Cephalotes jheringi

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Cephalotes jheringi
Scientific classification
Kingdom: Animalia
Phylum: Arthropoda
Class: Insecta
Order: Hymenoptera
Family: Formicidae
Subfamily: Myrmicinae
Tribe: Attini
Genus: Cephalotes
Species group: fiebrigi
Species: C. jheringi
Binomial name
Cephalotes jheringi
(Emery, 1894)

Cephalotes jheringi casent0173682 profile 1.jpg

Cephalotes jheringi casent0173682 dorsal 1.jpg

Specimen labels

Synonyms

Nothing is known about the biology of Cephalotes jheringi.

Identification

de Andrade and Baroni Urbani (1999) - A member of the fiebrigi clade differing from its next out group species, Cephalotes bohlsi, in the worker and in the soldier by the sculpture more impressed and by the denser foveae, and, from the next ingroup species, fossithorax, in the worker, soldier and gyne by the mesosoma and pedicel without short, truncate hairs. The jheringi material we examined suggests a high degree of variability of the three female castes.

Among workers, small specimens have the head narrower than long while the large ones have the head broader than long. The pronotal spines can be more or less pointed among specimens from the same nest. The type series of the unavailable name gaudens has the first pair of pronotal spines more salient and pointed than most other workers examined; a worker from Tintina (Santiago del Estero) has pronotal spines longer and less broad. The propodeum bears two or three pairs of spines, rarely four, in individuals of the same nest. The gastral lobes are normally pigmented but in the type series of gaudens and in the specimens from Tafi Viejo (Tucuman) the lobes can be either pigmented or semitransparent.

The soldiers exhibit a great variability in colour. The head can be yellow to brown and rarely black. Some specimens with light coloured head may have a black clypeus. The pronotum can vary from completely yellow to brown with different combinations thereof. Small soldiers have the gaster either entirely black or with a pair of orange-brown spots on the posterior border of the first tergite. Large soldiers regularly have two pairs of well defined spots on the first gastral tergite.

The gynes equally show a great degree of variation in colour. Specimens from Tafi Viejo (Tucuman) have black head with yellow-orange frontal carinae while the rest of the gynes examined have yellow-brown head with or without darker clypeus. The pronotum follows the same colour variation pattern as in the soldiers. The four gastral spots can be present or absent according to the specimen.

Keys including this Species

Distribution

Argentina, Brazil and Paraguay.

Latitudinal Distribution Pattern

Latitudinal Range: -22.809943° to -34.583333°.

   
North
Temperate 		North
Subtropical 		Tropical South
Subtropical 		South
Temperate

Distribution based on Regional Taxon Lists

Neotropical Region: Argentina, Brazil (type locality), Paraguay.

Distribution based on AntMaps

AntMapLegend.png

Distribution based on AntWeb specimens

Check data from AntWeb

Countries Occupied

Number of countries occupied by this species based on AntWiki Regional Taxon Lists. In general, fewer countries occupied indicates a narrower range, while more countries indicates a more widespread species.
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Estimated Abundance

Relative abundance based on number of AntMaps records per species (this species within the purple bar). Fewer records (to the left) indicates a less abundant/encountered species while more records (to the right) indicates more abundant/encountered species.
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Biology

Explore-icon.png Explore Overview of Cephalotes biology 
The biology of many Cephalotes species is not known. Ants in this genus are common in the New World tropics and subtropics and are especially abundant and diverse in the canopies of Neotropical forests. The majority of species are arboreal. Species that live in other strata inhabit smaller trees, bushes or grass stems. These noon-arboreal species, due to their accessibility, are among the better studied members of the genus. There are also species that can be found in downed wood but it is likely the wood housed the colony before it fell to the ground. Soil nests are not known for any species nor do most species appear to extensively excavate plant tissue. They nest instead in preformed cavities. Overall, ants in the genus utilize a wide range of plants. Some species are predictable in their plant use but none appear to have evolved specialized mutualisms with particular plant species.

Worker castes typically include two forms, a worker and soldier, but there are a few species that are monomorphic. The larger soldier caste typically has an enlarged head disk. In some species the head of the soldier is very different from the worker while in others these differences are less pronounced. Queens and soldiers tend to share similar head morphology. Soldiers use their heads to plug the nest entrance. This can be very effective in excluding potential intruders. Other morphological differences between the worker castes are present but these differences have not been studied as well as head moprhology.

