Stenamma brevicorne

From Smith (1957): Stenamma brevicorne is not only one of the most widely distributed but the oldest and perhaps best known of our North American species of Stenamma. Although apparently preferring to nest in wooded areas, the species is also found in meadows and other places. It usually nests in moderately dense to dense woods of such composition as maple, mixed oaks, oak-maple, red and white oaks, and beech-maple. Here the ants more commonly nest in the soil under stones, logs, humus, moss or other debris, as well as in the rotting wood of logs and stumps. Colonies are small, consisting of only a few dozen to a hundred or so adult individuals. The largest that has come to my attention was found at Tiffin, Ohio, on August 21, 1948, by Talbot and Headley and was composed of 105 workers, 8 alate females, and 12 males. Although no one has made a detailed study of Stenamma brevicorne, random observations indicate that this species is timid and sluggish, subterranean or hypogaeic. The workers are undoubtedly carnivorous, but may also be predaceous. Wheeler, so far as I am aware, was the first individual to advance the theory that, in some colonies at least, winged females and males overwinter in the parental nest, these sexual castes having been produced from late summer to early fall. None of the evidence I have seen disproves this, but no doubt the age of the colony has a great deal to do with whether or not sexual castes can be produced. As the colonies are not large, it is likely that there is only one mother queen per colony. This also seems to be the case in many, if not most of our own species. The overwintering sexual castes apparently start emerging from the parental colony in spring or early summer of the following year. It is believed that it is then that mating and the forming of new colonies are started. H. V. Weems, Jr., captured a copulating male and female on May 14, 1950. Since alate females have been captured at light traps and males taken while flying at dusk, it appears that both castes fly freely at night, like many other ants. Although altitudunal records are lacking for Stenamma brevicorne, it appears that this form lives in low lands or areas of only moderately high elevation.

Identification
The worker of Stenamma brevicorne may be distinguished by its moderately large eye, which usually bears 8-10 ommatidia in its greatest diameter; the strong mesoepinoraI impression which is frequently twice as long as deep; base of epinotum usually with a transverse welt following the mesoepinorai impression; the distinct spines which are approximately one-fourth to one-third the length of the base of the epinotum; petiolar node subconical, when viewed from behind; postpetiolar node, from above, stout, as broad as, or broader than long; head, thorax, petiole and postpetiole subopaque; promesonotum largely coarsely longitudinaIly rugulose; body usually brown with the gaster light at the base and apex and darker medianly. (Smith 1957)

Distribution
CANADA, USA. From Nova Scotia, Quebec, and Ontario south to Tennessee and west to Nebraska and Minnesota.

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

Missouri
Talbot (1965) offers the following observations about Stenamma brevicorne from Missouri:

Stenamma brevicorne, together with Lasius brevicornis, seemed the most typical ant of the low field habitat. It was common and widespread here (23 colonies on 18 plots), but had been collected only once before on the Reserve (at swamp edge). Three other species of Stenamma (Stenamma impar, Stenamma diecki, Stenamma schmittii) present on the Reserve did not live in the field.

Stenamma brevicorne workers foraged over the ground in cool damp weather, and the finding of workers just below the surface indicates that they also foraged below ground in the crumbly topsoil. Nest entrances were simply small openings hidden among leaves and grasses, and were not found unless workers were seen entering them. An attempt was made to collect whole colonies; therefore nests were dug completely when possible. This was not always a successful process, but the general pattern of nest structure seemed clear. Usually there was a chamber within 2 inches of the surface, which might be empty during dry weather. Some nests appeared to have no chambers in the crumbly surface soil but started with a chamber 4 inches down in more solid clay. No colony had more than 4 chambers, and most had only 2. Lowest chambers ranged in depth from 3 to 13 inches, with most at 6 inches. Large colonies dug down to soil moisture, while small colonies did not. Chambers were surprisingly large and were often only partly filled with brood. Those measured ranged from 1/2 X 1/2X 1/4 in. to 1 1/2 X 1 1/2 X 1 in. with most being larger than an inch in diameter and a half inch high. The range of size of the 10 most complete population counts is recorded in Table II. Eggs, larvae and worker pupae were present from before June 23, when the first colonies were dug, through the rest of the summer, although the September 10 colony had only one worker pupa. Twelve colonies which had winged adults or winged pupae averaged 20.6 alates per colony, of which 11.2 were females and 9.4 were males. Winged pupae were found in colonies from July 9 to August 11, and adults from August 10 to September 10 when observations ceased. It is known that S. brevicorne has flights in the spring (Smith, 1957; Kannowski, 1958). It may also fly in the late fall when conditions are right.

