Holcoponera striatula

A common, widespread species that exhibits wide variation in some characters. It commonly nests in twigs within leaf litter (Fernandes et al., 2018).

Identification
A member of the striatula subgroup (in the striatula species group). This widespread, common, and fairly variable ant has been the object of a lengthy synonymic list. Most of the names were based upon differences in size, gauge of costulation, and the shape of the subpetiolar process and the mesopisternal lobe. These size differences are all normal for the species. One can recognize coarsely costulate and finely costulate specimens, as well as intermediate forms. The finely costulate ants can be found in several places: S. Brazil, Argentina, Guiana, Costa Rica, Jamaica and Hispaniola. Occasional series can be found that have transverse costulae on part or all of the propodeal declivity; this condition is not frequent on the finely costulate forms. Several series from Bolivia have a long anterior slope of the node, which approaches the condition in pleurodon. The use of the subpetiolar process in the separation of species is of limited value as a variation from subquadrate to anteriorly projecting lobe can be found in series from the same population or even nest. Brown (1957:489) already discussed the futility of using the mesepisternal lobe. (Lattke 2004)

Distribution based on Regional Taxon Lists
Neotropical Region: Argentina, Barbados, Bolivia, Brazil, Colombia, Costa Rica, Dominican Republic, Ecuador, French Guiana , Greater Antilles, Guadeloupe, Guatemala, Guyana, Lesser Antilles, Martinique, Mexico, Panama, Paraguay, Puerto Rico, Suriname, Trinidad and Tobago, Uruguay, Venezuela.

Biology
Rosumek (2017) examined trophic and temporal niches of ants in a secondary Atlantic forest (Florianópolis, south Brazil; ), between December 2015 and January 2016. For this species: Gnamptogenys striatula (possibly a species complex) is a rare example of a Neotropical ant extensively studied in the laboratory, covering many aspects of its biology (e.g. Giraud et al., 2000; Kaptein et al., 2005). However, the only information available about its trophic niche in the wild comes from Lattke (1990), who reports remnants of several insect orders inside nests, and posteriorly called it “a generalist epigeic forager of humid forests” (Lattke, 1995). A recent account recorded it rarely on experimental vertebrate carcasses left to rot in a forest, predating the larvae and pupae of necrophagous insects (Paula et al., 2016).

At baits in this study, G. striatula  was observed using termites frequently, but crushed insects, feces and sucrose were important as well. Hence, the species will scavenge and consume sugar when given the opportunity, although they were not attracted to melezitose (an oligossacharide analog used as a proxy for honeydew). The relatively high use of feces, a less preferred resource overall, is a noteworthy feature that differentiated G. striatula from most other species of this study. It showed a moderate but significant preference for nocturnal foraging.

Lattke (1995) - A mite was found on the anterolateral petiolar side of one specimen from Beni, Bolivia.

De la Mora et al. (2016) found this species nesting in logs found in coffee farms and forests in the Soconusco region of Chiapas, Mexico.

Koch et al. (2018) sampled this species in Caryocar barsiliense trees, in southeastern Brazil cerrado, as part of a study examining species interactions in ant-plants.

Worker
Several gamergates reproduce in most colonies, while one or several dealate queens reproduce in others. Cuticular Hydrocarbons differ qualitatively and quantitatively between egg-laying and infertile workers, thus providing reliable information about current fertility (Lommelen et al. 2006).