The behavioral repertoire of Cephalotes varians has been examined in great detail (ethograms from Wilson 1976, Cole 1980 and Cole 1983). Soldiers do little else besides defend the nest. This specialized soldier behavior is presumed to be the norm for most species. An especially interesting behavior occurs when workers are dislodged from trees: they "fly" towards the tree, often grabbing the trunk well above the ground (video).

Mature nest size varies, by species, from less than a hundred to many thousands of workers. Available evidence suggests most species are monogynous. Queens may mate with multiple males.

The proventriculus of the Cephalotes is peculiar relative to other ants. The morphology of the structure suggests it serves as a powerful pump and filter. This does not appear to lead these ants to have a highly specialized diet as most species appear to be general scavengers. Foragers have been observed feeding on carrion, bird feces, extrafloral nectaries and even tending membracids. Pollen feeding has been observed in some species, and this is somewhat specialized for ants, but it is not evident that any species restricts its diet to this resource in any significant way. Evidence for pollen feeding in Cephalotes has accumulated, in part, via finding digested pollen grains seen in infrabucal pellets. It has been suggested that the morphology of the proventriculus is a specialization for processing pollen.

More research examining all aspects of the biology of Cephalotes is needed. Our present understanding of these ants is largely based on species that live in locations other than the forest canopy, which is where Cephalotes are most common and diverse. ‎

Castes

Worker

Images from AntWeb

Cephalotes jheringi casent0173683 head 1.jpgCephalotes jheringi casent0173683 profile 1.jpgCephalotes jheringi casent0173683 dorsal 1.jpgCephalotes jheringi casent0173683 label 1.jpg
Queen (alate/dealate). Specimen code casent0173683. Photographer April Nobile, uploaded by California Academy of Sciences. Owned by ALWC, Alex L. Wild Collection.

Queen

Images from AntWeb

Cephalotes jheringi casent0173684 head 1.jpgCephalotes jheringi casent0173684 profile 1.jpgCephalotes jheringi casent0173684 dorsal 1.jpgCephalotes jheringi casent0173684 label 1.jpg
Queen (alate/dealate). Specimen code casent0173684. Photographer April Nobile, uploaded by California Academy of Sciences. Owned by ALWC, Alex L. Wild Collection.

Nomenclature

The following information is derived from Barry Bolton's Online Catalogue of the Ants of the World.