This species generally inhabits open areas, with either a soil nest that opens underneath an object resting on the ground or within the litter horizon in debri or other objects where they can aggregate within chamber like structures.

Reproduction
Kannowski (1958) offers these detailed observations on a mating flight:

On the afternoon of May 2, 1955, I observed swarming of Stenamma brevicorne alates near "Blanchard's Pond" on the outskirts of Ann Arbor, Michigan. The pond, about an acre in size, occupied a shallow depression surrounded by a field which had not been cultivated in the last five years. Around the periphery of the pond were isolated clumps of shrubs and small trees, of which willows (Salix spp.) and hawthorns (Crataegus spp.) were the most numerous. About 150 yards to the northeast of the pond was a large oak-hickory woodland ("Eber White Woods").

I discovered the swarms about 2:45 p.m. It was a cloudy and moderately warm day (about 70 F.) with very little wind. The swarms had formed at the tops of 8 to 12 foot high hawthorns and willows at the edge of the pond. The hawthorns were in bloom. When I first sighted the swarms from a distance, I assumed that they were composed of bees attracted to the pollen of the flowers. Closer examination revealed that they were alate ants. The alates were not in contact with the trees, but always remained slightly above the tree tops. About a dozen such swarms were found at the tops of these trees. Usually, there was only one swarm per tree top, but several large trees had two swarms each.

Each swarm was composed of 15 to 50 males. These swarms were observed for an hour and a half, but no females were seen. At 4:15 p.m. it was necessary for me to leave the area, and the observations were terminated. No new swarms were seen to form during the time of observation. Each swarm hovered in approximately the same position during this time. The size varied from swarm to swarm, but each occupied an area that was roughly cuboidal in shape and measured from one to two feet on a side. Movement of an entire swarm seemed restricted to a distance of about one foot in any direction. Within each swarm the males were in continual motion without moving in any set direction.

Stenamma brevicorne colonies seldom comprise more than 200-300 individuals. Probably not more than a few dozen alates are produced each season in a single colony. Thus, the large number of males in these swarms a total of at least 250 indicates that they came from a number of colonies. I attempted to find the sources of these alates without success. I had previously found this species nesting only in wooded areas in southeastern Michigan. However, parts of the field near the pond were cursorily examined for Stenamma nests, but none was found. I next examined a small area in Eber White Woods where I had previously collected S. brevicorne. Again I found no evidence of the species. Nevertheless, I believe that these alates most likely came from colonies in Eber White Woods and/or in other nearby woods.

The occurrence of exclusively male swarms is significant. Perhaps they developed because only males took part in flights on this day. The first flights of the season of some species of ants are mostly, if not exclusively, male ones. Possibly this is also true of S. brevicorne. On the other hand, both sexes may have taken part in the flights with the males forming the swarms first and the females being attracted later in the day (The only female of S. brevicorne that I have taken in flight was collected over a lawn at the Edwin S. George Reserve, Livingston County, Michigan, about 6:15 p.m. on May 30, 1956). If the latter hypothesis has any validity, it is necessary to account for the activities of the females from the time of flight until the time they join the swarms.

The swarms were observed in an environment quite distinct from that in which nests of this species were located in this region. Evidently the primary attraction for these alates was the top of a tree, not the major environment. Perhaps swarms form at the tops of trees in any local environment. They would, however, be most noticeable where the trees were small and scattered. Additional swarms may form over the trees in the woodlands, but, because of the height of the trees and the dense foliage, they would be difficult to observe. If the alates were attracted to the tree-tops as prominences, the fact that the trees surrounding the pond were isolated may be important. In a more or less open environment any tree-top would likely be a conspicuous promontory. In a woods probably only those individual trees which towered above the general canopy level would attract the alates.

