Holotype. Madagascar. •1 major worker; Toamasina; Ankerana; -18.40829, 48.82107; alt. 750 m; 21 Jan 2012; B.L. Fisher et al. leg.; CASENT0275436 (CASC). Paratype. Madagascar. •1 w.; Toamasina; Ankerana; -18.4061, 48.82029; alt. 725 m; 16 Jan 2012; B.L. Fisher et al. leg.; BLF27931, CASENT0275480 (CASC).

Other material. Madagascar. –Fianarantsoa: •5w.; Forêt de Vevembe, 66.6 km 293° Farafangana; -22.791, 47.18183; alt. 600 m; 23 Apr 2006; B.L. Fisher et al. leg.; CASENT0108005, CASENT0108016, CASENT0108021, CASENT0108022, CASENT0108028 (CASC). –Toamasina: •2w.; Réserve Spéciale Ambatovaky, Sandrangato River; -16.77274, 49.26551; alt. 450 m; 20 Feb 2010; B.L. Fisher et al. leg.; CASENT0164424, CASENT0164471 (CASC). •3w., 2s.; Réserve Spéciale Ambatovaky, Sandrangato River; -16.7633, 49.26692; alt. 520 m; 22 Feb 2010; B.L. Fisher et al. leg.; CASENT0163714, CASENT0163724, CASENT0163831, CASENT0163832, CASENT0163976 (CASC). •1w.; Réserve Spéciale Ambatovaky, Sandrangato River; -16.81739, 49.29402; alt. 360 m; 25 Feb 2010; B.L. Fisher et al. leg.; CASENT0164250 (CASC). •2w.; Ankerana; -18.4061, 48.82029; alt. 725 m; 16 Jan 2012; B.L. Fisher et al. leg.; CASENT0275346, CASENT0275461 (CASC).

Major workers. Large species: HL: 1.67–1.91 (1.75), HW: 1.66–1.86 (1.71), WL: 1.23–1.3 (1.27); head in full-face view rectangular; in lateral view sub-rectangular with visible inner hypostomal teeth; antennal scrobes absent; sides of the head with dense, relatively long, erect pilosity; antennal sockets deep; frontal lobes distinct and lobe-like; inner hypostomal teeth distinct, closely spaced, triangular, with rounded apex and base slightly wider than apex; outer hypostomal teeth absent. Minor workers. Head shiny, with sparse, thick, irregular rugae, interspaces smooth or rugulose; frontal lobes present; antennal sockets deep; propodeal spines long and thin (PSL: 0.17–0.21 (0.19)).

Major workers. Measurements (N = 5): HL: 1.67–1.91 (1.75); HW: 1.66–1.86 (1.71); SL: 0.8–0.87 (0.85); EL: 0.15–0.16 (0.155); WL: 1.23–1.3 (1.27); PSL: 0.27–0.3 (0.28); MTL: 0.78–0.87 (0.81); PNW: 0.65–0.74 (0.69); PTW: 0.16–0.2 (0.17); PPW: 0.42–0.54 (0.46); CI: 97.1–99.4 (98.3); SI: 46.7–51.6 (48.9); PSLI: 14.8–17.7 (16.3); PPI: 36.1–39.8 (37.7); PNI: 38.9–42.8 (40.3); MTI: 46.9–47.8 (47.2). Head. In full-face view rectangular, anterior of eyes relatively straight, posterior of eyes slightly convex (Fig. 12B). In lateral view sub-rectangular; ventral and dorsal faces relatively flat; inner hypostomal teeth visible. Sides of the head with dense, relatively long, erect pilosity; whole head with moderately dense, long, suberect to erect pilosity. Antennal scrobes absent. Antennal sockets deep, smooth, or with a few rugae. Frontal lobes distinct and lobe-like. Occipital lobes with thick, sparse rugae; sculpture slightly weakening posteriorly. Whole head shiny, with sparse, thick, irregular rugae, interspaces smooth. Clypeus shiny, with a few longitudinal rugae; median notch present, wide, and shallow. Scape, when laid back, just reaching the midlength of head; pilosity decumbent to erect (Fig. 12B, D). Inner hypostomal teeth distinct, closely spaced, triangular, with rounded apex and base slightly wider than apex; outer hypostomal teeth absent (Fig. 84M). Mesosoma. In lateral view, promesonotum short, angular, and low, posterior mesonotum convex, with low tubercle-like projection; promesonotal groove absent; metanotal groove indistinct; propodeal spines long, massive basally, with acute apex; humeral area with short, triangular tubercles (Fig. 12D). Surface shiny, with thick, dense rugoreticulation, sculpture slightly weakening on dorsum. Pilosity moderately dense, long and erect (Fig. 12D, F). Petiole. Shiny, foveolate; peduncle relatively long, without horizontal lobes on its basal part; node, low and narrow, with flat apex, in rear view node dorsoventrally depressed; pilosity moderately dense and erect (Fig. 12D, F). Postpetiole. Foveolate; in dorsal view sides with short, acute, and triangular projections; pilosity long, moderately sparse and erect (Fig. 12D, F). Gaster. Shagreened on the whole surface; pilosity moderately dense, very long and erect (Fig. 12D, F). Colour. Unicolourous, dark brown (Fig. 12D, F).

Figure 12. 

Pheidole diakritos sp. nov., full-face view (A), profile (C), and dorsal view (E) of paratype minor worker (CASENT0275480) and full-face view (B), profile (D), and dorsal view (F) of holotype major worker (CASENT0275436).

Minor workers. Measurements (N = 10): HL: 0.68–0.78 (0.73); HW: 0.59–0.68 (0.64); SL: 0.73–0.83 (0.81); EL: 0.07–0.12 (0.1); WL: 0.89–1.04 (0.98); PSL: 0.17–0.21 (0.19); MTL: 0.56–0.66 (0.62); PNW: 0.44–0.52 (0.49); PTW: 0.08–0.13 (0.1); PPW: 0.12–0.19 (0.16); CI: 85.3–90.2 (87.8); SI: 121.9–129.7 (125.3); PSLI: 25.1–28.7 (26.4); PPI: 59.1–69.3 (64.6); PNI: 73.9–78.4 (76.0); MTI: 92.6–103.0 (97.1). Head. Occipital margin indistinctly concave; occipital carina indistinct, weakly developed (Fig. 12A). Pilosity sparse, long, suberect to erect; antennal sockets deep, smooth or with a few rugae; frontal lobes distinct and lobe-like. Whole head shiny, with sparse, thick, irregular rugae, interspaces smooth or with rugulae. Clypeus rugulose and shiny; median longitudinal carina absent; two lateral longitudinal carinae absent. Scape, when laid back, surpassing posterior head margin by two-fifths of its length; pilosity suberect to erect (Fig. 12A, C). Mesosoma. In lateral view, promesonotum convex; promesonotal groove present; metanotal groove present; humeral area with short triangular tubercles; propodeal spines moderately long, with base almost as wide as apex, apex acute (Fig. 12C). Pronotum and mesonotum with thick, dense rugoreticulation, sculpture slightly weakening on dorsum; katepisternum, anepisternum, and propodeum foveolate. Pilosity sparse, long, and erect (Fig. 12C, E). Petiole. Shiny; peduncle foveolate, moderately long and thin; node smooth, low, bulge-like; with few long, erect setae (Fig. 12C, E). Postpetiole. Sometimes partially foveolate; moderately short, low and slightly convex; with few long, erect setae at the anterior edge (Fig. 12C, E). Gaster. Pilosity sparse and erect (Fig. 12C, E). Colour. Brown to dark brown, legs brighter (Fig. 12C, E).