  • jheringi. Cryptocerus jheringi Emery, 1894c: 205, pl. 3, figs. 13, 14 (s.w.) BRAZIL (Rio Grande do Sul), PARAGUAY.
    • Type-material: 1 syntype soldier, syntype workers (number not stated, “several”).
    • [Note: De Andrade & Baroni Urbani, 1999: 626, cite 2w syntypes MSNG.]
    • Type-localities: soldier, workers Brazil: Rio Grande (do Sul) (H. von Ihering), 1 worker Paraguay (no further data) (L. Balzan).
    • Type-depository: MSNG.
    • [Also described as new by Emery, in von Jhering, 1894: 384 (footnote).]
    • [Misspelled as iheringii by Forel, 1895b: 141, and others.]
    • Kempf, 1958a: 54 (q.); De Andrade & Baroni Urbani, 1999: 632 (m.).
    • Combination in Paracryptocerus: Kusnezov, 1953b: 338;
    • combination in Paracryptocerus (Harnedia): Kempf, 1958a: 50;
    • combination in Zacryptocerus: Brandão, 1991: 386;
    • combination in Cephalotes: De Andrade & Baroni Urbani, 1999: 626.
    • Status as species: Forel, 1895b: 141; Santschi, 1912e: 529; Bruch, 1915: 530; Gallardo, 1915: 19; Emery, 1924d: 310; Borgmeier, 1927c: 118; Kusnezov, 1953b: 338; Kempf, 1958a: 50 (redescription); Kempf, 1972a: 178; Brandão, 1991: 386; Bolton, 1995b: 426; De Andrade & Baroni Urbani, 1999: 626 (redescription); Wild, 2007b: 31.
    • Senior synonym of ellenriederi: Kempf, 1958a: 50; Kempf, 1972a: 178; Brandão, 1991: 386; Bolton, 1995b: 426; De Andrade & Baroni Urbani, 1999: 627.
    • Senior synonym of peltatus: Kempf, 1958a: 50; Kempf, 1972a: 178; Brandão, 1991: 386; Bolton, 1995b: 426; De Andrade & Baroni Urbani, 1999: 627.
    • Material of the unavailable names gaudens, jocans referred here by Kempf, 1958a: 50; Kempf, 1972a: 178; Brandão, 1991: 386.
    • Distribution: Argentina, Brazil, Paraguay.
  • ellenriederi. Cryptocerus peltatus subsp. ellenriederi Forel, 1911e: 258 (s.w.q.) ARGENTINA (Santa Fé).
    • Type-material: syntype soldiers, syntype workers, syntype queen(s) (numbers not stated).
    • [Note: De Andrade & Baroni Urbani, 1999: 627, cite 2s, 3w, 1q syntypes (1s, 1w MHNG, 1s, 2w, 1q ZSBS.]
    • Type-locality: Argentina: Rosario di Santa Fé (von Ellenrieder).
    • Type-depositories: MACN, MHNG, ZSBS.
    • Combination in Paracryptocerus: Kusnezov, 1953b: 338.
    • Subspecies of peltatus: Forel, 1912e: 202; Forel, 1913l: 235; Forel, 1914d: 281; Bruch, 1914: 218; Bruch, 1916: 321; Santschi, 1922d: 255; Emery, 1924d: 310; Kusnezov, 1953b: 338.
    • Junior synonym of jheringi: Kempf, 1958a: 50; Kempf, 1972a: 178; Brandão, 1991: 386; Bolton, 1995b: 425; De Andrade & Baroni Urbani, 1999: 627.
  • peltatus. Cryptocerus peltatus Emery, 1896h: 633, fig. D (s.q.) PARAGUAY.
    • Type-material: 1 syntype soldier, 1 syntype queen.
    • Type-locality: Paraguay: San Salvator del Paraguay (J. Bohls).
    • Type-depository: MSNG.
    • Combination in Paracryptocerus: Kusnezov, 1953b: 338.
    • Status as species: Bruch, 1914: 218; Bruch, 1915: 530; Emery, 1924d: 310; Wheeler, W.M. 1942: 208.
    • Junior synonym of jheringi: Kempf, 1958a: 50; Kempf, 1972a: 178; Brandão, 1991: 386; Bolton, 1995b: 426; De Andrade & Baroni Urbani, 1999: 627.

Unless otherwise noted the text for the remainder of this section is reported from the publication that includes the original description.

Description

Worker

de Andrade and Baroni Urbani (1999) - Head almost as long as broad in small specimens and broader than long in large ones. Vertexal angles round or subtruncate and marginate; a very narrow lamellaceous border present in few specimens only. Vertexal margin concave. Frontal carinae not upturned over the eyes. Mandibles laterally carinate.

Mesosoma gently convex in side view. Scapular angles visible in dorsal view. Pronotum with a narrow lateral lamella with three pairs of irregular teeth, sometimes the first pair more pointed and the second and third pairs simply angulate. Sides of mesonotum with a pair of small teeth. Promesonotal suture superficially impressed. Propodeal suture more impressed laterally than medially. Propodeum with differentiate basal and declivous faces and with the sides narrowing backwards; sides of the basal face of the propodeum with two pairs of teeth, the first pair sometimes simply angulate and the second one larger; sides of the declivous face of the propodeum unarmed or armed with one or two pairs of denticles.

Petiole anteriorly truncate; its anterior border superficially concave medially. Petiolar spines arising from the anterior face of the petiole or slightly behind it, pointed and curved backwards. Postpetiole broader than the petiole; its lateral expansions variable in width, developed anteriorly, curved and pointed backwards.

Gaster suboval, with a pair of broad, anterolateral lobes with a thick, dark, marked margin not surpassing the stigma posteriorly; sometimes the lobes less thick and semitransparent.

Mid and hind femora not angulate; mid and hind basitarsi flat and with subparallel sides.

Sculpture. As in bohlsi except for the reticulation more impressed and for the foveae denser and larger.

Pilosity. As in bohlsi except the distribution of the hairs of type (2) which is denser on the sides of the frontal carinae and on the posterior borders of the gastral segments and the one of the hairs of type (3), rare to sparse on the whole first sternite.

Colour. Black. Frontal carinae, mesosomal and peduncular spines and border of the gastral lobes yellowish to light brown. Outer face of the tibiae brown.

