Cephalotes atratus

Within Cephalotes this species is the largest, one of the most abundant, and one of the most frequently encountered. The latter is due, in part, to the ability of C. atratus to thrive in urban areas. Mature colonies are comprised of several thousand individuals that can seemingly cover the trunk of the tree they inhabit when they are foraging.

This is one of the very few ants able to consume pollen, obtained by licking wind-transported pollen from leaves. The protein-rich internal part of the pollen is digested in the crop (foregut) and indigestible membranes are eliminated as 'infrabuccal pellets' through the mouth (Baroni Urbani & de Andrade 1997).

Identification
Note that nematode-infected workers have a red rather than black gaster (Poinar & Yanoviak, 2008).

Distribution based on Regional Taxon Lists
Neotropical Region: Argentina, Belize, Bolivia, Brazil, Colombia, Costa Rica, Ecuador, French Guiana, Guyana, Panama, Paraguay, Peru, Suriname, Trinidad and Tobago, Uruguay, Venezuela.

Biology
DaRocha et al. (2015) studied the diversity of ants found in bromeliads of a single large tree of Erythrina, a common cocoa shade tree, at an agricultural research center in Ilhéus, Brazil. Forty-seven species of ants were found in 36 of 52 the bromeliads examined. Bromeliads with suspended soil and those that were larger had higher ant diversity. Cephalotes atratus was found in 4 different bromeliads but was associated with twigs and bark cavities, rather than suspended soil or litter, of the plants.

Associations with other organisms
A known host for the fungi, , and  (Araujo et al., 2018; Shrestha et al., 2017), and the nematode  (Poinar & Yanoviak, 2008; Verble et al., 2012; Hughes et al., 2008). Note that in nematode-infected workers the gaster changes from black to red.

