Pheidole subarmata

Pheidole subarmata favors moist soil in open places, including habitats seriously disturbed by human activity—such as the edges of dirt roads, agricultural fields, coconut groves, secondary rainforest, city parks, and seashores. When exploring in the deeper rainforest is done, it is usually easy for the collector to add subarmata to his list by searching adjacent roads and fields. On St. Vincent, the indefatigable H. H. Smith (in Forel 1893j) found colonies of this species from sea level to 460 m. The nests are usually constructed in open soil or soil beneath rocks or sod, but also occasionally in pieces of rotting wood. When in the open they are marked by irregular accumulations of excavated soil. The galleries are irregular and difficult to follow by excavation, although I was able to do so in a Costa Rican banana plantation by digging a lateral pit and shaving the soil away from the side. The colonies are relatively small, with workers numbering probably only in the hundreds. On St. Vincent, H. H. Smith observed workers foraging on the ground and bushes. He found colony-founding queens in November. (Wilson 2003)

Identification
See the description in the nomenclature section.

Distribution
Pheidole subarmata is one of the most abundant and wide-ranging species of New World Pheidole, as evidenced by the heavy synonymic fouling cited above. It has been recorded from Veracruz, Mexico, south to Santa Catarina in southern Brazil, and widely through the West Indies, including the Bahamas, Cuba, Puerto Rico, and some of the Lesser Antilles, including St. Vincent and Grenada. It has also been found on Cocos Island off the Pacific Coast of Costa Rica. (Wilson 2003)

Distribution based on Regional Taxon Lists
Neotropical Region: Argentina, Bahamas, Barbados, Brazil, Colombia, Costa Rica, Cuba, Dominican Republic, French Guiana, Greater Antilles, Grenada, Guadeloupe, Guatemala, Haiti, Honduras, Mexico, Panama, Paraguay, Puerto Rico, St. Vincent, Trinidad and Tobago.

Biology
Wheeler (1905), Bahamas - Several soldiers and workers, taken in and about Nassau, N. P. (Queen's Staircase, Fort Charlotte, etc.), from small nests in grassy places and occasionally under stones.

Worker
Minor

Nomenclature

 *  subarmata. Pheidole subarmata Mayr, 1884: 37 (s.w.) FRENCH GUIANA. Senior synonym of borinquenensis, cornutula, dentimentum, elongatula, hondurensis, imbecillis, nassavensis, nefasta: Brown, 1981: 526. See also: Wilson, 2003: 759.
 * cornutula. Pheidole cornutula Emery, 1890b: 52 (footnote), pl. 5, figs. 14, 15 (s.) PARAGUAY. Forel, 1901e: 365 (w.); Santschi, 1923d: 62 (q.m.); Goni, Zolessi & Imai, 1983: 365 (k.). Junior synonym of subarmata: Brown, 1981: 526.
 * elongatula. Pheidole subarmata var. elongatula Forel, 1893g: 408 (s.w.q.m.) ANTILLES. Junior synonym of subarmata: Brown, 1981: 526.
 * nassavensis. Pheidole subarmata var. nassavensis Wheeler, W.M. 1905b: 92 (s.w.) BAHAMAS. Junior synonym of subarmata: Brown, 1981: 526.
 * imbecillis. Pheidole cornutula var. imbecillis Emery, 1906c: 151 (s.w.) ARGENTINA. Junior synonym of subarmata: Brown, 1981: 526.
 * borinquenensis. Pheidole subarmata var. borinquenensis Wheeler, W.M. 1908a: 133, pl. 12, figs. 19-21 (s.w.) PUERTO RICO. Junior synonym of subarmata: Brown, 1981: 526.
 * hondurensis. Pheidole (Elasmopheidole) hondurensis Mann, 1922: 25, fig. 12 (s.w.) HONDURAS. Junior synonym of subarmata: Brown, 1981: 526.
 * dentimentum. Pheidole cornutula var. dentimentum Santschi, 1929d: 285 (s.) BRAZIL. Junior synonym of subarmata: Brown, 1981: 526.
 * nefasta. Pheidole subarmata var. nefasta Santschi, 1929d: 285 (s.w.) BRAZIL. Junior synonym of subarmata: Brown, 1981: 526.

Description
From Wilson (2003): DIAGNOSIS Similar to Pheidole allarmata, Pheidole cramptoni, Pheidole hasticeps, Pheidole synarmata and Pheidole triplex, differing from them and other Pheidole by the following combination of traits.