Measurements (in mm) and indices: TL 3.36-5.36; HL 1.08-1.24; HW 1.10-1.44; EL 0.33-0.35; PW 0.99-1.24; PeW 0.50-0.61; PpW 0.63-0.69; HBaL 0.37-0.49; HBaW 0.10-0.13; CI 101.8-116.1; PI 111.1-116.1; PPeI 198.0-203.3; PPpI 157.1-179.7; HBaI 26.5-27.0.

Soldier

Kempf (1958) - Total length 5.5-7.9 mm (specimens under 6.0 mm are generally intermediates between this and the worker caste); head disc 1.50-2.07 mm. Black; yellowish-brown to testaceous: anterolateral portions of head disc, or nearly the entire head disc, both dorsally and the sides (in this case, there is usually a median black spot on the center of the disc, near the vertex of the clypeal triangle), anterolateral portions of pronotum, extensor face of tibiae; ferruginous: tip of mandibles and scapes, first funicular segment, the four apical tarsites. In darker specimens the yellow color may be substituted by ferruginous, and the ferruginous color by fuscous-ferruginous. A single soldier from Tafi Viejo, Tucuman, of an otherwise completely black series, has a quadrimaculate gaster.

Head disc subquadrate to subrectangular, completely marginate round its scarcely raised, somewhat crenulate border; the anterior corner rounded, the posterior corners either obliquely truncate or rounded. Floor of disc rather flat, slightly excavated anterolaterally. In profile, the sides of the head do not cover the dorsum of the cephalic disc, and the distance between the upper border of the eye and the rim of the disc is much less than the diameter of the eye. Sides of disc subparallel. Occipital border straight in smaller specimens, rounded in larger specimens. Occipital borders either bluntly rounded, or angulate or even subdentate. Floor of disc, sides of head, and occiput reticulate-punctate and foveolate in smaller specimens, whereas in larger specimens the foveolae are close together and the intervals form a network of raised rugosities. Lower face of head reticulate-punctate with sparse, elongate foveolae, and a few rugosities anterolaterally.

Thorax subopaque. Shoulders dentate. Sides of pronotum at the level of the usually vestigial, never strongly marked nor crested transverse carina, forming a blunt angle, the posterior corner of the pronotum being subrectangular. Mesonotum with a subtruncate projecting lateral lobe. Mesoepinotal suture distinct, scarcely impressed mesally. Sides of basal face of epinotum usually with two lateral, angulate or rounded and slightly projecting lobes and a stouter tooth at the posterior corner. Declivous face slightly excavated, its sides marginate. Mesopleural tooth absent. Sculpture, in general, as on head, but the foveolae are smaller, the intervals often shining. Laterotergite of pronotum longitudinally costate at the lower half. Pleura more or less rugose.

Peduncular segments usually equal in width (not always, sometimes the: postpetiole is distinctly broader!). Postpetiole greatly convex middorsally. Gaster elongate-ovate. Anterolateral lobes solid, submarginate, not crested. First tergite finely reticulate-punctate, the anterior fifth having fine longitudinal rugosities. Piligerous punctures scarcely impressed, inconspicuous.

Pilosity, in general, not differing from that of bivestitus, and fossithorax, with the exception of the following: Foveolae of head disc with short, thick, more or less decumbent hair, the free end of which does usually not project beyond the rim of the pit. Rim of cephalic disc with sparse, thick, subclavate, projecting setae. Scalelike hair of thorax, peduncle, and gaster appressed.

de Andrade and Baroni Urbani (1999) - Measurements (in mm) and indices: TL 6.20-7.98; HL 1.52-1.92; HW 1.68-2.00; EL 0.35-0;39; PW 1.48-1.80; PeW 0.58-0.75; PpW 0.61-0.80; HBaL 0.45-0.52; HBaW 0.15-0.17; CI 104.2-110.5; PI 111.1-113.5; PPeI 240.0-255.2; PPpI 225.0-242.6; HBal 32.7-33.3.

Queen

Kempf (1958) - Total length 8.4-8.9 mm; maximum length of head 1.85-2.03 mm; of thorax 2.46-2.53 mm. Black; the following yellowish-brown: anterolateral portion of head disc (variable as regards extension, occasionally the entire rim of disc and the sides may be included), a narrow, elongate spot on the shoulder, usually (not always, variation i.e. absence of spots noticed in female specimens of a colony that usually have them!) four spots on the first gastral tergite, extensor face of tibiae. Scape and tarsites more or less fuscous-ferruginous.