Images from AntWeb
Minor

Images from AntWeb
Major

Nomenclature

 * . Formica atrata Linnaeus, 1758: 581 (w.) South America (no state data).
 * Type-material: holotype (?) worker.
 * [Note: no indication of number of specimens is given.]
 * Type-locality: “America meridionali” (no further data) (Rolander).
 * Type-depository: ZMLS.
 * [Misspelled as atratulus by Santschi, 1929d: 301.]
 * Latreille, 1802c: 274 (q.); Spinola, 1851b: 47, Spinola, 1853: 63 (m.); Kempf, 1951: 116 (q.m.), Wheeler, G.C. & Wheeler, J. 1954b: 154 (l.).
 * Combination in Cryptocerus: Latreille, 1803: 280; Fabricius, 1804: 418; Latreille, 1809: 132; Leach, 1815: 147;
 * combination in Cryptocerus (Cephalotes): Wheeler, W.M. 1916c: 12;
 * combination in Cephalotes: Latreille, 1802a: 358; Emery, 1914c: 39; Santschi, 1916e: 381; Santschi, 1920f: 148; Smith, M.R. 1949c: 19.
 * Status as species: Linnaeus, 1767: 965; De Geer, 1773: 609; Fabricius, 1775: 395; Fabricius, 1782: 493; Retzius, 1783: 76; Gmelin, 1790: 2803; Christ, 1791: 516; Olivier, 1792: 500; Fabricius, 1793: 363; Latreille, 1802c: 272; Latreille, 1803: 280; Fabricius, 1804: 418; Gravenhorst, 1807: 287; Latreille, 1809: 132; Leach, 1815: 147; Latreille, 1817a: 526; Klug, 1824: 200; Lepeletier de Saint-Fargeau, 1835: 170; Guérin-Méneville, 1844a: 426; Spinola, 1851b: 47; Spinola, 1853: 63; Smith, F. 1853: 215; Smith, F. 1858b: 187; Mayr, 1862: 757; Smith, F. 1862b: 35; Smith, F. 1862d: 408; Roger, 1862c: 291; Mayr, 1863: 405; Roger, 1863b: 38; Mayr, 1865: 114; Emery, 1878a: ix (in list); Mayr, 1884: 37; Emery, 1890a: 68; Dalla Torre, 1893: 141; Emery, 1894e: 4; Forel, 1895b: 133; Forel, 1899c: 48; Forel, 1899d: 273; Forel, 1901h: 50; Wheeler, W.M. 1905b: 128; Emery, 1906c: 170; Forel, 1906d: 235; Forel, 1907e: 3; Wheeler, W.M. 1908a: 144; Forel, 1909a: 257, 266; Forel, 1912e: 199; Bruch, 1914: 218; Emery, 1914c: 39; Mann, 1916: 448; Wheeler, W.M. 1916c: 12; Wheeler, W.M. 1916d: 326; Crawley, 1916b: 371; Santschi, 1916e: 381; Luederwaldt, 1918: 40; Wheeler, W.M. 1918b: 26; Santschi, 1920f: 148; Forel, 1921b: 134; Wheeler, W.M. 1922c: 12; Wheeler, W.M. 1923a: 4; Emery, 1924d: 303; Wheeler, W.M. 1925a: 36; Borgmeier, 1927c: 114; Borgmeier, 1934: 103; Menozzi, 1935b: 197; Eidmann, 1936b: 81; Santschi, 1939f: 163; Wheeler, W.M. 1942: 207; Smith, M.R. 1949c: 19; Kempf, 1951: 114 (redescription); Kempf, 1958a: 10; Kempf, 1961b: 514; Kempf, 1970b: 335; Kempf, 1972a: 75; Zolessi, et al. 1988: 4; Brandão, 1991: 337; Bolton, 1995b: 140; De Andrade & Baroni Urbani, 1999: 111 (redescription); Wild, 2007b: 31; Bezděčková, et al. 2015: 115; Sandoval-Gómez & Sánchez-Restrepo, 2019: 904; Oliveira, Powell & Feitosa, 2021: 19.
 * Senior synonym of crassispina: Kempf, 1951: 115; Kempf, 1958a: 10; Kempf, 1972a: 75; Bolton, 1995b: 140; De Andrade & Baroni Urbani, 1999: 111.
 * Senior synonym of decemspinosus: Oliveira, Powell & Feitosa, 2021: 19 (by implication).
 * Senior synonym of dubitatus: Smith, F. 1858b: 188; Mayr, 1863: 405; Roger, 1863b: 38; Mayr, 1865: 114; Dalla Torre, 1893: 141; Forel, 1899c: 48; Emery, 1924d: 304; Borgmeier, 1927c: 114; Kempf, 1951: 115; Kempf, 1972a: 75; Bolton, 1995b: 140; De Andrade & Baroni Urbani, 1999: 111.
 * Senior synonym of erectus: De Andrade & Baroni Urbani, 1999: 112.
 * Senior synonym of marginatus: Oliveira, Powell & Feitosa, 2021: 19.
 * Senior synonym of nitidiventris: Kempf, 1951: 115; Kempf, 1972a: 75; Bolton, 1995b: 140; De Andrade & Baroni Urbani, 1999: 111.
 * Senior synonym of quadridens: Retzius, 1783: 76; Latreille, 1802c: 272; Fabricius, 1804: 419; Latreille, 1817a: 526; Klug, 1824: 200; Smith, F. 1853: 215; Smith, F. 1858b: 188; Mayr, 1863: 405; Roger, 1863b: 38; Mayr, 1865: 114; Dalla Torre, 1893: 141; Forel, 1895b: 133; Forel, 1899c: 48; Wheeler, W.M. 1908a: 144; Emery, 1924d: 303; Borgmeier, 1927c: 114; Kempf, 1951: 115; Kempf, 1958a: 10; Kempf, 1972a: 75; Bolton, 1995b: 140; De Andrade & Baroni Urbani, 1999: 111.
 * Senior synonym of rufiventris: Kempf, 1951: 115; Kempf, 1958a: 10; Kempf, 1972a: 75; Bolton, 1995b: 140; De Andrade & Baroni Urbani, 1999: 111.
 * Senior synonym of spissus: Oliveira, Powell & Feitosa, 2021: 19 (by implication).
 * Material of the unavailable name dehnowi referred here by Kempf, 1951: 115; Kempf, 1972a: 75; Bolton, 1995b: 140.
 * Distribution: Argentina, Belize, Bolivia, Brazil, Colombia, Costa Rica, Ecuador, French Guiana, Guyana, Panama, Paraguay, Peru, Suriname, Trinidad, Uruguay, Venezuela, Virgin Is.
 * crassispina. Cephalotes atratus st. crassispina Santschi, 1920f: 148 (w.) BRAZIL (Mato Grosso), ARGENTINA (Misiones, Formosa), PARAGUAY.
 * Type-material: syntype workers (number not stated).
 * [Note: De Andrade & Baroni Urbani, 1999: 111, report 7 syntypes.]
 * Type-localities: Brazil: Mato Grosso (F. Silvestri), Argentina: Misiones (E.R. Wagner), Argentina: Formosa (Joergensen), Paraguay: Sta Trinidad (Jorgensen).
 * Type-depository: NHMB.
 * Subspecies of atratus: Borgmeier, 1927c: 115.
 * Junior synonym of atratus: Kempf, 1951: 115; Kempf, 1958a: 10; Kempf, 1972a: 75; Bolton, 1995b: 140; De Andrade & Baroni Urbani, 1999: 111.