Major: head elongate; frontal lobes seen from the side triangular, pointing slightly downward; pronotum low and rounded, descending into the mesonotum in a weakly convex, continuous curve; propodeal spines very small; carinulae limited to the anterior half of the head, and those immediately mesad to the eyes reaching only to the posterior margins of the eyes; rest of head and body smooth and shiny.

Minor: occiput broad, lacking nuchal collar; propodeal spines reduced to denticles; except for circular carinulae of the antennal fossa, entire head and body smooth and shiny. I have confirmed most of the synonymies made by William L. Brown (1981) and listed above.

MEASUREMENTS (mm) Paralectotype major (Santa Catarina, Brazil): HW 0.94, HL 1.18, SL 0.50, EL 0.14, PW 0.54. Lectotype minor: HW 0.42, HL 0.48, SL 0.42, EL 0.08, PW 0.28. COLOR Major: reddish yellow to dark brown, including appendages; on some specimens at least, there is a diffuse round light brown spot on the vertex, as illustrated.

Minor: light yellowish brown, appendages yellow.



'''Figure. Upper: major. Lower: minor. COLOMBIA: Yumbo (compared with syntypes from Santa Catarina, Brazil, by E. O. Wilson.) Scale bars = 1 mm.'''

Etymology
L subarmata, presumably, less well armed, referring to the small propodeal spines. (Wilson 2003)

