Cephalotes pusillus

C. pusillus is one of the commonest and most widespread species of the genus, though its distribution is narrower than that of other common species like Cephalotes minutus and Cephalotes atratus. (de Andrade and Baroni Urbani 1999)

Identification
A member of the pusillus clade characterised in the worker and soldier by the less regular body foveae and by the narrow, semi-transparent gastral lamellae. (de Andrade and Baroni Urbani 1999)

Distribution based on Regional Taxon Lists
Neotropical Region: Argentina, Bolivia, Brazil, Colombia, Ecuador, French Guiana, Guyana, Paraguay, Peru, Suriname, Trinidad and Tobago, 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 pusillus was found in 5 different bromeliads but was associated with twigs and bark cavities, rather than suspended soil or litter, of the plants.

Koch et al. (2018) sampled this species in Caryocar barsiliense trees, in southeastern Brazil cerrado, as part of a study examining species interactions in ant-plants.

Spider Mimic and Predator
Aphantochilus rogersi is an an-mimicking spider that preys exclusively on cephalotine ants. The spiders oviposit in close proximity to nests of the model ant Cephalotes pusillus, and emergent spiderlings tend to remain in the vicinity of natal egg sacs. Females of A. rogersi actively defend their egg sacs against approaching workers of C. pusillus, but the latter may sometimes destroy the eggs. Feeding specialization on these ants is confirmed by more than 300 observations of young and adult A. rogersi carrying ant corpses in the field. Although A. rogersi possesses several behavioural traits which may reduce the risk of being injured by ants during subjugation, field and laboratory observations showed that social defence by C. pusillus may cause mutilation to the spiders. Tests in captivity revealed an ontogenetic change in the prey-capture techniques employed by A. rogersi. Early-instar spiderlings can apparently only seize the ant’s petiole tightly if they are able to approach the ant from the front. As the ant is paralysed, the spiderling positions itself vertically in relation to the substratum. Larger spiders, on the other hand, attack ants most frequently from behind, and seem better equipped to seize the ant’s petiole firmly with their larger chelicerae. Owing to their greater strength, late-instar spiders are able to lift the struggling ant aloft. The selection of a suitable oviposition site, the mother’s ability to defend herself and the eggs from nearby ants, and the capacity to capture and subdue ants safely from emergence to maturity, are regarded as crucial traits inherent in the mimetic and feeding specialization by A. rogersi. (Castanho & Oliveira, 1997)