 * decemspinosus. Cephalotes decemspinosus Santschi, 1920f: 148 (w.) FRENCH GUIANA.
 * Type-material: lectotype worker (by designtion of Kempf, 1951: 125), 9 paralectotype workers.
 * Type-locality: lectotype French Guiana (“Guyane”): St Jean du Maroni (E. Le Moult); paralectotypes with same data.
 * Type-depositories: MZSP (lectotype); NHMB (paralectotypes).
 * Status as species: Kempf, 1951: 125; Kempf, 1972a: 76; Bolton, 1995b: 140.
 * Junior synonym of marginatus: De Andrade & Baroni Urbani, 1999: 124.
 * Junior synonym of atratus: Oliveira, Powell & Feitosa, 2021: 19 (by implication as marginatus synonymised under atratus).
 * dubitatus. Cryptocerus dubitatus Smith, F. 1853: 216, pl. 20, fig. 1 (m.) BRAZIL (no state data).
 * Type-material: holotype male.
 * Type-locality: Brazil: (no further data) (H.W. Bates?).
 * Type-depository: unknown (holotype lost).
 * [Note: holotype not in BMNH or OXUM, according to De Andrade & Baroni Urbani, 1999: 111.]
 * Junior synonym of atratus: Smith, F. 1858b: 188; Mayr, 1863: 405; Roger, 1863b: 38; Mayr, 1865: 114; Dalla Torre, 1893: 141; Forel, 1899c: 48; Emery, 1924d: 304; Borgmeier, 1927c: 114; Kempf, 1951: 115; Kempf, 1972a: 75; Bolton, 1995b: 141; De Andrade & Baroni Urbani, 1999: 111.
 * erectus. Cephalotes atratus subsp. erectus Kempf, 1951: 124.
 * Type-material: 4 syntype workers.
 * Type-locality: Panama: Darien (F. Geay).
 * Type-depository: MNHN.
 * [First available use of Cephalotes atratus st. quadridens var. erecta Santschi, 1920f: 149 (w.q.) PANAMA; unavailable (infrasubspecific) name.]
 * Subspecies of atratus: Kempf, 1972a: 75; Bolton, 1995b: 141.
 * Junior synonym of atratus: De Andrade & Baroni Urbani, 1999: 112.
 * marginatus. Cryptocerus marginatus Fabricius, 1804: 419 (q.) GUYANA.
 * Type-material: holotype queen.
 * Type-locality: “America meridionali”: (no further data) (Dom. Smidt. Mus. Dom. Lund).
 * [Note: De Andrade & Baroni Urbani, 1999: 124, add that the holotype is labelled: “Essequibo Smidt Mus. J. Lund”; hence type-locality is Guyana: Essequibo.]
 * Type-depository: ZMUC.
 * De Andrade & Baroni Urbani, 1999: 124 (m.).
 * Combination in Cephalotes: Emery, 1924d: 304; De Andrade & Baroni Urbani, 1999: 124.
 * Junior synonym of atratus: Klug, 1824: 200; Smith, F. 1853: 215; Smith, F. 1858b: 188; Roger, 1862c: 291; Mayr, 1863: 405; Roger, 1863b: 38; Mayr, 1865: 114; Dalla Torre, 1893: 141; Forel, 1899c: 48; Emery, 1924d: 304; Borgmeier, 1927c: 114; Kempf, 1972a: 75; Bolton, 1995b: 141.
 * Status as species: De Andrade & Baroni Urbani, 1999: 124 (redescription); Sandoval-Gómez & Sánchez-Restrepo, 2019: 912.
 * Junior synonym of atratus: Oliveira, Powell & Feitosa, 2021: 19.
 * nitidiventris. Cephalotes atratus var. nitidiventris Santschi, 1920f: 148 (w.) FRENCH GUIANA.
 * Type-material: 3 syntype workers.
 * Type-localities: 2 workers French Guiana: Cayenne, 1905 (E. Grouvelle), 1 worker French Guiana: Rivière Lumier (F. Geay).
 * Type-depositories: MNHN, NHMB.
 * Junior synonym of atratus: Kempf, 1951: 115; Kempf, 1951: 115; Kempf, 1972a: 75; Bolton, 1995b: 141; De Andrade & Baroni Urbani, 1999: 111.
 * quadridens. Formica quadridens De Geer, 1773: 609, pl. 31, figs.17-20 (w.) SURINAME.
 * Type-material: syntype workers (number not stated).
 * Type-locality: Suriname: (no further data) (Rolander).
 * Type-depository: MNHN.
 * Combination in Cryptocerus: Fabricius, 1804: 419;
 * combination in Cephalotes: Santschi, 1919f: 44.
 * Status as species: Fabricius, 1804: 419; Santschi, 1919f: 44.
 * Subspecies of atratus: Santschi, 1920f: 149; Forel, 1922: 97; Borgmeier, 1927c: 115; Santschi, 1939f: 163.
 * Junior synonym of atratus: Retzius, 1783: 76; Latreille, 1802c: 272; Fabricius, 1804: 419; Latreille, 1817a: 526; Klug, 1824: 200; Smith, F. 1853: 215; Smith, F. 1858b: 188; Mayr, 1863: 405; Roger, 1863b: 38; Mayr, 1865: 114; Dalla Torre, 1893: 141; Forel, 1895b: 133; Forel, 1899c: 48; Wheeler, W.M. 1908a: 144; Emery, 1924d: 303; Borgmeier, 1927c: 114; Kempf, 1951: 115; Kempf, 1958a: 10; Kempf, 1972a: 75; Bolton, 1995b: 141; De Andrade & Baroni Urbani, 1999: 111.
 * rufiventris. Cryptocerus atratus ab. rufiventris Emery, 1894e: 4 (in text) (w.) ARGENTINA (Tucumán).
 * Type-material: holotype worker.
 * Type-locality: Argentina: Tucumán (A. Borelli).
 * Type-depository: MSNG.
 * As unavailable (infrasubspecific) name: Santschi, 1920f: 149 (in key).
 * Junior synonym of atratus: Kempf, 1951: 115; Kempf, 1958a: 10; Kempf, 1972a: 75; Bolton, 1995b: 141; De Andrade & Baroni Urbani, 1999: 111.
 * spissus. Cephalotes decemspinosus var. spissus Santschi, 1929d: 302 (w.) SURINAME.
 * Type-material: syntype workers (number not stated).
 * [Note: De Andrade & Baroni Urbani, 1999: 124, record 3 syntypes.]
 * Type-locality: Suriname (“Guyane hollandaise”): Lucie River, Gebrid, vii.,viii.1928 (Corporaal).
 * Type-depository: NHMB.
 * Junior synonym of decemspinosus: Kempf, 1951: 125; Kempf, 1972a: 76; Bolton, 1995b: 141.
 * Junior synonym of marginatus: De Andrade & Baroni Urbani, 1999: 124.
 * Junior synonym of atratus: Oliveira, Powell & Feitosa, 2021: 19 (by implication as marginatus synonymised under atratus).

