Nomamyrmex esenbeckii

Identification
Watkins (1977) - N. esenbecki. Workers: postoccipital sulcus distinct; dorsum of petiole with longitudinal rugae. Males: median border of head behind ocellar peduncle with a narrow lamella; first gastric tergite usually with longitudinal rugae; width of, blade of stipes about one-half its length; volsella blunt, gradually tapered or snout-shaped.

N. esenbecki s. str. Workers: transversely keeled posterior margin of mesonotum usually straight or slightly convex in the middle; anterior corners of postpetiole prominent and abruptly rounded causing the postpetiole to appear subquadrate with an anterior face as broad or broader than the petiole; entire body usually dark reddish brown with gaster sometimes slightly lighter. The mesonotal keel is occasionally slightly concave, but usually not in all specimens in a series. The posterior edge of the propodeum is often only slightly indented between the propodeal “teeth,” but a few specimens may be deeply indented (dorsal view). Males: setae on fifth gastric tergite broadly separated along the midline (i.e., the separation is about one half the width of the tergite near its base; distal portion of volsella distinctly snout-shaped, but the “snout” varies somewhat in detail.

N. esenbecki crassicornis. Workers: transversely keeled posterior margin of mesonotum usually straight or slightly convex in the middle; anterior corners of the postpetiole not prominent with the sides gradually curving inward to the anterior face which is slightly narrower than the petiole causing the postpetiole to appear trapezoidal; gaster usually yellowish brown, remainder of body reddish brown. A few workers from Panama have slightly concave mesonotal keels; however, these workers have postpetioles and yellowish brown gasters typical for crassicornis. Males: setae on fifth gastric tergite not distinctly separated along the midline; longitudinal rugae distinct on the second gastric tergite; apex of volsella slightly enlarged and rounded. The entire posterior half of the fifth tergite is usually covered with long thick setae; however, some specimens from Panama have the setae along the midline more confined to the posterior border and these setae are shorter than the more lateral setae. The distal portion of the volsella is slightly upturned with a gradually rounded apex in specimens from Panama, almost straight with a bluntly rounded apex in specimens from Trinidad, and somewhat more downturned in specimens from Peru.

Distribution based on Regional Taxon Lists
Nearctic Region: United States. Neotropical Region: Argentina, Bolivia, Brazil, Colombia, Costa Rica, Ecuador, French Guiana, Guatemala, Guyana, Honduras, Mexico , Panama, Paraguay, Peru, Suriname, Trinidad and Tobago, Venezuela.

Biology
Jack Longino: Nomamyrmex esenbeckii forages in columns during the day; I do not know if they forage at night. The workers are robust and their columns very conspicuous. In Costa Rica this species is uncommon relative to several species of Eciton and Labidus.

Nomamyrmex esenbeckii appears to be a specialist raider of Atta colonies (Swartz 1998 and included references). Swartz reviewed the earlier literature, in which workers reported Nomamyrmex raids on Atta colonies in Mexico and various sites in Brazil. She also observed and carefully documented a raid on an Atta cephalotes colony in Corcovado National Park, Costa Rica.

During work in Corcovado National Park in the early 1980's, I saw this species twice with prey and three times in the process of raiding ant colonies. Four of these observations involved attines. I observed Atta brood as prey in one column, and Acromyrmex brood as prey in another. I observed two attacks on mature Atta cephalotes colonies. Both these raids were observed mid-morning. In the first instance, Nomamyrmex were entering through a collapsed Atta chamber in the middle of a trail, taking away brood. I saw very little response on the Atta's part. There were two dead soldiers near the raiding column. In the second case, Atta workers were abundant on the surface of their mound, and had brought piles of brood to the surface of the largest, most active mounds. Nomamyrmex columns were carrying adult Atta soldiers and brood down into Atta nest entrances, suggesting a largely subterranean raid. I observed one raid on a Pheidole nest.

Jack Longino: I have once observed this species with prey. A column was entering and issuing from fissures in a trail, carrying abundant larvae and pupae of Atta, including pupae of fairly large workers.

