Temnothorax pergandei

This species nests in stumps, logs, nutshells and in the soil. It is widely distributed and is usually found in plant associations on higher dry sites but also nests in low flatwoods and hummocks, shaded deciduous forests or shaded prairie sites. It is active above ground throughout the year in Florida. (Mackay, 1993)

Identification
Prebus (2021) - Temnothorax pergandei can be separated from all other species in the salvini clade by the following character combination: promesonotum evenly convex; metanotal groove deeply impressed and propodeum depressed below the level of the promesonotum; propodeal spines present and typically represented by small teeth; in profile view, postpetiole evenly convex dorsally, with a small flat area posteriorly; in dorsal view, petiolar node 1.1 to 1.3 times as broad as caudal cylinder; setae on head, mesosoma, legs, waist segments and gaster erect, moderately long, abundant and tapering; integument variously colored: may be uniformly dark brown, uniformly yellow, or bicolored.

Similar species: This striking species is difficult to confuse with any other species in Temnothorax, except the closely related Temnothorax bison The presence of propodeal teeth, the structure of the postpetiole, which bulges anterodorsally in T. bison, and the relative width of the petiolar node in dorsal view (1.3 to 1.6 times as broad as the caudal cylinder in T. bison) will separate the two species.

Temnothorax pergandei is a highly morphologically variable species with a broad range, with a great deal of diversity in integument coloration, sculpturation, and propodeal spine length

Distribution
All elevations, eastern U.S.A. as far north as Michigan, as far west as Arizona, and south as far as Hidalgo, Mexico.

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

Habitat
From shaded portions of open sites to forests.

Abundance
Common.

Biology
Prebus (2021) - Temnothorax pergandei is often found in open, sparsely vegetated habitats throughout its range. With its rapid movements and constricted mesosoma, T. pergandei presents itself in the field as distinctly Pheidole-like, often foraging for small arthropods such as aphids and heteropterans during the diurnal hours (Wheeler, 1903b). Wheeler observed that workers of T. pergandei are territorial, showing aggression toward workers from other colonies. It often nests directly in the ground, with a very indistinct nest entrance that is typically only discoverable by following foraging workers back to it, much like other ground nesting Temnothorax. This is by no means the rule, though: nests have been collected under the bark of pine stumps (Smith, 1929), in rotten sticks on the ground (this study), or under moss mats and grass roots (Wheeler, 1903b). Emery (1895) states that Pergande collected T. pergandei in association with a Monomorium minimum Buckley nest, but this observation appears to be spurious; most collections of T. pergandei have been found apart from other ant species. Nests are small and monogynous, with 75 to 250 workers (Smith, 1929, Wesson, 1935; Heinze, Hoelldobler & Trenkle, 1995). Records of mating flights have been made from late spring in the vicinity of Austin, Texas, U.S.A. (Wheeler, 1903b), and early summer near Portal, Arizona, U.S.A. (Heinze, Hoelldobler & Trenkle, 1995). Gynes attract males with secretions from the poison gland, and typically mate with males only once in swarming flights (Heinze, Hoelldobler & Trenkle, 1995).

Wheeler (1903): "The type specimens of this fine species were taken by Mr. Pergande in a nest of Monomorium minutum Mayr var. minimum, and it was supposed that the Temnothoraxwas a guest in the nests-of the Monomorium, but Forel, who observed the species in North Carolina, showed that this was altogether an exceptional case. He found Temnothorax Pergandei living" in independent formicaries, in the moss of woods or in the earth of meadows, like the ordinary species of Temnothorax" (Ann. Soc. Ent, de Belg., Tome XLV, 1901,pp. 389-398). In Texas I have had ample opportunity to observe the habits of this ant, especially in the neighborhood of Austin, where it is found making its nests in very sparsely grassy spots among the mesquite and Opuntia thickets. The nests can be found only by carefully tracking foraging workers, as the entrance is a small hole often concealed under a dead twig or a tuft of grass roots. The colonies are hardly more populous than those of other species of Temnothorax. The winged forms appear during the last week in April and the first week in May. The workers run about on the soil in the hot sun as fierce hunters of small insects (Aphids, minute Heteroptera, etc). As they are extremely pugnacious even toward individuals of the same species from other nests, and as I have never found them nesting with Monomorium minimum, though this species is very common in the same localities, I believe, with Forel, that Pergande's observation must be quite exceptional or may even involve some misinterpretation."

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. Temnothorax pergandei was found twice, one colony with 76 workers and 1 queen and the other with 94 workers and 1 queen. While T. pergandei colonies were found with relative ease in the hickory nuts by MacGown (2006), the species was otherwise only rarely encountered in the Tombigbee National Forest when using pitfall and other collecting methods.

Ohio
Wesson and Wesson (1940) - Seen everywhere in dry, sunny fields and meadows, sometimes in dry open woods. Winged phases were taken from nests in early July.

Reproduction
Colonies are monogynous. Reproductives were collected in nests throughout the year, specifically on April 25, 1902 (Wheeler AMNH). June 23, 1941 (Buren USNM), November 18, 1932 (DeRead USNM) and December 4, 1928 (Smith MVIC). A flight occurred in Raleigh, NC on June 26, 1969 (specimens in NCSU). (Mackay 1993)

Nest in objects on the ground such as sticks and nuts (e.g., hickory or oak (MacGown, 2006)), in litter or in the ground.