Type Material


de Andrade and Baroni Urbani (1999):
 * Formica quadridens. Type locality: Surinam.
 * Cryptocerus dubitatus. Male. Type locality: Brazil. Type material presumably lost (neither in the BMNH nor in ).
 * Cryptocerus atratus ab. Rufiventris. Worker. Type locality: Tucuman (Argentina). Type material not available for the present dtudy.
 * Cephalotes atratus var. nitidiventris. Worker. Type locality: Cayenne and Lumier River (French Guyana). Type Material: 2 syntype workers labeled “America, Cayena 1905, E. Grouvelle” (sic) in, examined.
 * Cephalotes atratus stirps crassispina. Worker. Type localities: Mato Grosso (Brazil), Formosa (Misiones, Argentina), and Santa Trinidad (Paraguay). Type material 3 workers from Urucuru, Mato Grosso (Silvestri), 3 workers from Formosa (Argentina, Joergensen), 1 worker from Paraguay without further data, all in NHMB, examined.
 * Cephalotes marginatus. Gyne. Type locality: "in America meridionali", actually Essequibo (Guyana). Type material: Holotype winged gyne (wings detached) labelled "Essequibo, Smidt, Mus. J. Lund, Cryptocerus atratus (Fabr.) Linn., marginatus Fabr. ", in, examined.
 * Cephalotes decemspinosus Worker. Type locality: St. Jean du Maroni (French Guyana). Type material: 4 syntype workers from St. Jean du Maroni, French Guyana, Le Moult in (examined), lectotype worker, in Coll. Borgmeier, designated by Kempf, 1951 same data as the syntypes.
 * Cephalotes decemspinosus var. spissus (sic). Worker. Type locality: surroundings of the Lucie River (Surinam). Type material: 3 syntype workers labelled Suriname-Exped., Lucie-riv. -Gebied, VII -VIII. 1 926,, examined.

Worker
Kempf (1951) - Monomorphic. Length 8-14 mm. Black; the following fuscous ferruginous: two semitranslucid spots on each frontal carinae, tips of tarsi.

