Pheidole bessonii Forel, 1891

Salata, Sebastian & Fisher, Brian L., 2020, The ant genus Pheidole Westwood, 1839 (Hymenoptera: Formicidae) in Madagascar-taxonomic revision of the bessonii species-group, Zootaxa 4843 (1), pp. 1-64 : 17-21

Pheidole bessonii Forel, 1891

Figs 11 A–F, 27C, 28C

Pheidole oswaldi r. bessonii Forel, 1891: 176 , pl. 5, fig. 3 (w.).

Forel, 1905: 162 (s.).

Status as species: Dalla Torre, 1893: 88.

LECTOTYPE [designated here]: 1w., top specimen of the pin, Madagascar, Fianarantsoa, coll. Besson, ANTC3392, CASENT0101838 ( MHNG) . PARALECTOTYPES: 1w., bottom specimen, the same pin as lectotype; 3w., the same data as lectotype (CASENT0923201) .

Other material. Madagascar. Antananarivo: 11w., Ambohitantely 1410, -18.2253 47.28683, 1410 m, Fisher et al. leg. ( CASC) . Fianarantsoa: 1w., 9.0 km NE Ivohibe, -22.42667 46.93833, 900 m, Fisher et al. leg. ( CASC) ; 12w., Belle Vue trail, Ranomafana National Park , Fianarantsoa Prov., -21.2665 47.42017, 1020 m, R. Harin’Hala leg. ( CASC) ; 4w., dry wash, 1 km E of Isalo National Park Interpretive Center , Fianarantsoa Prov., -22.62667 45.35817, 885 m, Irwin et al. leg ( CASC) ; 5w., Fitovavy Fitovinany Region , District of Ifanadiana Belle vue area 1200 m S of Ranomafana National Park entrance, -21.2665 47.42017, 1018 m, M. Rin’ha leg.( CASC) ; 9w.,2s., Forêt d’Analalava , 29.6 km 280° W Ranohira, -22.59167 45.12833, 700 m, Fisher et al. leg. ( CASC) ; 4w., JIRAMA water works near river, Ranomafana National Park , Fianarantsoa Prov., -21.2485 47.45217, 690 m, R. Harin’Hala leg. ( CASC) ; 1w., Mampiarika IV Non Protected Area , 27.98 km SW Ambositra, -20.73528 47.08382, 1486 m, A. Ravelomanana leg. ( CASC) ; 8w., 4s., Parc National d’Isalo , 9.1 km 354° N Ranohira, -22.48167 45.46167, 725 m, Fisher et al. leg. ( CASC) ; 1w., Parc National d’Isalo, Ambovo Springs, 29.3 km 4° N Ranohira , -22.29833 45.35167, 990 m, Fisher et al. leg. ( CASC) ; 3w., 1s., Parc National d’Isalo, Sahanafa River , 29.2 km 351° N Ranohira, -22.31333 45.29167, 500 m, Fisher et al. leg. ( CASC) ; 1w., Parc National de Ranomafana, Vatoharanana River , 4.1 km 231° SW Ranomafana, -21.29 47.43333, 1100 m, Fisher et al. leg. ( CASC) ; 4w., R.S. Ivohibe 8.0 km E Ivohibe, -22.48333 46.96833, 1200 m, Fisher et al. leg. ( CASC) ; 2w., R.S. Ivohibe, 7.5 km ENE Ivohibe , -22.47 46.96, 900 m, Fisher et al. leg. ( CASC) ; 19w., radio tower, Ranomafana National Park , Fianarantsoa Prov., -21.25833 47.40717, 1130 m, Irwin et al. leg. ( CASC) ; 1s., Ranomafana National Park , Talatakely; Sahambavy ; Fianarantsoa Rural , -21.451179 47.3023894, 1139 m, Lee & Ribardo leg. ( CASC) ; 2w., stream area, 900 m E of Isalo National Park Interpretive Center , Fianarantsoa Prov., -22.62667 45.35817, 750 m, R. Harin’Hala leg. ( CASC) ; 12w., Vohiparara broken bridge, Fianarantsoa Prov., -21.22617 47.36983, 1110 m, R. Harin’Hala leg. ( CASC) ; 1w., Ampangabe I Non Protected Area , 21.4 km W Itremo, -20.61111 46.60688, 1414 m, A. Ravelomanana leg. ( CASC) ; 1w., Ampangabe II Non Protected Area , 21.29 km W Itremo, -20.61139 46.60809, 1402 m, A. Ravelomanana leg. ( CASC) ; 2w., Antapia III Non Protected Area , 26.43 km SW Ambositra, -20.72 47.08785, 1494 m, A. Ravelomanana leg. ( CASC) ; 1w., Antohatsahomby II Non Protected Area , 23.38 km NW Itremo, -20.55444 46.58438, 1640 m, A. Ravelomanana leg. ( CASC) ; 3w., Forêt d’Analalava , 29.6 km 280° W Ranohira, -22.59167 45.12833, 700 m, Fisher et al. leg. ( CASC) ; 1w., Mampiarika II Non Protected Area , 28.08 