Nomenclature

 * floridanus. Leptothorax (Dichothorax) floridanus Emery, 1895c: 324 (w.) U.S.A. Status as species: Wheeler, W.M. 1903c: 259; Wheeler, W.M. 1932a: 11. Subspecies of pergandei: Smith, M.R. 1929: 551; Creighton, 1950a: 260. Senior synonym of flavus and material of the unavailable name spinosus referred here: Creighton, 1950a: 260. Junior synonym of pergandei: Mackay, 1993a: 289.
 *  pergandei. Leptothorax (Dichothorax) pergandei Emery, 1895c: 323, pl. 8, fig. 13 (w.) U.S.A. Wheeler, W.M. 1903c: 257 (q.m.); Wheeler, G.C. & Wheeler, J. 1960b: 23 (l.). Combination in Temnothorax: Bolton, 2003: 272. Senior synonym of manni: Wesson, L.G. 1939: 180; of floridanus (and its junior synonym flavus): Mackay, 1993a: 289.
 * flavus. Leptothorax (Dichothorax) pergandei subsp. flavus Smith, M.R. 1929: 549 (w.q.) U.S.A. Junior synonym of floridanus: Creighton, 1950a: 260.
 * manni. Leptothorax (Dichothorax) manni Wesson, L.G. 1935: 208 (w.q.m.) U.S.A. [Unresolved junior primary homonym of manni Wheeler, W.M. 1914b: 53.] Junior synonym of pergandei: Wesson, L.G. 1939: 180.

Prebus (2021) - Temnothorax pergandei is a highly morphologically variable species with a broad range, with a great deal of diversity in integument coloration, sculpturation, and propodeal spine length (Fig. 130 & Fig. 131). These qualities have led to a profusion of names, with the morphologically similar but geographically distant Leptothorax floridanus appearing alongside the original description of T. pergandei. However, little sense has been made of the different forms, as there is often a great deal of intranidal and geographic variation (Creighton, 1950). Mackay (1993) synonymized T. floridanus under T. pergandei following a morphometric analysis of specimens throughout the range of both. I have found that the yellow form, formerly the subspecies T. pergandei flavus, is paraphyletic with respect to the typical dark form, as is the case with bicolored specimens. Most of the genetic structure that I have observed in the phylogeny of T. pergandei is explained by geography, with distinct Eastern, Central Great Plains, Northwestern and Southwestern clades, with the Eastern clade corresponding roughly to the Ozarks, the Appalachians and everything east of them. Whether these clades indicate distinct species is a question left to further analysis, but I cannot yet discern a distinct morphological pattern that supports separation.

Worker
Fusca, mandibulis, antennis (clava obscuriore), articulationibus pedum, tarsis et pedunculi segmenti 1. basi testaceis; nitida, metathorace opaco, pilosa, scapis pedibusque pube longa, obliqua vestitis; capite disperse punctato (punctis piligeris) praetereaque microscopice irregulariter aciculato, foveis antennalibus curvatim, genis longitrorsum rugosis, clypea media subtiliter, acute carinato, lateribus rugoso, mandibulis striatis, latiusculis, 5 dentatis; thoracis dorso profunde impresso, promesonoto convexo, nitido, disperse punctato et subtilissime aciculato, metanoto opaco, dentibus erectis armato, meso- et metapleuris longitrorsum rugosis et creberrime punctatis ,. abdominis pedunculi segment 1. antice longe petiolato, superne cum nodo transverso, medio impresso, 2. praecedente circiter dimidio latiore, transverse ovali. Long. 3-31/ 4 mm.

(Wheeler 1903) Length 2.5-3.25 mm.

Mandibles rather broad, 5-toothed. Clypeus moderately convex, broadly rounded in front, with a distinct median carina. Antennae 12-jointed, scape extending beyond the posterior angle of the head a distance fully equal to twice its breadth; first funicular joint as long as the three succeeding joints together; joints 3-8 nearly as long as broad; club 3-jointed, the two basal joints subequal, together shorter than the terminal joint. Thorax long, rather robust, not much wider in front than behind, without abrupt declivity at the juncture of the neck and pronotum, pro- and mesonotum convex; mesoepinotal constriction very deep and broad. Epinotal spines small, not longer than broad at the base, directed upward. Petiole from above nearly three times as long as its greatest width which is in the middle; in profile the node is low and rounded above, the anterior slope very long, at first nearly horizontal, then gradually as.cending, the posterior slope shorter, somewhat flattened; summit of node distinctly impressed or concave when seen from behind; ventral surface of peduncle with a long but not very prominent tooth. Postpetiole fully half again as broad as the petiole, a little broader than long, nearly square, its anterior angles prominent, its dorsal surface in profile very convex, especially in front. Gaster rather large, of the usual shape. Sting well developed. Legs robust.

Mandibles longitudinally striated. Clypeus smooth, especially behind, its sides longitudinally rugose. Head smooth and shining above and behind, sparsely punctate and irregularly and delicately reticulate. Antennal foveal with curved, parallel rugal; front and cheeks with straight rugal. Neck opaque and delicately rugose; pro- and mesonotum shining, very finely and irregularly reticulate, pleural, epinotum and mesoepinota,l constriction subopaque, coarsely reticulate-rugose; in the constriction and on the meso- and metapleural the rugae have a distinctly longitudinal trend. Declivous surface of epinotum smooth and shining. Petiole and postpetiole shining and finely reticulate above, more opaque and reticulate-rugose on the sides. Gaster smooth and shining, finely and irregularly reticulate.

Hairs white, long and very abundant, obtuse but not clavate, erect on the trunk, suberect on the legs and antennal. Black or rich dark-brown, mandibles, antennal, except. the club, frontal carinal, thorax, pedicel and legs yellowish-red or testaceous. In some specimens the dorsal surface of the thorax and nodes is blackened, while in others the whole of the thorax, nodes and legs is black except the peduncle of the petiole and the tarsi and joints of the legs, which are yellow.