Etymology. Greek for distinct [διακριτός], in reference to its being the only known species with distinct frontal lobes and deep antennal sockets.

Biology. The species was collected at elevation between 360–865 m, in rainforest, and in montane rainforest. Nesting preferences are unknown.

Holotype. Madagascar. •1 major worker; Fianarantsoa; Parc National d’Isalo, 9.1 km 354°N Ranohira; -22.48167, 45.46167; alt. 725 m; 31 Jan 2003; B.L. Fisher et al. leg.; BLF07348, CASENT0485691 (CASC). Paratypes. Madagascar. •5w.; same data as for holotype; CASENT0485689, CASENT0485690, CASENT0872151–CASENT0872153 (CASC).

Major workers. Head in full-face view oval, with anterior and posterior sides slightly convex; sides of the head with moderately dense, long, erect pilosity; occipital lobes and genae shiny, with thick, sparse, irregular rugae, interspaces with indistinct foveolae, sculpture weakening posteriorly; inner hypostomal teeth distinct, moderately high, closely spaced, triangular, with rounded apex and wide base; outer hypostomal approximately as high as inner hypostomal teeth, dentate, with very wide base, and upper half thin, rectangular with top pointed outward; mesosoma shiny, with fine, sparse rugoreticulation and indistinct, sparse foveolae, sculpture weakening on dorsum, anepisternum, katepisternum, and mesosoma smooth; gaster smooth; body orange to bright brown. Minor workers. Frons, genae, and malar area smooth and shiny; vertex smooth and shiny with few arcuate, interrupted rugae; promesonotum moderately high, short, evenly arched; propodeal spines very small, indistinct, triangular; mesosoma smooth and shiny, only dorsum with few transverse, thick rugulae.

Major workers. Measurements (N = 1): HL: 0.97; HW: 0.9; SL: 0.47; EL: 0.09; WL: 0.8; PSL: 0.1; MTL: 0.48; PNW: 0.48; PTW: 0.14; PPW: 0.34; CI: 92.5; SI: 52.6; PSLI: 10.7; PPI: 40.8; PNI: 53.1; MTI: 53.6. Head. In full-face view oval, with anterior and posterior sides slightly convex (Fig. 15B). In lateral view sub-oval; ventral and dorsal faces convex; dorsal face not depressed posteriorly; inner hypostomal teeth visible. Sides of the head with moderately dense, long, erect pilosity; whole head with dense, short, suberect to erect pilosity. Antennal scrobes indistinct and not delimited by carinulae, scrobe surface shiny, with sparse, thick, longitudinal rugae, interspaces smooth or indistinctly rugulose. Occipital lobes and genae shiny, with thick, sparse, irregular rugae, interspaces with indistinct foveolae, sculpture weakening posteriorly; frons and malar area with dense, thick and longitudinal rugae, interspaces smooth. Centre of clypeus smooth and shiny, lateral sides with longitudinal rugae; median notch present, wide, and indistinct; median longitudinal carina absent; lateral longitudinal carinae absent. Scape, when laid back, slightly exceeding the midlength of head; pilosity suberect to erect (Fig. 15B, D). Inner hypostomal teeth distinct, moderately high, closely spaced, triangular, with rounded apex and wide base; outer hypostomal approximately as high as inner hypostomal teeth, dentate, with very wide base, and upper half thin, rectangular with top pointed outward (Fig. 84Z). Mesosoma. In lateral view, promesonotum short, angular, and relatively high, posterior mesonotum relatively steep, tubercle-like projections absent or very indistinct; promesonotal groove absent; metanotal groove absent; propodeal spines minute, indistinct, triangular, with acute apex; humeral area laterally weakly produced (Fig. 15D). Surface shiny, with fine, sparse rugoreticulation and indistinct, sparse foveolae, sculpture weakening on dorsum, anepisternum, and katepisternum, mesosoma smooth. Pilosity moderately dense, long, and erect (Fig. 15D, F). Petiole. Shiny with fine, indistinct, and sparse foveolae; peduncle moderately short, with indistinct horizontal lobes on its basal part; node smooth, relatively low, triangular, with rounded apex, in rear view node straight; pilosity moderately sparse and erect (Fig. 15D, F). Postpetiole. Shiny, with fine, indistinct, and sparse foveolae; in dorsal view sides with short, acute, and triangular projections; pilosity long, moderately dense, and erect (Fig. 15D, F). Gaster. Smooth; pilosity dense, moderately long, and erect (Fig. 15D, F). Colour. Unicolourous, orange to bright brown; legs yellow; gaster brown (Fig. 15D, F).

Figure 15. 

Pheidole lutea sp. nov., full-face view (A), profile (C), and dorsal view (E) of paratype minor worker (CASENT0485689) and full-face view (B), profile (D), and dorsal view (F) of holotype major worker (CASENT0485691).

Minor workers. Measurements (N = 4): HL: 0.46–0.48 (0.47); HW: 0.41–0.43 (0.42); SL: 0.39–0.43 (0.42); EL: 0.06–0.08 (0.07); WL: 0.51–0.55 (0.53); PSL: 0.04–0.06 (0.05); MTL: 0.31–0.32 (0.32); PNW: 0.28–0.29 (0.28); PTW: 0.06–0.07 (0.07); PPW: 0.1–0.11 (0.1); CI: 88.7–90.3 (89.7); SI: 95.2–104.3 (100.0); PSLI: 9.4–12.4 (10.7); PPI: 61.6–71.2 (66.8); PNI: 65.6–68.4 (67.2); MTI: 74.8–76.8 (75.5). Head. Occipital margin straight or indistinctly concave; occipital carina absent (Fig. 15A). Pilosity moderately dense, relatively short, erect. Frons, genae, and malar area smooth and shiny; vertex smooth and shiny with few arcuate, interrupted rugae; antennal sockets with few thick, curved outward rugae. Clypeus with median longitudinal carina absent; two lateral longitudinal carinae absent. Scape, when laid back, surpassing the posterior head margin by one-fifth of its length; pilosity suberect (Fig. 15A, C). Mesosoma. In lateral view, promesonotum moderately high, short, evenly arched; promesonotal groove absent; metanotal groove absent; propodeal spines very small, indistinct, triangular, apex acute (Fig. 15C). Sculpture smooth and shiny, only dorsum with few transverse, thick rugulae. Pilosity moderately sparse, moderately long, and erect (Fig. 15C, E). Petiole. Peduncle moderately short and thin with ventral face slightly convex; node low, triangular, and small; with few moderately long, erect setae (Fig. 15C, E). Postpetiole. Short, low and slightly convex; with few moderately long, erect setae (Fig. 15C, E). Gaster. With moderately sparse, erect pilosity (Fig. 15C, E). Colour. Unicoloured, yellow (Fig. 15C, E).