km SW Ambositra, -20.73556 47.08384, 1464 m, A. Ravelomanana leg. ( CASC) ; 3w., Mampiarika III Non Protected Area , 28.93 km SW Ambositra, -20.73583 47.08399, 1487 m, A. Ravelomanana leg. ( CASC) ; 1w., 2s., Mampiarika IV Non Protected Area , 27.98 km SW Ambositra, - 20.73528 47.08382, 1486 m, A. Ravelomanana leg. ( CASC) ; 1w., Manandriana II Non Protected Area , 27.12 km SW Ambositra, -20.7325 47.09461, 1589 m, A. Ravelomanana leg. ( CASC) ; 1s., Parc National d’Isalo , 9.1 km 354° N Ranohira, -22.48167 45.46167, 725 m, Fisher et al. leg. ( CASC) ; 1w., Parc National d’Isalo, Sahanafa River , 29.2 km 351° N Ranohira, -22.31333 45.29167, 500 m, Fisher et al. leg. ( CASC) ; 1w., Soanierenana II Non Protected Area , 25.61 km SW Ambositra, -20.72194 47.10896, 1732 m, A. Ravelomanana leg. ( CASC) . Mahajanga: 9w., 5s., Forêt Ambohimanga, 26.1 km 314° Mampikony , -15.96267 47.43817, 250 m, Fisher et al. leg. ( CASC) ; 1w., Mahavavy River , 6.2 km 145° SE Mitsinjo, -16.05167 45.90833, 20 m, Fisher et al. leg. ( CASC) ; 4w., 8s., 1q., Parc National d’Ankarafantsika, Ampijoroa Station Forestière, 5.4 km 331° NW Andranofasika, -16.29889 46.813, 70 m, Fisher et al. leg. ( CASC) ; 2w., Parc National d’Ankarafantsika, Ampijoroa Station Forestière, 40 km 306° NW Andranofasika, -16.32083 46.81067, 130 m, Fisher et al. leg. ( CASC) ; 8w., Parc National d’Ankarafantsika, Forêt de Tsimaloto , 18.3 km 46° NE de Tsaramandroso, -16.22806, 47.14361, 135 m, Fisher et al. leg. ( CASC) ; 7w., 5s., Parc National de Baie de Baly , 12.4 km 337° NNW Soalala, -16.01 45.265, 10 m, Fisher et al. leg. ( CASC) ; 12w., 3s., Parc National de Namoroka , 16.9 km 317° NW Vilanandro, -16.40667 45.31, 100 m, Fisher et al. leg. ( CASC) ; 2w., 1s., Parc National de Namoroka , 9.8 km 300° WNW Vilanandro, -16.46667 45.35, 140 m, Fisher et al. leg. ( CASC) ; 1w., Parc National Tsingy de Bemaraha, 10.6 km ESE 123° Antsalova , -18.70944 44.71817, 150 m, Fisher et al. leg. ( CASC) ; 4w., 1s., Parc National Tsingy de Bemaraha, 2.5 km 62° ENE Bekopaka, Ankidrodroa River , -19.13222 44.81467, 100 m, Fisher et al. leg. ( CASC) ; 6w., 1s., Parc National Tsingy de Bemaraha, 3.4 km 93° E Bekopaka, Tombeau Vazimba , -19.14194 44.828, 50 m, Fisher et al. leg. ( CASC) ; 1w, 1s., Réserve d’Ankoririka , 10.6 km 13° NE de Tsaramandroso, -16.26722 47.04861, 210 m, Fisher et al. leg. ( CASC) ; 2w., 1s., Réserve Spéciale de Bemarivo , 23.8 km 223° SW Besalampy, -16.925 44.36833, 30 m, Fisher et al. leg. ( CASC) ; 1w., Réserve forestière Beanka , 50.2 km E Maintirano, -18.02649 44.05051, 250 m, Fisher et al. leg. ( CASC) . Toamasina: 1w., 1s., 6.3 km S Ambanizana, Andranobe , -15.6813 49.958, 25 m, Fisher et al. leg. ( CASC) . Toliara: 1w., 50 km N Morondava , - 20.06667, 44.58333, A. Pauly leg. ( CASC) ; 1w., 1s., Anosy Region, Parc National d’Andohahela, Forêt de Manatalinjo , -24.82505 46.57811, 90 m, Fisher et al. leg. ( CASC) ; 1w., Antafoky , -23.48778 44.0775, 60 m, Fisher et al. leg. ( CASC) ; 1w., 1s., Beza-Mahafaly , 27 km E Betioky, -23.65 44.63333, 135 m, Fisher et al. leg. ( CASC) ; 1s., Fiherenana , -23.17694 43.96083, 100 m, Fisher et al. leg. ( CASC) ; 1w., Fiherenana , -23.23528 43.87083, 50 m, Fisher et al. leg. ( CASC) ; 1w., Forêt Classée d’Analavelona , 29.2 km 343° NNW Mahaboboka, -22.675 44.19, 1100 m, Fisher et al. leg. ( CASC) ; 3w., 3s., Forêt de Beroboka , 5.9 km 131° SE Ankidranoka, -22.23306 43.36633, 80 m, Fisher et al. leg. ( CASC) ; 4w., Forêt de Kirindy , 15.5 km 64° ENE Marofandilia, -20.045 