Pogonomyrmex naegelii

A wide ranging, common South American Pogonomyrmex species.

Identification
Johnson (2015) - Worker Within the P. naegelii-group, the combination of: (1) approximately 8–10 coarse, longitudinal rugae between frontal lobes, (2) small lobe projecting dorsally from anterior margin of antennal fossa, (3) peduncle of petiole and anterior surface of petiolar node meet at an obtuse angle, (4) in dorsal view, posterior surface of petiolar node narrow, width similar to or slightly greater than distance between tips of superior propodeal spines, and (5) longest hairs on mesosoma rarely >0.7–0.8x MOD uniquely characterize this species.

Alate Queen This caste is diagnosed by: (1) caste-specific morphology of the mesosoma related to wing-bearing and presence of ocelli on head, (2) small size (HW < 1.35 mm), (3) petiolar node and postpetiole relatively narrow (petiolar node < 0.45 mm, postpetiole < 0.60 mm), (4) posterior surface of petiolar node rugoreticulate to vermiculate, and (5) mesoscutum and mesoscutellum rugoreticulate-vermiculate.

Ergatoid Queen This caste is diagnosed by: (1) ergatoid, with small ocelli on head, (2) small species (HW < 1.30 mm), (3) mesoscutum, mesoscutellum, and posterior surface of petiolar node rugoreticulate, (4) in profile, mesosomal outline discontinuous with a break between mesoscutellum and metanotum and between metanotum and propodeum.

Male This caste is diagnosed by: (1) first gastral tergum lacking striae, (2) small (HW < 1.10 mm; ML < 2.00 mm), (3) in dorsal view, posterior surface of petiolar node rugoreticulate, (4) in profile, petiolar node rounded, (5) pronotal sides rugoreticulate, (6) in profile, inferior propodeal spines moderately well-developed, broadly rounded, and (7) notauli present. Note that males are unknown for Pogonomyrmex abdominalis and Pogonomyrmex tenuipubens.

Pogonomyrmex naegelii co-occurs with the other two P. naegelii-group species. Pogonomyrmex naegelii can be distinguished from P. tenuipubens by: (1) the approximately 8–10 coarse longitudinal rugae between the frontal lobes, (2) a small lobe that project dorsally from the anterior margin of the antennal fossa, and (3) the longest hairs on the psammophore and mesosoma are coarse, their length >0.5x MOD. In P. tenuipubens: (1) there are 16–20 fine, weak, longitudinal rugae between the frontal lobes, (2) the anterior margin of the antenna fossa lacks a lobe-like projection, and (3) the psammophore and abundant hairs on the mesosoma are very short, delicate, length <0.2x MOD, except for one to few long, coarse hairs that are sometimes present on the posterior margin of head and pronotum. Pogonomyrmex naegelii can be distinguished from P. abdominalis by: (1) usually smaller (HW = 1.05–1.23 mm), (2) a small lobe that projects dorsally from anterior margin of antennal fossa, (3) peduncle of petiole and anterior surface of petiolar node meet at an obtuse angle, and (4) width of posterior surface of petiolar node is similar to or slightly greater than distance between tips of superior propodeal spines. In P. abdominalis: (1) the body is usually larger (HW = 1.14–1.33 mm; (2) no small lobe projects dorsally from anterior margin of antennal fossa, (3) the peduncle of petiole and anterior surface of petiolar node meet at or near a right angle, and (4) the posterior surface of petiolar node is distinctly wider than distance between tips of superior propodeal spines.

Distribution based on Regional Taxon Lists
Neotropical Region: Argentina, Bolivia, Brazil, Colombia, Guyana, Paraguay, Peru, Uruguay, Venezuela.