Nomenclature

 * . Gnamptogenys striatula Mayr, in Radoszkowsky, 1884: 32 (w.) FRENCH GUIANA.
 * Type-material: syntype workers (number not stated).
 * Type-locality: French Guiana (“Cayenne française”): (no further data) (Jelski).
 * Type-depositories: BMNH, NHMW.
 * Mayr, 1887: 542 (q.m.); Wheeler, G.C. & Wheeler, J. 1952a: 123 (l.).
 * Combination in Ectatomma (Holcoponera): Mayr, 1887: 541;
 * combination in Gnamptogenys: Brown, 1958g: 229;
 * combination in Holcoponera: Emery, 1911d: 40; Camacho, Franco, Branstetter, et al. 2022: 11.
 * Status as species: Mayr, 1887: 542; Dalla Torre, 1893: 26; Forel, 1895b: 113; Emery, 1896g: 44 (in key); Forel, 1901h: 81; Forel, 1909a: 253; Emery, 1911d: 40; Bruch, 1915: 528; Mann, 1916: 404; Wheeler, W.M. 1917g: 457; Gallardo, 1918b: 25 (redescription); Wheeler, W.M. 1922c: 3; Borgmeier, 1923: 56; Wheeler, W.M. 1925a: 4; Santschi, 1929h: 442 (redescription); Brown, 1956b: 490; Brown, 1958g: 229, 327; Kempf, 1970b: 325; Kempf & Lenko, 1976: 52; Zolessi, et al. 1988: 2; Lattke, 1990b: 20; Brandão, 1991: 345; Bolton, 1995b: 211; Lattke, 1995: 186; Lattke, et al. 2004: 348; Wild, 2007b: 27; Lattke, et al. 2007: 258 (in key); Lattke, et al. 2008: 98; Branstetter & Sáenz, 2012: 255; Bezděčková, et al. 2015: 111; Feitosa, 2015c: 98; Wetterer, et al. 2016: 11; Feitosa & Prada-Achiardi, 2019: 673; Lubertazzi, 2019: 119; Camacho, et al. 2020: 454 (in key); Camacho, Franco, Branstetter, et al. 2022: 11.
 * Senior synonym of angustiloba: Brown, 1958g: 229; Kempf, 1972a: 115; Bolton, 1995b: 211; Lattke, 1995: 186; Camacho, Franco, Branstetter, et al. 2022: 11.
 * Senior synonym of angustipleura: Brown, 1958g: 229; Kempf, 1972a: 115; Bolton, 1995b: 211; Lattke, 1995: 186; Camacho, Franco, Branstetter, et al. 2022: 11.
 * Senior synonym of antillana: Brown, 1956b: 490; Brown, 1958g: 300; Kempf, 1972a: 115; Bolton, 1995b: 211; Lattke, 1995: 186; Camacho, Franco, Branstetter, et al. 2022: 11.
 * Senior synonym of arcuata: Lattke, 1995: 186; Camacho, Franco, Branstetter, et al. 2022: 11.
 * Senior synonym of brasiliensis: Brown, 1958g: 229, 327; Kempf, 1972a: 115; Bolton, 1995b: 211; Lattke, 1995: 186; Camacho, Franco, Branstetter, et al. 2022: 11.
 * Senior synonym of calcarata: Brown, 1958g: 229; Kempf, 1972a: 115; Bolton, 1995b: 211; Lattke, 1995: 186; Camacho, Franco, Branstetter, et al. 2022: 11.
 * Senior synonym of curtulum: Lattke, 1995: 186; Camacho, Franco, Branstetter, et al. 2022: 11.
 * Senior synonym of isthmica: Lattke, 1995: 186; Camacho, Franco, Branstetter, et al. 2022: 11.
 * Senior synonym of mayri: Lattke, 1995: 186; Camacho, Franco, Branstetter, et al. 2022: 11.
 * Senior synonym of obscurum: Santschi, 1929h: 442; Kempf, 1972a: 115; Bolton, 1995b: 211; Lattke, 1995: 186; Camacho, Franco, Branstetter, et al. 2022: 11.
 * Senior synonym of paulina: Brown, 1958g: 229; Kempf, 1972a: 115; Bolton, 1995b: 211; Lattke, 1995: 186; Camacho, Franco, Branstetter, et al. 2022: 11.
 * Senior synonym of recta: Lattke, 1995: 186; Camacho, Franco, Branstetter, et al. 2022: 11.
 * Senior synonym of rustica: Lattke, 1995: 186; Camacho, Franco, Branstetter, et al. 2022: 11.
 * Senior synonym of simplicoides: Lattke, 1995: 186; Camacho, Franco, Branstetter, et al. 2022: 11.
 * Senior synonym of stolli: Lattke, 1995: 186; Camacho, Franco, Branstetter, et al. 2022: 11.
 * Senior synonym of vollenweideri: Brown, 1958g: 229; Kempf, 1972a: 115; Bolton, 1995b: 211; Lattke, 1995: 186; Camacho, Franco, Branstetter, et al. 2022: 11.