Prebus (2021) - (n = 20): SL = 0.543-0.708 (0.631); FRS = 0.213-0.283 (0.248); CW = 0.643-0.853 (0.726); CWb = 0.597-0.784 (0.663); PoOC = 0.262-0.307 (0.284); CL = 0.701-0.860 (0.766); EL = 0.165-0.227 (0.189); EW = 0.106-0.158 (0.132); MD = 0.141-0.199 (0.172); WL = 0.847-1.135 (1.003); SPST = 0.109-0.222 (0.177); MPST = 0.278-0.399 (0.325); PEL = 0.346-0.500 (0.403); NOL = 0.195-0.272 (0.226); NOH = 0.099-0.156 (0.128); PEH = 0.187-0.275 (0.234); PPL = 0.171-0.255 (0.224); PPH = 0.240-0.344 (0.294); PW = 0.413-0.557 (0.473); SBPA = 0.118-0.228 (0.194); SPTI = 0.134-0.264 (0.220); PEW = 0.158-0.225 (0.191); PNW = 0.178-0.262 (0.228); PPW = 0.257-0.372 (0.322); HFL = 0.609-0.798 (0.703); HFWmax = 0.137-0.186 (0.160); HFWmin = 0.051-0.068 (0.059); CS = 0.948-1.214 (1.046); ES = 0.226-0.306 (0.255); SI = 84-104 (95); OI = 23-26 (24); CI = 82-91 (87); WLI = 142-157 (151); SBI = 20-33 (29); PSI = 13-21 (18); PWI = 156-183 (169); PLI = 155-213 (181); NI = 144-245 (178); PNWI = 111-133 (120); NLI = 51-64 (56); FI = 239-315 (271).

In full-face view, head subquadrate, longer than broad (CI 82-91). Mandibles densely striate but shining and armed with five teeth: the apical-most well developed and acute, followed by a less developed preapical tooth and three equally developed smaller teeth. Anterior clypeal margin weakly convex medially. Antennal scapes moderately long: when fully retracted, surpassing the posterior margin of the head capsule by about the maximum width of the scape (SI 84-04). Antennae 12-segmented; antennal club of composed of three segments, with the apical-most segment about as long as the preceding two in combination. Frontal carinae short and apices directed laterally, extending past the antennal toruli by about the maximum width of the antennal scape. Compound eyes moderately protruding past the lateral margins of the head capsule. Lateral margin of head convex, forming a continuous arc from the mandibular insertions to the posterior margin of the head. Posterior head margin flat but rounding evenly into the lateral margins.

In profile view, compound eyes ovular and moderately large (OI 23-26), with 16 ommatidia in longest row. Pronotal declivity indistinct, neck and anterior face of pronotum forming a ~120° angle. Mesosoma with promesonotum forming an even convexity; metanotal groove deeply impressed, distinctly separating the promesonotum from the propodeum, which is depressed below the level of the promesonotum and dorsally convex. Promesonotal suture extending from the posterior margin of the procoxal insertion to the mesothoracic spiracle, which is moderately well developed, then continuing to the dorsal surface as a very weak sulcus. Propodeal spiracle well developed, directed posterolaterally, and separated from the propodeal declivity by about three spiracle diameters. Propodeal spines poorly developed and short (PSI 13-21), in the form of small triangular teeth, angled posterodorsally. Propodeal declivity flat, forming a ~120° angle with the base of the propodeal spines. Propodeal lobes rounded and weakly developed. Metapleural gland bulla large, extending from the metacoxal insertion three quarters of the way to the propodeal spiracle. Petiole moderately long (PLI 155-213), without tubercles anterodorsally. Subpetiolar process in the form of a small, triangular, blunt tooth, which continues as a low carina to the caudal cylinder; ventral margin of petiole posterior to the subpetiolar process strongly concave. Petiolar peduncle moderately long: comprising about half of the petiole. Petiolar node robust and cuneiform: transition between peduncle and node marked by a rounded angle of ~130°; anterior face forming a rounded ~90° angle with the posterior face; caudal cylinder long: about the as long as the maximum width of the antennal scape. Postpetiole weakly convex anteriorly, rounding evenly into the dorsal face, which bulges before flattening posteriorly; lobed ventrally.

In dorsal view, humeri weakly developed: evenly rounded and wider than the rest of the mesosoma, but not distinct from it; mesothoracic spiracles weakly protruding past the lateral margins of the mesosoma, visible as slight angles where the pronotum meets the mesonotum. Promesonotal suture visible as a weak sulcus. Metanotal strongly impressed, distinctly dividing the promesonotum from the propodeum. Propodeal spines very short, but broadly approximated basally and diverging apically, their apices separated from each other by about five times their length. Petiolar peduncle with spiracles protruding past the lateral margins, but not noticeably constricted anterior to them. Petiolar node, when viewed posterodorsally, trapezoidal and apically broadened, the apex flat; caudal cylinder narrower than the apex of the node. Postpetiole narrow (PWI 156-183) and subquadrate. Anterior margin of the postpetiole weakly convex and meeting the lateral margins at a ~95° angle; lateral margins parallel to the angulate posterior corners; posterior margin broadly concave. Metafemur moderately to strongly incrassate (FI 239-315).

