Pogonomyrmex mendozanus

A desert inhabitant that builds its nests in open areas with sandy soils.

Identification
Johnson (2015) - Worker Within the P. laticeps-group, the combination of: (1) head and mesosoma black, gaster dark ferruginous orange, and (2) head and mesosoma covered by fine, regular, incised rugae uniquely characterize this species.

Brachypterous Queen This caste is diagnosed by: (1) brachypterous with very small wings and small ocelli on head, (2) in dorsal view, mesoscutum poorly-developed, anterior margin barely surpassing humeral shoulders of pronotum, (3) pronotum well-developed, (4) in profile, the pronotum rises at an approximately 45° angle to meet the mesoscutum, (5) fine, regular, longitudinal rugae on head, mesoscutum, and mesoscutellum, and (6) head and mesosoma black; gaster dark ferruginous orange.

Male This caste is diagnosed by: (1) first gastral tergum lacking striae, (2) weakly regular to regular, subparallel rugae prominent on sides of head, pronotal sides, and mesopleura, (3) in profile, rugae forming circumocular whorls posterior to eyes, and (4) notauli present.

Pogonomyrmex mendozanus was misidentified as Pogonomyrmex pronotalis in a list of ant species in Ñancuñan Biosphere Reserve (Claver & Fowler, 1993) and in subsequent publications until it was described by Cuezzo & Claver (2009). However, these two species are easily separated with coloration being one of the few characters in common. In describing P. mendozanus, Cuezzo & Claver (2009) listed several characters to distinguish it from P. pronotalis. Sculpturing on the head and mesosoma is the best character: rugae on the cephalic dorsum of P. mendozanus are fine, very regular, and incised, whereas they are coarse and irregular on P. pronotalis. Lack of striae on the first gastral tergum combined with the fine, regular, incised sculpturing on the head and mesosoma and coloration pattern separate P. mendozanus from all South American congeners.

Distribution based on Regional Taxon Lists
Neotropical Region: Argentina.

Habitat
Pogonomyrmex mendozanus inhabits sites at elevations from 465–545 m. This species is common from southeastern San Juan to eastcentral Mendoza, and it appears to be restricted to the northern one-third of the Low Monte Desert ecoregion as defined by Olson et al. (2001). Interestingly, the distribution of P. mendozanus stops at the southern boundary of the High Monte Desert ecoregion. Field observations suggest that P. mendozanus is restricted to deep, loose, sandy soils (pers. obs.); nests are most common in open, disturbed areas such as roadsides (Pirk et al., 2004).

Biology
Cuezzo and Claver (2009) - We observed more than 20 nests of Pogonomyrmex mendozanus in natural areas. The nests were built in sandy soils, in bare or very sparsely vegetated areas. Nest mounds were 5 cm high and 12-15 cm in diameter and had one or more irregular entrances 1-2 cm in diameter. They were often located near the base of grasses. Excavated nests had a maximum depth of 45 cm and a diameter of 30 cm. Nest chambers with immature stages (pupae, larvae and eggs) were irregularly distributed throughout the nest, and the first chambers were found a few centimeters beneath soil surface. Seed chambers were found at depths of more than 30 cm, next to the nest chambers. The major foraging periods were during daytime from spring to fall. We observed an individual foraging pattern and no foraging trails. Despite high soil temperatures, ants were seen running away with the tip of the gaster curved forward below the mesosoma. In the Ñacuñán Biosphere Reserve, we observed that these ants mainly harvest caryopses from grasses (e.g. Trichloris crinita, Pappophorum caespitosum and Digitaria californica) and seeds of dicotyledoneous plants (e.g., Atriplex lampa and Verbena aspera) to a lesser degree. Workers were seen climbing up the stems of grasses to cut spikes and were also seen collecting diaspores from plant litter. Ants only accumulated seed parts or caryopses in their nest and discarded all the surrounding plant structures. Unlike other ant species of the genus, these ants leave no discarded plant material outside the nest. The workers were extremely aggressive when disturbed and their bite is painful.