Same diagnostic features as in soldier, excepting the following characteristics: Head disc distinctly more elongate. Shape of peduncular and epinotal teeth quite variable. Mesopleural tooth always present, but sometimes very small. Wings somewhat infuscated, veins brown. Venation of fore wing not significantly different from that of liogaster. Fore wing, when folded over the back, projects a little over the apex of the normally contracted gaster.

de Andrade and Baroni Urbani (1999) - Measurements (in mm) and indices: TL 7.32-9.04; HL 1.40-1.80; HW 1.48-1.72; EL 0.35-0.42; PW 1.44-1.68; PeW 0.59-0.63; PpW 0.68-0.87; HBaL 0.44-0.56; HBaW 0.16; CI 95.5-105.7; PI 102.4-102.8; PPeI 244.1-266.7; PPpI 193.1-211.8; HBaI 28.6-34.8.

Male

de Andrade and Baroni Urbani (1999) - Head (eyes included, mandibles excluded) about 1/3 broader than long; vertexal margin straight, superficially carinate and diverging towards the sides. Vertex convex. Ocelli and compound eyes protuberant. Frontal carinae diverging backwards and not reaching the posterior border of the eyes. Frons flat. Clypeus convex, with impressed posterior border. Mandibles superficially carinate laterally. Scapes thick, twice as long as the first funicular joint.

Mesosoma. Pronotum in dorsal view with nearly parallel sides; mesonotum convex; median Mayrian and parapsidal furrows weakly impressed. Propodeum with well differentiate basal and declivous faces; basal face gently convex, medially sloping posteriorly; its sides converging posteriorly towards the declivous face, the latter with lateral and median cannae.

Petiole and postpetiole convex dorsally. Petiole narrower than the postpetiole and with a node deeply concave anteriorly; petiolar sides slightly convex. Postpetiolar sides convex. Gaster almost as broad as the mesosoma.

Wings as in the gyne.

Legs stout.

Sculpture. Head dorsum minutely reticulate, with irregular, small foveae superimposed by sparse, irregular, longitudinal, thin rugosities, the rugosities transversal around the eyes. Ventral face of the head with the same sculpture as on the head dorsum but with denser foveae. Pronotum reticulate and with dense, irregular foveae. Mesonotum and scutellum superficially reticulate, with superimposed small, sparse and shallow foveae and with sparse, longitudinal rugosities. Basal face of the propodeum irregularly foveolate-rugulose. Declivous face of the propodeum superficially reticulate, shining and with few, short, longitudinal rugosities. Propleurae reticulate and with sparse, longitudinal, thin rugosities, denser on the ventral part. Mesopleurae with the same sculpture as on the mesonotum. Lower metapleurae superficially reticulate and longitudinally rugulose; upper metapleurae reticulate and irregularly rugulose. Peduncular segments superficially reticulate and longitudinally rugulose, the rugosities more irregular on the petiole. First gastral tergite deeply reticulate; its anterior half with longitudinal rugosities originating from the postpetiolar articulation. Remaining tergites and sternites with superficial reticulation. Coxae and femora superficially reticulate, remaining parts of the legs simply punctate.

Pilosity. Body with dense, long, suberect, flexuous hairs, sparser and subdecumbent on the gaster, rare on the femora, absent on the tibiae and tarsi. Femora, tibiae and tarsi with decumbent hairs much shorter than the flexuous ones. Colour. Black. Coxae and femora brown, distal parts of the legs lighter.

Measurements (in mm) and indices: TL 6.32; HL 0.92; HW 1.12; EL 0.47; PW 1.04; PeW 0.59; PpW 0.69; HBaL 0.61; HBaW 0.14; CI 121.7; PI 107.7; PPeI 176.3; PPpI 150.7; HBaI 22.9.

Type Material

de Andrade and Baroni Urbani (1999):

Worker and soldier. Type locality: Rio Grande do Sul (Brazil). Type material: 2 syntype workers labelled “Rio Grande, Ihering”, in Museo Civico di Storia Naturale, Genoa, examined. Note: A syntype of jheringi presumably part of the material originally examined by Emery and preserved in the Musee d'Histoire Naturelle Genève is actually pallidicephalus.

Cryptocerus peltatus. Soldier, gyne. Type locality: San Salvador (Paraguay). Type material: 1 soldier (without gaster) and 1 gyne, syntypes, labelled “Paraguay, Bohls”, in MCSN, examined.