Sculpture: median clypeal carina present, extending posteriorly to the level of the anterior margins of the antennal toruli, and flanked on either side by one weaker carina. Lateral clypeal lobes with additional, weaker carinae; ground sculpture smooth and shining on the posterior half of the median clypeal lobe, and weakly areolate anteriorly. Antennal scapes shining through weak coriarious ground sculpture. Cephalic dorsum predominantly weakly coriarious; concentric costulae surrounding the antennal insertions, extending from the apices of the frontal carinae, to the anterior margins of the lateral clypeal lobes. Lateral surfaces of head predominantly weakly coriarious, with strong costulae between the compound eye and the mandibular insertion. Ventral surface of head smooth and shining. Pronotal neck areolate. Lateral surface of the pronotum and the surface surrounding the propodeal spiracle weakly coriarious. Lateral face of the propodeum, meso- and metapleurae with strong costulae. Dorsal surface of promesonotum weakly coriarious; dorsal surface of propodeum areolate. Femora smooth and shining. Petiole shining ventrally through weak coriarious sculpture; dorsum of petiolar peduncle and lateral face of petiolar node with weak costulae; petiolar node otherwise weakly coriarious. Postpetiole weakly coriarious. First gastral tergite and sternite weakly coriarious, without spectral iridescence.

Setae: antennal scapes and funiculi with long, subdecumbent pilosity. Dorsum of the head, pronotum, waist segments, and gaster with abundant, erect, flexuous, tapering setae, the longest of which are slightly longer than the length of the compound eye. The head bears ~36, mesosoma ~32, petiole 18, postpetiole ~28, and first gastral tergite ~66 setae. Pubescence is long, coarse, difficult to distinguish from the setae, and present over the entire body except for the ventral surface of the petiole.

Color: predominantly medium brown, with mandibles and tarsi testaceous yellow; masticatory margins of the mandibles dark brown.

Queen
Wheeler (1903) - Length 3.5-4 mm.

Apart from the usual sexual characters, differs from the worker in having the lower surfaces of the head, thorax, pedicel and the legs more yellowish. The smooth surfaces of the body are hardly reticulate. Mesonotum adorned with a median brown blotch on its anterior half and a large comma-shaped spot on either parapsis. Scutellum, epinotum and pleurm more or less spotted with brown, and a small black spot at the insertion of the forewing. Femora and tibim infuscated in the middle. Antennal club infuscated. Wings milky-hyaline, with colorless veins and stigma. Concavity at smnmit of petiolar node very distinct; postpetiole nearly twice as broad as long. Epinotal spines distinctly shorter than broad at their bases.

Prebus  (2021) - (n = 9): SL = 0.633-0.728 (0.680); FRS = 0.270-0.327 (0.301); CW = 0.848-0.963 (0.906); CWb = 0.755-0.868 (0.816); PoOC = 0.289-0.339 (0.315); CL = 0.821-0.943 (0.880); EL = 0.244-0.284 (0.267); EW = 0.173-0.222 (0.198); MD = 0.142-0.195 (0.171); WL = 1.405-1.554 (1.488); SPST = 0.203-0.274 (0.240); MPST = 0.352-0.458 (0.407); PEL = 0.453-0.515 (0.492); NOL = 0.212-0.319 (0.262); NOH = 0.135-0.181 (0.162); PEH = 0.293-0.362 (0.328); PPL = 0.229-0.291 (0.259); PPH = 0.370-0.439 (0.406); PW = 0.786-0.903 (0.835); SBPA = 0.341-0.405 (0.377); SPTI = 0.307-0.439 (0.370); PEW = 0.233-0.277 (0.258); PNW = 0.256-0.328 (0.295); PPW = 0.384-0.465 (0.429); HFL = 0.793-0.896 (0.852); HFWmax = 0.160-0.185 (0.177); HFWmin = 0.063-0.085 (0.072); CS = 1.166-1.340 (1.256); ES = 0.343-0.395 (0.366); SI = 79-89 (83); OI = 27-32 (29); CI = 90-96 (93); WLI = 178-188 (183); SBI = 41-49 (46); PSI = 14-18 (16); PWI = 161-170 (166); PLI = 175-223 (191); NI = 143-233 (163); PNWI = 107-119 (114); NLI = 46-63 (53); FI = 209-283 (248).

In full-face view, head subquadrate, slightly longer than broad (CI 90-96). Mandibles densely striate but shining and armed with five teeth: the apical-most well developed, followed by a less developed preapical tooth and three equally developed smaller teeth. Anterior clypeal margin flat medially. Antennal scapes moderately long: when fully retracted, just reaching the posterior margin of the head capsule (SI 79-89). Antennae 12-segmented; antennal club composed of three segments, with the apical-most segment as long as the preceding two in combination. Frontal carinae moderately long, directed laterally at their apices, and extending past the antennal toruli by about three times the maximum width of the antennal scape. Compound eyes moderately protruding past the lateral margins of the head capsule. Lateral margin of head evenly convex, converging from below the compound eyes to the mandibular insertions. Posterior head margin flat, rounding evenly into the lateral margins.