Latin for yellow, in reference to body colouration of major and minor workers.

The species was collected at elevation 725 m, in gallery forest. Nest was located in rotten log.

Pheidole lutea sp. nov. is most similar to P. ranohirensis sp. nov. and P. voasara sp. nov. Major workers. Pheidole lutea sp. nov. differs from those both taxa by surface of genae and antennal scrobes never foveolate, outer hypostomal teeth approximately as high as inner hypostomal teeth, with very wide base, and upper half thin, rectangular with top pointed outward, and by smooth anepisternum, katepisternum, and mesosoma. Minor workers. Pheidole lutea sp. nov. differs from P. ranohirensis sp. nov. in evenly arched, high and short promesonotum. Minor workers of P. voasara sp. nov. are unknown.

Holotype. Madagascar. •1 major worker; Fianarantsoa; Parc National d’Isalo, Sahanafa River, 29.2 km 351°N Ranohira; -22.31333, 45.29167; alt. 500 m; 10 Feb 2003; B.L. Fisher et al. leg.; BLF07681, CASENT0490851, top specimen (CASC). Paratypes. Madagascar. •6w., 5s.; same data as for holotype; CASENT0490850, CASENT0490852–CASENT0490854, CASENT0490856, CASENT0872083, CASENT0872206–CASENT0872214 (CASC).

Other material. Madagascar. –Antsiranana: •1s.; Ampasindava, Forêt d’Ambilanivy, 3.9 km 181°S Ambaliha; -13.79861, 48.16167; alt. 600 m; 4 Mar 2001; Fisher et al. leg.; CASENT0464406 (CASC). –Fianarantsoa: •9w., 8s., 2q.; Parc National d’Isalo, Sahanafa River, 29.2 km 351°N Ranohira; -22.31333, 45.29167; alt. 500 m; 10 Feb 2003; Fisher et al. leg.; CASENT0031826, CASENT0490740, CASENT0490756, CASENT0490757, CASENT0490761, CASENT0490763, CASENT0490768, CASENT0490772, CASENT0490864, CASENT0490865 (CASC). –Mahajanga: •2s.; Forêt de Tsimembo, 11.0 km 346°NNW Soatana; -18.99528, 44.4435; alt. 50 m; 21 Nov 2001; Fisher et al. leg.; CASENT0483413, CASENT0483517 (CASC). •1s.; Parc National d’Ankarafantsika, Forêt de Tsimaloto, 18.3 km 46°NE de Tsaramandroso; -16.22806, 47.14361; alt. 135 m; 2 Apr 2001; Fisher et al. leg.; CASENT0432165 (CASC). •3w., 3s.; Parc National de Namoroka, 16.9 km 317°NW Vilanandro; -16.40667, 45.31; alt. 100 m; 12-Nov-2002; Fisher et al. leg.; CASENT0038806, CASENT0038833, CASENT0038875, CASENT0023575, CASENT0023577, CASENT0023583 (CASC). •1w., 1s.; Parc National Tsingy de Bemaraha, 10.6 km ESE 123° Antsalova; -18.70944, 44.71817; alt. 150 m; 16 Nov 2001; Fisher et al. leg.; CASENT0078424, CASENT0078429 (CASC). •1s.; Parc National Tsingy de Bemaraha, 3.4 km 93°E Bekopaka, Tombeau Vazimba; -19.14194, 44.828; alt. 50 m; 6 Nov 2001; Fisher et al. leg.; CASENT0477494 (CASC). •4w., 1s.; Réserve Spéciale de Bemarivo, 23.8 km 223°SW Besalampy; -16.925, 44.36833; alt. 30 m; 19 Nov 2002; Fisher et al. leg.; CASENT0022564, CASENT0022764, CASENT0022772, CASENT0022775, CASENT0022787 (CASC).

Major workers. Head in full-face view oval, with anterior and posterior sides slightly convex; sides of the head with dense, long, erect pilosity; occipital lobes and genae shiny, with thick, sparse, irregular rugae, interspaces with fine but distinct foveolae, sculpture weakening posteriorly; inner hypostomal teeth distinct, moderately high, closely spaced, triangular, with rounded apex and narrow base; outer hypostomal approximately smaller and thinner than inner hypostomal teeth, with moderately wide base, dentate; mesosoma shiny, foveolate with additional sparse and thick rugae on promesonotal dorsum and propodeum, sculpture slightly weakening on dorsum; anepisternum and mesosoma with indistinct sculpture or smooth; body reddish brown. Minor workers. Frons, genae, and malar area smooth and shiny; vertex smooth and shiny with few arcuate, interrupted rugae; promesonotum low, short, slightly convex, with relatively steep posterior declivity; propodeal spines very small, indistinct, triangular; mesosoma smooth and shiny, only dorsum with few transverse, thick rugulae.

Major workers. Measurements (N = 10): HL: 0.92–1.08 (0.97); HW: 0.83–0.99 (0.87); SL: 0.46–0.57 (0.49); EL: 0.08–0.11 (0.1); WL: 0.76–0.93 (0.8); PSL: 0.09–0.12 (0.11); MTL: 0.42–0.53 (0.45); PNW: 0.46–0.51 (0.49); PTW: 0.12–0.15 (0.13); PPW: 0.33–0.37 (0.35); CI: 89.1–92.7 (90.9); SI: 51.9–59.0 (55.6); PSLI: 9.4–13.2 (11.3); PPI: 35.9–44.1 (39.8); PNI: 51.4–59.4 (55.9); MTI: 49.5–53.5 (51.6). Head. In full-face view oval, with anterior and posterior sides slightly convex (Fig. 16B). In lateral view sub-oval; ventral and dorsal faces convex; dorsal face not depressed posteriorly; inner hypostomal teeth visible. Sides of the head with dense, long, erect pilosity; whole head with dense, short, suberect to erect pilosity. Antennal scrobes indistinct and not delimited by carinulae; scrobe surface shiny, with sparse, thick, longitudinal rugae, interspaces with fine but distinct foveolae. Occipital lobes and genae shiny, with thick, sparse, irregular rugae, interspaces with fine but distinct foveolae, sculpture weakening posteriorly; frons with dense, thick, and longitudinal rugae, interspaces with fine but distinct foveolae; malar area with dense, thick, longitudinal rugulae, interspaces smooth. Centre of clypeus smooth and shiny, lateral sides with longitudinal rugae; median notch present, wide, and indistinct; median longitudinal carina absent; lateral longitudinal carinae absent. Scape, when laid back, slightly exceeding the midlength of head; pilosity suberect to erect (Fig. 16B, D). Inner hypostomal teeth distinct, moderately high, closely spaced, triangular, with rounded apex and narrow base; outer hypostomal teeth approximately smaller and thinner than inner hypostomal teeth, with moderately wide base, dentate (Fig. 85T). Mesosoma. In lateral view, promesonotum short, angular, and relatively high, posterior mesonotum relatively steep, with small tubercle-like projections; promesonotal groove absent; metanotal groove absent; propodeal spines minute, triangular, with acute apex; humeral area laterally weakly produced (Fig. 16D). Surface shiny, foveolate with additional sparse and thick rugae on promesonotal dorsum and propodeum, sculpture slightly weakening on dorsum; anepisternum and mesosoma with indistinct sculpture or smooth. Pilosity sparse, long, and erect (Fig. 16D, F). Petiole. Shiny with fine and sparse foveolae; peduncle moderately long, with indistinct horizontal lobes on its basal part; node smooth, relatively low, triangular, with rounded apex, in rear view node dorsoventrally concave; pilosity moderately sparse and erect (Fig. 16D, F). Postpetiole. Shiny, with fine and sparse foveolae; in dorsal view postpetiole very wide, almost semi-oval; pilosity long, moderately sparse and erect (Fig. 16D, F). Gaster. Shiny and finely shagreened; pilosity dense, moderately long and erect (Fig. 16D, F). Colour. Unicolourous, reddish brown; legs dark yellow; gaster brown (Fig. 16D, F).