Habitat
Pogonomyrmex naegelii inhabits sites at elevations from 0–1115 m, and it is the most widespread and commonly collected Pogonomyrmex in South America. It is known from all countries except Chile, Ecuador, Suriname, and French Guiana. Pogonomyrmex naegelii occurs in numerous ecoregions, but appears to be absent from deserts, rainforests, and high-elevation areas. (Johnson 2015)

Biology
Johnson (2015) - Pogonomyrmex naegelii is a solitary forager that has a generalist and season-dependent diet that consists of grass seeds, and to a lesser extent nongrass species, various plant parts, and insects; ants and termites are the primary animal prey. The diet is primarily granivorous during the dry/cold season, whereas arthropods are collected to a greater extent during the warm/wet season (Belchior, Del-Claro, & Oliveira, 2012). Nests range from a cryptic entrance to a 10 cm tumulus, and they sometimes have a secondary entrance. Nests in the cerrado savanna of Brazil are relatively shallow, reaching about 70 cm below the soil surface and have 5–10 chambers (Belchior et al., 2012). Colonies are relatively small, ranging from 166–580 workers (n = 3 nests) (Belchior et al., 2012). Nests had one queen (n = 3) (Belchior et al., 2012), and the author has found multiple dealate queens in several nests. In the latter case, it is unknown if these queens were reproductive, especially given that colonies of the related North American species P. pima sometimes have numerous unmated dealate queens that forage (see Johnson et al., 2007).

Collection dates for sexuals range from 3–25 December. Mating flights have not been observed, but they likely occur during austral summer based on dates on which sexuals have been collected. Pogonomyrmex naegelii has both alate and ergatoid queens. From collections to date, the author has not seen any series in which both queen phenotypes were collected in the same colony, similar to observations for Pogonomyrmex pima (Johnson et al., 2007). One alate queen had eight ovarioles (unpub. data).

Nomenclature

 *  naegelii. Pogonomyrmex naegelii Forel, in Emery, 1878a: x (w.) BRAZIL. Mayr, 1887: 612 (q.m.); Wheeler, G.C. & Wheeler, J. 1972b: 237 (l.). Combination in P. (Ephebomyrmex): Wheeler, W.M. 1902c: 390; in Ephebomyrmex: Kempf, 1972a: 106; in Pogonomyrmex: Bolton, 1995b: 341. Senior synonym of rupununi, venezuelensis: Kempf, 1960f: 428. See also: Forel, 1886b: xli; Gallardo, 1932b: 109.
 * rupununi. Pogonomyrmex (Ephebomyrmex) venezuelensis var. rupununi Weber, 1943b: 71 (w.) GUYANA. Junior synonym of naegelii: Kempf, 1960f: 428.
 * venezuelensis. Pogonomyrmex (Ephebomyrmex) venezuelensis Weber, 1943b: 69, fig. 2 [fig. is incorrectly captioned as Leptothorax anduzei] (w.) VENEZUELA. Junior synonym of naegelii: Kempf, 1960f: 428.

Johnson (2015):

Worker
Johnson (2015) - Lectotype (n = 31 + 1 paralectotype). HL 1.15 (1.05–1.23); HW 1.15 (1.06–1.20); MOD 0.24 (0.22–0.28); OMD 0.28 (0.21–0.31); SL 0.84 (0.80–0.94); PNW 0.78 (0.69–0.84); HFL 1.15 (0.95–1.20); ML 1.34 (1.20–1.46); PW 0.30 (0.28–0.36); PPW 0.45 (0.40–0.49). Indices: SI 73.04 (68.97–85.45); CI 100.00 (91.53–106.25); OI 20.87 (20.17–24.30); HFI 100.00 (84.82–103.64).

Head subquadrate to quadrate (CI = 91.53–106.25), widest just posterior to eye; posterior margin flat to weakly concave. Longitudinal rugae on cephalic dorsum prominent, usually moderately to coarsely rugoreticulate especially medial to eyes and near posterior margin; approximately 8–10 coarse longitudinal rugae between frontal lobes; in full-face view, medial rugae not diverging toward posterior corners of head. In profile, area posterior to eyes moderately to strongly rugoreticulate. Cephalic interrugae moderately to strongly granulate, dull to weakly shining; vertex rugose to rugoreticulate. Anterior margin of clypeus flat to weakly concave, dorsal surface with numerous subparallel, longitudinal rugae; small lobe projects dorsally from anterior margin of antennal fossa. Mandible with six teeth; mandibular dorsum coarsely rugose. Up to several moderately long, curved, bristle-like, yellow-brown to brownish hairs project from anterior margin of clypeus and basolateral margin of mandibles. MOD ranging from 0.19–0.25x HL. In profile, eyes situated anterior to middle of head, OMD = 0.84–1.25x MOD. Antennal scapes moderately long (SI = 70.83–85.45), failing to reach vertex by 1.0–1.5x length of basal funicular segment; entire scape with strong longitudinal striae, dull to weakly shining. Basal flange of scape flattened, well-developed with carinate margin. Psammophore poorly-developed, consisting of short to medium-length hairs scattered across ventral side of head.