 * Senior synonym of wasmanni Lattke, 1995: 186; Camacho, Franco, Branstetter, et al. 2022: 11.
 * Senior synonym of wheeleri: Lattke, 1995: 186; Camacho, Franco, Branstetter, et al. 2022: 11.
 * Material of the unavailable name hybrida referred here by Brown, 1958g: 229; Kempf, 1972a: 115; Bolton, 1995b: 211; Lattke, 1995: 186.
 * Distribution: Argentina, Bolivia, Brazil, Colombia, Costa Rica, Dominican Republic, Ecuador, El Salvador, French Guiana, Guatemala, Guyana, Honduras, Jamaica, Lesser Antilles, Martinique, Mexico, Nicaragua, Panama, Paraguay, Peru, Puerto Rico, Suriname, Trinidad, Uruguay, Venezuela.
 * angustiloba. Holcoponera striatula var. angustiloba Emery, 1911d: 41.
 * Type-material: syntype workers (number not stated).
 * Type-locality: Brazil: São Paulo (von Ihering).
 * Type-depositories: MHNG, NHMB.
 * [First available use of Ectatomma (Holcoponera) striatulum subsp. obscurum var. angustiloba Forel, 1908c: 341 (w.) BRAZIL (São Paulo); unavailable (infrasubspecific) name.]
 * Forel, 1911e: 254 (q.); Gallardo, 1918b: 23 (m.).
 * As unavailable (infrasubspecific) name: Forel, 1911e: 254.
 * Subspecies of striatula: Emery, 1911d: 41; Luederwaldt, 1918: 34; Santschi, 1919f: 37; Borgmeier, 1923: 56.
 * Subspecies of brasiliensis Santschi, 1929d: 275; Santschi, 1929h: 453 (redescription).
 * Junior synonym of striatula: Brown, 1958g: 229; Kempf, 1972a: 115; Bolton, 1995b: 208; Lattke, 1995: 186; Camacho, Franco, Branstetter, et al. 2022: 11.
 * angustipleura. Holcoponera striatula var. angustipleura Emery, 1911d: 41.
 * Type-material: syntype workers (number not stated).
 * Type-locality: Brazil: Pará (Goeldi).
 * Type-depositories: MHNG, NHMB.
 * [First available use of Ectatomma (Holcoponera) striatulum subsp. obscurum var. angustipleura Forel, 1908c: 342 (w.) BRAZIL (Pará); unavailable (infrasubspecific) name.]
 * Subspecies of striatula: Emery, 1911d: 41; Borgmeier, 1923: 56; Eidmann, 1936a: 33.
 * Status as species: Santschi, 1929h: 456 (redescription).
 * Junior synonym of striatula: Brown, 1958g: 229; Kempf, 1972a: 115; Bolton, 1995b: 208; Lattke, 1995: 186; Camacho, Franco, Branstetter, et al. 2022: 11.
 * antillana. Holcoponera striatula var. antillana Santschi, 1929h: 444, figs. 16, 21 (w.m.) MARTINIQUE, JAMAICA.
 * Type-material: syntype workers, syntype males (numbers not stated).
 * Type-localities: Martinique (no further data), Jamaica: Kingstown (Forel), Jamaica: Port Antonio (Wright).
 * Type-depositories: MCZC, NHMB (possibly also in IEUB).
 * Junior synonym of striatula: Brown, 1956b: 490; Brown, 1958g: 300; Kempf, 1972a: 115; Bolton, 1995b: 208; Lattke, 1995: 186; Camacho, Franco, Branstetter, et al. 2022: 11.
 * arcuata. Holcoponera regularis var. arcuata Santschi, 1929h: 457 (w.) BRAZIL (Rio de Janeiro, Minas Gerais).
 * Type-material: holotype worker.
 * Type-locality: Brazil: Rio de Janeiro, Passo Quatro (Zikan).
 * Type-depository: NHMB.
 * Replacement name for Holcoponera regularis Santschi, 1929d: 275 [Junior secondary homonym of Gnamptogenys regularis Mayr, 1870b: 965.]
 * [Note: arcuata junior synonym of regularis Santschi, 1929d: 275 (synonymy by Brown, 1958g: 227); hence arcuata first available replacement name (Bolton, 1995b: 208).]
 * Combination in Gnamptogenys: Brown, 1958g: 227.
 * Status as species: Brown, 1958g: 227; Kempf, 1972a: 112; Bolton, 1995b: 208.
 * Junior synonym of striatula: Lattke, 1995: 186; Camacho, Franco, Branstetter, et al. 2022: 11.