In profile view, compound eyes ovular and large (OI 27-32), with 20 ommatidia in longest row. Mesoscutum rounded evenly anteriorly, not quite covering the dorsal surface of the pronotum, and weakly convex dorsally. Mesoscutellum on the same level of the mesoscutum, and evenly convex dorsally. Posterior margin of metanotum extending slightly past the posterior margin of the mesoscutum. Propodeal spiracle well developed, directed posterolaterally, and separated from the propodeal declivity by about two and a half spiracle diameters. Propodeal spines short and stout (PSI 14-18), represented by posterodorsally directed, blunt, triangular teeth, the bases of which are attached to the propodeal declivity by flanges. Propodeal declivity straight and flat, forming a ~120° angle with the base of the propodeal spines. Propodeal lobes rounded and weakly developed. Metapleural gland bulla large, extending from the metacoxal insertion three quarters of the way to the propodeal spiracle. Petiole moderately long (175-223), without tubercles anterodorsally. Subpetiolar process in the form of a small, triangular, blunt tooth, which continues as a low carina to the caudal cylinder; ventral margin of petiole posterior to the subpetiolar process strongly concave. Petiolar peduncle moderately long: comprising about half the length of the petiole. Petiolar node robust and cuneiform: transition between peduncle and node marked by a rounded angle of ~130°; anterior face forming a rounded ~90° angle with the posterior face; caudal cylinder long: about the as long as the maximum width of the antennal scape. Postpetiole weakly convex anteriorly, rounding evenly into the dorsal face, which bulges before flattening posteriorly; lobed ventrally.

In dorsal view, mesoscutum not fully covering pronotum anteriorly; humeri evenly rounded laterally. Propodeal teeth diverging apically, their apices separated from each other by about six times their length. Petiolar peduncle with spiracles protruding past the lateral margins, but not noticeably constricted anterior to them. Petiolar node, when viewed posterodorsally, trapezoidal and apically broadened, the apex weakly emarginate; caudal cylinder narrower than the apex of the node. Postpetiole narrow (PWI 161-170), and subquadrate. Anterior margin of postpetiole weakly convex, with corners marked by rounded angles as it transitions to the lateral margins, which are parallel to the angulate posterior corners; posterior margin broadly concave. Metafemur weakly to moderately incrassate (FI 209-283).

Sculpture: median clypeal carina present, extending from the anterior margin nearly to the level of the anterior margins of the antennal insertions; flanked by weaker, indistinct carinae near the lateral margins of median clypeal lobe. Lateral clypeal lobes with additional weaker carinae; ground sculpture smooth and shining on the median clypeal lobe medially but becoming weakly areolate-coriarious laterally. Antennal scapes weakly coriarious. Cephalic dorsum predominantly costulate over smooth and shining sculpture, with moderately strong striae flanking the frontal carinae; a median strip of smooth sculpture extends from the frontal triangle to the median ocellus, then expands to encompass the middle of the ocellar triangle and extends to the posterior head margin; concentric costulae surrounding the antennal insertions, extending from the frontal carinae to the anterior margin of the lateral clypeal lobes. Lateral surfaces of head with weak coriarious sculpture posterior to the compound eye, denser rugose sculpture surrounding the compound eye, extending between the compound eye and the mandibular insertion. Ventral surface of head weakly costulate. Pronotal neck areolate. Lateral face of the pronotum weakly coriarious, with costulae on the ventral third. Anepisternum smooth and shining on its anterior half, but weakly coriarious on its posterior half; katepisternum predominantly

Male
Wheeler (1903) - Length 2-2.25 mm.

Head a little longer than broad; cheeks very short, eyes and ocelli prominent. Mandibles overlapping, 4-toothed. Clypeus convex. Antennae 13-jointed, scape about as long as the five succeeding joints, first. funicular joint about as long as the second and third together; joints 3-8 slerider, cylindrical, subequal, twice as long as broad; club 4-jointed, the three basal joints subequal, together as long as the terminal joint. Thorax long, mesonotum rounded, projecting forward, so that the head is scarcely visible when the insect is viewed from above. Epinotum with a pair of slight projections in the place of the spines. Petiole long and slender, like that of the worker but with lower node. Postpetiole as long as broad, squar.e from above, in profile with the node highest in the middle of the segment and rounded. Gaster and legs of the usual conformation.

Mandibles and clypeus subopaque, the latter coarsely rugose. Head shining, very irregularly and sparsely reticulate, in.front of the ocelli with a few shallow foveolae. Pronotum subopaque, reticulate; mesonotum very smooth and shining, finely reticulate. Scutellum and epinotum more opaque, disk of former coarsely reticulate, sides of both longitudinally rugulose. Petiole and postpetiole opaque, more shining above, delicately corrugated. Gaster very smooth and shining.

Hairs white, rather sparse and long, even on the legs and antennal scape, where they are reclinate but not appressed.

Black. Mandibles, antennae and legs white except the following portions, which are infuscated or blackened: Edges and teeth of mandibles, terminal joint of antennae, basal two-thirds of coxae, middle of femora and tibiae and last tarsal joint. Wings like those of the female.

Prebus  (2021) - (n = 2): SL = 0.261–0.324 (0.293); FRS = 0.133–0.151 (0.142); CW = 0.552–0.592 (0.572); CWb = 0.466–0.475 (0.471); PoOC = 0.251–0.266 (0.259); CL = 0.561–0.603 (0.582); EL = 0.240–0.267 (0.254); EW = 0.196–0.209 (0.203); MD = 0.038–0.041 (0.040); WL = 1.062–1.169 (1.116); SPST = 0.188–0.216 (0.202); MPST = 0.310–0.378 (0.344); PEL = 0.334–0.335 (0.335); NOL = 0.194–0.258 (0.226); NOH = 0.076–0.078 (0.077); PEH = 0.160–0.166 (0.163); PPL = 0.202–0.215 (0.209); PPH = 0.216–0.260 (0.238); PW = 0.600–0.616 (0.608); SBPA = 0.224–0.257 (0.241); SPTI = 0.224–0.257 (0.241); PEW = 0.163–0.182 (0.173); PNW = 0.142–0.197 (0.170); PPW = 0.255–0.268 (0.262); HFL = 0.819–0.891 (0.855); HFWmax = 0.093–0.098 (0.096); HFWmin = 0.051; CS = 0.747–0.777 (0.762); ES = 0.338–0.372 (0.355); SI = 56–68 (62); OI = 45–48 (47); CI = 79–83 (81); WLI = 228–246 (237); SBI = 48–54 (51); PSI = 18; PWI = 140–164 (152); PLI = 156–165 (161); NI = 249–339 (294); PNWI = 87–108 (98); NLI = 58–77 (68); FI = 182–192 (187).