Head subopaque; finely reticulate-punctate. Mandibles densely and finely rugose and punctured. Frontal carinae slightly diverging caudad, scarcely upturned laterad, vestigially crenulate anteriorly with only a few projecting setae, remaining part not crenulate and without setae. Occiput continuous with upper surface of head, inclined ventrad; its angles with a pair of spines. Vertex without spines nor swellings. Lower occipital border straight, carinulate. Upper surface of head and cheeks covered with sparse oval foveolae, containing a short, scalelike, silvery, decumbent hair. Lower surface of head more coarsely reticulate-rugose and foveolate. Apical segment of maxillary palpi somewhat longer than the preceding.

Thorax subopaque. Shoulders obtusely dentate. Scapular spines strong, obliquely inclined backward, and laterad, subacuminate, the apical half more or less recurved, with a small tooth projecting from the anterior face at the bent. Median pronotal spines usually minute, vestigial or absent. Promesonotum moderately longitudinally convex, immarginate laterad. Promesonotal suture distinct. Mesonotum flat, with a minute tubercle laterally toward the posterior angle. Mesoepinotal suture distinct and impressed. The basal face of the epinotum slightly inclined forward, immarginate laterad, slightly constricted in front of the oblique, more or less divergent, straight to slightly curved epinotal spines. Entire thorax finely reticulate-punctate, with sparse setigerous foveolae, crowded and more deeply impressed on pronotum, very sparse on the pleura, leaving free spaces, which arc mostly longitudinally rugose. Mesopleura with a small tooth above the anterior corner of the mid coxae. Declivous face mostly without coarse sculpture. Outer surface of fore coxae transversely striated. Legs without foveolae, but with appressed setae. Oblique setae on apical end of femora, tibiae and on tarsi. Petiole shorter than broad, subcuboidal, denesely foveolate above and on the sides, with a small sidewards projecting tubercle or tooth on each anterior corner. Ventral face strongly keeled, ending anteriorly in an obtuse tooth. Postpetiole more than 1.5 times as broad as long. Sides with a conspicuous, anteriorly angulate, posteriorly rounded swelling, the sides converging caudad. Upper face flat, with a pair of small tubercles on the anterior border. Ventral face with a large, truncate, rounded or bifid lobe, projecting ventrad and somewhat cehalad. Sculpture as on petiole.

Gaster elliptical, subopaque to subfulgid, finely reticulate-punctate, covered with larger, elongate, shallow grooves, containing a decumbent hair. Posterior half of first sternites and the exposed portion of the tergites and sternites 2-4 with sparse, suberect, pale-brownish hairs.

de Andrade and Baroni Urbani (1999) - Measurements (in mm) and indices: TL 7.64-15.10; HL 1.60-3.30; HW 2.08-4.50; EL 0.48-0.80; PW 1.88-4.80; PeW 0.50-1.00; PpW 0.59-1.08; HBaL 1.32-2.30; HBaW 0.34-0.51; CI 130.0-136.4; PI 93.7-112.9; PPel 349.3-480.0; PPpl 317.9-444.4; HBal 22.2-25.7.

Queen
Kempf (1951) - Length up to 23 mm. Somewhat similar to the worker. Black, subopaque. Head somewhat broader than long. Mandibles strongly rugose. Clypeus much broader than long, subtriangular. Frontal carinae distinctly converging anteriorly, not parallel not upturned laterally, slightly crenulate on anterior third, with a few projecting setae. Vertex with a pair of obtuse swellings, just in front of the subtruncate occiput. Occipital angles bituberculate. Cheeks immarginate below. culpture as in worker. Thorax subopaque. Pronotum immarginate cephalad and laterad. Shoulders low, bluntly angulate. Scapular spines short, stout, acute. Transverse crest of pronotum vestigial, not carinate, two median pronotal spines short, obtuse. Mesopleura with a small tooth above the mid coxa. Epinotum narrow, immarginate, laterad, spines short, icrassate at base, apices slightly diverging, their interapical distance less than 1/2 of the maximum width of thorax. Declivous face twice as long as basal face of epinotum. Sculpture as on head. Basitarsus of mid and hind leg flattened and broadened, not as thin as in worker. Petiole short, about twice as broad as long, the upper face presenting a transverse keel. Postpetiole longitudinally, greatly convex, with a large, blunt, rounded swelling laterad, about twice as broad as long. Ventral lobe less conspicuous than in worker. Both peduncular segments coarsely and rather densely foveolate. Gaster as in worker, more elongate, sides subparallel, the microsculpture less prominent; first tergite rather fulgid discad, sparsely foveolate, the foveolae smaller than on thorax and on head. Wings infumated. Fore wing with fuscous stigma and a dark streak along the subcostal vein. Marginal cell closed and appendiculate. Submarginal cell shorter than in Zacryptocerus clypeatus. Transverse cubital vein absent, the cubital vein touching the marginal cell. Second abscissa of subcostal vein shorter than the section of basal vein that encloses the submarginal cell. Occasionally stubs projecting from the cubital vein into the ubmarginal cell and the apical field.

de Andrade and Baroni Urbani (1999) - Measurements (in mm) and indices: TL 15.06-19.50; HL 2.76-3.65; HW 3.32-4.50; EL 0.64-0.80; PW 2.60-4.50; PeW 0.96-1.40; PpW 1.16-1.60; HBaL 1.76-2.00; HBaW 0.486-0.60; CI 120.0-126.5; PI 93.3-127.7; PPeI 270.8-321.4; PPpI 224.1-281.2; HBaI 27.3-30.0.