Johnson (2015) - Pogonomyrmex mendozanus is one of the more well-studied species of Pogonomyrmex in Argentina. This species harvests the seeds of various grass species and nongrass species to a lesser extent (Pirk & Lopez de Casenave, 2006, 2010, 2011; Pirk et al., 2007; Pirk et al., 2009; Pol & Lopez de Casenave, 2004; Pol et al., 2011), and it is a solitary forager that recruits nestmates to high-density seed patches (Pol et al., 2015). Five colonies averaged 108–186 foragers, which consisted of an estimated 10–13% of all workers in the nests (Nobua-Behrmann et al., 2013). Nests have a tumulus that ranges up to 40 cm in diameter; a midden of seed chaff sometimes surrounds the nest. Colonies of P. mendozanus are relatively small: two excavated colonies in Reserva Ñancuñan contained an average of 615 workers plus 292 larvae and pupae (Nobua Behrmann et al., 2010). Colony size was similar in another study, but varied by grazing intensity: colonies in lightly grazed areas contained 731 ± 249 (n = 12; mean ± SD) individuals including brood (535 ±105 of which were workers), while those in heavily grazed areas contained 557 ± 325 (n = 12) individuals including brood (382 ± 230 of which were workers). All colonies contained one reproductive queen (n = 19) (R.G. Pol, pers. comm.). Worker dry mass averaged 3.87 mg, while that of brachypterous queens averaged 9.16 mg; the mean queen to worker dry mass ratio was 2.37 (n = 2 colonies) (R.A. Johnson, unpub. data).

Reproduction
Collection dates for sexuals range from 28 December to 15 March, and mating flights have been observed on 21 January, 14–15 February, and 15 March. Flights occur during early to late afternoon on days following rain. Mating involves the brachypterous queens leaving their natal nest to mate at aggregations in low-growing vegetation at the top of or near their natal nest; aggregations are often small and contain approximately 20–30 individuals, but over 100 brachypterous queens have been observed on vegetation outside their nests (pers. obs.; R.G. Pol, pers. comm.). Queens mate with multiple males with an average (± SD) effective mating frequency of 8.75 ± 3.26 (range = 4–16; n = 24) (Pol et al., 2008). After mating, queens leave the aggregation to initiate a nest using independent colony founding (R.G. Pol, pers. comm.), which is an unusual behavior for brachypterous queens (see Johnson, 2010). No information is available, but these brachypterous queens are probably obligate foragers (see Peeters et al., 2012).

Foraging
Pol et al. (2015) - Field experiments studying foraging were conducted in the Monte desert, Argentina. P. mendozanus display an intermediate foraging strategy in which workers are typically solitary foragers but recruit nestmates to high-density seed patches when they become available. The sum of their foraging efforts, in terms of the area covered, suggests their nest surroundings are continuously and thoroughly explored for food.

Diet
Dietary preferences of these seed harvesting ants have been studied in the Monte Desert (Biosphere Reserve of Ñacuñán ), Argentina (see Pirk et. al 2011). In choice tests this species preferred seeds of grass species to those of forbs and shrub.

Nomenclature

 *  mendozanus. Pogonomyrmex mendozanus Cuezzo & Claver, 2009: 101, figs. 1, 2, 5, 6, 7-11 (w.q.m.) ARGENTINA.

Worker
Holotype (Paratype variations in parentheses, n=20, notation: minimum - maximun). HL: 1.775 (1.5-2.0); HW: 1.9 (1.625-2.075); SL: 1.35 (1.15-1.425); DFL: 0.775 (0.725-0.85); EL: 0.4 (0.325-0.425); OCD: 0.525 (0.45-0.625); PW: 1.175 (1.125-1.375); WL: 2.075 (2.0- 2.525); CI: 107 (103-108); SI: 71 (70-71).

Head, mesosoma, petiole and postpetiole black to blackish brown, gaster yellow to reddish brown. Dorsal surface of body with short and whitish setae, sparsely distributed. Fine and very regular rugae present in dorsal surface of head, diverging towards occipital corners. Area between rugae, on dorsal surface of head, shagreened. Head: slightly wider than long (CI= 103-108). Posterior margin slightly concave with longitudinal rugae, without transverse rugae. Mandible strongly striated with six teeth. Anterior clypeal margin convex, bidentate. Clypeal area, between frontal lobes, with six-eigth rugae always present. Eye convex, protruding from side of head, centered on midpoint of side of head, without hairs between ommatidia. Scape not reaching posterior margin of head, but surpassing midpoint between posterior edge of eye and occipital margin. Scape covered by longitudinal rugae moderately to weakly developed. Psammophore well developed. Palpal formula: 4:3.