Worker
Minor

Nomenclature

 * . Cryptocerus pusillus Klug, 1824: 202 (w.) BRAZIL (Pará, Rio de Janeiro), GUYANA.
 * Type-material: syntype workers (number not stated).
 * Type-localities: Brazil: Pará (“ex Museo de Hoffmannsegg”), Brazil: Rio de Janeiro (“sent by Dr de Olfers”), Guyana: Demerara (“from Hemmerich”).
 * Type-depository: MNHU (perhaps also in MRSN).
 * Emery, 1890b: 75 (s.q.); Emery, 1896h: 636 (m.); Wheeler, G.C. & Wheeler, J. 1954b: 156 (l.).
 * Combination in Cryptocerus (Paracryptocerus): Emery, 1915i: 192;
 * combination in Paracryptocerus: Kempf, 1951: 176;
 * combination in Paracryptocerus (Paracryptocerus): Kempf, 1958a: 14;
 * combination in Zacryptocerus: Brandão, 1991: 388;
 * combination in Cephalotes: Baroni Urbani, 1998: 316; De Andrade & Baroni Urbani, 1999: 172.
 * Status as species: Guérin-Méneville, 1844a: 426; Smith, F. 1853: 220; Smith, F. 1858b: 190; Smith, F. 1862d: 410; Mayr, 1863: 406; Roger, 1863b: 38; Dalla Torre, 1893: 144; Forel, 1895b: 134; Emery, 1896h: 636; Forel, 1905b: 158; Emery, 1906c: 170; Forel, 1906d: 234; Forel, 1907e: 3; Forel, 1908c: 354; Forel, 1908e: 69; Forel, 1909a: 257; Forel, 1912e: 201; Bruch, 1914: 218; Wheeler, W.M. 1916c: 12; Wheeler, W.M. 1916d: 326; Mann, 1916: 450; Crawley, 1916b: 371; Santschi, 1916e: 383; Luederwaldt, 1918: 41; Wheeler, W.M. 1918b: 26; Santschi, 1921h: 127; Wheeler, W.M. 1922c: 11; Wheeler, W.M. 1923a: 4; Emery, 1924d: 307; Wheeler, W.M. 1925a: 36; Borgmeier, 1927c: 116; Menozzi, 1935b: 197; Eidmann, 1936b: 83; Wheeler, W.M. 1942: 209; Kempf, 1951: 176 (redescription); Kempf, 1958a: 14; Kempf, 1961b: 514; Kempf, 1972a: 179; Brandão, 1991: 388; Bolton, 1995b: 427; De Andrade & Baroni Urbani, 1999: 172 (redescription); Wild, 2007b: 32; Bezděčková, et al. 2015: 116; Sandoval-Gómez & Sánchez-Restrepo, 2019: 913.
 * Senior synonym of brevispinosa: Kempf, 1951: 176; Kempf, 1958a: 14; Kempf, 1972a: 180; Brandão, 1991: 388; Bolton, 1995b: 427; De Andrade & Baroni Urbani, 1999: 172.
 * Senior synonym of caustica: Emery, 1924d: 307; Borgmeier, 1927c: 116; Kempf, 1951: 176; Kempf, 1958a: 14; Kempf, 1972a: 180; Brandão, 1991: 388; Bolton, 1995b: 427; De Andrade & Baroni Urbani, 1999: 172.
 * Senior synonym of elongatus: Roger, 1862c: 297; Mayr, 1863: 406; Roger, 1863b: 38; Dalla Torre, 1893: 144; Forel, 1895b: 134; Emery, 1924d: 307; Borgmeier, 1927c: 116; Kempf, 1951: 176; Kempf, 1972a: 180; Brandão, 1991: 388; Bolton, 1995b: 427; De Andrade & Baroni Urbani, 1999: 172.
 * Senior synonym of obtusus: Forel, 1911e: 262; Emery, 1924d: 307; Borgmeier, 1927c: 116; Kempf, 1951: 176; Kempf, 1972a: 180; Brandão, 1991: 388; Bolton, 1995b: 427; De Andrade & Baroni Urbani, 1999: 172.
 * Material of the unavailable name rufescens referred here by Kempf, 1951: 177; Kempf, 1958a: 14; Kempf, 1972a: 180; Brandão, 1991: 388.
 * Distribution: Argentina, Bolivia, Brazil, Colombia, Ecuador, Guyana, Paraguay, Peru, Trinidad, Venezuela.
 * brevispinosa. Cryptocerus pusillus var. brevispinosa Santschi, 1921h: 127, fig. 2 (w.) ARGENTINA (Misiones, Córdoba), PARAGUAY, BRAZIL (Rio de Janeiro).
 * Type-material: syntype workers (number not stated).