Kjetil Aasen, a student working at La Selva Biological Station during the summer of 1997, made the following observation of a Nomamyrmex raid on Atta cephalotes:

On the evening on my last day at La Selva, I went to take a look at the Atta cephalotes colony between the bridge and the library and I was surprised to find a fierce battle in progress. It was dark and it was raining and thousands of Atta soldiers and Nomamyrmex esenbeckii were fighting in the area surrounding the main entrances of the Atta nest. They were all emerging from two closely spaced nest entrances and most of the Nomamyrmex were in the mandibles of some large leafcutter. In most cases, the Nomamyrmex were caught by their antenna, but they still managed to sting Atta, and they seemed to be very good at this because there were many dead or dying leafcutters on the battlefield. I did not see many dead Nomamyrmex.

O'Donnell et al. (2020) report this species foraging both day and night.

Brown & Fenner (1998) report this species conducting raids on the nests of Camponotus cingulatus, Camponotus crassus and Camponotus rufipes at Itatiaia, Brazil.

Castes
Primarily wingless queen described by Borgmeier 1958

Nomenclature

 * . Labidus esenbeckii Westwood, 1842: 75, pl. 20, fig. 4 (m.) BRAZIL (Goiás).
 * Type-material: holotype male.
 * Type-locality: Brazil: Rio Vendinha, 10.ix.1828, in the evening (W. Burchell).
 * Type-depository: OXUM.
 * Combination in Eciton (Labidus): Emery, 1890b: 39; Santschi, 1920d: 364; Creighton, 1950a: 64;
 * combination in E. (Holopone): Santschi, 1925b: 11;
 * combination in E. (Nomamyrmex): Borgmeier, 1936: 56;
 * combination in Nomamyrmex: Borgmeier, 1953: 14.
 * Status as species: Smith, F. 1859b: 7; Roger, 1863b: 41; Mayr, 1863: 424; Emery, 1890b: 39; Dalla Torre, 1893: 2; Emery, 1894k: 46; Forel, 1895b: 119; Forel, 1899c: 25; Emery, 1900a: 177 (in key); Forel, 1901h: 47; Wheeler, W.M. 1908e: 409 (redescription); Emery, 1910b: 23; Wheeler, W.M. 1910g: 562; Forel, 1911c: 288; Mann, 1916: 422; Santschi, 1920d: 364; Borgmeier, 1923: 43; Borgmeier, 1936: 56; Creighton, 1950a: 64; Smith, M.R. 1951a: 779; Borgmeier, 1953: 14; Borgmeier, 1955: 139 (redescription); Borgmeier, 1958: 201; Kempf, 1970b: 322; Kempf, 1972a: 164; Kempf & Lenko, 1976: 46; Watkins, 1976: 7 (in key); Watkins, 1977a: 204; Watkins, 1982: 215 (in key); Bolton, 1995b: 292; Palacio, 1999: 161 (in key); Wild, 2007b: 26, 43; Branstetter & Sáenz, 2012: 254; Guénard & Economo, 2015: 226; Shattuck & Cover, 2016: 19; Palacio, 2019: 626.
 * Senior synonym of afflictus: Borgmeier, 1955: 139; Kempf, 1972a: 164; Bolton, 1995b: 292.
 * Senior synonym of crassicornis: Wild, 2007b: 43.
 * Senior synonym of crudele: Borgmeier, 1955: 139; Kempf, 1972a: 164; Bolton, 1995b: 292.
 * Senior synonym of fimbriatum: Borgmeier, 1953: 14; Borgmeier, 1955: 139; Kempf, 1972a: 164; Bolton, 1995b: 292.
 * Senior synonym of hirsutipodex: Wild, 2007b: 43 (by implication).
 * Senior synonym of indomabile: Borgmeier, 1955: 139; Kempf, 1972a: 164; Bolton, 1995b: 292.
 * Senior synonym of interrupta: Borgmeier, 1953: 14; Borgmeier, 1955: 139; Kempf, 1972a: 164; Bolton, 1995b: 292.
 * Senior synonym of intrepidum: Wild, 2007b: 43 (by implication).
 * Senior synonym of mordax: Shattuck & Cover, 2016: 19.