 * brasiliensis. Holcoponera brasiliensis Emery, 1902a: 181 (w.q.m.) BRAZIL (Santa Catarina, Rio de Janeiro).
 * Type-material: syntype workers, syntype queens, syntype males (numbers not stated).
 * Type-localities: Brazil: Santa Catarina (no collector’s name), Brazil: Rio de Janeiro (no collector’s name).
 * Type-depository: MSNG.
 * Wheeler, G.C. & Wheeler, J. 1952a: 124 (l.).
 * Status as species: Emery, 1906c: 113; Emery, 1911d: 40; Bruch, 1914: 213; Gallardo, 1918b: 23 (redescription); Wheeler, W.M. 1922c: 3; Borgmeier, 1923: 55; Santschi, 1929h: 451 (redescription); Kusnezov, 1953b: 336.
 * Junior synonym of striatula: Brown, 1958g: 229, 327; Kempf, 1972a: 115; Bolton, 1995b: 208; Lattke, 1995: 186; Camacho, Franco, Branstetter, et al. 2022: 11.
 * calcarata. Holcoponera brasiliensis var. calcarata Santschi, 1929h: 452, fig. 28 (w.) BRAZIL (São Paulo).
 * Type-material: 2 syntype workers.
 * Type-locality: Brazil: São Paulo, Iguape (B.S. Pessoa).
 * Type-depository: NHMB.
 * Subspecies of brasiliensis: Santschi, 1931e: 273; Santschi, 1933e: 106.
 * Junior synonym of striatula: Brown, 1958g: 229; Kempf, 1972a: 115; Bolton, 1995b: 208; Lattke, 1995: 186; Camacho, Franco, Branstetter, et al. 2022: 11.
 * curtulum. Ectatomma (Holcoponera) curtulum Emery, 1896g: 47, pl. 1, fig. 8 (w.) COSTA RICA.
 * Type-material: syntype workers (number not stated).
 * Type-locality: Costa Rica: Alajuela (A. Alfaro).
 * Type-depositories: MSNG, NHMB.
 * Forel, 1899c: 7 (q.).
 * Combination in Holcoponera: Emery, 1911d: 40;
 * combination in Gnamptogenys: Brown, 1958g: 228.
 * Status as species: Forel, 1899c: 7; Emery, 1911d: 40; Bruch, 1914: 213; Santschi, 1929h: 469 (redescription); Wheeler, W.M. 1933a: 62; Brown, 1958g: 228, 320; Kempf, 1972a: 112; Bolton, 1995b: 209.
 * Junior synonym of striatula: Lattke, 1995: 186; Camacho, Franco, Branstetter, et al. 2022: 11.
 * isthmica. Holcoponera wasmanni var. isthmica Santschi, 1929h: 467 (w.) PANAMA.
 * Type-material: 3 syntype workers.
 * Type-locality: Panama: Tabogila, 19.xi.1911 (W.M. Wheeler).
 * Type-depository: NHMB.
 * Junior synonym of wasmanni: Brown, 1958g: 230; Kempf, 1972a: 116; Bolton, 1995b: 209.
 * Junior synonym of striatula: Lattke, 1995: 186; Camacho, Franco, Branstetter, et al. 2022: 11.
 * mayri. Holcoponera brasiliensis var. mayri Santschi, 1929h: 453 (w.) COSTA RICA.
 * Type-material: 2 syntype workers.
 * Type-locality: Costa Rica: (no further data).
 * Type-depositories: NHMB, NHMW.
 * Junior synonym of wheeleri: Brown, 1958g: 230; Kempf, 1972a: 116; Bolton, 1995b: 209.
 * Junior synonym of striatula: Lattke, 1995: 186; Camacho, Franco, Branstetter, et al. 2022: 11.
 * obscurum. Ectatomma (Holcoponera) obscurum Emery, 1896g: 48 (w.) BRAZIL (Pará).
 * Type-material: holotype worker.
 * Type-locality: Brazil: Pará (no collector’s name).
 * Type-depository: MSNG.
 * Combination in Holcoponera: Emery, 1911d: 41.
 * Subspecies of striatula: Forel, 1908c: 341; Forel, 1911e: 254; Emery, 1911d: 41; Luederwaldt, 1918: 34; Borgmeier, 1923: 56; Donisthorpe, 1927b: 385.
 * Junior synonym of striatula: Santschi, 1929h: 442; Kempf, 1972a: 115; Bolton, 1995b: 210; Lattke, 1995: 186; Camacho, Franco, Branstetter, et al. 2022: 11.
 * paulina. Ectatomma (Holcomyrmex) curtulum var. paulina Forel, 1908c: 342 (w.) BRAZIL (São Paulo).
 * [Note: Holcomyrmex here is a misspelling of Holcoponera.]
 * Type-material: syntype workers (number not stated).
