Brachymyrmex heeri

 Brachymyrmex heeri nests under stones and other objects, often in disturbed areas, as well as in bamboo (Fagundes et al., 2010).

Identification
Small workers (1.2 to 2mm) with nine segmented antenna. Color varies from yellow to brown.

Distribution
Central America, South America and the Caribbean. Also introduced in a few locations in Europe and in the Galapagos Islands.

Distribution based on Regional Taxon Lists
Neotropical Region: Argentina, Bolivia, Brazil , Colombia, Costa Rica, Ecuador, French Guiana, Galapagos Islands, Guatemala, Guyana, Haiti, Lesser Antilles, Mexico, Panama, Paraguay, Puerto Rico, Uruguay, Venezuela. Palaearctic Region: Switzerland, Ukraine.

Biology
Ortiz-Sepulveda et al. (2019) - Brachymyrmex heeri was described from specimens collected from a tropical-orchid greenhouse in Switzerland (Forel 1874). The ants were found climbing on various plants and associated with aphids. No nest was located but nesting information was included with the descriptions of two species that are now synonyms of B. heeri. A nest of B. giardi var. cordobensis (Santschi 1929) was found at the base of a tree and B. physogaster (Kusnezov 1960) was collected from shaded, loamy soil.

The description of the now synonymized B. giardi var. cordobensis (Santschi 1929) noted physogastric specimens and “worker-queens” with a strongly expanded gaster. Kusnezov (1960) labeled physogastric specimens (of B. physogaster) we have assigned to B. heeri as honey pot workers. These were observed hanging from the ceiling of the chambers of the nest, similarly to Myrmecocystus species. De Zolessi et al. (1978) also reported repletes of Brachymyrmex melensis (= Brachymyrmex giardi) were observed hanging from the roof together with more typical B. melensis workers.

Our examination of Santschi's B. heeri specimens, and other atypical-morphed specimens of this species, have been exclusively limited to a robust, worker-like morph with an expanded gaster. Beyond their ocelli, these ants had no morphological features reminiscent of a queen. Compared to the more typical Brachymyrmex workers in their colonies, the worker-like form exhibited a larger body size, a subquadrate head, a posterior cephalic margin that is slightly concave to almost flat, scapes that barely reach the posterior margin of the head, three ocelli, eyes with approximately 11 ommatidia along their maximal diameter, a deep metanotal groove that is wider than the diameter of the metathoracic spiracles, and a strongly expanded gaster. To exclude the possibility that the museum samples we studied reflect a mixture of two species, we sequenced specimens of both morphs, and found them to be genetically very similar. In sum, B. heeri appear to have dimorphic workers, as opposed to a worker-queen intercaste, but more research (with more collections and nest observations) is needed.

The wide distribution of B. heeri suggests this may be a species complex. Further study is needed, with more sampling, examining the morphological and molecular variation of B. heeri across its range. Likewise, it would be interesting to study the biology of the variable morphs known from some colonies. This would ideally include examining the morph's distribution within colonies, variation across the range of B. heeri and within colonies, their behavior, and the underlying developmental processes that produce this caste.

It is not known why physogastry would be present in some Brachymyrmex species but not others. Kusnezov (1960) suggested it may relate to a trophobiotic lifestyle, whereas others suggested it is an adaptation to periodically arid conditions or food scarcity (Forel 1902; Wheeler 1910). However, Creighton (1950) argued exactly the opposite by suggesting that physogastry may develop when a xerophyte species encounters and adapts to less arid and more resource-rich habitats. It is interesting that the habitat from which B. physogaster was reported by Kusnezov (1960) does not appear to confirm the “scarcity hypothesis.” Regardless, these curious circumstances currently only allow for speculation. More specimens, and additional observations of nests with worker-like morphs, would be needed to assess if there any ecological correlates associated with this physogastric, worker-like Brachymyrmex caste.

Greater Antilles
In Haiti this species was "found to be very common beneath stones, boards etc. on the shores of the salty Lake Assuei (Haiti)" (Wheeler and Mann 1914).

Wheeler (1908) found "several small colonies under stones" in Puerto Rico and in the Bahamas (Wheeler 1905) discovered "a single colony, including several winged females, of the typical form of this species.under a stone on a key in the Southern Bight, Andros Island, May 23 (1904)."

Costa Rica
Longino 2004 - This is a common species of synanthropic habitats in Costa Rica. It can be found in small parks in the middle of San José, in hotel landscaping, along road edges, in scrubby second growth vegetation, and in pastures. It occurs in almost any bioclimatic region: dry Guanacaste lowlands, wet Atlantic slope lowlands, Central Valley urban areas, and roads and pastures near Monteverde cloud forest. Nests are often under stones on the ground but also occur in cavities in low vegetation. Colonies are polygynous, with multiple dealate queens often occurring together in nests. Workers are omnivorous and opportunistic foragers.