 * Senior synonym of terrificum: Borgmeier, 1953: 14; Borgmeier, 1955: 139; Kempf, 1972a: 164; Bolton, 1995b: 292.
 * Senior synonym of wilsoni: Smith, M.R., 1951a: 779; Shattuck & Cover, 2016: 19.
 * Distribution: Argentina, Belize, Bolivia, Brazil, Colombia, Costa Rica, French Guiana, Guatemala, Guyana, Honduras, Mexico, Panama, Paraguay, Peru, Suriname, Trinidad, U.S.A., Venezuela.
 * afflictus. Acanthostichus afflictus Gallardo, 1919a: 239, fig. 3 (m.) ARGENTINA (Formosa).
 * Type-material: 2 syntype males.
 * Type-locality: Argentina: Formosa, Rio Pilcomayo, vii.1907, no. 7653 (no collector’s name).
 * Type-depository: MACN.
 * Combination in Eciton: Emery, 1920a: 60.
 * Junior synonym of esenbeckii: Borgmeier, 1955: 139; Kempf, 1972a: 164; Bolton, 1995b: 292.
 * crassicornis. Eciton crassicornis Smith, F. 1855c: 163, pl. 13, figs. 1, 2 (w.) BRAZIL (Amazonas).
 * Type-material: syntype workers (number not stated).
 * Type-locality: Brazil: Villa Nova (H.W. Bates).
 * Type-depository: BMNH.
 * Borgmeier, 1958: 202 (q.).
 * Combination in E. (Labidus): Emery, 1910b: 23;
 * combination in E. (Holopone): Santschi, 1925b: 12;
 * combination in E. (Nomamyrmex): Borgmeier, 1936: 55;
 * combination in Nomamyrmex: Borgmeier, 1953: 16.
 * Status as species: Smith, F. 1858b: 151; Mayr, 1863: 409; Roger, 1863b: 36; Mayr, 1865: 77 (in key); Norton, 1868b: 45; Mayr, 1886b: 115 (in key); Emery, 1890a: 55; Emery, 1890b: 38; Dalla Torre, 1893: 2; Emery, 1894c: 179; Emery, 1894k: 45; Forel, 1895b: 141; Emery, 1896h: 625; Forel, 1899c: 24; Emery, 1900a: 185; Forel, 1901c: 124; Forel, 1906d: 249; Forel, 1908b: 40; Forel, 1908c: 346; Forel, 1908e: 64; Forel, 1909a: 254; Emery, 1910b: 23; Mann, 1916: 422; Wheeler, W.M. 1916d: 324; Luederwaldt, 1918: 37; Santschi, 1920d: 367; Wheeler, W.M. 1922c: 1; Borgmeier, 1923: 42; Wheeler, W.M. 1925a: 2; Santschi, 1929a: 414; Borgmeier, 1936: 55; Borgmeier, 1948b: 460; Creighton, 1950a: 63; Smith, M.R. 1951a: 779; Borgmeier, 1953: 16.
 * Subspecies of esenbeckii: Borgmeier, 1955: 149 (redescription); Borgmeier, 1958: 202; Kempf, 1972a: 164; Watkins, 1977a: 205; Bolton, 1995b: 292.
 * Junior synonym of esenbeckii: Wild, 2007b: 43.
 * Material of the unavailable name furiosum referred here by Borgmeier, 1955: 139; Kempf, 1972a: 164.
 * crudele. Eciton (Holopone) crassicorne subsp. crudele Santschi, 1929a: 414, figs. c, g (w.) PARAGUAY, BOLIVIA.
 * Type-material: holotype worker.
 * Type-locality: Paraguay: (no further data) (Fiebrig).
 * [Note: Santschi also includes non-type material from Bolivia: Caranditi (Lizer & Deletang).]
 * Type-depository: NHMB.
 * Subspecies of crassicorne: Santschi, 1929f: 85.
 * Junior synonym of esenbeckii: Borgmeier, 1955: 139; Kempf, 1972a: 164; Bolton, 1995b: 292.
 * fimbriatum. Eciton (Labidus) fimbriatum Santschi, 1920d: 363, figs. 1, 2 (m.) FRENCH GUIANA.
 * Type-material: 8 paratype males.
 * Type-locality: French Guiana: St Jean du Maroni (Le Moult).
 * Type-depositories: MNHN, NHMB.
 * Status as species: Eidmann, 1936a: 29.
 * Junior synonym of esenbeckii: Borgmeier, 1953: 14; Borgmeier, 1955: 139; Kempf, 1972a: 164; Bolton, 1995b: 292.
 * hirsutipodex. Eciton (Labidus) esenbecki st. hirsutipodex Santschi, 1920d: 364, fig. 2a (m.) MEXICO (Yucatan).
 * Type-material: holotype male.
