Megalomyrmex modestus

Large polygynous colonies (>2000 workers) have been collected nesting in or under logs and in diffuse soil at the base of trees. Longino (2010) reported a colony ~ 20 cm deep soil on a creek embankment. In a Costa Rican population where Apocephalus phorid flies (B. Brown det.) were seen attacking workers, individuals walked around with their gasters tucked under their bodies. It is difficult to bait this species with Keebler Pecan Sandies as they are mainly subterranean and nocturnal; if the bait is placed on top of a soil nest they will emerge and bury the cookies. Colonies construct soil galleries on the side and at the base of trees when tending immature white flies (Aleyrodoidea: Aleyrodidae). Megalomyrmex modestus was the most frequently collected Megalomyrmex species during Malaise trapping in La Selva, Costa Rica, with well over fifty specimens recovered. (Boudinot et al. 2013)

Identification
Boudinot et al. (2013) - Worker uniquely identified among Central American Megalomyrmex by the following combination: (1) mandible with 5 robust, equally-spaced, subequal triangular teeth; (2) eye small (EL < 0.25); (3) dorsal face of mandible striate; (4) clypeus with triangular median process; (5) disc of katepisternum roughened in ventral half by striations and weak carinulae; (6) petiolar peduncle long and thin. Queen similarly identifiable as worker, but alate, with larger eyes, and less-robust petiolar and postpetiolar nodes. Male uniquely identified among Central American Megalomyrmex by either of the following: (1) postpetiolar helcium laterally compressed; (2) telomere with a dorsomedian dentiform process. Otherwise recognized by the following combination: (1) forewing 1m-cu present; (2) scape length less than antennomeres 2–4; (3) occipital carina visible in full-face view; (4) mesosoma robust.

Workers, alate queens, and males of Megalomyrmex modestus have relatively dense, short ocular setae. Meglomyrmex modestus workers are set off from the Central American leoninus- and modestus-group species by several traits unique in this two groups: (1) body comparatively small (ML 1.40–1.65 vs. ML 1.83–2.71); (2) eyes relatively small (EI 20–21 vs. EI 25–33); (3) dorsal mandibular face striate (vs. smooth); (4) anterior clypeal margin strongly angulate medially (vs. weakly dentate or convex); (5) antennal club distinctly broader than preceeding antennomeres of funiculus (vs. subequal in width to preceeding antennomeres); (6) gastral shoulders strong (vs. weak or indistinct); petiolar peduncle long and thin (vs. comparatively shorter and stout); and body completely pale yellow (vs. head and mesosoma orange and/or gaster black). However, given these distinct characters, it is notable that M. modestus shares with Megalomyrmex wallacei (another modestus group species) a notch between the basalmost and basal mandibular margins.

Distribution
Sea-level to 940 m elevation.

Distribution based on Regional Taxon Lists
Neotropical Region: Colombia, Costa Rica, Ecuador, French Guiana, Panama, Venezuela.

Biology
Longino (2010): Forel (1899) reported that the type specimens were collected by A. Alfaro from a rotten log. Kugler collected specimens in Colombia, from a “large nest in rotten log, soil - rocks, under forest” (reported in Brandão 1990).

In Costa Rica, M. modestus is found in mature wet forest of the Atlantic slope. Although it occurs at sea level, it is most abundant at middle elevations, 500–800 m. Nests are subterranean and very large. From these nests, inconspicuous covered galleries extend large distances up trees and under liana stems, and workers may be found under epiphyte mats in the canopy. Workers tend Coccoidea under the shelters. Workers are occasionally collected at baits on the forest floor, but in general they are not often seen as exposed foragers.

At Refugio Eladio, on the Río Peñas Blancas, I found a nest on the bank of a small stream. An area of excavated soil about 2m across covered the ground. There were hundreds of small entrances, and excavations in the surface soil turned up only scattered workers, and no brood. As I dug deeper into the center of the nest (more than 20cm deep) workers became more dense and I began finding adult males and some brood. At a depth of about arm's length workers were very dense, brood more abundant, and two dealate queens were turned up. The soil was very friable, and I could not discern any gallery or chamber structure. Given that I was digging in only a small part of the colony, and turned up two queens, the colony must have had many queens. Around the periphery of the colony there were abundant Coccoidea (Pseudococcidae?) in the soil.

Brandão (2003) - Three workers of this species, from Rio Toro Amarillo, near Guapiles, Alajuela province, belonging to the MZSP collection, were erroneously listed in the 1990 revision as gynes. Also from Costa Rica, I received from the LACM two workers of M. modestus collected in La Selva, Heredia province (10°26’N, 83°59’W), March 1974 and August 4, 1974 (# 0515-1740) in pitfall traps set at the forest litter by Talbot & Van Devender. From the INBC collection, I received several workers of M. modestus collected at Río Peñas Blancas, Alajuela province (10°19’N, 84°43’W), 800-950 m, by J. Longino (different acc. numbers and dates), including a gyne and a male (# 2019) collected in April 27, 1988. J. Longino also sent me two M. modestus workers from Reserva Biologica Hitgi-Cerere, Limón province, Costa Rica (09°40’N, 83°02’W), 200 m of altitude, collected in August 29, 1985, # 942-S.