Mesosoma: profile interrupted by a strong ruga placed between mesonotum and propodeum. Pronotal collar smooth and shiny. Dorso-lateral angle of pronotum poorly developed, rounded. From side view, pronotal profile higher than mesonotum. Dorsal surface of pronotum covered with thick rugae, conspicuous and extensive. Humerus angular, with tubercle rounded. Propodeal rugae fine, regular and transverse. Procoxa with transverse rugae strongly developed. Dorsal propodeal spine well developed, slender and acutely pointed, more than twice as long as ventral spine, connected by conspicuous keel to contralateral tooth.

Petiole: peduncle smooth and shining with a very weak ventral process. Petiolar node elongated, more than twice longer than wide. Dorsal face with transversal rugae.

Postpetiole: dorsal surface with tranversal rugae. Ventral process strongly developed and covered with transverse rugae.

Gaster: smooth and shiny with sparse piligerous punctulae. First segment without rugae.

Johnson (2015) - (n = 16). HL 1.65–2.04; HW 1.87–2.28; MOD 0.37–0.49; OMD 0.47–0.67; SL 1.25–1.59; PNW 1.20–1.41; HF 1.79–2.28; ML 2.16–2.58; PW 0.49–0.67: PPW 0.65–0.82. Indices: SI 65.45–75.66; CI 102.50–113.33; OI 20.27–23.04; HFI 94.14–106.95. See also Cuezzo & Claver (2009).

Head subquadrate to wider than long (CI = 102.50–113.33), widest just posterior to eyes; posterior margin flat to weakly concave in full-face view. Cephalic dorsum with fine, regular, incised, longitudinal rugae; in full-face view, medial rugae diverging weakly toward posterior corners of head. In profile, rugae posterior to eyes converging toward vertex. Cephalic interrugae appearing as furrows, weakly to moderately coriarious, weakly shining; vertex rugose. Anterior margin of clypeus weakly to moderately concave; dorsal surface with numerous subparallel, longitudinal rugae. Mandible with six teeth; mandibular dorsum coarsely rugose. Numerous long, curved, bristle-like, yellowish to brownish hairs project from anterior margin of clypeus and basolateral margin of mandibles. MOD ranging from 0.20–0.25x HL. In profile, eyes situated near middle of head, OMD = 1.24–1.43x MOD. Antennal scapes relatively long (SI = 65.45–75.66), reaching vertex or failing to reach vertex by less than length of basal funicular segment; scapes often with moderately coarse longitudinal striae. Basal flange of scape flattened with carinate margin. Psammophore well-developed.

Promesonotal profile flattened, propodeum descending; all mesosomal surfaces with prominent subparallel/parallel rugae similar to that on cephalic dorsum. In profile and dorsal views, humeral shoulders of pronotum angulate, distinctly elevated above medial portion of pronotum. Dorsum of promesonotum with longitudinal rugae that diverge anterad toward humeral shoulders of pronotum; anterior margin of pronotum with transverse rugae that traverse posteroventrally or obliquely on pronotal sides. Mesopleura with transverse rugae, those near dorsal margin often traversing posterodorsally. Dorsum and sides of propodeum with transverse rugae. Superior propodeal spines long, acuminate; spines longer than distance between their bases. Inferior propodeal spines absent or reduced to indistinct broadly rounded processes. Propodeal spiracles narrowly ovate facing posterad. Interrugae on mesosoma weakly coriarious, weakly shining to shining. Legs moderately coriarious, weakly shining.