Mesosomal profile convex; all mesosomal surfaces rugoreticulate to vermiculate. Mesoepinotal sulcus sometimes weakly to moderately impressed. Dorsum of promesonotum and sides of pronotum rugoreticulate to vermiculate. Mesopleura with highly irregular rugae angling posterodorsally to rugoreticulate-vermiculate. Dorsum and sides of propodeum with irregular to very irregular, transverse rugae to occasionally rugoreticulate-vermiculate. Superior propodeal spines moderately long, acuminate, connected by well-defined keel; spine length approximately 0.7–0.8x distance between their bases. Inferior propodeal spines well-developed, acuminate, length approximately 0.5–1.0x that of superior spines, base wider than length of superior spines; inferior and superior spines connected by crest that defines the propodeal declivity. Propodeal spiracles ovoid to circular facing posterad. Interrugae on mesosoma weakly to strongly granulate, dull to weakly shining to smooth and strongly shining. Legs moderately coriarious, weakly shining.

Peduncle of petiole about as long as petiolar node, anteroventral margin with weakly to strongly-developed triangular process. In profile, petiolar node asymmetrical with anterior surface shorter than posterior surface; apex weakly rounded to subangulate; anterior surface meeting peduncle of petiole at an obtuse angle. In dorsal view, petiolar node longer than wide, widest near middle, narrowing to spatulate to subangulate anterior margin; maximal width of posterior surface similar to slightly greater than distance between tips of superior propodeal spines; posterior surface and sides strongly rugoreticulate-vermiculate, interrugae weakly to moderately granulate, weakly shining. Dorsum of postpetiole convex in profile; in dorsal view, postpetiole widest at or near posterior margin, narrowing near middle to convex anterolateral margin; maximal width about equal to length; dorsum and sides moderately to coarsely rugoreticulate or with several irregular, longitudinal to oblique rugae, interrugae moderately to strongly granulate, dull to weakly shining. Ventral process of postpetiole large, bulbous, height similar to dorsal portion of postpetiole. First gastral tergum moderately to strongly coriarious, weakly shining, to smooth and strongly shining; anterior margin to anterior one-half often with weak to moderately strong longitudinal striae.

Erect yellow-brown to brownish pilosity moderately abundant on head, similar in length, mostly short, often with one or more longer hairs that approximate MOD. Moderately abundant suberect to semidecumbent pilosity on scape, abundant decumbent hairs on funicular segments. Legs with moderately abundant semidecumbent, brownish setae. Mesosoma, petiolar node, postpetiole, gastral terga with moderately dense, erect setae, mostly similar in length, longest approximately 0.7–0.8x MOD; hairs on propodeum less dense. Concolorous tan to tannish-brown with darker to blackish gaster.

Queen
Alate (n = 11). HL 1.15–1.29; HW 1.15–1.26; MOD 0.24–0.32; OMD 0.23–0.30; SL 0.85–0.98; PNW 0.87–1.01; HFL 1.09–1.30; ML 1.56–1.75; PW 0.34–0.44; PPW 0.50–0.60. Indices: SI 73.91–78.86; CI 95.93–107.83; OI 20.69–25.40; HFI 87.90–108.47. Ergatoid (n = 6). HL 1.12–1.27; HW 1.17–1.27; MOD 0.23–0.29; OMD 0.23–0.28; SL 0.83–0.94; PNW 0.89–0.99; HFL 1.09–1.22; ML 1.45–1.60; PW 0.39–0.41; PPW 0.56–0.63. Indices: SI 65.35–77.78; CI 94.49–110.71; OI 18.55–23.93; HFI 89.76–99.19.

Ergatoid; small species (HW = 1.17–1.27 mm), head subquadrate to wider than long (CI = 94.49–110.71), posterior margin flat to weakly concave. Longitudinal rugae on cephalic dorsum prominent medially, becoming rugoreticulate laterally; interrugae weakly to moderately coriarious, weakly shining. Mandible with six teeth, dorsal surface coarsely rugose. Psammophore poorly-developed, consisting of numerous short hairs scattered across ventral side of head.

