Pheidole biconstricta

P. biconstricta is a conspicuous ant in much of the tropical forests of the New World. It forms large colonies, with populations possibly in the tens of thousands, that nest in rotting logs and stumps on the forest floor. John T. Longino (1997) reports that in Costa Rica, “Workers are aggressive, and forage day or night. Large numbers of minor and major workers may be observed swarming out from nests and retrieving live insect prey, with a behavior reminiscent of army ants. Workers also tend Homoptera, and visit extrafloral nectar sources. Colonies may build scattered carton shelters on low vegetation, and tend membracids and other Homoptera beneath them. Workers may aggressively defend extrafloral nectar sources (e.g. Passiflora shoots), driving away herbivores and other ants. Colonies use carton construction to form baffles in rotten wood, and galleries running up tree trunks. At Rara Avis, workers were observed tending large riodinid larvae under carton galleries. Founding queens are found under loose bark of dead wood, in dead branches, and very commonly under epiphyte mats on recently fallen trees.” (Wilson 2003)

Identification
See the description in the nomenclature section.

Distribution
Widespread and locally abundant, occurring mostly in tropical moist forests from Guatemala to Brazil and Bolivia; present in Trinidad but absent from Tobago and the rest of the West Indies. Ranges to at least 1500 m in Costa Rica and to 2500 m in Colombia. (Wilson 2003)

Distribution based on Regional Taxon Lists
Neotropical Region: Belize, Bolivia, Brazil, Colombia, Costa Rica, Ecuador, Guatemala, Honduras, Mexico , Nicaragua, Panama, Peru, Trinidad and Tobago, Venezuela.

Worker
Minor

Biology
Charles Kugler (1979d) has described the capture of live insect prey by “gang-pulling, and the hypertrophial pygidial glands, which secrete a viscous gumming agent and irritant when smeared on enemies. Another behavior unusual for Pheidole is the lifting of the gaster toward the enemy, making release of the toxin material more effective. Alarm pheromones also emanate from the same gland.”

Nomenclature

 *  biconstricta. Pheidole biconstricta Mayr, 1870a: 399 (s.w.) COLOMBIA. Senior synonyn of bicolor, burtoni, festata, holmgreni, hybrida, lallemandi, rubicunda, surda and material of the unavailable names angustella, fuscata, regina referred here: Wilson, 2003: 143..
 * bicolor. Pheidole biconstricta r. bicolor Emery, 1890b: 50 (s.w.) COSTA RICA. [Also described as new by Emery, 1894k: 55.] Raised to species: Dalla Torre, 1893: 88. Subspecies of biconstricta: Forel, 1899c: 64; Emery, 1922e: 98. Junior synonym of biconstricta: Wilson, 2003: 143.
 * rubicunda. Pheidole biconstricta r. rubicunda Emery, 1890b: 50 (s.w.) COSTA RICA. [Also described as new by Emery, 1894k: 55.] Raised to species: Dalla Torre, 1893: 96. Subspecies of biconstricta: Forel, 1899c: 64. Junior synonym of biconstricta: Wilson, 2003: 143.
 * hybrida. Pheidole biconstricta subsp. hybrida Emery, 1894c: 154 (s.w.) BOLIVIA. Borgmeier, 1934: 97 (m.). Junior synonym of biconstricta: Wilson, 2003: 143.
 * lallemandi. Pheidole radoszkowskii r. lallemandi Forel, 1901c: 133 (s.w.) COLOMBIA. Subspecies of biconstricta: Forel, 1901e: 362; Forel, 1912f: 222. Junior synonym of biconstricta: Wilson, 2003: 143.
 * surda. Pheidole biconstricta var. surda Forel, 1912f: 222 (s.w.) COSTA RICA. Junior synonym of biconstricta: Wilson, 2003: 143.
 * burtoni. Pheidole (Pheidole) biconstricta subsp. burtoni Mann, 1916: 436, pl. 3, fig. 24 (s.) BRAZIL. Junior synonym of biconstricta: Wilson, 2003: 143.
 * festata. Pheidole holmgreni subsp. festata Wheeler, W.M. 1925a: 20 (s.w.) BOLIVIA. Junior synonym of biconstricta: Wilson, 2003: 143.
 * holmgreni. Pheidole holmgreni Wheeler, W.M. 1925a: 18 (s.w.) BOLIVIA. Junior synonym of biconstricta: Wilson, 2003: 143.
 * inermis. Pheidole inermis Mayr, 1870b: 984 (s.w.) MEXICO.
 * Species status: Wilson, 2003: 146.
 * Junior synonym of biconstricta: Longino, 2019: 27.