In full-face view, head elongate and subovate, longer than broad (CI 79–83). Mandibles very weakly striate, shining, and armed with five teeth: the apical-most well developed, followed by a smaller preapical tooth and three roughly equally developed smaller teeth. Anterior clypeal margin entire and weakly convex. Antennal scapes short: when fully retracted, failing to reach the posterior margin of the head capsule by about two times the maximum width of the scape (SI 56–68). Antennae 13-segmented; antennal club composed of four segments, with the apical-most segment slightly longer than the preceding two in combination. Frontal carinae short, extending past the antennal toruli by two times the maximum width of the antennal scape. Compound eyes strongly protruding past the lateral margins of the head capsule. Margin between the anterior margin of the compound eye and the mandibular insertions straight. Posterior head margin flat convex, evenly rounding into the lateral margins. In profile view, compound eyes ovular and large (OI 45–48), with 30 ommatidia in the longest row. Mesoscutum bulging anteriorly, covering the dorsal surface of the pronotum, and evenly convex dorsally. Mesoscutellum on the same level of the mesoscutum and evenly rounded dorsally. Posterior margin of metanotum extending beyond the posterior margin of scutellum. Propodeal spiracle well developed, directed laterally, and separated from the propodeal declivity by about four spiracle diameters. Propodeal spines absent but indicated by angulate flanges on the dorsal and declivitous faces of the propodeum. Propodeal lobes rounded and weakly developed. Metapleural gland bulla large, extending three quarters of the way between the insertion of the metacoxa and the propodeal spiracle. Petiole moderately long (PLI 156–165), without anterodorsally. Subpetiolar process absent; ventral margin of petiole concave. Petiolar peduncle short: comprising about a third of the total length of the petiole. Petiolar node low and cuneiform, the convergence of the anterior and posterior faces marked by a rounded angle. Postpetiole evenly rounded anterodorsally, flattened dorsally, and with a lobed, concave ventral surface. In dorsal view, mesoscutum covering pronotum anteriorly, humeri not visible. Petiolar peduncle with spiracles strongly protruding past the lateral margins, the peduncle broadened where they arise. Petiolar node slightly wider than the peduncle; petiole narrowing posterior to the node, before widening again to the caudal cylinder, which is about the same width as the node. Postpetiole narrow (PWI 140–164) and campaniform. Anterior margin of postpetiole convex, with the anterior corners evenly rounding into the lateral margins, which are parallel to the angulate posterior corners; posterior margin of postpetiole broadly concave. Metafemur not incrassate (FI 182–192).

Sculpture: median clypeal carina absent. Lateral clypeal lobes with weak, indistinct carinae, ground sculpture weakly coriarious. Antennal scapes smooth and shining. Head predominantly weakly areolate-coriarious, with a central strip of smooth and shining sculpture extending from the frontal triangle to the median ocellus. Lateral surface of head weakly coriarious. Ventral surface of head smooth and shining anteriorly, transversely rugulose posteriorly. Pronotal neck areolate. Lateral surfaces of pronotum, anepisternum, katepisternum, predominantly smooth and shining, but with anterior third of pronotum densely rugose. Metapleuron and lateral face of propodeum longitudinally rugose. Dorsally, mesoscutum weakly coriarious, the Mayrian furrows moderately well impressed. Mesoscutellum smooth and shining medially, becoming coriarious laterally. Femora smooth and shining. Petiole costulate ventrally; lateral faces of the peduncle and node longitudinally rugose, the posterodorsal surface of the petiole smooth and shining. Dorsal surface of postpetiole shining. First gastral tergite and sternite weakly coriarious. Setae: antennal scapes and funiculi with long, subdecumbent pilosity. Dorsum of the head, pronotum, waist segments, and gaster with abundant, erect, flexuous, tapering setae, the longest of which are about half the width of the compound eye. Pubescence is long, coarse, difficult to distinguish from the setae, and present over the entire body except for the ventral surface of the petiole.

Color: predominantly medium brown, with mandibles, genae, antennal scapes, antennal funiculus, and tarsi testaceous yellow; masticatory margins of the mandibles dark brown.

Type Material
Prebus (2021) - Syntype workers. Washington, D.C., U.S.A. One syntype worker here designated lectotype.

Lectotype worker of Leptothorax (Dichothorax) pergandei: U.S.A.: District of Columbia: Washington, D.C.: September 1910, [no collector data], no. 54078 U.S.N.M. cotype, (USNMENT00922524).

Paralectotype workers: same data as lectotype: 1 worker (USNMENT00922508) [USNM] 1 worker (USNMENT00922509) [USNM] 1 worker (USNMENT00922515) [USNM] 1 worker (USNMENT00922517) [USNM] 1 worker (USNMENT00922518) [USNM] 1 worker (USNMENT00922510) [USNM] 1 worker (USNMENT00922511) [USNM] 1 worker (USNMENT00922522) [USNM] 1 worker (USNMENT00922523) [USNM] 1 worker (USNMENT00922525) [USNM] 1 worker (USNMENT00532783) [USNM] 1 worker (images of CASENT0904767 examined on antweb. org).

Etymology
Patronym, in honor of Theodore Pergande, a myrmecologist who sent his extensive collections to Forel and Emery.