Nomenclature

 * . Brachymyrmex heeri Forel, 1874: 89, 91, figs. 16, 20 (w.) SWITZERLAND (hothouse in botanical garden, Zurich).
 * [Note: description repeated in Latin, p. 92.]
 * Forel, 1876: 52 (q.m.).
 * Status as species: Forel, 1876: 52; Emery, 1878b: 47; Emery & Forel, 1879: 466; Forel, 1881: 3; André, 1882b: 214 (in key); Mayr, 1886d: 431; Cresson, 1887: 257; Dalla Torre, 1893: 174; Forel, 1895b: 106; Forel, 1899c: 123; Wheeler, W.M. 1904e: 304; Wheeler, W.M. 1905b: 132; Wheeler, W.M. 1906e: 350; Forel, 1907e: 9; Wheeler, W.M. 1908a: 153; Wheeler, W.M. 1911a: 29; Forel, 1912i: 62; Wheeler, W.M. & Mann, 1914: 44; Forel, 1915d: 49 (in key); Wheeler, W.M. 1916m: 591; Mann, 1920: 431; Wheeler, W.M. 1921f: 166; Wheeler, W.M. 1922c: 15; Wheeler, W.M. 1923c: 5; Santschi, 1923b: 664; Emery, 1925b: 42; Stärcke, 1926: 118 (in key); Borgmeier, 1927c: 141; Menozzi, 1927c: 268; Menozzi & Russo, 1930: 166; Menozzi, 1931b: 274; Aguayo, 1932: 224; Santschi, 1933e: 122; Eidmann, 1936b: 92; Smith, M.R. 1937: 866; Santschi, 1939e: 319; Wheeler, W.M. 1942: 253; Kusnezov, 1953b: 339; Kempf, 1961b: 522; Bernard, 1967: 280 (redescription); Kempf, 1970b: 340; Kempf, 1972a: 39; Alayo, 1974: 26 (in key); Bolton, 1995b: 82; Wetterer & Wetterer, 2004: 215; Wild, 2007b: 27; Branstetter & Sáenz, 2012: 255; Borowiec, L. 2014: 24 (see note in bibliography); Radchenko, 2016: 348; Fernández & Ortiz-Sepúlveda, 2019: 728; Lubertazzi, 2019: 78; Ortiz-Sepúlveda, et al. 2019: 503 (redescription).
 * Senior synonym of cordobensis: Ortiz-Sepúlveda, et al. 2019: 503.
 * Senior synonym of goeldii: Ortiz-Sepúlveda, et al. 2019: 503.
 * Senior synonym of physogaster: Ortiz-Sepúlveda, et al. 2019: 503.
 * cordobensis. Brachymyrmex giardi var. cordobensis Santschi, 1929d: 309 (w.) ARGENTINA (Córdoba).
 * Subspecies of giardi: Kempf, 1972a: 39; Bolton, 1995b: 82.
 * Junior synonym of heeri: Ortiz-Sepúlveda, et al. 2019: 503.
 * goeldii. Brachymyrmex goeldii Forel, 1912i: 65 (w.) BRAZIL (São Paulo).
 * Status as species: Luederwaldt, 1918: 48; Santschi, 1923b: 663; Emery, 1925b: 42; Borgmeier, 1927c: 141; Kempf, 1972a: 39; Bolton, 1995b: 82.
 * Junior synonym of heeri: Ortiz-Sepúlveda, et al. 2019: 503.
 * physogaster. Brachymyrmex physogaster Kusnezov, 1960c: 382, figs. 1-4 (w.) ARGENTINA (Salta).
 * Status as species: Bolton, 1995b: 82.
 * Junior synonym of heeri: Ortiz-Sepúlveda, et al. 2019: 503.