Male
Kempf (1951) - Length 13.2 mm. Median head length 1.53 mm. Weber's length of thorax 4.05 mm. Black: the following brunneous: antennae, coxae, trochanters, base of femora. Orange-brown: gaster, femora, tibiae, tarsi.

Head, without mandibles much broader than long (103 : 63). Mandibles finely rugulose and punctate; chewing border with distinct apical and preapical tooth. Clypeus transverse, anterior face truncate, subperpendicular to upper surface of head, posterior half at right angles to it; horizontal. Anterior border straight, posterior border marked by a deep transverse depression in front of the antennal sockets. Entire clypeus finely shagreened. Frontal carinae scarcely expanded, divergent caudad, fading out before the posterior border of eye. Antennal scrobes indistinct. Ocelli on vertex, conspicuous, subequal to diameter of second funicular segment. First funicular segment shorter than broad. Second funicular segment longer than the following, except of the apical segment.

Thorax subopaque, almost twice as long as maximum width (83 : 47). Pronotum immarginate laterad, with a minute tooth on the anterior angle, and a longer blunt tooth somewhat behind; rugose-reticulate and foveolate. Scutum convex with deeply impressed Y-shaped Mayrian furrows, antero-mesal part deeply and densely foveolate, postero-lateral portion more sparsely and shallowly foveolate. Scutellum similarly sculptured with anterolateral lobes almost constricted off by a deep transverse furrow immediately behind the anterior border. Basal face of epinotum shorter than declivous face, transversely convex, immarginate laterally, posterior corners with a sharp carinule, reticulate-rugose and foveolate. Declivous face finely reticulate, subfulgid, with a few faint, distinct rugulae. Sides of mesothorax finely shagreened anteriorly, sparsely, shallowly foveolate and rugose posteriorly. Basitarsi elongate, compressed.

Petiole subopaque, slightly broader than long, anterior face truncate, excavate, the sides somewhat converging behind, finely shagreened above, finely longitudinally rugose laterad, and reticulate-rugose antero-laterad. Postpetiole broader than long, sides greatly convex, subtuberculate, sculptured as petiole.

Gaster subopaque, short, fusiform, finely shagreened. Subgenital plate transverse, apical border rounded.

Wings leathery, infumated. Veins brunneous. Fore wings with fuscous stigma, and fuscous streak along the subcostal vein. Marginal cell closed and appendiculate. Discoidal cell extremely small. Transverse cubital vein present, cubital vein not touching the marginal cell.

Pilosity gold-brown. Antennae with dense fine pubescence. Long dense hair on head and thorax; sparse, appressed on first gastral tergite, erect on apical border of the gastral tergites and sternites.

Certain features are highly variable, even among males of the same nest. Among those are the occipital spines, which may form mere perpendicular carinule on each side, or even be quite obsolete. The same is true for the carinules on the posterior corner of the epinotum, which may be occasionally tooth-like. The sculpture in some instances is coarser than in the specimen described above. The following variability occurs in the fore wing: discoidal cell always small, sometimes minute, open or even completely absent. Stubs projecting into submarginal cell from various places of the recurrent and the cubital veins.

de Andrade and Baroni Urbani (1999) - Measurements (in mm) and indices: TL 9.78-13.21; HL 1.28-1.68; HW 1.68-2.16; EL 0.53-0.70; PW 1.60-2.24; PeW 0.72-0.92; PpW 0.88-0.96; HBaL 1.32-1.84; HBaW 0.18-0.26; CI 128.3-131.9; PI 96.4-105.0; PPeI 227.3-272.2; PPpI 208.3-233.3; HBaI 13.3-15.1.

Worker
de Andrade and Baroni Urbani (1999) - Moderately variable in size. Head with the sides (frontal carinae) subparallel, very weakly crenulate only on the anterior fourth and upturned laterally. Vertexal border continuing the frontal carinae and sloping ventrally, bearing two pairs of lateral spines, the distal pair ca. 1/2 longer than the proximal one, very weakly carinate to straight posteriorly. Eyes ca. at 6/7 of head length posteriorly and protruding from the ventral border of the antennal scrobes.