Figure 16. 

Pheidole ranohirensis sp. nov., full-face view (A), profile (C), and dorsal view (E) of paratype minor worker (CASENT0490856) and full-face view (B), profile (D), and dorsal view (F) of holotype major worker (CASENT0490851).

Minor workers. Measurements (N = 10): HL: 0.43–0.47 (0.45); HW: 0.39–0.42 (0.41); SL: 0.41–0.44 (0.42); EL: 0.07–0.09 (0.08); WL: 0.5–0.56 (0.53); PSL: 0.05–0.07 (0.06); MTL: 0.3–0.35 (0.32); PNW: 0.26–0.28 (0.27); PTW: 0.05–0.07 (0.06); PPW: 0.1–0.14 (0.12); CI: 86.1–95.8 (89.9); SI: 101.7–112.2 (104.8); PSLI: 11.9–15.2 (13.4); PPI: 47.3–59.8 (52.2); PNI: 65.4–72.7 (67.5); MTI: 74.3 –85.7 (79.0). Head. Occipital margin straight or indistinctly concave; occipital carina absent (Fig. 16A). Pilosity moderately dense, long, erect. Frons, genae, and malar area smooth and shiny; vertex smooth and shiny with few arcuate, interrupted rugae; antennal sockets with few thick, curved outward rugae. Clypeus with median longitudinal carina absent; two lateral longitudinal carinae absent. Scape, when laid back, surpassing the posterior head margin by one-fifth of its length; pilosity suberect (Fig. 16A, C). Mesosoma. In lateral view, promesonotum low, short, slightly convex, with relatively steep posterior declivity; promesonotal groove absent; metanotal groove absent; propodeal spines very small, indistinct, triangular, apex acute (Fig. 16C). Sculpture smooth and shiny, only dorsum with few transverse, thick rugulae. Pilosity sparse, moderately short, and erect (Fig. 16C, E). Petiole. Peduncle short and thin with ventral face slightly convex; node low, triangular, and small; with few short, erect setae (Fig. 16C, E). Postpetiole. Short, low, and slightly convex; with few short, erect setae (Fig. 16C, E). Gaster. With sparse, erect pilosity (Fig. 16C, E). Colour. Unicolourous, yellow (Fig. 16C, E).

From the type locality.

The species was collected at elevation between 30–600 m, in gallery forest, in tropical dry forest, in rainforest. Nests were located in rotten logs and in dead twigs above ground.

Pheidole ranohirensis sp. nov. is most similar to P. lutea sp. nov. and P. voasara sp. nov. Major workers. Pheidole ranohirensis sp. nov. differs from P. lutea sp. nov. by surface of genae and antennal scrobes foveolate, dentate outer hypostomal teeth, which are smaller and thinner than inner hypostomal teeth and is never pointed outward, and by anepisternum, katepisternum, and mesosoma never entirely smooth; from P. voasara sp. nov. in indistinct or partially smooth sculpture on anepisternum and mesosoma, foveolate frons, and dentate outer hypostomal teeth, which are smaller and thinner than inner hypostomal teeth. Minor workers. Pheidole ranohirensis sp. nov. differs from P. lutea sp. nov. in promesonotum low, short, slightly convex, and with relatively steep posterior declivity. Minor workers of P. voasara sp. nov. are unknown.

Holotype. Madagascar. •1 major worker; Toliara; Réserve Spéciale d’Ambohijanahary, Forêt d’Ankazotsihitafototra, 35.2 km 312°NW Ambaravaranala; -18.26667, 45.40667; alt. 1050 m; 13 Jan 2003; B.L. Fisher et al. leg.; BLF07018, CASENT0050060 (CASC).

Madagascar. –Fianarantsoa: •1s.; Parc National d’Isalo, 9.1 km 354°N Ranohira; -22.48167, 45.46167; alt. 725 m; 27 Jan 2003; Fisher et al. leg.; CASENT0036511 (CASC). –Toliara: •2s.; Réserve Spéciale d’Ambohijanahary, Forêt d’Ankazotsihitafototra, 34.6 km 314°NW Ambaravaranala; -18.26, 45.41833; alt. 1100 m; 16 Jan 2003; Fisher et al. leg.; CASENT0029300, CASENT0029748 (CASC). •1s.; Réserve Spéciale d’Ambohijanahary, Forêt d’Ankazotsihitafototra, 35.2 km 312°NW Ambaravaranala; -18.26667, 45.40667; alt. 1050 m; 13 Jan 2003; Fisher et al. leg.; CASENT0028086 (CASC).

Major workers. Head, in full-face view, oval, with anterior and posterior sides slightly convex; sides of the head with sparse, long, erect pilosity; occipital lobes and genae shiny, with sparse, indistinct and irregular rugae, interspaces with fine but distinct foveolae, sculpture weakening posteriorly; inner hypostomal teeth distinct, moderately high, closely spaced, triangular, with rounded apex and wide base, tops directed inward; outer hypostomal bigger and wider than inner hypostomal teeth, with wide base, lobe-like; mesosoma shiny, shiny, foveolate with additional sparse to moderately dense, and moderately thick, rugae, sculpture slightly weakening on dorsum; upper part of mesosoma with indistinct sculpture; body yellowish brown.