All mesosomal surfaces rugoreticulate. In profile, mesoscutum and mesoscutellum flattened, mesoscutellum angled upward posterad; metanotum discontinuously connected to mesoscutellum and propodeum. Superior and inferior propodeal spines moderately well-developed, similar in length. Peduncle of petiole long, anteroventral margin with small to moderately well-developed acuminate triangular process. In profile, petiolar node asymmetrical with anterior surface notably shorter than posterior surface, apex angulate. Postpetiole slightly wider than long. Posterior surface of petiolar node coarsely rugoreticulate, dorsum of postpetiole finely rugoreticulate; interrugae weakly to moderately punctate, weakly shining. First gastral tergum weakly to strongly coriarious, dull to weakly shining with faint to moderately strong longitudinal striae near base that sometimes extend over anterior one-half of tergum. Most body surfaces with abundant suberect to erect, brownish hairs that are similar in length, longest approximately 0.5–0.7x MOD. Body concolorous tannish-brown to orangish-brown, posterior gastral terga often darker.

Male
(n = 12). HL 0.89–1.10; HW 0.84–1.08; MOD 0.32–0.42; OMD 0.06–0.18; SL 0.14–0.23; HFL 1.03–1.40; ML 1.51–1.95; PW 0.33–0.43; PPW 0.44–0.58. Indices: SI 14.00–22.12; CI 94.38–109.18; OI 33.98–42.00; HFI 106.80–136.00.

Type Material
Syntypes examined: 1 worker, 2 workers , 1 worker , BRAZIL, Rio de Janeiro (Carlos Naegeli leg.); MSNG worker here designated LECTOTYPE [CASENT0280989].

Etymology
The specific epithet, naegelii (Latinization of Naegeli), is derived from Carlos Naegeli, who collected the syntype workers.