 * Type-locality: Brazil: São Paulo (von Ihering).
 * Type-depository: MHNG.
 * Combination in Holcoponera: Emery, 1911d: 40.
 * Subspecies of curtula: Emery, 1911d: 40; Forel, 1912c: 33; Borgmeier, 1923: 55.
 * Subspecies of brasiliensis: Santschi, 1929h: 456 (redescription).
 * Junior synonym of striatula: Brown, 1958g: 229; Kempf, 1972a: 115; Bolton, 1995b: 210; Lattke, 1995: 186; Camacho, Franco, Branstetter, et al. 2022: 11.
 * recta. Holcoponera emeryi var. recta Santschi, 1929h: 465 (w.) BOLIVIA.
 * Type-material: 3 syntype workers.
 * Type-locality: Bolivia: Tumufasa (= Tumupasa) (W.M. Mann).
 * Type-depositories: MSNG, NHMB.
 * [Note: specimens from the original series that are present in MCZC and USNM (Brown, 1958g: 320) were not included among the 3 specimens on which the taxon was based. Santschi, 1929h: 465, records that the 3 workers came to him from MSNG, and Baroni Urbani, 1973b: 127, notes that only 1 syntype is retained in NHMB.]
 * Junior synonym of pleurodon: Brown, 1958g: 229, 320; Kempf, 1972a: 114; Bolton, 1995b: 210.
 * Junior synonym of striatula: Lattke, 1995: 186; Camacho, Franco, Branstetter, et al. 2022: 11.
 * regularis. Holcoponera regularis Santschi, 1929d: 275 (w.) BRAZIL (Minas Gerais, Rio de Janeiro).
 * Type-material: 7 syntype workers.
 * Type-localities: 4 workers Brazil: Rio de Janeiro, Passo Quatro (Zikan), 1 worker Brazil: Rio de Janeiro, Petropolis (no collector’s name), 2 workers Brazil: Paraná, Cascatinha (Borgmeier).
 * Type-depository: NHMB.
 * [Note: Baroni Urbani, 1973b: 128, records the 4 Passo Quatro specimens in NHMB; location of remainder unknown.]
 * [Also described as new by Santschi, 1929h: 457.]
 * [Junior secondary homonym of Gnamptogenys regularis Mayr, 1870b: 965.]
 * Replacement name: Holcoponera regularis var. arcuata Santschi, 1929h: 457.
 * [Note: arcuata junior synonym of regularis Santschi (synonymy by Brown, 1958g: 227, 327); hence arcuata first available replacement name.]
 * rustica. Holcoponera rustica Santschi, 1929h: 446, figs. 11, 18 (w.) PARAGUAY.
 * Type-material: holotype worker.
 * Type-locality: Paraguay: (no further data) (Forel).
 * Type-depository: NHMB.
 * Combination in Gnamptogenys: Brown, 1958g: 229.
 * Status as species: Brown, 1958g: 229; Kempf, 1972a: 114; Bolton, 1995b: 210.
 * Junior synonym of striatula: Lattke, 1995: 186; Camacho, Franco, Branstetter, et al. 2022: 11.
 * simplicoides. Holcoponera striatula var. simplicoides Emery, 1911d: 41.
 * Type-material: syntype workers (number not stated).
 * Type-locality: Brazil: São Paulo (von Ihering).
 * Type-depositories: MHNG, NHMB.
 * [First available use of Ectatomma (Holcoponera) striatulum subsp. obscurum var. simplicoides Forel, 1908c: 341 (w.) BRAZIL (São Paulo); unavailable (infrasubspecific) name.]
 * Santschi, 1929h: 455 (m.).
 * Combination in Gnamptogenys: Brown, 1958g: 229.
 * Subspecies of striatula: Emery, 1911d: 41; Forel, 1912c: 33; Luederwaldt, 1918: 34; Borgmeier, 1923: 56.
 * Subspecies of brasiliensis: Santschi, 1929h: 454 (redescription).
 * Status as species: Brown, 1958g: 229; Kempf, 1972a: 114; Bolton, 1995b: 210.
 * Junior synonym of striatula: Lattke, 1995: 186; Camacho, Franco, Branstetter, et al. 2022: 11.
 * stolli. Ectatomma (Holcoponera) curtulum var. stolli Forel, 1899c: 7 (w.m.) GUATEMALA.
 * Type-material: syntype workers, syntype males (numbers not stated).
 * Type-locality: Guatemala: Retalhuleu (Stoll).
 * Type-depositories: MHNG, NHMB.