44.66222, 100 m, Fisher et al. leg. ( CASC) ; 7w., Forêt de Kirindy , 15.5 km 64° ENE Marofandilia, -20.06855 44.6595667, 30 m, Fisher et al. leg. ( CASC) ; 4w., Forêt de Mahavelo, Isantoria River , -24.75833 46.15717, 110 m, Fisher et al. leg. ( CASC) ; 1w., Forêt de Mahavelo, Isantoria River , 5.5 km 37° NE Ifotaka, -24.75361 46.1515, 115 m, Fisher et al. leg. ( CASC) ; 3w., 3s., Forêt de Mite , 20.7 km 29° WNW Tongobory, -23.52417 44.12133, 75 m, Fisher et al. leg. ( CASC) ; 1w., Forêt Ivohibe 55.0 km N Tolagnaro, -24.569 47.204, 200 m, Fisher et al. leg. ( CASC) ; 1w., Forêt Vohidava 88.9 km N Amboasary, -24.24067 46.28783, 500 m, Fisher et al. leg. ( CASC) ; 1w., 1s., Forêt Vohidava 89.2 km N Amboasary, -24.239 46.28233, 850 m, Fisher et al. leg. ( CASC) ; 2w., Ivahona , -23.45591 46.17376, 820 m, Fisher et al. leg. ( CASC) ; 1w., Makay Mts. , -21.21836 45.3106, 510 m, Fisher et al. leg. ( CASC) ; 2w., Makay Mts. , -21.30997, 45.12946, 590 m, Fisher et al. leg. ( CASC) ; 3w., Manderano , -23.52722 44.0875, 70 m, Fisher et al. leg. ( CASC) ; 1w., Manombo , -22.81092 43.7344, 177 m, Fisher et al. leg. ( CASC) ; 1w., Manombo , -22.8123 43.73932, 165 m, Fisher et al. leg. ( CASC) ; 1w., Parc National d’Andohahela , Col du Sedro, 3.8 km 113° ESE Mahamavo, 37.6 km 341° NNW Tolagnaro, -24.76389 46.75167, 900 m, Fisher et al. leg. ( CASC) ; 6w., Parc National d’Andohahela, Forêt d’Ambohibory , 1.7 km 61° ENE Tsimelahy, 36.1 km 308° NW Tolagnaro, -24.93 46.6455, 300 m, Fisher et al. leg. ( CASC) ; 3w., Parc National d’Andohahela, Forêt de Manatalinjo, 33.6 km 63° ENE Amboasary, 7.6 km 99° E Hazofotsy , -24.81694 46.61, 150 m, Fisher et al. leg. ( CASC) ; 1w., Parc National de Kirindy Mite , 16.3 km 127° SE Belo sur Mer, -20.79528 44.147, 80 m, Fisher et al. leg. ( CASC) ; 4w., Parc National de Zombitse, 19.8 km 84° E Sakaraha , -22.84333 44.71, 770 m, Fisher et al. leg. ( CASC) ; 11w., 5s., 3m., Réserve Privé Berenty, Forêt de Bealoka, Mandraré River , 14.6 km 329° NNW Amboasary, -24.95694 46.2715, 35 m, Fisher et al. leg. ( CASC); 5w., Réserve Spéciale d’Ambohijanahary, Forêt d’Ankazotsihitafototra , 34.6 km 314° NW Ambaravaranala, -18.26 45.41833, 1100 m, Fisher et al. leg. ( CASC) ; 2w., Réserve Spéciale d’Ambohijanahary, Forêt d’Ankazotsihitafototra , 35.2 km 312° NW Ambaravaranala, -18.26667 45.40667, 1050 m, Fisher et al. leg. ( CASC) ; 1w., Sakaraha , -22.91233 44.53283, 470 m, Fisher et al. leg. ( CASC) ; 1w., Sept Lacs , -23.52833, 44.15556, 80 m, Fisher et al. leg. ( CASC) ; 1w., southern Isoky-Vohimena Forest , 59 km NE Sakaraha, -22.46667 44.85, 730 m, Fisher et al. leg. ( CASC) ; 14w., 9s., Forêt Classée d’Analavelona , 29.2 km 343° NNW Mahaboboka, -22.675 44.19, 1100 m, Fisher et al. leg. ( CASC) ; 1w., Forêt Classée d’Analavelona , 29.4 km 343° NNW Mahaboboka, - 22.675 44.18667, 1050 m, Fisher et al. leg. ( CASC) ; 1s., Forêt Vohidava 89.2 km N Amboasary, -24.239 46.28233, 850 m, Fisher et al. leg. ( CASC) ; 2w., Makay Mts. , -21.21761, 45.33917, 500 m, Fisher et al. leg. ( CASC) ; 2w., Makay Mts. , -21.22336 45.32628, 480 m, Fisher et al. leg. ( CASC) ; 4w., Makay Mts. , -21.30997 45.12946, 590 m, Fisher et al. leg. ( CASC) ; 2w., 2m., Makay Mts. , -21.31664 45.1296, 620 m, Fisher et al. leg. ( CASC) ; 3w., 2s., Parc National de Zombitse, 19.8 km 84° E Sakaraha , -22.84333 44.71, 770 m, Fisher et al. leg. ( CASC) ; 1w., Réserve Privé Berenty, Forêt de Malaza, Mandraré River , 8.6 km 314° NW Amboasary, -25.00778 46.306, 40 m, Fisher et al. leg. ( CASC) .