References based on Global Ant Biodiversity Informatics

 * Belchior C., K. Del-Claro, and P. S. Oliveira. 2012. Seasonal patterns in the foraging ecology of the harvester ant Pogonomyrmex naegelii (Formicidae, Myrmicinae) in a Neotropical savanna: Daily rhythms, shifts in granivory and carnivory, and home range. Arthropod-Plant Interactions 6: 571582.
 * 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.
 * Caldart V. M., S. Iop, J. A. Lutinski, and F. R. Mello Garcia. 2012. Ants diversity (Hymenoptera, Formicidae) of the urban perimeter of Chapecó county, Santa Catarina, Brazil. Revista Brasileira de Zoociências 14 (1, 2, 3): 81-94.
 * Canepuccia A. D., F. Hidalgo, J. L. Farina, F. Cuezzo, and O. O. Iribarne. 2016. Environmental harshness decreases ant β-diversity between salt marsh and neighboring upland environments. Wetlands DOI 10.1007/s13157-016-0777-0.
 * Cardoso, D.C., T.G. Sobrinho and J.H. Schoereder. 2010. Ant community composition and its relationship with phytophysiognomies in a Brazilian Restinga. Insectes Sociaux 57:293-301
 * Clemes Cardoso D., and J. H. Schoereder. 2014. Biotic and abiotic factors shaping ant (Hymenoptera: Formicidae) assemblages in Brazilian coastal sand dunes: the case of restinga in Santa Catarina. Florida Entomologist 97(4): 1443-1450.
 * Clemes Cardoso D., and M. Passos Cristiano. 2010. Myrmecofauna of the Southern Catarinense Restinga sandy coastal plain: new records of species occurrence for the state of Santa Catarina and Brazil. Sociobiology 55(1b): 229-239.
 * Coelho M. S., G. W. Fernandes, J. C. Santos, and J. H. C. Delabie. 2009. Ants (Hymenoptera: Formicidae) as bioindicators of land restoration in a Brazilian Atlantic forest fragment. Sociobiology 54(1): 51-63.
 * Costa-Milanez C. B., G. Lourenco-Silva, P. T. A. Castro, J. D. Majer, and S. P. Ribeiro. 2014. Are ant assemblages of Brazilian veredas characterised by location or habitat type? Braz. J. Biol. 74(1): 89-99.
 * 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.
 * Diehl-Fleig E. 2014. Termites and Ants from Rio Grande do Sul, Brazil. Sociobiology (in Press).
 * Emery C. 1906. Studi sulle formiche della fauna neotropica. XXVI. Bullettino della Società Entomologica Italiana 37: 107-194.
 * Escalante Gutiérrez J. A. 1993. Especies de hormigas conocidas del Perú (Hymenoptera: Formicidae). Revista Peruana de Entomología 34:1-13.
 * Fernández F., and E. E. Palacio. 1995. Hormigas de Colombia IV: nuevos registros de géneros y especies. Caldasia 17: 587-596.
 * Forel A. 1908. Ameisen aus Sao Paulo (Brasilien), Paraguay etc. gesammelt von Prof. Herm. v. Ihering, Dr. Lutz, Dr. Fiebrig, etc. Verhandlungen der Kaiserlich-Königlichen Zoologisch-Botanischen Gesellschaft in Wien 58: 340-418.
 * Forel A. 1911. Ameisen des Herrn Prof. v. Ihering aus Brasilien (Sao Paulo usw.) nebst einigen anderen aus Südamerika und Afrika (Hym.). Deutsche Entomologische Zeitschrift 1911: 285-312.
 * Gallardo A. 1932. Las hormigas de la República Argentina. Subfamilia Mirmicinas, segunda sección Eumyrmicinae, tribu Myrmicini (F. Smith), género Pogonomyrmex Mayr. Anales del Museo Nacional de Historia Natural de Buenos Aires 37: 89-170.
 * Iop S., V. M. Caldart, J. A. Lutinski, and F. R. Mello Garcia. 2009. Formigas urbanas da cidade de Xanxerê, Santa Catarina, Brasil. Biotemas 22(2): 55-64.
 * Johnson R. A. 2015. A taxonomic revision of South American species of the seed-harvester ant genus Pogonomyrmex (Hymenoptera: Formicidae). Part I. Zootaxa 4029(1):1-142
 * Johnson R. Personnal Database. Accessed on February 5th 2014 at http://www.asu.edu/clas/sirgtools/resources.htm
 * Johnson Robert. 2014. List of South American species of Pogonomyrmex. Accessed on February 5th 2014 at http://www.asu.edu/clas/sirgtools/pogonomyrmex/SOUTHAMERICANPOGOS.htm
