). Master's Thesis, Department of Biology and the faculty of the Graduate of Nebraska at Omaha, 72 pages.
 * 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
 * Parys K. A., M. L. Gimmel, and S. J. Johnson. 2013. Checklist of Insects Associated with Salvinia minima Baker in Louisiana, USA. Check List 9(6): 14881495.
 * Procter W. 1938. Biological survey of the Mount Desert Region. Part VI. The insect fauna. Philadelphia: Wistar Institute of Anatomy and Biology, 496 pp.
 * 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
 * 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.
 * Rowles, A.D. and J. Silverman. 2009. Carbohydrate supply limits invasion of natural communities by Argentine ants. Oecologia 161(1):161-171
 * Sanders, C.J. 1972. Seasonal and daily activity patterns of Carpenter ants (Camponotus spp.) in Northwestern Ontario (Hymenoptera: Formicidae). Canadian Naturalist 104:1681-1687
 * Sargent J. M., Benson. E.P., Zungoli. P. A. and Bridges. W. C. 2002. Carpenter Ant (Hymenoptera: Formicidae) Fauna of South Carolina. J. Agric. Urban Entomol. 18: 227-236
 * 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. 1934. A list of the ants of South Carolina. Journal of the New York Entomological Society 42: 353-361.
 * Smith M. R. 1934. Dates on which the immature or mature sexual phases of ants have been observed (Hymen.: Formicoidea). Entomological News 45: 247-251.
 * Smith M. R. 1935. A list of the ants of Oklahoma (Hymen.: Formicidae) (continued from page 241). Entomological News 46: 261-264.
 * Smith M. R. 1936. A list of the ants of Texas. Journal of the New York Entomological Society 44: 155-170.
 * Sturtevant A. H. 1931. Ants collected on Cape Cod, Massachusetts. Psyche (Cambridge) 38: 73-79
 * 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.
 * 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.
 * 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.
 * Varady-Szabo, H. 2004. Spiders and Ants Associated with Fallen Logs in Forillon National Park of Canada, Quebec. Masters Thesis, MgGill University, Montreal, Canada
 * Viereck H. L. 1903. Hymenoptera of Beulah, New Mexico. [part]. Trans. Am. Entomol. Soc. 29: 56-87.
 * 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
 * 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 E. W. Wheeler. 1944. Ants of North Dakota. North Dakota Historical Quarterly 11:231-271.
 * 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. 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. 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. 1910. The North American ants of the genus Camponotus Mayr. Annals of the New York Academy of Sciences 20: 295-354.
 * 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. 1916. Formicoidea. Formicidae. Pp. 577-601 in: Viereck, H. L. 1916. Guide to the insects of Connecticut. Part III. The Hymenoptera, or wasp-like insects, of Connecticut. Connecticut State Geological and Natural History Survey. Bulletin 22: 1-824.
 * 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.
 * Yitbarek S., J. H. Vandermeer, and D. Allen. 2011. The Combined Effects of Exogenous and Endogenous Variability on the Spatial Distribution of Ant Communities in a Forested Ecosystem (Hymenoptera: Formicidae). Environ. Entomol. 40(5): 1067-1073.
 * Young J., and D. E. Howell. 1964. Ants of Oklahoma. Miscellaneous Publication. Oklahoma Agricultural Experimental Station 71: 1-42.
 * Young, J. and D.E. Howell. 1964. Ants of Oklahoma. Miscellaneous Publications of Oklahoma State University MP-71
 * Zettler J. A., M. D. Taylor, C. R. Allen, and T. P. Spira. 2004. Consequences of Forest Clear-Cuts for Native and Nonindigenous Ants (Hymenoptera: Formicidae). Ann. Entomol. Soc. Am. 97(3): 513-518.