Peduncle of petiole slightly shorter than petiolar node, anteroventral margin usually flat, lacking tooth or lobe. In profile, petiolar node asymmetrical with anterior surface shorter than posterior surface; apex weakly angulate to rounded. In dorsal view, petiolar node longer than wide, widest near spatulate anterior margin; posterior surface with numerous fine, transverse to arcuate, wavy striae that curve posteroventrally on sides. Dorsum of postpetiole convex in profile; in dorsal view, widest near posterior margin, narrowing anterad; maximal width about equal to length; dorsum and sides with numerous transverse, wavy striae that are finer, denser than those on posterior surface of petiolar node. Interrugae on posterior surface of petiolar node and dorsum of postpetiole weakly to moderately punctate or coriarious, weakly shining to shining. First gastral tergum weakly to moderately coriarious, weakly shining to shining.

Erect white pilosity moderately abundant on head, variable in length, longest hairs not exceeding MOD. Moderately abundant suberect to semidecument pilosity on scape, abundant decumbent hairs on funicular segments. Legs with moderately abundant suberect white setae. Mesosoma, petiolar node, postpetiole with moderately dense, erect white setae, often similar in length, longest on pronotum, none >MOD; gastral terga with moderately dense pilosity, only those on posterior gastral terga approaching MOD. Head, mesosoma black; petiolar node dark orangish-brown to orangish-black; postpetiole, gaster lighter orangish-brown; mandibles, circumference of eyes, tarsi often dark reddish-brown.

Queen
n=4. HL: 2.075-2.3; HW: 2.43-2.73; ML: 1.03-1.28; EL: 0.48-0.53; EW: 0.2-0.28; SL: 1.55-1.65; DFL: 0.88-1.00; OCD: 0.6-0.7; PW: 1.45-1.58; WL: 2.4-2.73; CI:117-119; MI: 0.42-0.47; OI: 0.04-0.05; SI: 60-64.

Same pattern of color and sculpture as worker.

Head: Mandible striated with only six teeth. Scape short, not reaching posterior cephalic margin, covered with very fine rugae. Occipital margin slightly concave in middle.

Mesosoma: Similar to worker in color and sculpture. All four examined specimens are brachypterous. Forewing with only two longitudinal dark veins and no cells or pterostigma.

Petiole and Postpetiole: Dorsal and lateral faces transversely striated.

Gaster: First gastral segment without sculpture, covered by scattered and appressed hairs.

Johnson (2015) - (n = 3). HL 2.07–2.53; HW 2.44–2.81; MOD 0.50–0.53; OMD 0.55–0.81; SL 1.58–1.74; PNW 1.54–1.68; HFL 2.28–2.48; ML 2.59–2.76; PW 0.70–0.77; PPW 1.00–1.05. Indices: SI 61.92–66.13; CI 111.07–119.81; OI 18.86–20.56; HFI 88.26–95.16. See also Cuezzo & Claver (2009).

Male
N=1. HL: 1.2; HW: 1.48; ML: 2.5; EL: 0.55; EW: 0.18; SL: 0.45; PW: 1.4; WL: 2.5; CI: 123; MI: 169; OI: 0.07; SI: 30.

Pattern of color similar to that of worker and queen.

Head: subquadrate in frontal view, with posterior margin convex, with its maximum width after compound eyes. Dorsal surface of body with short and whitish setae, sparsely distributed. Fine and very regular rugae present in dorsal surface of head, diverging towards occipital corners. Vertex covered with transverse rugae. Compound eye well developed, convex, occupying more than one-third of lateral cephalic margin. Clypeus without smooth areas, all its dorsal surface covered by longitudinal rugae. Mandible with rugae on the outermost margin and with only four teeth. Scape shorther than combined length of first two flagellar segments, without rugae and with short, erect whitish hairs.

Mesosoma: in profile strongly sculptured with longitudinal rugae. Fore and hind wings well developed. Fore wing with two closed cubital cells and one discoidal cell. Hind wing with only two closed basal cells and a narrow costal cell. Hamulus with 10 hooks. Fore and hind wing disc covered with short hairs. Anterior surface of first coxa with long hairs. Coxa I shining. Propodeum without teeth, spines or tubercles in lateral view.

Petiole: Dorsal face areolate in lateral view, with very small ventral process below peduncle.

Postpetiole: Dorsal face with few rugae.