Nomenclature

 * . Aphaenogaster brevicornis Mayr, 1886d: 447 (w.q.) U.S.A. (Virginia).
 * Type-material: syntype workers (number not stated), 1 syntype queen.
 * Type-locality: U.S.A.: Virginia (T. Pergande).
 * Type-depositories: NHMW, USNM.
 * Emery, 1895c: 299 (m.); Smith, M.R. 1957b: 167 (m.); Hauschteck, 1962: 219 (k.).
 * Combination in Stenamma: Emery, 1895c: 299.
 * Status as species: Cresson, 1887: 260; Dalla Torre, 1893: 100; Emery, 1895c: 299; Forel, 1901e: 347; Wheeler, W.M. 1903d: 166 (in key); Wheeler, W.M. 1904e: 302; Wheeler, W.M. 1905f: 382; Wheeler, W.M. 1906b: 4; Wheeler, W.M. 1910g: 565; Wheeler, W.M. 1916m: 585; Emery, 1921f: 54; Menozzi, 1932b: 311; Cole, 1936a: 36; Wing, 1939: 162; Wesson, L.G. & Wesson, R.G. 1940: 93; Buren, 1944a: 284; Creighton, 1950a: 136; Smith, M.R. 1951a: 794; Smith, M.R. 1957b: 163 (redescription); Smith, M.R. 1958c: 116; Smith, M.R. 1967: 352; Francoeur, 1977b: 206; Smith, D.R. 1979: 1358; Allred, 1982: 506; DuBois & LaBerge, 1988: 142; Wheeler, G.C., et al. 1994: 304; Bolton, 1995b: 393; Coovert, 2005: 40; Ellison, et al. 2012: 319.
 * Senior synonym of neoarcticum: Mayr, 1887: 628; Dalla Torre, 1893: 100; Emery, 1895c: 299; Creighton, 1950a: 136; Smith, M.R. 1957b: 165; Smith, M.R. 1958c: 116; Smith, D.R. 1979: 1358; Bolton, 1995b: 393.
 * Distribution: Canada, U.S.A.
 * neoarcticum. Stenamma neoarcticum Mayr, 1886d: 454 (w.q.m.) U.S.A. (New Hampshire, Virginia, California).
 * Type-material: 2 syntype queens, 2 syntype males.
 * [Note: 2 syntype workers (1 New Hampshire (T. Pergande), 1 Virginia (T. Pergande)) were excluded from the original type-series by Mayr, 1887: 628, and Emery, 1895c: 299, who identified them as specimens of S. brevicorne (Mayr), and not conspecific with the queens and males.]
 * Type-locality: U.S.A.: California, Oct. (by restriction of Creighton, 1950a: 138).
 * Type-depositories: NHMW, USNM.
 * [Misspelled as nearcticum by Emery, 1895c: 299, Wheeler, W.M. 1903d: 166, and others.]
 * Emery, 1895c: 299 (w.).
 * Subspecies of westwoodii: Emery, 1895c: 299.
 * Status as species: Cresson, 1887: 261; Dalla Torre, 1893: 121; Forel, 1901e: 347; Wheeler, W.M. 1903d: 166; Wheeler, W.M. 1910g: 565; Wheeler, W.M. 1917a: 519; Emery, 1921f: 54; Creighton, 1950a: 138; Smith, M.R. 1951a: 795.
 * Junior synonym of brevicorne: Smith, M.R. 1957b: 165; Smith, M.R. 1958c: 116; Smith, D.R. 1979: 1358; Bolton, 1995b: 393.
 * [Note: these synonymies with brevicorne all appear to relate to the excluded workers, not to the genuine syntype queens and males of neoarcticum. Snelling, R.R. 1973c: 25, identifies the queen and male syntypes of neoarcticum as synonymous with occidentale Smith, M.R., but gives occidentale priority, whereas neoarcticum should be the senior synonym.]