 * Combination in Holcoponera: Emery, 1911d: 40.
 * Subspecies of curtulum: Emery, 1911d: 40; Santschi, 1929h: 470 (redescription).
 * Junior synonym of curtulum: Brown, 1958g: 228, 320; Kempf, 1972a: 112; Bolton, 1995b: 211.
 * Junior synonym of striatula: Lattke, 1995: 186; Camacho, Franco, Branstetter, et al. 2022: 11.
 * vollenweideri. Holcoponera curtula var. vollenweideri Forel, 1912c: 33 (w.) ARGENTINA (no province data).
 * Type-material: holotype (?) worker.
 * [Note: no indication of number of specimens is given.]
 * Type-locality: Argentina: (no further data) (Vollenweider).
 * Type-depository: MHNG.
 * Subspecies of curtulum: Santschi, 1916b: 510; Gallardo, 1918b: 24; Santschi, 1919f: 37.
 * Junior synonym of angustiloba: Santschi, 1929d: 275; Santschi, 1929h: 453.
 * Junior synonym of striatula: Brown, 1958g: 229; Kempf, 1972a: 115; Bolton, 1995b: 210; Lattke, 1995: 186; Camacho, Franco, Branstetter, et al. 2022: 11.
 * wasmanni. Holcoponera wasmanni Santschi, 1929h: 466, fig. 42 (w.) BRAZIL (Pará).
 * Type-material: holotype worker.
 * Type-locality: Brazil: Pará (Schmalz).
 * Type-depository: MSNG.
 * Combination in Gnamptogenys: Brown, 1958g: 230.
 * Status as species: Brown, 1958g: 230; Kempf, 1972a: 116; Bolton, 1995b: 211.
 * Junior synonym of striatula: Lattke, 1995: 186; Camacho, Franco, Branstetter, et al. 2022: 11.
 * wheeleri. Holcoponera rustica st. wheeleri Santschi, 1929h: 448, figs. 9, 19 (w.) COSTA RICA.
 * Type-material: 3 syntype workers.
 * [Note: Santschi described this taxon from 3 workers, which Baroni Urbani, 1973b: 128, recorded as present in NHMB. Therefore the specimen in MCZC, noted by Lattke, 1995: 149, may be from the same series but is not a syntype.]
 * Type-locality: Costa Rica: Porto Limon, 24.xi.1928 (W.M. Wheeler).
 * Type-depository: NHMB.
 * Wheeler, G.C. & Wheeler, J. 1952a: 124 (l.).
 * Combination in Gnamptogenys: Brown, 1958g: 230.
 * Status as species: Brown, 1958g: 230; Kempf, 1972a: 116; Bolton, 1995b: 211.
 * Junior synonym of striatula: Lattke, 1995: 186; Camacho, Franco, Branstetter, et al. 2022: 11.

Taxonomic Notes
Lattke (1995): Diagnosis - Pronotal costulae semicircular; dorsoposterior mesosomal costulae longitudinal and slightly diverging caudad; mesometanotum sometimes with semiciruclar costulae; triangular metacoxal teeth; body dark brown.

Specimens determined as G. curtula by Brown for the 1958 revision are G. striatula, and so are several specimens determined as H. wasmanni var. isthmica. Specimens labelled as type series of H. emeryi recta were studied in the USNM and found to be conspecific with striatula and not with G. pleurodon. I was able to examine two G. striatula syntypes in the BMNH. Bill Brown generously shared the notes he took during his study of the following Santschi types in 1963: H. regularis, H. regularis v. arcuata, H. rustica, and H. wheeleri. He concluded that they are all the same species. He also wrote that these specimens plus additional types and specimens he studied (H. striatula v. antillana, v. angustipleura, H. wasmanni, H. wasmanni v. isthmica, H. brasiliensis v. calcarata, v. mayri, H. curtula v. paulina (det. Santschi)) perhaps could be separated into two species using the subpetiolar process. But as already has been mentioned, this process has proven to be an unreliable character for species determination in this group.

References based on Global Ant Biodiversity Informatics

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