Mesosoma relatively slender. Pronotum making an angle of ca. 90° with the mesonotum, straight and inclined antero-ventrally at about 45° on the longitudinal body axis. Posterior pronotal third with two pairs of spines, the external pair almost continuing laterally the pronotal surface and long as half of the maximum head width; the median pair, perpendicular to the pronotum, is regularly much longer than in atratus, ca. 1/4 as long as the external pair. Promesonotal suture well marked. Mesonotum straight and flat in profile. Propodeal suture deeply impressed. Propodeum with basal and declivous faces subequal in length and forming an angle of ca. 90°. Basal face practically flat, passing into the sides with a round angle. Declivous face very weakly concave. Propodeal spines only slightly shorter than the external pronotal ones, obliquely pointing backwards. Metapleural-propodeal suture oblique and deeply impressed laterally.

Petiole with slightly concave anterior and slightly convex posterior faces, bearing two small, dorso-lateral denticles. Postpetiole subequal to petiole in width, narrower posteriorly, with a very short, perpendicular anterior face and nearly flat dorsal one; the angle between the anterior and superior faces bears a pair of short spines, slightly longer than the petiolar ones and directed upwards; ventrally, another pair of spines obliquely directed forwards, subequal to the dorsal ones in length but thinner.

Gaster elongate, incised at the articulation with the postpetiole and marginate anteriorly.

Mid and hind tibiae strongly compressed laterally anq with marginate external borders.

Pilosity and sculpture. Whole body, except the spines and the appendages covered by dense, superficial, oval pits. The pits on the head, mesosoma and pedicel bear a thick, truncate hair and can be interpreted as sensilla auricillica. The pits on the gaster are more superficial, each pit bearing simply pointed hairs (in atratus the deeper pits are generally limited to the head and pronotum). A few suberect, thick bristles on the anterior pair of vertexal spines, on both pairs of pronotal spines and on the pedicel. Numerous yellowish standing hairs on the posterior border of all gastral segments and at least on the distal half of the first gastral sternite. Legs with few, yellowish, subdecumbent or decumbent hairs, particularly on the coxae and on the tibiae.

Colour black moderately shining except the frontal carinae, ferrugineous.

Measurements (in mm) and indices: TL 9.60-17.00; HL 2.04-3.50; HW 2.60-5.20; EL 0.53-0.80; PW 2.60-5.00; PeW 0.68-1.08; PpW 0.75-1.28; HBaL 1.60-2.48; HBaW 0.38-0.60; CI 127.1-148.6; PI 88.2-110.6; PPeI 382.3-555.5; PPpI 346.7-510.2; HBaI 23.6-24.8.

Queen
de Andrade and Baroni Urbani (1999) - Head slightly wider than long, with the sides converging anteriorly and weakly crenulate only on their anterior fifth. Clypeus broadly concave. Vertexal border weakly concave, nearly straight. Abroad median dorsal depression of the head capsule at about 2/3 of its length. Frontal line reaching the median depression or continuing through it up to the median ocellus. Two pairs of lateral vertical spines subequal in size and disposed perpendicularly to the vertexal angles; a third pair of smaller, blunt, vertexal spines behind and between the pair ocelli. Eyes, in dorsal view, not covered by the frontal carinae. Ocelli deeply impressed and subequal in distance each one to the other two. Mandibles robust, serially toothed and transversely impressed at mid length. Antennal scape thick; funicular joints gradually increasing in size from the second to the last.

Mesosoma, in dorsal view, remarkably broadened at the height of the mesopleurae, with flat mesonotum and scutellum, both higher than the pronotum and propodeum in profile. Humeral spines developed and pointing perpendicularly to the major body axis, followed by a pair of minute denticles caudally. Medial pronotal spines much broader and shorter, obtuse to pointed. Propodeum with basal face gently sloping backwards, shorter than the declivous face, which is nearly straight. Propodeal spines robust, their dorsal surface complanary with the basal face of the propodeum, long 1/2 of their distance at the apex, diverging caudally.

Petiole sessile, slightly lower than the postpetiole, with anterior face flat and posterior face only slightly shorter than the anterior one, bearing only two weak dorsal protuberances and no spines. Postpetiole slightly broader than petiole, twice broader than long, bearing two more prominent dorsal protuberances and a broad ventral lamella.

Gaster incised at the articulation with the postpetiole, not marginate; in dorsal view only the first and part of the second tergites visible.

Legs with mid and hind tibiae compressed laterally and distally marginate on the external face.