Worker
Smith (1957) - Length 2.7-3.8 mm. Funicular segments 2-6 as broad as, or broader than long; the four apical segments of the funiculus distinctly enlarged but not forming a well defined club; the length of the club exceeding the length of the remainder of the funiculus. Eye moderately large for a Stenamma, oblong, usually with approximately 8-10 ommatidia in its greatest diameter. The promesonotum, in profile, forming a rather long, uninterrupted arch in which the anterior half of the arch is distinctly more convex than the posterior half. Mesoepinotal impression usually strong and approximately twice as long as deep. Epinotum lower than the mesonotum and usually with a prominent, transverse welt near the base following the mesoepinotal impression. Base and declivity of the epinotum meeting to form a pair of short, but distinct, ruberculate spines which are approximately one-fourth to one-third the length of the base of the epinotum. Petiolar node viewed posteriorly with dorsally converging sides and truncate or feebly rounded superior border. Postpetiolar node viewed from above, stout, of variable shape, usually subspherical but occasionally subrectangular or transversely elliptical.

Frontal area and much of the clypeus smooth. Frontal region bearing fine, longitudinal and posteriorly diverging striae. Cheeks mostly longitudinally rugulose-punctate. Promesonotum usually largely, longitudinally rugulose, with the anterior border of the pronotum transversely rugulose. Dorsal surface of the epinorum with variable sculpture but quite commonly irregularly rugulose or rugulose-reticulate. Sides of thorax mostly rather coarsely, longitudinally rugulose with the exception of the lower part of the mesopleuron which is commonly reticulate-punctate. Base of gaster with distinct but short, longitudinal rugulae which are apparently never as long as the postpetiole. Petiolar and postpetiolar nodes mostly punctate or rugulose-punctate.

Head (with the exceptions of the mandibles, clypeus and frontal area), thorax, petiole and postpetiole, subopaque. Gaster smooth and shining. Propleuron occasionally somewhat shining.

Body with rather abundant, suberect to erect, grayish or light yellowish hairs of variable length. Head, and especially the gaster, usually more hairy: than the thorax. Hairs on appendages abundant but not always very closely appressed.

Head, thorax, petiole and postpetiole dark brown; antennae and legs light brown or yellowish brown. Gaster brown or blackish with the apex and the base lighter thus causing the dark area to appear as a more or less distinct, transverse band.

The head varies very noticeably in shape; with some individuals it is slender, very distinctly longer than broad whereas in other individuals the head is almost square but never quite as broad as long. The number of ommatidia in the greatest diameter of the eye is also subject to considerable variation, usually there are approximately 8-10 but one individual had only 4 or 5. Although it is common for the epinotum to bear a transverse welt near the base, the welt may be obscure or even lacking in some individuals. The postpetiolar node although usually subspherical may sometimes be transversely elliptical or even subrectangular. The sculpture on the promesonotum is usually largely longitudinally rugulose, occasionally there are individuals in which the rugulae have an arched or oblique trend or even an irregular, indescribable pattern. The color of the head, thorax, petiole and postpetiole is quite commonly reddish brown but often may be a light brown. Although the gaster is usually darker in the middle than at the base and tip there are individuals in which the base may be as dark as the median area.

Queen
Smith (1957) - Similar to the worker except as described below.

Length 3.4-4.2 mm. Ocelli small, yellowish, not noticeably protruding above the general surface of the head, the anterior ocellus located from 0.3-0.4 mm posterior to the frontal area. Greatest diameter of the eye ranging from 0.2-0.25 mm and with 14-18 ommatidia. Antennal scape failing to attain the posterior border of the head by its greatest diameter or less, funicular segments 2-7 as broad as, or broader than long; last segment of the club not as long as the combined lengths of the three preceding segments. Thorax 1.35-1.6 mm in length from the anterior border of the pronotal collar to the apices of the epinotal spines, widest anterior to the aniculations of the anterior wings where it measures 0.65-0.75 mm. Epinotal spines, in profile, variable in size and shape, usually stout and prominent and ranging from 0.10-0.15 mm in length; from above, the spines are approximately one-fourth to one-third as long as their interapical distance. Anterior wing grayish to yellowish depending upon the light, normally containing a closed cubital and a closed discoidal cell and a well developed stigma; without vein rom but with MB present, occasionally one or both of the anterior wings may have a partly or fully closed second cubital cell. MB is seldom absent but an occasional individual may lack this vein in one or both anterior wings. Upon infrequent occasions the discoidal cell may be only partly closed in one or both wings. Petiolar peduncle, in profile, often rather short and stout, the node subangular, slightly higher than long; postpetiole, in profile, approximately as high as long. Postpetiolar node, from above, convex, wider than long.