Major workers. Measurements (N = 10): HL: 1.05–1.2 (1.16); HW: 0.96–1.11 (1.06); SL: 0.46–0.52 (0.5); EL: 0.09–0.13 (0.11); WL: 0.79–0.89 (0.85); PSL: 0.12–0.15 (0.13); MTL: 0.46–0.52 (0.49); PNW: 0.46–0.56 (0.53); PTW: 0.13–0.17 (0.15); PPW: 0.3–0.42 (0.38); CI: 90.9–94.2 (92.0); SI: 45.3–49.6 (46.9); PSLI: 10.2–13.0 (11.5); PPI: 32.7–43.3 (39.2); PNI: 47.3–52.3 (49.6); MTI: 44.4–48.5 (46.3). Head. In full-face view oval, with anterior and posterior sides slightly convex (Fig. 17C). In lateral view sub-oval; ventral and dorsal faces convex; dorsal face not depressed posteriorly; inner hypostomal teeth visible. Sides of the head with moderately dense, long, erect pilosity; whole head with moderately dense, short, suberect to erect pilosity. Antennal scrobes indistinct and not delimited by carinulae; scrobe surface shiny, with sparse, thick, longitudinal to irregular rugae, interspaces with fine but distinct foveolae. Occipital lobes and genae shiny, with sparse, indistinct and irregular rugae, interspaces with fine but distinct foveolae, sculpture weakening posteriorly; frons with dense, thick, and longitudinal rugae, interspaces smooth; malar area with dense, thick, longitudinal rugulae, interspaces smooth. Centre of clypeus smooth and shiny, lateral sides with longitudinal rugae; median notch present, wide and indistinct; median longitudinal carina absent; lateral longitudinal carinae absent. Scape, when laid back, reaching the midlength of head; pilosity suberect to erect (Fig. 17B–C). Inner hypostomal teeth distinct, moderately high, closely spaced, triangular, with rounded apex and wide base, tops directed inward; outer hypostomal teeth larger and wider than inner, with wide base, lobe-like (Fig. 85Z). Mesosoma. In lateral view, promesonotum short, angular, and high, posterior mesonotum relatively steep, with small tubercle-like projections; promesonotal groove absent; metanotal groove absent; propodeal spines small, triangular, with acute apex; humeral area laterally weakly produced (Fig. 17B). Surface shiny, foveolate with additional sparse to moderately dense, and moderately thick rugae, sculpture slightly weakening on dorsum; upper part of mesosoma with indistinct sculpture. Pilosity moderately sparse, long, and erect (Fig. 17A, B). Petiole. Shiny and with foveolae; peduncle moderately long, with distinct horizontal lobes on its basal part; node relatively low, triangular, with rounded apex, in rear view node dorsoventrally concave; pilosity moderately dense and erect (Fig. 17A, B). Postpetiole. Shiny and with foveolae, dorsum partially smooth; short and flat; in dorsal view very wide, almost semi-oval; pilosity long, moderately dense, and erect (Fig. 17A, B). Gaster. Shiny and finely shagreened on the basal part of first tergite; pilosity dense, moderately long and erect (Fig. 17A, B). Colour. Unicolourous, orange to yellowish brown; gaster brown; legs dark yellow (Fig. 17A, B).

Figure 17. 

Pheidole voasara sp. nov., dorsal view (A), profile (B), and full-face view (C) of holotype major worker (CASENT0050060).

Minor workers. Unknown.

Malagasy for orange, in reference to bright body colouration.

The species was collected at elevation between 725–1100 m, in gallery forest, and in montane rainforest. Nesting preferences unknown.

Pheidole voasara sp. nov. is most similar to P. lutea sp. nov. and P. ranohirensis sp. nov. Major workers. Pheidole voasara sp. nov. differs from P. lutea sp. nov. by surface of genae and antennal scrobes foveolate, lobe-like outer hypostomal teeth, inner hypostomal teeth pointed inward, and by anepisternum, katepisternum, and mesosoma never smooth; from P. ranohirensis sp. nov. in distinct and never smooth sculpture on anepisternum and mesosoma, frons never foveolate, and lobe-like outer hypostomal teeth, which are bigger and wider than inner hypostomal teeth. Minor workers. Unknown.

Holotype. Madagascar. •1 major worker; Antananarivo; Navoatra I Non-Protected Area, 7.64 km NW Arivonimamo; -18.97806, 47.11929; alt. 1373 m; 6 May 2010; Andrianjaka & Ravelomanana leg.; CASENT0204235 (CASC). Paratype. Madagascar. •1w.; same data as for holotype; ARA0915, CASENT0204254 (CASC).

Madagascar. –Fianarantsoa: •3w.; Ampangabe I Non-Protected Area, 21.4 km W Itremo; -20.61111, 46.60688; alt. 1414 m; 21 Mar 2010; Andrianjaka & Ravelomanana leg.; CASENT0211553, CASENT0213731, CASENT0213781 (CASC). •1w.; Ampangabe V Non-Protected Area, 21.37 km W Itremo; -20.61361, 46.60799; alt. 1449 m; 22 Mar 2010; Andrianjaka & Ravelomanana leg.; CASENT0207631 (CASC). •2w.; Ampangabe VI Non-Protected Area, 21.16 km W Itremo; -20.61444, 46.6104; alt. 1379 m; 21 Mar 2010; Andrianjaka & Ravelomanana leg.; CASENT0164859, CASENT0236320 (CASC). •1w.; Ampotoampoto I National Parc, 8.02 km NW Ilakaka; -22.62833, 45.18859; alt. 917 m; 26 Feb 2010; Andrianjaka & Ravelomanana leg.; CASENT0207300 (CASC). •3w.; Antohatsahomby I Non-Protected Area, 22.77 km NW Ambatofinandrahana; -20.55056, 46.58562; alt. 1550 m; 15 Mar 2010; Andrianjaka & Ravelomanana leg.; CASENT0207419, CASENT0207420, CASENT0207442 (CASC). •1w.; Antohatsahomby I Non-Protected Area, 22.77 km NW Ambatofinandrahana; -20.55056, 46.58562; alt. 1550 m; 15 Mar 2010; Andrianjaka & Ravelomanana leg.; CASENT0207433 (CASC). •1w.; Antohatsahomby II Non-Protected Area, 23.38 km NW Itremo; -20.55444, 46.58438; alt. 1640 m; 15 Mar 2010; Andrianjaka & Ravelomanana leg.; CASENT0211933 (CASC). •2s.; Parc National d’Isalo, 9.1 km 354°N Ranohira; -22.48167, 45.46167; alt. 725 m; 27 Jan 2003; Fisher et al. leg.; CASENT0036594, CASENT0036601 (CASC). –Mahajanga: •2s.; Parc National d’Ankarafantsika, Ampijoroa Station Forestière, 5.4 km 331°NW Andranofasika; -16.29889, 46.813; alt. 70 m; 26 Mar 2001; Fisher et al. leg.; CASENT0469065, CASENT0469084 (CASC). •1w.; Réserve forestière Beanka, 52.7 km E Maintirano; -18.0622, 44.52587; alt. 300 m; 24 Oct 2009; Fisher et al. leg.; CASENT0157649 (CASC). –Toliara: •10w.; Makay Mts.; -21.30997, 45.12946; alt. 590 m; 3 Dec 2010; Fisher et al. leg.; CASENT0205797, CASENT0205801, CASENT0205807, CASENT0205808, CASENT0205812, CASENT0205824, CASENT0205827, CASENT0205832, CASENT0205984 (CASC). •1s.; Réserve Spéciale Kalambatritra; -23.4185, 46.4583; alt. 1365 m; 8 Feb 2009; Fisher et al. leg.; CASENT0149710 (CASC).