From Venezuela, John Lattke kindly sent me two samples of M. modestus workers (24 an 13 workers respectively) from, Ayan Tepui, 1500 m of altitude (ca. Salto Angel) Estado Bolivar (5°57’N, 62°30’W), collected in November 19, 1984 by J. Lattke & K. Jaffé (leg. # 611), and southwestern Kamarcabarai Tepui, 1800 m, 50°53’N, 62°01’W, collected in May 24, 1986 by J. Lattke (leg. # 853).

From Instituto Humboldt, I recently studied four workers collected in October 28, 2000 at Parque Nacional Farallones de Cali, Embalses de Alto Anchicaya Valle del Cauca, Colombia (03 26’N, 76°48’W), 650 m of altitude (W1), by S. Barrio and others. From the same locality and collection (altitude 900 m), I studied another sample of nine workers collected using Malaise traps in November 8 to 21, 2000 by S. Sarria (samples M. 1105, 1111, and 1114). From the IHVL collection, I studied a very tiny dealated gyne collected in Reserva Nacional La Planada, Nariño, 1850 m, in July to August 1995, by C. Estrada. The sculpturation on the head disk and mandibles is very much effaced, while in the mesopleura and over the propodeum the sculpture is much stronger than in the other studied gynes. However, this Colombian specimen agrees in all other characters with M. modestus, and could represent a microgyne, expected in a genus like Megalomyrmex where female reproductives vary in size and development from typical alate gynes. A worker from the same sample agrees with other M. modestus workers. I also studied another worker of the same species, from the same locality (01 09’N, 77°68’W), collected by F. Escobar in 1994 (leg. # 39).

Nomenclature

 * . Megalomyrmex modestus Emery, 1896g: 94 (w.) COSTA RICA.
 * Type-material: lectotype worker (by designation of Brandão, 1990b: 425), 1 paralectotype worker.
 * Type-locality: lectotype Costa Rica: Suerre, nr Jiménez (A. Alfaro); paralectotype with same data.
 * Type-depository: MSNG.
 * Brandão, 1990b: 426 (q.m.).
 * Status as species: Forel, 1899c: 58; Wheeler, W.M. 1909b: 236 (in key); Emery, 1922e: 190; Wheeler, W.M. 1925a: 33 (in key); Ettershank, 1966: 105; Kempf, 1972a: 140; Brandão, 1990b: 425 (redescription); Brandão, 1991: 355; Bolton, 1995b: 249; Brandão, 2003: 148; Longino, 2010: 46; Boudinot, et al. 2013: 50; Fernández & Serna, 2019: 806.
 * Distribution: Colombia, Costa Rica, Panama, Venezuela.

Worker
Brandão (1990) - Mandibles striate; clypeus with anterior medium denticle; antennal club with 4 segments; frontal suture not impressed; occipital margin not raised; pronotum dorsal profile round; promesonotal suture not impressed dorsally, mesosternum and metasternum with sharp paired teeth: propodeum faces meeting in an obtuse angle in lateral view, sometimes forming a carina: declivity smooth; dorsal profile of petiole round in frontal view: ventral face of postpetiole without process.

Pilosity: small erect hairs (ca. .1 mm) uniformly distributed on the whole body, but dorsal aspect of thorax, petiole and postpetiole, apex of nodes, and gaster, with long hairs (ca. .2mm).

Color: bright brownish-yellow.

Queen
Brandão (1990) - Differing from workers in superficial sculptures covering most of head capsule; anterior ocellus similar in size to the posterior ones; pterothorax with areas of striation at margins of pronotum and mesonotum; paraptera with longitudinal rugosities on lateral areas; mesonotum with notaulus and parapsidal sutures; petiolar spiracles laterally produced; ventral face of petiole with a longitudinal translucid flange; ventral face of postpetiole with an acute tooth.

Male
Boudinot et al. (2013) - (n=2): HW 0.87–0.89, HL 0.83, SL 0.27, EL 0.43–0.45, ML 1.60–1.64, CI 105–107, SI 33, EI 49–51.