 * Kamura, C.M., M.S.C. Morini, C.J. Figueiredo, O.C. Bueno, and A.E.C. Campos-Farinha. 2007. Comunidades de formigas (Hymenoptera: Formicidae) em um ecossistema urbano próximo à Mata Atlântica. Brazilian Journal of Biology 67(4): 635-641
 * Kempf W. W. 1960. Miscellaneous studies on Neotropical ants (Hymenoptera, Formicidae). Studia Entomologica (n.s.)3: 417-466.
 * Kempf, W.W. 1972. Catalago abreviado das formigas da regiao Neotropical (Hym. Formicidae) Studia Entomologica 15(1-4).
 * Kusnezov N. 1951. El género Pogonomyrmex Mayr (Hym., Formicidae). Acta Zoologica Lilloana 11: 227-333.
 * Kusnezov N. 1953. La fauna mirmecológica de Bolivia. Folia Universitaria. Cochabamba 6: 211-229.
 * Kusnezov N. 1958. Lilidris metatarsalis gen. et spec. nov. (Hym. Formicidae). Acta Zoologica Lilloana 15: 189-193.
 * Kusnezov N. 1978. Hormigas argentinas: clave para su identificación. Miscelánea. Instituto Miguel Lillo 61:1-147 + 28 pl.
 * Lattke J. E. 1991. Una nueva especie de Pogonomyrmex Mayr de selva húmeda tropical (Hymenoptera: Formicidae). Revista de Biología Tropical 38: 305-309.
 * Luederwaldt H. 1918. Notas myrmecologicas. Rev. Mus. Paul. 10: 29-64.
 * Lutinski J. A., B. C. Lopes, and A. B. B.de Morais. 2013. Diversidade de formigas urbanas (Hymenoptera: Formicidae) de dez cidades do sul do Brasil. Biota Neotrop. 13(3): 332-342.
 * Lutinski J. A., F. R. Mello Garcia, C. J. Lutinska, and S. Iop. 2008. Ants diversity in Floresta Nacional de Chapecó in Santa Catarina State, Brazil. Ciência Rural, Santa Maria 38(7): 1810-1816.
 * Lutinski J. A., and F. R. Mello Garcia. 2005. Análise faunística de Formicidae (Hymenoptera: Apocrita) em ecossistema degradado no município de Chapecó, Santa Catarina. Biotemas 18(2): 73 - 86.
 * Maciel L., J. Iantas, F. C. Gruchowski-W, and D. R. Holdefer. 2011. INVENTORY OF FAUNA OF ANTS (HYMENOPTERA: FORMICIDAE) IN ECOLOGICAL SUCCESSION FLORISTIC ENVIRONMENT IN THE CITY OF UNION OF VICTORIA, PARANA, BRAZIL. Biodiversidade Pampeana Pucrs, Uruguiana 9(1): 38-43.
 * Maravalhas J., and H. L. Vasconcelos. 2014. Revisiting the pyrodiversitybiodiversity hypothesis: long-term fire regimes and the structure of ant communities in a Neotropical savanna hotspot. Journal of Applied Ecology 51: 1661-1668.
 * Menozzi C. 1926. Neue Ameisen aus Brasilien. Zoologischer Anzeiger. 69: 68-72.
 * Nascimento Santos M., J. H. C. Delabie, and J. M. Queiroz. 2019. Biodiversity conservation in urban parks: a study of ground-dwelling ants (Hymenoptera: Formicidae) in Rio de Janeiro City. Urban Ecosystems https://doi.org/10.1007/s11252-019-00872-8
 * Neves F. S., K. S. Queiroz-Dantas, W. D. da Rocha, and J. H. C. Delabie. 2013. Ants of Three Adjacent Habitats of a Transition Region Between the Cerrado and Caatinga Biomes: The Effects of Heterogeneity and Variation in Canopy Cover. Neotrop Entomol 42: 258268.
 * Osorio Rosado J. L, M. G. de Goncalves, W. Drose, E. J. Ely e Silva, R. F. Kruger, and A. Enimar Loeck. 2013. Effect of climatic variables and vine crops on the epigeic ant fauna (Hymenoptera: Formicidae) in the Campanha region, state of Rio Grande do Sul, Brazil. J Insect Conserv 17: 1113-1123.
 * Pacheco R., and H. L. Vasconcelos. 2007. Invertebrate conservation in urban areas: ants in the Brazilian Cerrado. Landscape and Urban Planning 81: 193199.
 * Pacheco R., and H. L. Vasconcelos. 2012. Subterranean Pitfall Traps: Is ItWorth Including Them in Your Ant Sampling Protocol? Psyche doi:10.1155/2012/870794
 * 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.
 * Pires de Prado L., R. M. Feitosa, S. Pinzon Triana, J. A. Munoz Gutierrez, G. X. Rousseau, R. Alves Silva, G. M. Siqueira, C. L. Caldas dos Santos, F. Veras Silva, T. Sanches Ranzani da Silva, A. Casadei-Ferreira, R. Rosa da Silva, and J. Andrade-Silva. 2019. An overview of the ant fauna (Hymenoptera: Formicidae) of the state of Maranhao, Brazil. Pap. Avulsos Zool. 59: e20195938.