Geographic range. Madagascar, widely distributed on the island except Antsiranana.

Diagnosis. Major workers. Head in full-face view sub-rectangular, not or slightly widening posteriorly, lateral margin slightly convex with sparse, long, decumbent to subdecumbent pilosity; medial frons with moderately thick to thin and dense costulae that are on the posterior part; interspaces between costulae smooth to indistinctly punctate; frons laterally with sparser and thinner rugocostulae; interspaces between rugocostulae distinctly punctate; occipital lobes punctate, sometimes with additional network of sparse and thin rugae, sculpture weakening posteriorly; scape, when laid back, exceeding the midlength of head by two-fifths of its length; inner hypostomal tooth absent or indistinct, bulge-like; outer hypostomal tooth lobe-like, high and wide, directed posteriorly; median tooth absent; promesonotum short, angular, and moderately low; posterior mesonotum moderately steep; mesonotal process distinct and tubercle-like; mesosoma microreticulate; promesonotum with additional thin, sparse, and transverse rugae; gaster shagreened; body brown to blackish brown. Minor workers. Occiput elongated; nuchal collar distinct; head sculpture smooth; anterior frons with sparse puncta; scape, when laid back, exceeding the posterior head margin by two-fifths of its length; promesonotum low, long, and arched; promesonotal groove present; metanotal groove distinct; propodeal spines very small with relatively wide base; promesonotum mostly smooth with sparse microreticulae laterally; mesopleuron and propodeum with sparse microreticulae; body bright brown to brown.