Taxonomic Notes
Longino (2019) - Pheidole inermis is a version of P. biconstricta that lacks any trace of a propodeal spine. There is continuous variation in the size of the propodeal spine in P. biconstricta, and in some cases it is reduced to a minute denticle. DNA barcoding data places three specimens that lack the propodeal spine among P. biconstricta specimens. I treat P. inermis as an extreme of variation within P. biconstricta.

Description
From Wilson (2003): I have examined the types of all the biconstricta related forms (Pheidole biconstricta Mayr 1870a: 399. Syn.: Pheidole biconstricta subsp. bicolor Emery 1890c: 50, n. syn.; Pheidole biconstricta r. rubicunda Emery 1890c: 50, n. syn.; Pheidole biconstricta rubicunda var. fuscata Emery 1890c: 51 (unavailable name, quadrinomial); Pheidole biconstricta subsp. hybrida Emery 1894d: 154, n. syn.; Pheidole radoszkowskii r. lallemandi Forel 1901d: 133, n. syn.; Pheidole biconstricta bicolor var. regina Forel 1908c: 52, n. syn.; Pheidole biconstricta hybrida var. angustella Forel 1912g: 222 (unavailable name, quadrinomial); Pheidole biconstricta var. surda Forel 1912g: 222, n. syn; Pheidole biconstricta subsp. burtoni Mann 1916: 436, n. syn. (provisional); Pheidole holmgreni Wheeler 1925a: 18, n. syn.; Pheidole holmgreni festata Wheeler 1925a: 20, n. syn.).

What I have regarded here as the single species biconstricta is highly variable in details of size, sculpturing, and color, both locally and geographically, with general and overlapping intergradation. Closer studies with more material may well reveal biconstricta to be a complex of sibling species, to which at least some of the names will apply, but for the time being I have chosen the more conservative arrangement, that is, recognition of a single, very variable species.

A member of the biconstricta group distinguished as follows.

Major: large, with well-developed propodeal spines and prominent rounded humeral angles; head dorsal surface foveolate, space between eye and antennal fossa rugoreticulate; first 2 gastral tergites mostly shagreened and opaque; pilosity sparse on head, moderate on rest of body; body color reddish yellow (“orange”) to dark reddish brown, usually a lighter shade; one variant (bicolor, possibly a distinct species) has a contrasting paler gaster.

Minor: head conspicuously narrowed, with nuchal collar; mesonotal convexity well-developed, leaning forward; propodeal spines short, thick, and erect; occiput, mesothorax, and propodeum foveolate and opaque; anterior half of first gastral tergite shagreened and opaque.

MEASUREMENTS (mm)
 * Lectotype major: HW 1.62, HL 1.70, SL 1.08, EL 0.22, PW 0.82.
 * Paralectotype minor: HW 0.78, HL 0.94, SL 1.12, EL 0.16, PW 0.54.

COLOR Major: yellowish brown (possibly faded).

Minor: light reddish brown; otherwise, see Diagnosis above.

P. inermis MEASUREMENTS (mm)
 * Lectotype major: HW 1.84, HL 1.72, SL 1.06, EL 0.24, PW 0.84.
 * Paralectotype minor: HW 0.80, HL 0.94, SL 1.10, EL 0.14, PW 0.54.

COLOR Major: most of body light reddish brown, with mandibles, postpetiole, and gaster medium reddish brown.

Minor: gaster medium brown, rest of body light reddish brown.

'''Figure. Upper: lectotype, major (with 2 hypostomal teeth; a 4-toothed variant is also shown). Lower: paralectotype, minor. Scale bars = 1 mm.''' '''Figure. Upper: lectotype, major. Lower: paralectotype, minor. Scale bars = 1 mm.'''