 * Type-localities: Argentina: Misiones, vic. San Ignacio, Villa Lutecia (E.R. Wagner), Argentina: Cerro Cora (von Steiger), Argentina: Córdoba (C. Bruch), Paraguay: (no further data) (Fiebrig), Brazil: Rio de Janeiro, Alto da Boa Vista (E.R. Wagner).
 * Type-depository: NHMB.
 * Subspecies of pusillus: Borgmeier, 1927c: 116.
 * Junior synonym of pusillus: Kempf, 1951: 176; Kempf, 1958a: 14; Kempf, 1972a: 180; Brandão, 1991: 388; Bolton, 1995b: 424; De Andrade & Baroni Urbani, 1999: 172.
 * caustica. Formica caustica Kollar, in Pohl & Kollar, 1832: 17, fig. 12 (w.) BRAZIL (no state data).
 * Type-material: holotype (?) worker.
 * [Note: no indication of number of specimens is given.]
 * Type-locality: Brazil: (no further data).
 * Type-depository: unknown.
 * [Note: neither Pohl nor Kollar are mentioned in Horn & Kahle, 1935, 1936.]
 * Combination in Cryptocerus: Guérin-Méneville, 1844a: 426.
 * Status as species: Smith, F. 1853: 222; Smith, F. 1858b: 191; Mayr, 1862: 764; Smith, F. 1862d: 410; Mayr, 1863: 405; Roger, 1863b: 38; Emery, 1878a: ix (in list).
 * Junior synonym of minutus: Emery, 1890b: 74; Dalla Torre, 1893: 143; Forel, 1895b: 134; Forel, 1899c: 50.
 * Junior synonym of pusillus: Emery, 1924d: 307; Borgmeier, 1927c: 116; Kempf, 1951: 176; Kempf, 1958a: 14; Kempf, 1972a: 180; Brandão, 1991: 388; Bolton, 1995b: 424; De Andrade & Baroni Urbani, 1999: 172.
 * [Note: De Andrade & Baroni Urbani, 1999: 172, 194, enter caustica as a junior synonym under both pusillus and minutus. It rests here as a junior synonym of pusillus, though its true identity may never be known.]
 * elongatus. Cryptocerus elongatus Klug, 1824: 214 (q.) BRAZIL (Pará, Rio de Janeiro), GUYANA.
 * Type-material: syntype queens (number not stated).
 * Type-localities: Brazil: Pará (“sent by Sieber, ex Mus. Com. De Hoffmannsegg”), Brazil: Rio de Janeiro (“sent by Sello”), Guyana: Demerara (“from Hemmerich”).
 * Type-depository: MNHU.
 * Smith, F. 1860c: 75 (w.); Smith, F. 1862d: 409 (s.).
 * Status as species: Guérin-Méneville, 1844a: 426; Smith, F. 1853: 218; Smith, F. 1858b: 190; Smith, F. 1860c: 75; Smith, F. 1862d: 409.
 * Junior synonym of pusillus: Roger, 1862c: 297; Mayr, 1863: 406; Roger, 1863b: 38; Dalla Torre, 1893: 144; Forel, 1895b: 134; Emery, 1924d: 307; Borgmeier, 1927c: 116; Kempf, 1951: 176; Kempf, 1972a: 180; Brandão, 1991: 388; Bolton, 1995b: 425; De Andrade & Baroni Urbani, 1999: 172.
 * obtusus. Cryptocerus obtusus Smith, F. 1858b: 191 (w.) BRAZIL (Pará).
 * Type-material: holotype (?) worker.
 * [Note: no indication of number of specimens is given.]
 * Type-locality: Brazil: Santarem (H.W. Bates).
 * [Note: De Andrade & Baroni Urbani, 1999: 172, record a second syntype (in ZSBS) from Amazonas, but this locality is not mentioned in the original description.]
 * Type-depository: BMNH.
 * Status as species: Smith, F. 1862d: 410; Mayr, 1862: 764; Mayr, 1863: 406; Roger, 1863b: 38; Dalla Torre, 1893: 143; Forel, 1895b: 134.
 * Junior synonym of pusillus: Forel, 1911e: 262; Emery, 1924d: 307; Borgmeier, 1927c: 116; Kempf, 1951: 176; Kempf, 1972a: 180; Brandão, 1991: 388; Bolton, 1995b: 426; De Andrade & Baroni Urbani, 1999: 172.