 * Ribas C. R., R. R. C. Solar, R. B. F. Campos, F. A. Schmidt, C. L. Valentim, and J. H. Schoereder. 2012. Can ants be used as indicators of environmental impacts caused by arsenic? Insect Conserv 16: 413421.
 * Rosa da Silva R. 1999. Formigas (Hymenoptera: Formicidae) do oeste de Santa Catarina: historico das coletas e lista atualizada das especies do Estado de Santa Catarina. Biotemas 12(2): 75-100.
 * Rosado J. L. O., M. G. de Gonçalves, W. Dröse, E. J. E. e Silva, R. F. Krüger, R. M. Feitosa, and A. E. Loeck. 2012. Epigeic ants (Hymenoptera: Formicidae) in vineyards and grassland areas in the Campanha region, state of Rio Grande do Sul, Brazil. Check List, Journal of species lists and distribution 8(6): 1184-1189.
 * Santoandre S., J. Filloy, G. A. Zurita, and M. I. Bellocq. 2019. Ant taxonomic and functional diversity show differential response to plantation age in two contrasting biomes. Forest Ecology and Management 437: 304-313.
 * Santschi F. 1912. Quelques fourmis de l'Amérique australe. Revue Suisse de Zoologie 20: 519-534.
 * Santschi F. 1925. Fourmis des provinces argentines de Santa Fe, Catamarca, Santa Cruz, Córdoba et Los Andes. Comunicaciones del Museo Nacional de Historia Natural "Bernardino Rivadavia" 2: 149-168.
 * Silva F. H. O., J. H. C. Delabie, G. B. dos Santos, E. Meurer, and M. I. Marques. 2013. Mini-Winkler Extractor and Pitfall Trap as Complementary Methods to Sample Formicidae. Neotrop Entomol 42: 351358.
 * Suguituru S. S., D. R. de Souza, C. de Bortoli Munhae, R. Pacheco, and M. S. de Castro Morini. 2011. Diversidade e riqueza de formigas (Hymenoptera: Formicidae) em remanescentes de Mata Atlântica na Bacia Hidrográfica do Alto Tietê, SP. Biota Neotrop. 13(2): 141-152.
 * Suguituru S. S., M. Santina de Castro Morini, R. M. Feitosa, and R. Rosa da Silva. 2015. Formigas do Alto Tiete. Canal 6 Editora 458 pages
 * Ulyssea M.A., C. E. Cereto, F. B. Rosumek, R. R. Silva, and B. C. Lopes. 2011. Updated list of ant species (Hymenoptera, Formicidae) recorded in Santa Catarina State, southern Brazil, with a discussion of research advances and priorities. Revista Brasileira de Entomologia 55(4): 603-611.
 * Viana-Silva F. E. C., and C. M. Jacobi. 2012. Myrmecofauna of Ironstone Outcrops: Composition and Diversity. Neotrop Entomol 41: 263271.
 * 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).
 * Weber N. A. 1943. New ants from Venezuela and neighboring countries. Bol. Entomol. Venez. 2: 67-78.
 * Wheeler W. M. 1925. Neotropical ants in the collections of the Royal Museum of Stockholm. Arkiv för Zoologi 17A(8): 1-55.
 * Wild, A. L. "A catalogue of the ants of Paraguay (Hymenoptera: Formicidae)." Zootaxa 1622 (2007): 1-55.
 * Zolessi L. C. de, Y. P. Abenante, and M. E. de Philippi. 1988. Lista sistematica de las especies de Formicidos del Uruguay. Comun. Zool. Mus. Hist. Nat. Montev. 11: 1-9.
 * Zolessi L. C. de; Y. P. de Abenante, and M. E. Philippi. 1989. Catálogo sistemático de las especies de Formícidos del Uruguay (Hymenoptera: Formicidae). Montevideo: ORCYT Unesco, 40 + ix pp.
 * da Silva Araujo, M., Castro Della Lucia, T.M., DA VEIGA, Clayton E y CARDOSO DO NASCIMENTO, Ivan. 2004. Efeito da queima da palhada de cana-de-açúcar sobre comunidade de formicídeos. Ecol. austral. 14(2): 191-200.
 * da Silva de Oliveira A. B., and F. A. Schmidt. 2019. Ant assemblages of Brazil nut trees Bertholletia excelsa in forest and pasture habitats in the Southwestern Brazilian Amazon. Biodiversity and Conservation 28(2): 329-344.
 * de Zolessi, L.C., Y.P. de Abenante and M.E. Philippi. 1987. Lista sistemática de las especies de formícidos del Uruguay. Comunicaciones Zoologicas del Museo de Historia Natural de Montevideo 11(165):1-9
 * de Zolessi, L.C., Y.P. de Abenante and M.E. Phillipi. 1989. Catalago Systematico de las Especies de Formicidos del Uruguay (Hymenoptera: Formicidae). Oficina Regional de Ciencia y Technologia de la Unesco para America Latina y el Caribe- ORCYT. Montevideo, Uruguay