References based on Global Ant Biodiversity Informatics

 * Ahuatzin D. A., E. J. Corro, A. Aguirre Jaimes, J. E. Valenzuela Gonzalez, R. Machado Feitosa, M. Cezar Ribeiro, J. Carlos Lopez Acosta, R. Coates, W. Dattilo. 2019. Forest cover drives leaf litter ant diversity in primary rainforest remnants within human-modified tropical landscapes. Biodiversity and Conservation 28(5): 1091-1107.
 * Brandao, C.R.F. 1991. Adendos ao catalogo abreviado das formigas da regiao neotropical (Hymenoptera: Formicidae). Rev. Bras. Entomol. 35: 319-412.
 * Branstetter M. G. and L. Sáenz. 2012. Las hormigas (Hymenoptera: Formicidae) de Guatemala. Pp. 221-268 in: Cano E. B. and J. C. Schuster. (eds.) 2012. Biodiversidad de Guatemala. Volumen 2. Guatemala: Universidad del Valle de Guatemala, iv + 328 pp
 * Brown W. L., Jr. 1981. Preliminary contributions toward a revision of the ant genus Pheidole (Hymenoptera: Formicidae). Part I. J. Kans. Entomol. Soc. 54: 523-530.
 * Calixto J. M. 2013. Lista preliminar das especies de formigas (Hymenoptera: Formicidae) do estado do Parana, Brasil. Universidad Federal do Parana 34 pages.
 * Castano-Meneses, G., M. Vasquez-Bolanos, J. L. Navarrete-Heredia, G. A. Quiroz-Rocha, and I. Alcala-Martinez. 2015. Avances de Formicidae de Mexico. Universidad Nacional Autonoma de Mexico.
 * 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.
 * Dattilo W. et al. 2019. MEXICO ANTS: incidence and abundance along the Nearctic-Neotropical interface. Ecology https://doi.org/10.1002/ecy.2944
 * Del Toro, I., M. Vázquez, W.P. Mackay, P. Rojas and R. Zapata-Mata. Hormigas (Hymenoptera: Formicidae) de Tabasco: explorando la diversidad de la mirmecofauna en las selvas tropicales de baja altitud. Dugesiana 16(1):1-14.
 * Deyrup M., L. Davis, and S. Buckner. 1998. Composition of the ant fauna of three Bahamian islands. Proceedings of the seventh symposium on the natural history of the Bahamas. 23-32. Bahamian Field Station, San Salvador, Bahamas
 * Donoso D. A. 2014. Assembly mechanisms shaping tropical litter ant communities. Ecography 37 doi: 10.1111/j.1600-0587.2013.00253.x
 * Emery C. 1890. Studii sulle formiche della fauna neotropica. Bull. Soc. Entomol. Ital. 22: 38-8
 * Emery C. 1894. Estudios sobre las hormigas de Costa Rica. Anales del Museo Nacional de Costa Rica 1888-1889: 45-64.
 * Favretto M. A., E. Bortolon dos Santos, and C. J. Geuster. 2013. Entomofauna from West of Santa Catarina State, South of Brazil. EntomoBrasilis 6 (1): 42-63.
 * Fernandes, P.R. XXXX. Los hormigas del suelo en Mexico: Diversidad, distribucion e importancia (Hymenoptera: Formicidae).
 * Fernández, F. and S. Sendoya. 2004. Lista de las hormigas neotropicales. Biota Colombiana Volume 5, Number 1.
 * Field Museum Collection, Chicago, Illinois (C. Moreau)
 * Fleck M. D., E. Bisognin Cantarelli, and F. Granzotto. 2015. Register of new species of ants (Hymenoptera: Formicidae) in Rio Grande do Sul state. Ciencia Florestal, Santa Maria 25(2): 491-499.
 * Fontanla Rizo J.L. 1997. Lista preliminar de las hormigas de Cuba. Cocuyo 6: 18-21.
 * Fontenla J. L., and J. Alfonso-Simonetti. 2018. Classification of Cuban ants (Hymenoptera: Formicidae) into functional groups. Poeyana Revista Cubana de Zoologia 506: 21-30.
 * Fontenla Rizo J. L. 1997. Lista preliminar de las hormigas de Cuba (Hymenoptera: Formicidae). Cocuyo 6: 18-21.
 * Forel A. 1901. Variétés myrmécologiques. Annales de la Société Entomologique de Belgique 45: 334-382.
 * Forel A. 1908. Fourmis de Costa-Rica récoltées par M. Paul Biolley. Bulletin de la Société Vaudoise des Sciences Naturelles 44: 35-72.
 * Forel A. 1909. Ameisen aus Guatemala usw., Paraguay und Argentinien (Hym.). Deutsche Entomologische Zeitschrift 1909: 239-269.
 * Forel A. 1912. Formicides néotropiques. Part III. 3me sous-famille Myrmicinae (suite). Genres Cremastogaster et Pheidole. Mémoires de la Société Entomologique de Belgique. 19: 211-237.
 * Franco W., N. Ladino, J. H. C. Delabie, A. Dejean, J. Orivel, M. Fichaux, S. Groc, M. Leponce, and R. M. Feitosa. 2019. First checklist of the ants (Hymenoptera: Formicidae) of French Guiana. Zootaxa 4674(5): 509-543.
 * Galkowski C. 2016. New data on the ants from the Guadeloupe (Hymenoptera, Formicidae). Bull. Soc. Linn. Bordeaux 151, 44(1): 25-36.
 * Houadria M., A. Salas-Lopez, J. Orivel, N. Bluthgen, and F. Menzel. 2015. Dietary and temporal niche differentiati on in tropical ants—can they explain local ant coexistence? Biotropica 47(2): 208-217.
 * Kempf W. W. 1961. A survey of the ants of the soil fauna in Surinam (Hymenoptera: Formicidae). Studia Entomologica 4: 481-524.
 * Kempf W. W. 1978. A preliminary zoogeographical analysis of a regional ant fauna in Latin America. 114. Studia Entomologica 20: 43-62.