Description. Major workers. Measurements (n=10): HL: 1.78–2.13 (1.97); HW: 1.74–2.01 (1.89); SL: 1.13–1.2 (1.17); EL: 0.22–0.27 (0.25); WL: 1.61–1.78 (1.68); PSL: 0.26–0.34 (0.3); MTL: 1.24–1.38 (1.3); PNW: 0.66–0.75 (0.7); PTW: 0.22–0.31 (0.28); PPW: 0.62–0.81 (0.7); CI: 99.7–106.5 (104.3); SI: 57.8–65.4 (61.7); PSLI: 13.9–16.6 (15.3); PPI: 32.1–47.0 (40.4); PNI: 34.2–39.1 (37.0); MTI: 64.8–74.8 (69.1).

Head. In full-face view sub-rectangular, slightly widening posteriorly with sides slightly convex ( Fig. 11B ); in lateral view sub-oval, not depressed posteriorly; ventral and dorsal margins convex; inner hypostomal tooth not visible. Sides of head with sparse, long, decumbent to subdecumbent pilosity; head dorsum with relatively dense, long, decumbent to erect pilosity. Medial frons with moderately thick to thin and dense costulae on the posterior part; interspaces between costulae smooth to indistinctly punctate. Frons laterally with sparser and thinner rugocostulae; interspaces between rugocostulae distinctly punctate. Occipital lobes punctate, sometimes with additional network of sparse and thin rugae; sculpture weakening posteriorly. Gena with dense, moderately thick to thin costulae; interspaces between costulae distinctly punctate. Sides posterolateral from eyes densely punctate to microreticulate, sometimes with additional network of thin and sparse rugulae. Center of clypeus shiny and with a few rugulae, lateral sides with distinct rugulae; median notch present, moderately wide, and shallow; median longitudinal carina present; lateral longitudinal carinae present. Scape, when laid back, exceeding the midlength of head by two-fifths of its length; pilosity subdecumbent to erect ( Fig. 11B, D ). Inner hypostomal tooth absent or indistinct, bulge-like; outer hypostomal tooth lobe-like, high, and wide,, with top directed posteriorly; median tooth absent ( Fig. 27C ).

Mesosoma. In lateral view, promesonotum short, angular, and moderately low; posterior mesonotum moderately steep; mesonotal process distinct and tubercle-like; promesonotal groove weakly developed; metanotal groove present; propodeal spines moderately long with wide base and acute top; humeral tubercle weakly produced ( Fig. 11D ). Surface shiny and microreticulate; promesonotum with additional thin, sparse, and transverse rugae. Pilosity moderately dense, long, and erect ( Fig. 11D, F ). Gaster. Dull and shagreened; pilosity moderately sparse, long, and erect ( Fig. 11D, F ). Color. Brown to blackish brown, antennae and legs yellowish brown, mandibles ochraceous ( Fig. 11D, F ).