Type Material


de Andrade and Baroni Urbani (1999):

Worker. Type locality: South America. Type material: Syntype workers presumably in the Museum fur Naturkunde der Humboldt-Universitat Berlin (Roger, 1862: 297), and in the Museo regionale di Scienze Naturali, Torino (Casolari & Casolari Moreno, 1980: 84) (not available for the present study).

Cryptocerus elongates Gyne. Type locality: South America. Type material: a gyne presumably in the Museum fur Naturkunde der Humboldt-Universitat Berlin (Roger, 1862: 297) (not available for the present study).

Formica caustica Worker. Type locality: Brazil. Type material not available for the present study. Synonymy by De Dalla Torre, 1893: 143. Note: synonym of minutus in Emery (1890) and Forel (1895).

Cryptocerus obtusus Soldier. Type locality: Santarem (Brazil). Type material: a syntype worker labelled "Santarem, 54 63, 1 74, C. obtusus (type) Sm.", other label (probably more recent): "pusillus, Rio", examined; a syntype worker labelled "Amazonas, H. W. Bates" in the (Forel, 1911b: 262), examined.

Cryptocerus pusillus var. brevispinosa Worker. Type localities: S. Ignacio, Villa Lutecia, Cerro Cora in the govern. of Misiones, Cordoba (Argentina), Paraguay, Alto da Boa Vista (Rio de Janeiro, Brazil). Type material: 3 syntype workers from Cordoba, Misiones and Paraguay in (examined).

Worker
Kempf (1951) - Length 4.4 mm. Median head length 1.14 mm; Weber's length of thorax 1.29 mm. Black; the following ferruginous: apex of mandibles, anterior portion of frontal carinae, apex of last funicular segment, apex of femora and basitarsi, distal half of the following tarsal segments, anterolateral border of first gastral tergite, apex of lateral projections of peduncular segments.

Head subopaque, about as long as broad; interocular distance subequal to maximum length of head (57:58). Sides of head subparallel, posterior margin between the internal corners of the short, obliquely truncate, infuscated occipital lamellae evenly curved and emarginate. Apical border of occipital lamellae straight. Cheeks strongly marginate beneath, densely covered with flat, silvery hairs, which lie in grooves on the posterior portion of the vertex. Eyes small, their maximum diameter less than 1 /3 of the median head length.

Thorax opaque. Anterior border arcuate, shoulders angulate. Pronotum somewhat expanded laterally behind into a narrow, denticulate crest, with two short teeth on each side, the posterior tooth distinctly shorter than the anterior; posterior corner of pronotum rectangular and projecting. Promesonotal suture distinct. Mesonotum on each side with a very small, subrectangular tooth. Mesoepinotal suture vestigial, obsolete mesad. Basal face of epinotum in the same plane as mesonotum, transverse, very short, with a short lateral tooth, longer than the second pronotal tooth, and a very long, subcylindrical posterior spine arising from the posterior corner, diverging about 45° degrees from the longitudinal axis. Dorsum of thorax moderately both longitudinally and transversely convex. Declivous face not excavate mesad, nor crested laterad. The entire thorax finely reticulate-punctate, with rather dense, sharply impressed, more or less elongate foveolae, each containing an appressed silvery, elongate glistening scale. Declivous face without nacrosculpture and without scales. Laterotergite of pronotum and mesopleura more or less regularly rugose and sparsely scaled.

Petiole opaque, transverse, with a very small somewhat recurved lateral denticule. Postpetiole opaque, as broad as petiole, with a broad, apically truncate, posteriorly subdentate-projection on each side, curving slightly forward. Both peduncular segments finely reticulatepunctate with sparse, glistening appressed hairs.

Gaster opaque, elliptical, emarginate anteriorly mesad. First gastral tergite with a very narrow, not distinctly set off nor crested antero-Iateral border. Tergites and sternites finely and rather sharply reticulate-punctate. Scales of gaster sparser, more slender than on head and thorax, not lying in distinctly excavated foveolae. Erect pile confined to mandibles and to the 2-4th tergites and the sternites of the gaster.

de Andrade and Baroni Urbani (1999) - Measurements (in mm) and indices: TL 3.84-5.52; HL 0.98-1.32; HW 1.24-1.76; EL 0.30-0.36; PW 0.92-1.38; PeW 0.37-0.57; PpW 0.44-0.66; HBaL 0.46-0.58; HBaW 0.10-0.15; CI 121.6-133.3; PI 127.5-134.8; PPeI 200.0-259.4; PPpI 204.2-209.1; HBaI 20.4-25.9.

Soldier
Kempf (1951) - Length 7.5 mm. Median head length 1 .95 mm; Weber's length of thorax 2.12 mm. Black; the following ferruginous: the four apical tarsal segments of the fore legs and the two apical segments of the mid and hind legs. Tips of last funicular segments orange-brown.