 * Kempf, W.W. 1972. Catalago abreviado das formigas da regiao Neotropical (Hym. Formicidae) Studia Entomologica 15(1-4).
 * Kusnezov N. 1952. El género Pheidole en la Argentina (Hymenoptera, Formicidae). Acta Zoologica Lilloana 12: 5-88.
 * Kusnezov N. 1978. Hormigas argentinas: clave para su identificación. Miscelánea. Instituto Miguel Lillo 61:1-147 + 28 pl.
 * LaPolla, J.S. and S.P. Cover. 2005. New species of Pheidole (Hymenoptera: Formicidae) from Guyana, with a list of species known from the country. Tranactions of the American Entomological Society 131(3-4):365-374
 * Landero-Torres I., I. Madrid-Neco, J. E. Valenzuela-Gonzalez, M. E. Galindo-Tovar, O. R. Leyva-Ovalle, J. Murguia-Gonzalez, H. E. Lee-Espinosa, and M. A. Garcia-Martinez. 2014. Myrmecofuana from three ornamental agroecosystems with different management and a forest remnant in Ixtaczoquitlan, Veracruz, Mexico. Southwestern Entomologist 39(4): 783-795.
 * Lapolla, J. S., and S. P. Cover. "New species of Pheidole (Hymenoptera : Formicidae) from Guyana, with a list of species known from the country." Transactions of the American Entomological Society 131, no. 3-4 (2005): 365-374.
 * Leponce M., J. H. C. Delabie, J. Orivel, J. Jacquemin, M. Calvo Martin, and A. Dejean. 2019. Tree-dwelling ant survey (Hymenoptera, Formicidae) in Mitaraka, French Guiana, in Touroult J. (ed.), “Our Planet Reviewed” 2015 large-scale biotic survey in Mitaraka, French Guiana. Zoosystema 41 (10): 163-179.
 * Longino J. T. 2013. Ants of Nicargua. Consulted on 18 Jan 2013. https://sites.google.com/site/longinollama/reports/ants-of-nicaragua
 * Longino J. T. L., and M. G. Branstetter. 2018. The truncated bell: an enigmatic but pervasive elevational diversity pattern in Middle American ants. Ecography 41: 1-12.
 * Longino J. T., and R. K. Colwell. 2011. Density compensation, species composition, and richness of ants on a neotropical elevational gradient. Ecosphere 2(3): 16pp.
 * Maes, J.-M. and W.P. MacKay. 1993. Catalogo de las hormigas (Hymenoptera: Formicidae) de Nicaragua. Revista Nicaraguense de Entomologia 23.
 * Mirmecofauna de la reserva ecologica de San Felipe Bacalar
 * Morrison L. W. 1998. A review of Bahamian ant (Hymenoptera: Formicidae) biogeography. Journal of Biogeography 25: 561-571.
 * 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
 * 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.
 * Ottonetti L., L. Tucci, F. Frizzi, G. Chelazzi, and G. Santini. 2010. Changes in ground-foraging ant assemblages along a disturbance gradient in a tropical agricultural landscape. Ethology Ecology & Evolution 22: 7386.
 * 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.
 * Smith M. R. 1937. The ants of Puerto Rico. Journal of Agriculture of the University of Puerto Rico 20: 819-875.
 * Smith M. R. 1942. The relationship of ants and other organisms to certain scale insects on coffee in Puerto Rico. Journal of Agriculture of the University of Puerto Rico 26: 21-27.
 * Solomon, S.E. and A.S. Mikheyev. 2005. The ant (Hymenoptera: Formicidae) fauna of Cocos Island, Costa Rica. Florida Entomologist 88(4):415-423
 * 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
 * Torres J.A. 1984. Niches and Coexistence of Ant Communities in Puerto Rico: Repeated Patterns. Biotropica 16(4): 284-295.
 * Torres, Juan A. and Roy R. Snelling. 1997. Biogeography of Puerto Rican ants: a non-equilibrium case?. Biodiversity and Conservation 6:1103-1121.
 * Vittar, F. 2008. Hormigas (Hymenoptera: Formicidae) de la Mesopotamia Argentina. INSUGEO Miscelania 17(2):447-466
 * Vásquez-Bolaños M. 2011. Lista de especies de hormigas (Hymenoptera: Formicidae) para México. Dugesiana 18: 95-133
 * Wheeler W. M. 1905. The ants of the Bahamas, with a list of the known West Indian species. Bulletin of the American Museum of Natural History 21: 79-135.
 * Wheeler W. M. 1908. The ants of Jamaica. Bulletin of the American Museum of Natural History 24: 159-163.
 * Wheeler W. M. 1908. The ants of Porto Rico and the Virgin Islands. Bulletin of the American Museum of Natural History 24: 117-158.
 * Wheeler W. M. 1922. The ants of Trinidad. American Museum Novitates 45: 1-16.
 * Wheeler, William Morton. 1911. Additions to the Ant-Fauna of Jamaica. Bulletin American Museum of Natural History. 30:21-29.
 * Wheeler, William Morton. 1923. Report on the Ants. The University of Iowa Studies in Natural History. 10(3):3-9.
 * Wild, A. L. "A catalogue of the ants of Paraguay (Hymenoptera: Formicidae)." Zootaxa 1622 (2007): 1-55.
 * Zelikova, T.J. and M.D. Breed. 2008. Effects of habitat disturbance on ant community composition and seed dispersal by ants in a tropical dry forest in Costa Rica. Journal of Tropical Ecology 24:309-316
 * 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 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