Major workers. Head elongate; sides of the head with very sparse, moderately short, erect pilosity; frons smooth, with few short, thick, longitudinal rugae on the anterior part; genae shiny and smooth; inner hypostomal teeth distinct, small, closely spaced, triangular, with rounded apex; outer hypostomal teeth weakly developed, dentate; body yellow to orange; mesosoma smooth, sometimes with indistinct and sparse foveolae or rugulae on propodeum and promesonotum. Minor workers. Body yellow; head shiny and smooth, only lateral sides of frons with short, indistinct, longitudinal rugulae; propodeal spines short and triangular; mesosoma smooth and shiny.

Major workers. Measurements (N = 10): HL: 1.09–1.45 (1.21); HW: 0.86–1.14 (0.94); SL: 0.46–0.58 (0.5); EL: 0.12–0.15 (0.13); WL: 0.84–1.02 (0.91); PSL: 0.14–0.19 (0.17); MTL: 0.49–0.6 (0.53); PNW: 0.5–0.65 (0.55); PTW: 0.15–0.19 (0.17); PPW: 0.3–0.41 (0.34); CI: 74.5–80.3 (77.8); SI: 50.2–56.7 (53.1); PSLI: 12.9–15.6 (14.0); PPI: 43.3–54.9 (49.3); PNI: 56.6–61.4 (58.0); MTI: 52.4 –59.9 (57.1). Head. In full-face view longer than wide, anterior of eyes straight, posterior of eyes straight and slightly convex (Fig. 18B). In lateral view sub-oval; ventral and dorsal faces convex; dorsal face not depressed posteriorly; inner hypostomal teeth invisible. Sides of the head with very sparse, moderately short, erect pilosity; whole head with moderately dense, very short, suberect to erect pilosity. Antennal scrobes very weakly impressed and not delimited, scrobe surface shiny, with thick, longitudinal, short, and interrupted rugae; interspaces distinctly rugo-foveolate. Occipital lobes shiny, with thick, irregular rugae, interspaces rugo-foveolate, rugae fading anteriorly; frons smooth, with few short, thick, longitudinal rugae on the anterior part; genae shiny and smooth; malar area with thin, longitudinal, moderately dense rugae, interspaces smooth or with indistinct rugulae. Centre of clypeus shiny and smooth, lateral sides with longitudinal rugulae; median notch present, moderately wide and shallow; median longitudinal carina absent; lateral longitudinal carinae indistinct. Scape, when laid back, not reaching the midlength of head; pilosity subdecumbent to erect (Fig. 18B, D). Inner hypostomal teeth distinct, small, closely spaced, triangular, with rounded apex; outer hypostomal teeth weakly developed, dentate (Fig. 85M). Mesosoma. In lateral view, promesonotum relatively low and arched, posterior mesonotum convex, without tubercle-like projections; promesonotal groove absent; metanotal groove absent; propodeal spines small, triangular, with sharp apex and wide base; humeral area laterally weakly produced (Fig. 18D). Surface shiny and smooth, sometimes with indistinct and sparse foveolae or rugulae on propodeum and promesonotum. Pilosity moderately dense, very long and suberect to erect (Fig. 18D, F). Petiole. Shiny; peduncle moderately long, finely foveolate, without horizontal lobes on its basal part; node smooth, relatively low, triangular, with rounded apex, in rear view node slightly concave; pilosity moderately dense and erect (Fig. 18D, F). Postpetiole. Shiny; in dorsal view sides with moderately short, acute, and triangular projections; pilosity long, moderately dense and erect (Fig. 18D, F). Gaster. Shiny and smooth; pilosity dense, short, and erect (Fig. 18D, F). Colour. Yellow to orange; malar area, lower frons and gaster dark yellow to yellowish brown (Fig. 18D, F).

Figure 18. 

Pheidole navoatrensis sp. nov., full-face view (A), profile (C), and dorsal view (E) of paratype minor worker (CASENT0204254) and full-face view (B), profile (D), and dorsal view (F) of holotype major worker (CASENT0204235).

Minor workers. Measurements (N = 5): HL: 0.45–0.49 (0.47); HW: 0.41–0.45 (0.43); SL: 0.39–0.43 (0.41); EL: 0.09–0.11 (0.1); WL: 0.53–0.59 (0.54); PSL: 0.06–0.09 (0.07); MTL: 0.33–0.36 (0.35); PNW: 0.27–0.3 (0.28); PTW: 0.06–0.08 (0.07); PPW: 0.1–0.12 (0.11); CI: 88.7–90.7 (89.7); SI: 90.7–99.1 (95.6); PSLI: 12.6–17.5 (15.1); PPI: 61.5–75.5 (66.2); PNI: 64.0–68.5 (66.7); MTI: 78.0–84.3 (80.7). Head. Occipital margin straight or indistinctly concave; occipital carina absent (Fig. 18A). Pilosity moderately dense, long, suberect. Whole head shiny and smooth, only lateral sides of frons with short, indistinct, longitudinal rugulae; antennal sockets with sparse carinae curved outward. Clypeus with median longitudinal carina absent; two lateral longitudinal carinae absent. Scape, when laid back, reaching posterior head margin; pilosity suberect to erect (Fig. 18A, C). Mesosoma. In lateral view, promesonotum convex; promesonotal groove absent; metanotal groove indistinct; propodeal spines short and triangular, apex acute (Fig. 18C, E). Sculpture smooth and shiny. Pilosity moderately dense, long, and erect (Fig. 18C, E). Petiole. Peduncle short and thin with ventral face slightly convex; node globular; with few moderately long, erect setae (Fig. 18C, E). Postpetiole. Short, low, and convex; with few moderately long, erect setae (Fig. 18C, E). Gaster. With moderately dense, erect pilosity (Fig. 18C, E). Colour. Unicolourous, yellow (Fig. 18C, E).

From the type locality.

The species was collected at elevation between 70–1640 m, in dry forest on sandy soil, in Uapaca woodland, in savannah woodland, in tropical dry forest, in gallery forest, in shrubland. Nests were located in soil and under stones.

Pheidole navoatrensis sp. nov. is most similar to P. typhlos sp. nov. Major workers. Pheidole navoatrensis sp. nov. differs from P. typhlos sp. nov. in well-developed eyes, never smooth occipital lobes, small inner hypostomal teeth and weakly developed outer hypostomal teeth, and posterior mesonotum never convex. Minor workers. Pheidole navoatrensis sp. nov. differs from P. typhlos sp. nov. in convex promesonotum, big eyes, and shallow metanotal groove.

Holotype. Madagascar. •1 major worker; Antananarivo; Ankalalahana; -19.00716, 47.1124; alt. 1370 m; 3 Jan 2013; B.L. Fisher et al. leg.; BLF31313, CASENT0303345 (CASC). Paratype. Madagascar. •1 w.; same data as for holotype; CASENT0923187 (CASC).