Fore wings infuscate with very dark R+ Sc connected with a marked pterostigma. Rsf5 connected with R1. R1 almost reaching or entirely reaching and outlining the wing border. A, Cu-A, Cu-A1 and Mf4 well marked and nearly reaching the wing border. Hind wings with well marked R, M+CuA, 1A, CuA, r-m and M. Distal parts of 1A, CuA and M obsolete and nearly reaching the wing border.

Sculpture and pilosity as in the worker but more uniform over the whole body, with the pits more deeply impressed on the pedicel. Abundant, short, truncate, yellowish hairs on the terminal parts of the gastral segments and at least on the distal half of the first gastral sternite.

Colour uniformly black, more shining on the gaster, with a ferrugineous strip parallel to the frontal carinae where the integument is thinner and semi-transparent.

Measurements (in mm) and indices: TL 19.60-20.30; HL 3.28-3.70; HW 4.35-4.90; EL 0.83-0.88; PW 4.60-4.90; PeW 1.44-1.72; PpW 1.60-1.64; HBaL 2.00-2.20; HBaW 0.60-0.61; CI 127.0-132.6; PI 89.8-97.9; PPeI 315.8-331.1 ; PPpI 287.5-306.2; HBaI 27.3-30.5.

Male
de Andrade and Baroni Urbani (1999) - Head much broader than long, with the cheeks strongly converging anteriorly. Vertexal margin prolonging backwards into a neck. Vertexal angles diverging towards the posterior border of the eyes, with a median pair of obtuse, triangular denticles sometimes connected through a carina to the external pair denticles close to the eyes. Frontal carinae more or less raised over the head dorsum. Frontal line marked and reaching the median ocellus posteriorly. Clypeus moderately convex anteriorly and remarkably raised in the middle, its medial part much higher than the mandibular plane. Eyes strongly protruding on the sides of the head which is much narrower in front than behind the eyes. Ocelli delimiting a subtriangular area adjecting dorsally over the already prominent frontal area. Antennal scape thick, ca. two times longer than broad and three times longer than the first funicular joint; second funicular joint ca. 1.5 times longer than the scape; third funicular joint slightly shorter; remaining antennomeres thickening from the base to the apex. Mandibles elongate and weakly carinate laterally.

Mesosoma elongate, in dorsal view the sides continuously arched from the anterior pronotal border to the propodeum. Pronotal sides diverging backwards, sometimes with a pair of minute denticles. Mayrian furrows deeply impressed. Mesonotum and scutellum gently convex in sides view. Propodeum with basal and declivous faces making an angle of ca. 100° and separate by a ridge bearing, at its sides, two small teeth. Petiole and postpetiole subequal in length and width, with their maximum width anteriorly. Petiolar node subrectangular; its anterior face truncate and laterally delimited by a pair of denticles. Postpetiole subexagonal. Gaster elongate; its maximum width subequal to the maximum width of the thorax.

Sculpture. Head punctate and covered by irregular foveae and short rugosities; the foveae and the rugosities broader on the posterior part of the head dorsum and on the whole ventral part of the head. Mesosoma punctate and densely covered by foveae more regular than those on the ventral part of the head. Anterior half of the basal face of the propodeum with the same type of sculpture as on the posterior part of the head dorsum; posterior half of the basal face and anterior part of the declivous face with longitudinal rugosities, the rugosities transversal on the posterior part of the declivous face. Pleurae punctate and with longitudinal rugosities on the pro- and lower metapleurae, the rugosities thicker on the lower metapleurae. Meso and upper metapleurae covered by irregular foveae, sparser on the upper mesopleurae. Pedicel punctate and with very thin, irregular, longitudinal rugosities more impressed on the sides. First gastral tergite reticulate; remaining tergites and sternites superficially reticulate and shining. Legs punctate; the punctures more impressed on the tibiae.

Pilosity. Head and mesosoma covered by long, thick, pointed, erect, golden hairs. The same hairs, but rarer, on the sides of the petiole and postpetiole. Gaster, dorsally, only with few short hairs at the end of each segment, practically bald; ventrally, numerous, long, golden hairs as in the worker and in the gyne. Antennae only pubescent. Legs with similar but shorter and appressed, golden hairs.

Colour. Head, mesosoma, petiole and postpetiole, coxae and trochanters black. Gaster and remaining parts of the legs yellowish-brown; antennae brown and opaque.

Measurements (in mm) and indices: TL 11.34-13.88; HL 1.64-1.76; HW 1.80-2.16; EL 0.62-0.72; PW 1.72-2.16; PeW 0.84-0.96; PpW 0.98-1.08; HBaL 1.80-2.08; HBaW 0.25-0.30; CI 109.7-122.7; PI 98.1-104.6; PPeI 179.2-257.1; PPpI 172.0-204.1; HBaI 11.5-15.9.

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