Head Antenna with 13 antennomeres, none of which are kinked, nor forming a club; scape length shorter than eye length. Entire antenna brown, except apex of terminal antennomere which is pale. Palpal formula 4,3. Mandible subfalcate; masticatory margin with three triangular teeth; apical tooth about twice the length of the subbasal tooth; subbasal tooth subequal in size to basal tooth from which it is separated by a short diastema. Dorsal face of mandible striate, at least at midlength. Minimum distance between lateral ocellus and compound eye about three lateral ocellus lengths. Compound eyes with dense, short ocular setae. Occipital carina visible in full-face view. Mesosoma Mesosoma robust. Notauli absent. Parapsidal lines weak. Foraminal carina present. Metasternum with a triangular process. Basitarsi tubular. Pterostigma elongated apically. Forewing crossvein 1m-cu present; submarginal cell 1 one fourth as wide as long; terminal abscissa of M branches from Rs at or just basad 2r-rs. Metasoma Basipetiolar carina arc-shaped; weak; most distinct laterally, obsolescent medially. Ventrolateral longitudinal carina of petiole present. Petiolar spiracle in anterior fourth. Posterior margin of petiole with distinct girdling carinae; postpetiole without. Postpetiolar helcium strongly compressed laterally. Postpetiolar spiracle placed about midlength of the lateral tergal margin. Sternum of postpetiole produced anteriorly; this process angular, sloping gently to posterior margin. Genitalia Abdominal sternum IX lateral margins subparallel, arching apically to posterior margin which is ventromedially produced as a relatively strongly sclerotized apical triangle; this triangle is keeled ventrally and produced apically; lateral faces of this apical triangle concave. Telomere triangular, elongate, with apex acutely angled; telomere with medially-directed dentiform process (hence “shouldered”); medial face of telomere sclerotized, arched; ventral margin with two anteriorly-directed sclerotized denticles. Cuspis with posterodorsal process; apical margin weakly convex. Digitus narrow; dorsal margin explanate, with posterodorsal margin subangular; ventral margin parabolic; apex triangular; ventral margin slightly obscuring apex of cuspal posterodorsal process. Valviceps about half as tall as long, subrectangular; dorsal margin curving smoothly through apical margin to meet the sclerotized ventral margin; basal two thirds of ventral margin linear; penisvalvar teeth short, truncate, close-set.

Type Material
Boudinot et al. (2013) - Lectotype worker (designated by Brandão 1990: 425): COSTA RICA: Suerre [near Jimenez, Limón Prov.] (not examined).

References based on Global Ant Biodiversity Informatics

 * Boudinot B. E., T. P. Sumnicht, and R. M. M. Adams. 2013. Central American ants of the genus Megalomyrmex Forel (Hymenoptera: Formicidae): six new species and keys to workers and males. Zootaxa 3732(1): 1-82.
 * Brandao, C.R.F. 1991. Adendos ao catalogo abreviado das formigas da regiao neotropical (Hymenoptera: Formicidae). Rev. Bras. Entomol. 35: 319-412.
 * Brandão C. R. F. 1990. Systematic revision of the Neotropical ant genus Megalomyrmex Forel (Hymenoptera: Formicidae: Myrmicinae), with the description of thirteen new species. Arquivos de Zoologia (São Paulo) 31: 411-481
 * Brandão C. R. F. 2003. Further revisionary studies on the ant genus Megalomyrmex Forel (Hymenoptera: Formicidae). Papeis Avulsos de Zoologia (São Paulo) 43: 145-159
 * Emery C. 1896. Studi sulle formiche della fauna neotropica. XVII-XXV. Bullettino della Società Entomologica Italiana 28: 33-107.
 * Fernández F., E. E. Palacio, W. P. Mackay, and E. S. MacKay. 1996. Introducción al estudio de las hormigas (Hymenoptera: Formicidae) de Colombia. Pp. 349-412 in: Andrade M. G., G. Amat García, and F. Fernández. (eds.) 1996. Insectos de Colombia. Estudios escogidos. Bogotá: Academia Colombiana de Ciencias Exactas, Físicas y Naturales, 541 pp
 * Fernández, F. and S. Sendoya. 2004. Lista de las hormigas neotropicales. Biota Colombiana Volume 5, Number 1.
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 * INBio Collection (via Gbif)
 * Longino J. T. 2010. A taxonomic review of the ant genus Megalomyrmex Forel (Hymenoptera: Formicidae) in Central America. Zootaxa 2720: 35-58
 * 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., J. Coddington, and R. K. Colwell. 2002. The ant fauna of a tropical rain forest: estimating species richness three different ways. Ecology 83: 689-702.
 * 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.
 * Longino J. et al. ADMAC project. Accessed on March 24th 2017 at https://sites.google.com/site/admacsite/
 * Siqueira de Castro F., A. B. Gontijo, P. de Tarso Amorim Castro, and S. Pontes Ribeiro. 2012. Annual and Seasonal Changes in the Structure of Litter-Dwelling Ant Assemblages (Hymenoptera: Formicidae) in Atlantic Semideciduous Forests. Psyche doi:10.1155/2012/959715
 * Siqueira de Castro F., A. B. Gontijo, W. Duarte da Rocha, and S. Pontes Ribeiro. 2011. As comunidades de formigas de serapilheira nas florestas semidecíduas do Parque Estadual do Rio Doce, Minas Gerais. MG.BIOTA, Belo Horizonte 3(5): 5-24.
 * Vieira J. M. 2005. Nuevos registros de Megalomyrmex Forel, 1884 (Hymenoptera: Formicidae) para Ecuador, con la descripción del macho de M. glaesarius Kempf, 1970. Boletín de la Sociedad Entomológica Aragonesa 36: 81-84.