Madagascar. – Antananarivo: •2w., 2s.; Andohony I Non-Protected Area, 22.62 km SW Antsirabe; -20.06784, 46.99068; alt. 1451 m; 6 Mar 2012; Andrianjaka & Ravelomanana leg.; CASENT0302331, CASENT0302483 (CASC). •2w.; Antaponimanadala I Non-Protected Area, 6.59 km E Manalalondo; -19.25528, 47.1771; alt. 1984 m; 13 May 2010; Andrianjaka & Ravelomanana leg.; CASENT0211687, CASENT0211787 (CASC). •3s.; Antaponimanadala III Non-Protected Area, 6.55 km E Manalalondo; -19.25583, 47.17751; alt. 1987 m; 17 May 2010; Andrianjaka & Ravelomanana leg.; CASENT0236559, CASENT0284933, CASENT0284998 (CASC). •2w., 1s., 1m.; Antaponimanadala IV Non-Protected Area, 6.66 km E Manalalondo; -19.25361, 47.17634; alt. 1951 m; 15 May 2010; Andrianjaka & Ravelomanana leg.; CASENT0228718, CASENT0228735 (CASC). •1w.; Beapombo II Non-Protected Area, 22.65 km SW Antsirabe; -20.07022, 47.00555; alt. 1689 m; 28 Feb 2012; Andrianjaka & Ravelomanana leg.; CASENT0292480 (CASC). •1w.; Navoatra II Non-Protected Area, 7.54 km NW Arivonimamo; -18.97889, 47.11975; alt. 1357 m; 6 May 2010; Andrianjaka & Ravelomanana leg.; CASENT0203015 (CASC). –Fianarantsoa: •3w., 1s.; Ambinanindranomena Non-Protected Area, 39.16 km SE Ambalavao; -21.96077, 47.29125; alt. 1002 m; 3 Feb 2012; Andrianjaka & Ravelomanana leg.; CASENT0285095, CASENT0289395, CASENT0289462, CASENT0289567 (CASC). •3w., 1s.; Ampanenitra Non-Protected Area, 41.19 km SE Ambalavao; -21.9652, 47.31001; alt. 1010 m; 8 Feb 2010; Andrianjaka & Ravelomanana leg.; CASENT0289604, CASENT0289694, CASENT0289797, CASENT0292637 (CASC). •4w.; Ampangabe IV Non-Protected Area, 21.37 km W Itremo; -20.61278, 46.60774; alt. 1417 m; 21 Mar 2010; Andrianjaka & Ravelomanana leg.; CASENT0236453, CASENT0236497, CASENT0236524, CASENT0236525 (CASC). •1w., 1s.; Mampiarika III Non-Protected Area, 28.93 km SW Ambositra; -20.73583, 47.08399; alt. 1487 m; 1 Feb 2010; Andrianjaka & Ravelomanana leg.; CASENT0165724, CASENT0165857 (CASC). •1w.; Manandriana I Non-Protected Area, 27.11 km SW Ambositra; -20.73194, 47.09413; alt. 1590 m; 9 Feb 2010; Andrianjaka & Ravelomanana leg.; CASENT0167038 (CASC). •1s., 1m.; Manandriana III Non-Protected Area, 27.25 km SW Ambositra; -20.73333, 47.09391; alt. 1578 m; 8 Feb 2010; Andrianjaka & Ravelomanana leg.; CASENT0206047, CASENT0210577 (CASC). •1w.; Soanierenana I Non-Protected Area, 25.33 km SW Ambositra; -20.72139, 47.10994; alt. 1723 m; 6 Feb 2010; Andrianjaka & Ravelomanana leg.; CASENT0207108 (CASC). •3s.; Soanierenana III Non-Protected Area, 25.25 km SW Ambositra; -20.72194, 47.11019; alt. 1707 m; 5 Feb 2010; Andrianjaka & Ravelomanana leg.; CASENT0213472, CASENT0213840, CASENT0213844 (CASC). •1w., 1s., 1m.; Soanierenana IV Non-Protected Area, 25.22 km SW Ambositra; -20.72389, 47.10705; alt. 1736 m; 5 Feb 2010; Andrianjaka & Ravelomanana leg.; CASENT0168320, CASENT0168431 (CASC). –Toliara: •1w.; Réserve Spéciale Kalambatritra; -23.4185, 46.4583; alt. 1365 m; 8 Feb 2009; B.L. Fisher et al. leg.; CASENT0149620 (CASC).

Major workers. Eyes small and reduced; body yellow; head rectangular, anterior of eyes relatively straight, posterior of eyes slightly convex; sides of the head with dense, relatively long, erect pilosity; occipital lobes and genae shiny and smooth, sometimes with indistinct, very sparse, longitudinal rugulae; frons and malar area shiny, with thick, sparse, short, and interrupted rugae, interspaces smooth; propodeal spines absent or weakly developed, lobe-like; mesosoma shiny, with fine, sparse to moderately sparse rugoreticulation, sculpture on dorsum weaker or with smooth patches; inner hypostomal teeth indistinct, very low, lobe-like, with rounded apex; outer hypostomal teeth distinct, dentate, and with rounded tops directed outward. Minor workers. Eyes small and reduced; body yellow; head and mesosoma shiny and smooth; propodeal spines indistinct, triangular.

Major workers. Measurements (N = 10): HL: 0.72–1.07 (0.93); HW: 0.6–0.83 (0.74); SL: 0.36–0.48 (0.43); EL: 0.05–0.07 (0.06); WL: 0.7–0.88 (0.8); PSL: 0.08–0.12 (0.1); MTL: 0.37–0.46 (0.43); PNW: 0.38–0.53 (0.47); PTW: 0.11–0.16 (0.14); PPW: 0.23–0.32 (0.28); CI: 76.1–82.9 (80.1); SI: 55.8–61.4 (58.4); PSLI: 9.7–12.2 (11.0); PPI: 45.2–54.6 (49.1); PNI: 60.0–65.5 (63.2); MTI: 54.7–62.9 (58.0). Head. In full-face view elongate, anterior of eyes relatively straight, posterior of eyes slightly convex (Fig. 19B). In lateral view sub-oval; ventral and dorsal faces moderately convex; inner hypostomal teeth invisible. Sides of the head with dense, relatively long, erect pilosity; whole head with moderately dense, short, suberect to erect pilosity. Antennal scrobes absent. Occipital lobes and genae shiny and smooth, sometimes with indistinct, very sparse, longitudinal rugulae; frons and malar area shiny, with thick, sparse, short and interrupted rugae, interspaces smooth. Clypeus shiny and smooth, with a few longitudinal rugae on the lateral sides; median notch present, wide, and shallow; median longitudinal carina indistinct; lateral longitudinal carinae absent. Scape, when laid back, reaching slightly over the midlength of head; pilosity decumbent to erect (Fig. 19B, D). Inner hypostomal teeth indistinct, very low, lobe-like, with rounded apex; outer hypostomal teeth distinct, dentate, and with rounded tops directed outward (Fig. 85P). Mesosoma. In lateral view, promesonotum short, angular, and low, posterior mesonotum convex, with low tubercle-like projection; promesonotal groove present; metanotal groove absent; propodeal spines absent or weakly developed, lobe-like; humeral area with short triangular tubercles (Fig. 19D). Surface shiny, with fine, sparse to moderately sparse rugoreticulation, sculpture on dorsum weaker or with smooth patches. Pilosity moderately dense, long, and erect (Fig. 19D, F). Petiole. Peduncle relatively long, without horizontal lobes on its basal part; node moderately high and globular, with convex apex, in rear view node concave; pilosity moderately dense and erect (Fig. 19D, F). Postpetiole. Smooth and shiny; in dorsal view sides with short, acute, and tubercle-like projections; pilosity long, moderately dense and erect (Fig. 19D, F). Gaster. Smooth and shiny; pilosity dense, short and erect (Fig. 19D, F). Colour. Yellow, malar area and lower part of frons dark yellow to bright brown (Fig. 19D, F).