Description
Ouvriere: Longueur 1, 2 a 2mm

Tout le corps court, large, trapu. Mandibules munies de cinq dents (les autres especes en ont quatre). Pas trace d'ocelles (les autres especes ont trois ocelles). Yeux composes d'environ 35 facettes chacun ; ils sont situes legerement en avant du milieu des bords lateraux de la tete. Palpes maxillaires longs; le second et le troisieme article sont plus longs que les quatre autres. Les palpes labiaux sont assez longs aussi; leurs quatre articles sont egaux entre eux. Chaperon (c.) en forme de capuchon (comme chez le Brachymyrmex patagonicus), sans carene, recouvrant un peu les mandibules, fortement voute de droite a gauche, moins fortement d'avant en arriere: Il est arrondi posterienrement, parfois aussi un peu echancre au milieu de son bord posterieur ; il ne se prolonge pas ou presque pas entre les articulations des antennes (entre les aretes frontales). Aire frontale distincte, triangulaire, souvent un peu arrondie a son angle posterieur. Sillon frontal distinct. Aretes frontales courtes, divergentes, a peine recourbees. La tete, un peu aplatie en dessus, basse en arriere, est echancree a son bord posterieur. Premier article du fouet des antennes un peu plus long que les deux suivants ensemble ; dernier (8m.) article du fouet aussi long que les trois qui le precedent a. la fois (fg.) Thorax tres court, surtout le mesonotum et le pronotum qui sont voutesis et forment ensemble seulement les 2/5 de sa longueur totale. Un sillon transversal enfonce, assez etroit, separe le mesonotum du metanotum; ce sillon ne renferme pas de petite spirale (il en renferme une chez le Brachymyrmex tristis). La face basale du metanotum est extremement courte, convexe (mtb); sa face declive, longue, large et plate (mtd), va en pente douce et egale jusqu'au pedicule. Eperons des pattes anterieures tres forts. Ecaille encore plus fortement inclinee en avant que chez la Plagiolepis pygmaea (e). Premier segment de l'abdomen prolonge en avant et recouvrant l'ecaille ; abdomen grand. D'un jaune plus on moins brunatre, grisatre ou rougeatre. Dessus de la tete et de l'abdomen plus fonce. Tout le corps luisant; chaperon et face declive du metanotum lisses; quelques rugosites assez grossieres sur les joues; tout le reste tres finement rugueux ponctue. Pubescence grisatre, assez forte sur l'abdomen, un peu moins forte sur la tete, plus faible sur le thorax, les pattes et les antennes, nulle sur le chaperon et sur la face declive du metanotum. Poils epars, un peu partout, sauf sur les pattes et sur les antennes. Les deux stigmates posterieurs sont gros et tres distincts.

Ortiz-Sepulveda et al. (2019) - Lectotype and paralectotypes (n = 5). HL1 0.39–0.43; HL2 0.27–0.30; HL3 0.10; HW 0.35–0.41; SL 0.37–0.39; EL 0.10–0.11; WL 0.39–0.45; PnL 0.10; PnW 0.25–0.31; ML 0.08–0.12; MW 0.16–0.20; Indices CI 90.00–100.00; SI1 90.48–111.11; SI2 126.67–142.86; OI1 23.81–27.78; OI2 22.72–25.00. Additional material (n = 5). HL1 0.41–0.60; HL2 0.27–0.39; HL3 0.10–0.13; HW 0.40– 0.66; SL 0.36–0.52; EL 0.09–0.19; WL 0.35–0.68; PnL 0.11– 0.16; PnW 0.27–0.45; ML 0.09–0.21; MW 0.18–0.35; Indices CI 93.75–110.81; SI1 78.05–93.62; SI2 114.29–135.71; OI1 21.74–29.27; OI2 16.22–28.00.

Head. Slightly longer than wide in full face view; posterior cephalic margin slightly concave. Clypeus with a rounded anterior margin and five long, erect hairs of which a single, usually conspicuous hair is near the anterior margin, two hairs are in mediolateral position, and two more near the toruli; other hairs on the clypeus are markedly shorter and appressed or decumbent. Toruli surpassing the posterior clypeal margin in oblique anterodorsal view. The scapes surpass the posterior margin of the head by a length smaller than the maximal diameter of the eye; they bear decumbent hairs. Ocelli absent. Eyes are positioned on the cephalic midline and have 6–7 ommatidia along their maximal diameter.

Mesosoma. With several decumbent hairs and usually two erect hairs on the pronotum and two on the mesonotum, but sometimes those on the mesonotum or on both are absent. The mesonotum is inflated and bulges dorsally above the pronotum in lateral view. Metanotal groove absent or narrower than the diameter of the metanotal spiracles. Metathoracic spiracles in dorsolateral position, not protruding, and touching the propodeal suture. Dorsum of the propodeum convex and shorter than the propodeal slope. Propodeal spiracles circular, positioned on the posterior propodeal margin, slightly posterior of the middle of the propodeal slope. Legs with appressed hairs. Petiole short and inclined forward.

Gaster. With dense pubescence and scattered long erect hairs at the edges of the segments. Some specimens of B. heeri resemble the regular worker in head and mesosoma, but they have a strongly expanded gaster (physogastry).

Color and sculpture. Head and gaster smooth, dorsum of the mesosoma with imbricate sculpture, body opaque and yellowish, sometimes with a somewhat darker gaster.

Type Material
Ortiz-Sepulveda et al. (2019) - Lectotype worker (: USNMENT00757169) and paralectotype workers, males, queen (MHNG: USNMENT00757167–00757171, USNMENT00758116– 00758120); here designated): 15 workers, three males, one queen [examined]. SWITZERLAND: Zurich: Serra des orchidiées.

The lectotype is designated here as the ant in the middle of their holder MHNG: USNMENT00757169, whereas the other specimens are paralectotypes.

The types (Forel 1874) were found in the "Jardin botanique de Zurich, dans la serre des Orchidees tropicales - the Botanical Garden of Zurich, in the greenhouse of tropical orchids." The ants were apparently brought to Europe in tropical plant material, which led to the odd circumstance of a Neotropical ant species having Switzerland as its type locality.

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