References based on Global Ant Biodiversity Informatics

 * Annotated Ant Species List Ordway-Swisher Biological Station. Downloaded at http://ordway-swisher.ufl.edu/species/os-hymenoptera.htm on 5th Oct 2010.
 * Atchison R. A., J. Hulcr, and A. Lucky. 2018. Managed fire frequency significantly influences the litter arthropod community in longleaf pine flatwoods. Environmental Entomology 20(10): 1-11.
 * Buren W. F. 1944. A list of Iowa ants. Iowa State College Journal of Science 18:277-312
 * Callcott A. M. A., D. H. oi, H. L. Collins, D. F. Williams, and T. C. Lockley. 2000. Seasonal Studies of an Isolated Red Imported Fire Ant (Hymenoptera: Formicidae) Population in Eastern Tennessee. Environmental Entomology, 29(4): 788-794.
 * 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
 * Carroll T. M. 2011. The ants of Indiana (Hymenoptera: Formicidae). Master's Thesis Purdue university, 385 pages.
 * Coovert G. A. 2005. The Ants of Ohio (Hymenoptera: Formicidae). Ohio Biological Survey, Inc. 15(2): 1-207.
 * Coovert, G.A. 2005. The Ants of Ohio (Hymenoptera: Formicidae) Ohio Biological Survey Bulletin New Series Volume 15(2):1-196
 * Cover S. P., and R. A. Johnson. 20011. Checklist of Arizona Ants. Downloaded on January 7th at http://www.asu.edu/clas/sirgtools/AZants-2011%20updatev2.pdf
 * Dash S. T. and L. M. Hooper-Bui. 2008. Species diversity of ants (Hymenoptera: Formicidae) in Louisiana. Conservation Biology and Biodiversity. 101: 1056-1066
 * Dattilo W. et al. 2019. MEXICO ANTS: incidence and abundance along the Nearctic-Neotropical interface. Ecology https://doi.org/10.1002/ecy.2944
 * Dean D. A., and S. R. Dean. 2018. A survey of the ant fauna and seasonal alate nuptial flights at two locations in South-Central Texas. Southwestern Entomologist 43(3): 639-647.
 * Deyrup M. 1998. Smithistruma memorialis (Hymenoptera: Formicidae), a new species of ant from the Kentucky Cumberland Plateau. Entomological News 109: 81-87.
 * Deyrup M., C. Johnson, G. C. Wheeler, J. Wheeler. 1989. A preliminary list of the ants of Florida. Florida Entomologist 72: 91-101
 * Deyrup, M. and J. Trager. 1986. Ants of the Archbold Biological Station, Highlands County, Florida (Hymenoptera: Formicidae). Florida Entomologist 69(1):206-228
 * Deyrup, M. and S. Cover. 2004. A new species of the ant genus Leptothorax from Florida, with a key to the Leptothorax of the southeast (Hymenoptera: Formicidae). Floridae Entomologist 87(1):51-59
 * 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: 153-772
 * DuBois M. B. 1985. Distribution of ants in Kansas: subfamilies Ponerinae, Ecitoninae, and Myrmicinae (Hymenoptera: Formicidae). Sociobiology 11: 153-773
 * DuBois M. B. 1985. Distribution of ants in Kansas: subfamilies Ponerinae, Ecitoninae, and Myrmicinae (Hymenoptera: Formicidae). Sociobiology 11: 153-774
 * DuBois M. B. 1985. Distribution of ants in Kansas: subfamilies Ponerinae, Ecitoninae, and Myrmicinae (Hymenoptera: Formicidae). Sociobiology 11: 153-775
 * DuBois M. B. 1985. Distribution of ants in Kansas: subfamilies Ponerinae, Ecitoninae, and Myrmicinae (Hymenoptera: Formicidae). Sociobiology 11: 153-776
 * DuBois M. B. 1985. Distribution of ants in Kansas: subfamilies Ponerinae, Ecitoninae, and Myrmicinae (Hymenoptera: Formicidae). Sociobiology 11: 153-777
 * DuBois M. B. 1985. Distribution of ants in Kansas: subfamilies Ponerinae, Ecitoninae, and Myrmicinae (Hymenoptera: Formicidae). Sociobiology 11: 153-778
 * DuBois M. B. 1985. Distribution of ants in Kansas: subfamilies Ponerinae, Ecitoninae, and Myrmicinae (Hymenoptera: Formicidae). Sociobiology 11: 153-779
 * DuBois M. B. 1985. Distribution of ants in Kansas: subfamilies Ponerinae, Ecitoninae, and Myrmicinae (Hymenoptera: Formicidae). Sociobiology 11: 153-780
 * DuBois M. B. 1985. Distribution of ants in Kansas: subfamilies Ponerinae, Ecitoninae, and Myrmicinae (Hymenoptera: Formicidae). Sociobiology 11: 153-781
 * DuBois M. B. 1985. Distribution of ants in Kansas: subfamilies Ponerinae, Ecitoninae, and Myrmicinae (Hymenoptera: Formicidae). Sociobiology 11: 153-782
 * DuBois M. B. 1985. Distribution of ants in Kansas: subfamilies Ponerinae, Ecitoninae, and Myrmicinae (Hymenoptera: Formicidae). Sociobiology 11: 153-783
 * DuBois M. B. 1985. Distribution of ants in Kansas: subfamilies Ponerinae, Ecitoninae, and Myrmicinae (Hymenoptera: Formicidae). Sociobiology 11: 153-784
 * DuBois M. B. 1985. Distribution of ants in Kansas: subfamilies Ponerinae, Ecitoninae, and Myrmicinae (Hymenoptera: Formicidae). Sociobiology 11: 153-788
 * 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
 * Forster J.A. 2005. The Ants (hymenoptera: Formicidae) of Alabama. Master of Science, Auburn University. 242 pages.
 * Frye J. A., T. Frye, and T. W. Suman. 2014. The ant fauna of inland sand dune communities in Worcester County, Maryland. Northeastern Naturalist, 21(3): 446-471.