Scutellum and much of the mesonotum largely longitudinally rugulose-punctate. Pronotum, exclusive of the collar, largely transversely rugulose-punctulate. Area above the epinotal spines transversely rugulose with punctulate interspaces, the sculpturing often extending well down below the spines, occasionally however the area below the spines may be smooth or almost so.

Head, thorax, petiole, and postpetiole ranging from a light brown or reddish brown through dark brown to almost blackish brown.

Variation occurs mostly in the length, proportion, and color of the body. The variation of the wings is apparently less than other types of variation; it commonly consists in a partly or fully dosed extra cubital cell, lack of Mf3 or in the presence of a partly dosed discoidal cell.

Male
Smith (1957) - Length 3-3.5 mm. Ocelli distinct but not prominent or protruding above the general surface of the head, the anterior ocellus approximately 0.05 mm in width and located 0.2-0.25 mm posterior to the frontal area. Frontal area small but distinct, longer than broad. Frontal carinae short, 0.15-0.20 mm in length, subparallel, with scarcely any lobes. Eye large and prominent, 0.25-0.30 mm through its greatest diameter. Antennal scape (excluding the pedicel) approximately 0.28-0.37 mm in length, approximately as long as the combined lengths of the first four to six funicular segments; last funicular segment approximately as long as the combined lengths of the two preceding segments. Middle of the dorsal surface of the clypeus with a flattened area or impression that is longitudinal in direction. Mandible rather small, subtriangular. Thorax 1.1-1.5 mm in length from the anterior border of the pronotal collar to the apices of the epinotal spines, greatest width 0.6-0.75 mm just anterior to the articulations of the anterior wings. Mayrian furrows well developed, the parapsidal sutures present but best seen only in certain lights. In profile, base of epinotum depressed or flattened but the surface not horizontal, the base and declivity meeting on each side to form a broad tubercle or angle. Petiole, in profile, peduriculate, with a low not strongly convex node, which is approximately as long as the peduncle. Postpetiole, in profile, larger than the petiolar node, approximately as high as long. From above, postpetiolar node as broad as, or broader than long, usually with the appearance of being broader than long.

Head varying from rather densely and finely punctulate to reticulate-punctulate, the front with fine longitudinal striae; base of epinotum and sides of the petiole and postpetiole largely punctulate. Mesonotum and scutellum largely longitudinally rugulose-reticulate and punctate or punctulate, the area between the Mayrian furrows less heavily sculptured and therefore more shining than the mesonotum. Mesopleuron usually more weakly sculptured than the side of the epinotum which is irregularly rugulose-reticulate and punctulate. Gaster, and dorsal surface of the postpetiolar node, smooth and shining.

Pilosity light yellowish or grayish, moderately abundant, consisting of hairs of variable length which are suberect to erect.

Body brown to brownish black; mandibles, antennae, legs and apex of gaster light brown to yellowish, usually distinctly lighter than the remainder of the body.

Variations occur in the size of the male as well as in the proportions of the body. Individuals have been noted with the following abnormalities in the venation of the anterior wings: one or both wings with either a partly or else a fully developed extra cubital cell; also one or both wings lacking a dosed discoidal cell or else having only a partly dosed discoidal cell.

Etymology
Morphological. Presumably brevi L., short and corn L., horn, are a reference to the spines on the propodeum.

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 * 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, G.C., J. Wheeler and P.B. Kannowski. 1994. CHECKLIST OF THE ANTS OF MICHIGAN (HYMENOPTERA: FORMICIDAE). Great Lakes Entomologist 26:1:297-310
 * Wing M. W. 1939. An annotated list of the ants of Maine (Hymenoptera: Formicidae). Entomological News 50:161-165.