Head subopaque, somewhat shorter than wide. Frontal carinae strongly converging anteriorly, prolonged behind above the eyes in the form of a raised carinule, fading out before reaching the occipital corner. Upper surface of head distinctly convex, vertex with a pair of faint, not tuberculate median swellings on the posterior border. Occiput truncate, immarginate above; occipital angles form a scarcely projecting blunt, strong tubercle. Cheeks not marginate beneath in front; a vestigial margination beneath the eye is present. Eyes very little convex. Upper and lower surface of head finely and shallowly reticulate, the upper surface with rather dense, small foveolae in which a minute, not shiny, scale is visible. foveolae slightly more crowded, longer on occiput. Still larger and more deeply impressed on occiput and lower surface of head, containing a short decumbent, somewhat glistening seta.

Thorax subopaque. Anterior border arcuate, shoulder subangulate. Sides of pronotum diverging until reaching the lateral end of the transverse crest, having before the crest a stout, apically rounded tooth, projecting obliquely forwars and laterad. Transverse pronotal crest broadly interrupted mesad, and obsolescent towards the sides. Lateral border of pronotum angulate at the crest, and converging posteriorly behind the crest. Promesonotum, in profile, greatly convex. Promesonotal suture more or less distinct laterad, vestigial mesad. Sides of mesonotum subparallel with a small posterior, acute, lateral tooth. Mesoepinotal suture impressed and distinct. Basal face of epinotum transversely convex, with a small, blunt tooth antero- Iaterad, and a much longer and stronger, blunt spine on the posterior corner, slightly more divergent than in worker. Entire thorax finely and sharply reticulate-punctate. Dorsum of thorax and laterotergite of pronotum covered with rather dense, elongate deeply impressed setiferous foveolae. declivous face and posterior portion of the sides of the thorax with more or less vestigial, irregular, fine rugosities.

Petiole opaque, transverse, its sides subparallel, with a minute denticule. Postpetiole opaque, slightly broader than petiole, with a tapering, plate-like lateral projection, the apex of which is rounded in front, sub dentate behind, curving slightly forward. Both segments finely reticulate-punctate, covered with vestigial foveolae and rugosities.

Gaster subopaque, elliptical, finely reticulate-punctate, not foveolate, deeply emarginate in front mesad. First gastral tergite anterolaterally sharply marginate, not forming a lamellate border.

All foveolae bear a decumbent seta, which is slightly flattened and glistening on thorax and peduncle. Erect setae on the terminal tergitcs and on the sternites of the gaster. First gastral tergite covered with minute, decumbent setae laterad, almost scaleless discad.

de Andrade and Baroni Urbani (1999) - Measurements (in mm) and indices: TL 6.44-7.76; HL 1.60-1.90; HW 2.24-2.36; EL 0.45-0.48; PW 1.80-2.04; PeW 0.62-0.78; PpW 0.71-0.89; HBaL 0.64-0.66; HBaW 0.18-0.19; CI 124.2-130.0; PI 115.5-115.7; PPeI 261.5-290.3; PPpI 229.2-253.5; HBaI 28.1-28.8.

Queen
Kempf (1951) - Length 10.5 mm. Median head length 2.13 mm; interocular width 2.27 mm; Weber's length of thorax 3. 1 7 mm. Black; tip of last funicular segment fuscous ferruginous. In general resembling minutus, from which it differs by the appreciably larger size, the black coloration of the appendages, the lack of the yellow maculae on the first gastral tergite. The head is shorter than wide, the upper face subopaque, greatly convex, with a distinct carinule laterad, below the eyes. Occiput subtruncate, immargin ate above. Pronotum with a distinct transverse crest. Sides of petiole without a spine, or tooth; subparallel. Sides of thorax more extensively foveolate.

de Andrade and Baroni Urbani (1999) - Measurements (in mm) and indices: TL 9.77-11.16; HL 1.76-1.96; HW 2.10-2.36; EL 0.45-049; PW 1.96-2.26; PeW 0.87-0.91; PpW 0.99-1.12; HBaL 0.72-0.75; HBaW 0.19; CI 119.3-127.3; PI 103.7-107.1; PPeI 222.7-248.3; PPpI 198.0-201.8; HBaI 25.3-26.4.

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