Etymology
L biconstricta, constricted (pinched) twice, once in front of the mesonotum and once behind it. (Wilson 2003)

L inermis, unarmed, referring to the absence of propodeal spines. (Wilson 2003)

References based on Global Ant Biodiversity Informatics

 * Borgmeier T. 1934. Contribuição para o conhecimento da fauna mirmecológica dos cafezais de Paramaribo, Guiana Holandesa (Hym. Formicidae). Archivos do Instituto de Biologia Vegetal (Rio de Janeiro) 1: 93-111.
 * Dattilo W. et al. 2019. MEXICO ANTS: incidence and abundance along the Nearctic-Neotropical interface. Ecology https://doi.org/10.1002/ecy.2944
 * Davidson, D.W. 2005. Ecological stoichiometry of ants in a New World rain forest. Oecologia 142:221-231
 * Delabie J. H. C., R. Céréghino, S. Groc, A. Dejean, M. Gibernau, B. Corbara, and A. Dejean. 2009. Ants as biological indicators of Wayana Amerindian land use in French Guiana. Comptes Rendus Biologies 332(7): 673-684.
 * Emery C. 1894. Studi sulle formiche della fauna neotropica. VI-XVI. Bullettino della Società Entomologica Italiana 26: 137-241.
 * Escalante Gutiérrez J. A. 1993. Especies de hormigas conocidas del Perú (Hymenoptera: Formicidae). Revista Peruana de Entomología 34:1-13.
 * Fernandes I., and J. de Souza. 2018. Dataset of long-term monitoring of ground-dwelling ants (Hymenoptera: Formicidae) in the influence areas of a hydroelectric power plant on the Madeira River in the Amazon Basin. Biodiversity Data Journal 6: e24375.
 * Fernández, F. and S. Sendoya. 2004. Lista de las hormigas neotropicales. Biota Colombiana Volume 5, Number 1.
 * Field Museum Collection, Chicago, Illinois (C. Moreau)
 * 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.
 * INBio Collection (via Gbif)
 * 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. 1972. Catalago abreviado das formigas da regiao Neotropical (Hym. Formicidae) Studia Entomologica 15(1-4).
 * Kusnezov N. 1953. La fauna mirmecológica de Bolivia. Folia Universitaria. Cochabamba 6: 211-229.
 * 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
 * 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.
 * Lapolla, J.S., T. Suman, J. Soso-Calvo and T.R. Schultz. 2006. Leaf litter ant diversity in Guyana. Biodiversity and Conservation 16:491510
 * Longino J. T. 2019. Pheidole (Hymenoptera, Formicidae) of Middle American wet forest. Zootaxa 4599: 1-126
 * 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. et al. ADMAC project. Accessed on March 24th 2017 at https://sites.google.com/site/admacsite/
 * Lopes M. C., G. P. A. Lamarre, C. Baraloto, P. V. A. Fine, A. Vincentini, and F. B. Baccaro. 2019. The Amazonas-trap: a new method for sampling plant-inhabiting arthropod communities in tropical forest understory. Entomologia Experimentalis et Applicata https://doi.org/10.1111/eea.12797
 * Maes, J.-M. and W.P. MacKay. 1993. Catalogo de las hormigas (Hymenoptera: Formicidae) de Nicaragua. Revista Nicaraguense de Entomologia 23.
 * Mann W. M. 1916. The Stanford Expedition to Brazil, 1911, John C. Branner, Director. The ants of Brazil. Bulletin of the Museum of Comparative Zoology 60: 399-490
 * 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.
 * Radoszkowsky O. 1884. Fourmis de Cayenne Française. Trudy Russkago Entomologicheskago Obshchestva 18: 30-39.
 * Robson Simon Ant Collection, 05-Sept-2014
 * Solomon, S.E. and A.S. Mikheyev. 2005. The ant (Hymenoptera: Formicidae) fauna of Cocos Island, Costa Rica. Florida Entomologist 88(4):415-423
 * Wheeler W. M. 1925. Neotropical ants in the collections of the Royal Museum of Stockholm. Arkiv för Zoologi 17A(8): 1-55.
 * Wilson, E.O. 2003. Pheidole in the New World: A Dominant, Hyperdiverse Genus. Harvard University Press
 * 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.