Cryptocerus peltatus Ellenriederi. Worker, soldier, gyne. Type locality: Rosario de Santa Fe (Argentina). Type material: 1 worker, 1 soldier (syntypes) in Musee d'Histoire Naturelle Genève, 2 workers, 1 soldier, 1 gyne (syntypes), in Zoologische Staatssammlung, Munich, all labelled “Cr. peltatus Em, r. Ellenriederi, type, Forel, Rosario de Santa Fe, in Weidenstamm”, examined.

References

References based on Global Ant Biodiversity Informatics

  • Bollazzi M. 2000. Nuevos taxa para los Formicidae Uruguayos (Hymenoptera: Formicidae). Nacional , Segundo Encuentro de Jóvenes Biólogos, Elio Garcia-Austt , Montevideo, Uruguay , 2000, 77
  • Brandao, C.R.F. 1991. Adendos ao catalogo abreviado das formigas da regiao neotropical (Hymenoptera: Formicidae). Rev. Bras. Entomol. 35: 319-412.
  • Bruch C. 1916. Contribución al estudio de las hormigas de la provincia de San Luis. Revista del Museo de La Plata 23: 291-357.
  • Claver S., S. L. Silnik, and F. F. Campon. 2014. Response of ants to grazing disturbance at the central Monte Desert of Argentina: community descriptors and functional group scheme. J Arid Land 6(1): 117?127.
  • Cuezzo, F. 1998. Formicidae. Chapter 42 in Morrone J.J., and S. Coscaron (dirs) Biodiversidad de artropodos argentinos: una perspectiva biotaxonomica Ediciones Sur, La Plata. Pages 452-462.
  • Emery C. 1894. Studi sulle formiche della fauna neotropica. VI-XVI. Bullettino della Società Entomologica Italiana 26: 137-241.
  • Emery C. 1896. Formiciden, gesammelt in Paraguay von Dr. J. Bohls. Zoologische Jahrbücher. Abteilung für Systematik, Geographie und Biologie der Tiere 9: 625-638.
  • Fernández, F. and S. Sendoya. 2004. Lista de las hormigas neotropicales. Biota Colombiana Volume 5, Number 1.
  • Forel A. 1911. Die Ameisen des K. Zoologischen Museums in München. Sitzungsber. Math.-Phys. Kl. K. Bayer. Akad. Wiss. Münch. 11: 249-303.
  • Forel A. 1913. Fourmis d'Argentine, du Brésil, du Guatémala & de Cuba reçues de M. M. Bruch, Prof. v. Ihering, Mlle Baez, M. Peper et M. Rovereto. Bulletin de la Société Vaudoise des Sciences Naturelles. 49: 203-250.
  • Gallardo A. 1915. Observaciones sobre algunas hormigas de la República Argentina. Anales del Museo Nacional de Historia Natural de Buenos Aires 27: 1-35.
  • Kempf, W.W. 1972. Catalago abreviado das formigas da regiao Neotropical (Hym. Formicidae) Studia Entomologica 15(1-4).
  • Pignalberi C. T. 1961. Contribución al conocimiento de los formícidos de la provincia de Santa Fé. Pp. 165-173 in: Comisión Investigación Científica; Consejo Nacional de Investigaciones Científicas y Técnicas (Argentina) 1961. Actas y trabajos del primer Congreso Sudamericano de Zoología (La Plata, 12-24 octubre 1959). Tomo III. Buenos Aires: Librart, 276 pp.
  • Santschi F. 1912. Quelques fourmis de l'Amérique australe. Revue Suisse de Zoologie 20: 519-534.
  • Vittar, F. 2008. Hormigas (Hymenoptera: Formicidae) de la Mesopotamia Argentina. INSUGEO Miscelania 17(2):447-466
  • Vittar, F., and F. Cuezzo. "Hormigas (Hymenoptera: Formicidae) de la provincia de Santa Fe, Argentina." Revista de la Sociedad Entomológica Argentina (versión On-line ISSN 1851-7471) 67, no. 1-2 (2008).
  • Wild, A. L.. "A catalogue of the ants of Paraguay (Hymenoptera: Formicidae)." Zootaxa 1622 (2007): 1-55.
  • de Andrade, M.L. & C. Baroni Urbani. 1999. Diversity and Adaptation in the ant genus Cephalotes, past and present. Stuttgarter Beitrage zur Naturkunde Serie B 271. 893 pages, Stuttgart
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