 * General D. M., and L. C. Thompson. 2011. New Distributional Records of Ants in Arkansas for 2009 and 2010 with Comments on Previous Records. Journal of the Arkansas Academy of Science 65: 166-168.
 * General D., and L. Thompson. 2008. Ants of Arkansas Post National Memorial: How and Where Collected. Journal of the Arkansas Academy of Science 62: 52-60.
 * General D.M. & Thompson L.C. 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.
 * 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
 * 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.
 * Hill J.G. & Brown R. L. 2010. The Ant (Hymenoptera: Formicidae) Fauna of Black Belt Prairie Remnants in Alabama and Mississippi. Southeastern Naturalist. 9: 73-84
 * Ipser R. M. 2004. Native and exotic ants (Hymenoptera: Formicidae) of Georgia: Ecological Relationships with implications for development of biologically-based management strategies. Doctor of Philosophy thesis, University of Georgia. 165 pages.
 * 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. 2015. Checklist of the ants (Hymenoptera: Formicidae) of Ohio. Conference: Ohio Natural History Conference, At Columbus OH
 * Ivanov, K. 2019. The ants of Ohio (Hymenoptera, Formicidae): an updated checklist. Journal of Hymenoptera Research 70: 65–87.
 * Ivanov K., L. Hightower, S. T. Dash, and J. B. Keiper. 2019. 150 years in the making: first comprehensive list of the ants (Hymenoptera: Formicidae) of Virginia, USA. Zootaxa 4554 (2): 532–560.
 * Johnson C. 1986. A north Florida ant fauna (Hymenoptera: Formicidae). Insecta Mundi 1: 243-246
 * Johnson R. Personnal Database. Accessed on February 5th 2014 at http://www.asu.edu/clas/sirgtools/resources.htm
 * Jusino-Atresino R., and S. A. Phillips, Jr. 1992. New ant records for Taylor Co., Texas. The Southern Naturalist 34(4): 430-433.
 * Lubertazzi D. and Tschinkel WR. 2003. Ant community change across a ground vegetation gradient in north Floridas longleaf pine flatwoods. 17pp. Journal of Insect Science. 3:21
 * 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 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
 * MacKay W. P. 1993. The status of the ant Leptothorax pergandei Emery (Hymenoptera: Formicidae). Sociobiology 21: 287-297.
 * MacKay W. P., S. P. Cover, J. Heinze, and B. Hölldobler. 1995. Range extensions for the ant Leptothorax pergandei (Hymenoptera: Formicidae): a mesic forest species discovered in the Chihuahuan Desert. Proceedings of the Entomological Society of Washington 97: 888.
 * 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.
 * Moreau C. S., M. A. Deyrup, and L. R. David Jr. 2014. Ants of the Florida Keys: Species Accounts, Biogeography, and Conservation (Hymenoptera: Formicidae). J. Insect Sci. 14(295): DOI: 10.1093/jisesa/ieu157
 * 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
 * Roeder K. A., and D. V. Roeder. 2016. A checklist and assemblage comparison of ants (Hymenoptera: Formicidae) from the Wichita Mountains Wildlife Refuge in Oklahoma. Check List 12(4): 1935.
 * Smith M. R. 1918. A key to the known species of South Carolina ants, with notes (Hym.). Entomological News 29: 17-29.
 * Smith M. R. 1934. Dates on which the immature or mature sexual phases of ants have been observed (Hymen.: Formicoidea) (continued from page 251). Entomological News 45: 264-267.
 * Smith M. R. 1935. A list of the ants of Oklahoma (Hymen.: Formicidae). Entomological News 46: 235-241.
 * Smith M. R. 1962. A new species of exotic Ponera from North Carolina (Hymenoptera, Formicidae). Acta Hymenopterologica 1: 377-382.
 * 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.
 * Trager, J. and C.Johnson. 1985. A slave-making ant in Florida: Polyergus lucidus with observations on the natural history of its host Formica archboldi (Hymenoptera: Formicidae). The Florida Entomologist 68(2):261-266.
 * 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. 1948. A Preliminary Key to the Worker Ants of Alachua County, Florida. The Florida Entomologist 30(4): 57-67
 * Van Pelt A. F. 1956. The ecology of the ants of the Welaka Reserve, Florida (Hymenoptera: Formicidae). American Midland Naturalist 56: 358-387
 * 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
 * Warren, L.O. and E.P. Rouse. 1969. The Ants of Arkansas. Bulletin of the Agricultural Experiment Station 742:1-67
 * Wetterer, J.K. and J.A. Moore. 2005. Red Imported Fire Ants (Hymenoptera: Formicidae) at Gopher Tortoise (Testudines: Testudinidae) Burrows. The Florida Entomologist 88(4):349-354
 * Wheeler G. C., and J. Wheeler J. 1989. A checklist of the ants of Oklahoma. Prairie Naturalist 21: 203-210.
 * 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. 1904. The ants of North Carolina. Bulletin of the American Museum of Natural History. 20: 299-306.
 * Wheeler, G.C. and J. Wheeler. 1985. A checklist of Texas ants. Prairie Naturalist 17:49-64.
 * 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