 * Type-locality: Mexico: Merida (no collector’s name).
 * Type-depository: NHMB.
 * Junior synonym of crassicornis: Borgmeier, 1955: 139; Kempf, 1972a: 164; Bolton, 1995b: 292.
 * Junior synonym of esenbeckii: Wild, 2007b: 43 (by implication).
 * indomabile. Eciton (Holopone) crassicorne var. indomabile Santschi, 1930e: 82 (w.) BRAZIL (Pernambuco).
 * Type-material: 19 syntype workers.
 * Type-locality: Brazil: Pernambuco, Olinda (Reichensperger).
 * Type-depository: NHMB.
 * Subspecies of crassicornis: Santschi, 1931c: 270.
 * Junior synonym of esenbeckii: Borgmeier, 1955: 139; Kempf, 1972a: 164; Bolton, 1995b: 292.
 * interrupta. Eciton fimbriatum var. interrupta Santschi, 1923c: 249 (m.) BRAZIL (Minas Gerais).
 * Type-material: holotype male.
 * Type-locality: Brazil: Minas Gerais, Piracicabo (Luja).
 * Type-depository: NHMB.
 * Junior synonym of esenbeckii: Borgmeier, 1953: 14; Borgmeier, 1955: 139; Kempf, 1972a: 164; Bolton, 1995b: 292.
 * intrepidum. Eciton (Holopone) crassicorne var. intrepidum Santschi, 1929a: 414, figs. b, f (w.) VENEZUELA.
 * Type-material: syntype workers (number not stated).
 * Type-locality: Venezuela: between Carupano and Cariaco, 1896 (no collector’s name).
 * Type-depository: unknown.
 * [Note: Baroni Urbani, 1977e: 68-71, does not record the presence of type-material of this taxon in NHMB.]
 * Junior synonym of crassicornis: Borgmeier, 1955: 139; Kempf, 1972a: 164; Bolton, 1995b: 292.
 * Junior synonym of esenbeckii: Wild, 2007b: 43 (by implication).
 * mordax. Eciton (Holopone) crassicorne st. mordax Santschi, 1929a: 415, figs. d, h (w.) MEXICO (Morelos).
 * Type-material: syntype workers (number not stated).
 * Type-locality: Mexico: Cuernavaca (A. Forel).
 * Type-depository: NHMB.
 * Watkins, 1977a: 210 (m.).
 * Combination in Nomamyrmex: Borgmeier, 1953: 13.
 * Subspecies of crassicorne: Menozzi, 1935b: 189.
 * Junior synonym of wilsoni: Borgmeier, 1955: 139; Smith, M.R. 1958c: 108; Kempf, 1972a: 165; Smith, D.R. 1979: 1329.
 * Subspecies of esenbeckii: Watkins, 1977a: 205; Brandão, 1991: 361; Bolton, 1995b: 292.
 * Junior synonym of esenbeckii: Shattuck & Cover, 2016: 19.
 * terrificum. Eciton (Holopone) fimbriatum var. terrificum Santschi, 1930e: 82 (m.) BRAZIL (no state data).
 * Type-material: holotype male.
 * Type-locality: Brazil: (no further data), 23.ix.1929 (Reichensperger).
 * Type-depository: NHMB.
 * Junior synonym of esenbeckii: Borgmeier, 1953: 14; Borgmeier, 1955: 139; Kempf, 1972a: 164; Bolton, 1995b: 292.
 * wilsoni. Eciton (Labidus) esenbecki st. wilsoni Santschi, 1920d: 366 (m.) U.S.A. (Texas).
 * Type-material: holotype male.
 * Type-locality: U.S.A.: Texas, Las Borragas, nr Brownsville, vi. (C. Schaeffer).
 * Type-depository: BMNY.
 * [Note: holotype may now be in AMNH, or if retained by Wheeler, in MCZC.]
 * Combination in E. (Nomamyrmex): Smith, M.R. 1951a: 779;
 * combination in Nomamyrmex: Borgmeier, 1953: 13.
 * Subspecies of esenbeckii: Borgmeier, 1955: 147; Smith, M.R. 1958c: 108; Kempf, 1972a: 165; Watkins, 1977a: 205; Smith, D.R. 1979: 1329; Watkins, 1985: 481; Bolton, 1995b: 292.
 * Junior synonym of esenbeckii: Smith, M.R. 1951a: 779; Shattuck & Cover, 2016: 19.

References based on Global Ant Biodiversity Informatics

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