Gaster: with only 5 visible tergites and 6 sternites. Pigostyle short, but present. Genital capsule strongly sclerotized. Paramere without a differentiated cuspis. Ventral border of aedeagus in lateral view with row of minute teeth.

Johnson (2015) - (n = 12). HL 1.27–1.67; HW 1.44–1.68; MOD 0.46–0.56; OMD 0.23–0.40; SL 0.37–0.56; HFL 1.53–2.19; ML 2.47–2.91; PW 0.50–0.60; PPW 0.70–0.79. Indices: SI 23.42–34.15; CI 98.16–113.39; OI 29.34–33.94; HFI 106.25–133.54. See also Cuezzo & Claver (2009).

Type Material
Holotype worker  (1w), Argentina,   Provincia   de   Mendoza,   Santa Rosa, Ñacuñán, 12-II-1997, Silvia Claver coll. (deposited at IADIZA, Mendoza, Argentina). Paratypes  (n=   106   workers,   1   male,   4 queens) from same locality and collector as the holotype, several samples collected with different  data:  5w,  03-XII-1981;  9w,  16-III-1982;  67w,  26-III-98;  9w,  12-II-1997;  11w, 20-II-1997;  5w, 12-VI-1997;  Paratypes will be  deposited  at  the  following  institutions: 56w, 1m and 2q at IADIZA, 10w at MZSP, 5w at ICN, 5w at  MIZA, 30w and 2q at IFML.



Etymology
The specific epithet, mendozanus (Latinization of Mendoza), is derived from the type locality occurring in Mendoza Province, Argentina.

Determination Clarifications
Pogonomyrmex mendozanus was misidentified as Pogonomyrmex pronotalis in a list of ant species in Ñancuñan Biosphere Reserve (Claver & Fowler, 1993) and in subsequent publications until it was described by Cuezzo & Claver (2009).

References based on Global Ant Biodiversity Informatics

 * Behrmann N. B. E., F. A. Milesi, J. Lopez de Casenave, R. G. Pol, and B. Pavan. 2010. Colony size and composition in three Pogonomyrmex ant species (Hymenoptera: Formicidae) in the central Monte desert, Argentina. Rev. Soc. Entomol. Argent. 69 (1-2): 117-122.
 * Claver S., S. L. Silnik, and F. F. Campon. 2014. Response of ants to grazing disturbance at the central Monte Desert of Argentina: community descriptors and functional group scheme. J Arid Land 6(1): 117?127.
 * Cuezzo F., and S. Claver. 2009. Two new species of the ant genus Pogonomyrmex (Hmyenoptera: Formicidae) from Argentina. Revista de la Sociedad Entomológica Argentina 68: 97-106.
 * Johnson R. A. 2015. A taxonomic revision of South American species of the seed-harvester ant genus Pogonomyrmex (Hymenoptera: Formicidae). Part I. Zootaxa 4029(1):1-142
 * Johnson Robert. 2014. List of South American species of Pogonomyrmex. Accessed on February 5th 2014 at http://www.asu.edu/clas/sirgtools/pogonomyrmex/SOUTHAMERICANPOGOS.htm
 * Nobua Behrmann B. E., F. A. Milesi, J. Lopez de Casenave, R. G. Pol, and B. Pavan. 2010. Colony size and composition in three Pogonomyrmex ant species (Hymenoptera: Formicidae) in the Central Monte desert, Argentina. Rev. Soc. Entomol. Argent. 69 (1-2): 117-122.
 * Ulyssea M. A., L. P. Prado, C. R. F. Brandao. 2015. Type specimens of the traditional Myrmicinae (Hymenoptera: Formicidae) ant tribes deposited in the Museu de Zoologia da Universidade de Sao Paulo, Brazil: Adelomyrmecini, Basicerotini, Blepharidattini, Crematogastrini, Formicoxenini, Lenomyrmecini, Myrmicini, Phalacromyrmecini, Pheidolini, Stegomyrmecini, Stenammini and Tetramoriini. Papeis Avulsos de Zoologia, Museu de Zoologia da Universidade de Sao Paulo 55(12): 175-204.