Description. Minor workers. Measurements (n=10): HL: 0.69–0.82 (0.77); HW: 0.52–0.61 (0.57); SL:0.9– 1.09 (1.0); EL: 0.13–0.19 (0.16); WL: 0.96–1.12 (1.04); PSL: 0.07–0.12 (0.1); MTL: 0.78–0.95 (0.88); PNW: 0.33–0.41 (0.37); PTW: 0.09–0.12 (0.1); PPW: 0.17–0.21 (0.18); CI: 131.4–137.6 (133.5); SI: 169.3–183.2 (175.0); PSLI: 10.2–15.3 (13.3); PPI: 48.7–60.0 (55.3); PNI: 61.0–68.7 (65.2); MTI: 144.7–158.1 (152.8).

Head. In full-face view oval, posterior region elongated; nuchal collar distinct ( Fig. 11A ). Pilosity relatively sparse, moderately long, subdecumbent to erect. Sculpture shiny and smooth; anterolateral frons with sparse puncta. Clypeus with median longitudinal carina absent; two lateral longitudinal carinae absent. Scape, when laid back, exceeding the posterior head margin by two-fifths of its length; pilosity dense, subdecumbent to erect ( Fig. 11A, C ).

Mesosoma. In lateral view, promesonotum low, long, and arched; promesonotal groove present; metanotal groove distinct; propodeal spines very small, triangular, and with relatively wide base ( Fig. 11C ). Promesonotum mostly smooth with sparse microreticulae on lateral sides; katepisternum, anepisternum, and propodeum with sparse microreticulae. Pilosity very sparse, moderately long, and erect ( Fig. 11C, E ). Gaster. With sparse and erect pilosity ( Fig. 11C, E ). Color. Bright brown to brown, legs, mandibles, and antennae yellowish ( Fig. 11C, E ).

Biology. The species was collected between 30–1732 m elevation, in Uapaca woodland, montane rainforest, tropical dry forest, savanna and woodland, gallery forest and rainforest. Nests were located in rotten logs, in the soil, and under stones. Workers were collected from sifted litter.

Comments. Pheidole bessonii is probably one of the most widespread Pheidole species known from Madagascar. Morphologically it is most similar to P. voreios and P. atsirakambiaty , and its major workers can be separated from both species by punctate occipital lobes (which sometimes have additional thin and very sparse rugulae that arch posterolaterally), and sparsely rugocostulate lateral frons with distinctly punctate interspaces between rugocostulae. In contrast, majors of P. voreios and P. atsirakambiaty always have distinctly rugulate to costulate occipital lobes and frons. Minors of P. bessonii can be separated from P. voreios and P. atsirakambiaty by the less elongated posterior part of the neck, a mostly smooth promesonotum with sparse microreticulae on its sides, and very small propodeal spines. Pheidole voreios and P. atsirakambiaty have a more elongated posterior part of the neck, mostly or entirely sculptured promesonotum, and larger propodeal spines. In addition, majors and minors of P. bessonii can be distinguished from P. voreios by the brown body.

Figures

FIGURE 11. Pheidole bessonii, full-face view (A), profile (C), and dorsal view (E) of minor worker (CASENT0496974) and full-face view (B), profile (D), and dorsal view (F) of major worker (CASENT0496976).

FIGURE 27. Major worker, hypostomal tooth. Pheidole antsahabe (A). Pheidole atsirakambiaty (B). Pheidole bessonii (C). Pheidole clara (D). Pheidole decollata (E). Pheidole flammea (F). Pheidole flavodepressa (G). Pheidole grallatrix (H). Pheidole madecassa (I). Pheidole mantadioflava (J). Pheidole maro (K). Pheidole oswaldi (L). Pheidole ovalinoda (M). Pheidole similis (N). Pheidole tenebrovulgaris (O). Pheidole uranus (P). Pheidole voreios (Q). Pheidole zirafy (R).