Figure 19. 

Pheidole parviocula sp. nov., full-face view (A), profile (C), and dorsal view (E) of paratype minor worker (CASENT0923187) and full-face view (B), profile (D), and dorsal view (F) of holotype major worker (CASENT0303345).

Minor workers. Measurements (N = 10): HL: 0.45–0.51 (0.49); HW: 0.4–0.45 (0.44); SL: 0.34–0.37 (0.35); EL: 0.04–0.06 (0.05); WL: 0.5–0.58 (0.55); PSL: 0.05–0.07 (0.06); MTL: 0.25–0.31 (0.29); PNW: 0.26–0.3 (0.29); PTW: 0.07–0.08 (0.07); PPW: 0.11–0.12 (0.11); CI: 86.4–92.8 (89.6); SI: 76.9–85.4 (81.1); PSLI: 9.3–13.5 (11.8); PPI: 60.2–72.4 (65.1); PNI: 59.1–70.1 (65.9); MTI: 62.4–71.5 (66.9). Head. In full-face view rectangular, posterior and anterior of eyes slightly convex, occipital margin straight or indistinctly convex; occipital carina absent (Fig. 19A). Pilosity moderately dense, short, subdecumbent to suberect. Whole head shiny and smooth, only antennal sockets with sparse carinae curved outward. Clypeus with median longitudinal carina absent; two lateral longitudinal carinae absent. Scape, when laid back, reaching posterior head margin; pilosity suberect to erect (Fig. 19A, C). Mesosoma. In lateral view, promesonotum convex; promesonotal groove present; metanotal groove indistinct; propodeal spines indistinct, triangular, apex acute (Fig. 19C). Sculpture smooth and shiny. Pilosity moderately sparse, moderately long and erect (Fig. 19C, E). Petiole. Peduncle moderately short and thin; node globular and moderately high; with few long, erect setae (Fig. 19C, E). Postpetiole. Moderately short, low, and slightly convex; with few long, erect setae at the anterior edge (Fig. 19C, E). Gaster. Pilosity moderately dense, erect, and moderately long (Fig. 19C, E). Colour. Unicolourous, yellow (Fig. 19C, E).

Latin for small eyes.

The species was collected at elevation between 1002–1987 m, in savannah grassland, in Uapaca woodland, in savannah woodland, and in grassland. Nests were located in soil, under stones, and in dead tree stumps.

This species is most similar to P. typhlos sp. nov. Major workers. Pheidole parviocula sp. nov. can be distinguished from P. typhlos sp. nov. by dense, relatively long pilosity on sides of the head, small, lobe-like inner hypostomal teeth and weakly developed or absent propodeal spines. Minor workers. Pheidole parviocula sp. nov. can be distinguished from P. typhlos sp. nov. by convex shape of promesonotum convex, presence of promesonotal groove and weakly developed, small, triangular, apex acute.

Holotype. Madagascar. •1 major worker; Antsiranana; Galoko chain, Mont Galoko; -13.58487, 48.71818; alt. 520 m; 19 Feb 2013; B.L. Fisher et al. leg.; BLF30851, CASENT0302953 (CASC). Paratype. Madagascar. •1 w.; same data as for holotype; CASENT0923217 (CASC).

Madagascar. – Antananarivo: •6s.; Galoko chain, Mont Galoko; -13.58487, 48.71818; alt. 520 m; 16 Feb 2013; B.L. Fisher et al. leg.; CASENT0305030 (CASC). •1w.; Galoko chain, Mont Galoko; -13.5888, 48.72864; alt. 980 m; 22 Feb 2013; B.L. Fisher et al. leg.; CASENT0304748 (CASC).

Major workers. Eyes small and reduced; body yellow; head rectangular, anterior of eyes relatively straight, posterior of eyes slightly convex; sides of the head with sparse, short, erect pilosity; occipital lobes, genae, and posterior part of frons shiny and smooth, sometimes with indistinct, very sparse, longitudinal rugulae; anterior part of frons and malar area shiny, with thick, very sparse, short, and interrupted rugae, interspaces smooth; propodeal spines short, triangular, with acute top; pronotum smooth and shiny, sometimes on lateral sides with indistinct and sparse rugulae; mesonotum and propodeum shiny, with fine, sparse to moderately sparse rugoreticulation; katepisternum indistinctly foveolate; anepisternum with thick rugoreticulation; inner hypostomal teeth distinct, moderately high, narrow, triangular, with rounded apex; outer hypostomal teeth distinct, triangular, and with rounded tops directed outward; outer hypostomal teeth are slightly lower than inner hypostomal teeth. Minor workers. Eyes small and reduced; body yellow; head and mesosoma shiny and smooth, lateral sides of frons with short, indistinct, longitudinal rugulae; propodeal spines short and triangular.

Major workers. Measurements (N = 8): HL: 1.2–1.34 (1.28); HW: 0.96–1.07 (1.01); SL: 0.49–0.52 (0.5); EL: 0.08–0.12 (0.1); WL: 0.79–0.91 (0.84); PSL: 0.15–0.19 (0.16); MTL: 0.48–0.52 (0.51); PNW: 0.48–0.55 (0.51); PTW: 0.14–0.16 (0.15); PPW: 0.37–0.43 (0.4); CI: 75.1–82.0 (79.5); SI: 48.0–52.3 (49.8); PSLI: 11.9–14.3 (12.7); PPI: 36.3–39.9 (37.9); PNI: 47.4–55.3 (50.6); MTI: 46.9–52.9 (49.8). Head. In full-face view elongate, anterior of eyes relatively straight, posterior of eyes slightly convex (Fig. 20B). In lateral view sub-oval; ventral and dorsal faces relatively flat; inner hypostomal teeth invisible. Antennal scrobes absent. Sides of the head with sparse, short, erect pilosity; whole head with dense, short, suberect to erect pilosity. Occipital lobes, genae, and posterior part of frons shiny and smooth, sometimes with indistinct, very sparse, longitudinal rugulae; anterior part of frons and malar area shiny, with thick, very sparse, short, and interrupted rugae, interspaces smooth. Clypeus shiny and smooth, with a few indistinct, longitudinal rugae on the lateral sides; median notch present, wide, and shallow; median longitudinal carina absent; lateral longitudinal carinae absent. Scape, when laid back, reaching the midlength of head; pilosity decumbent to erect (Fig. 20B, D). Inner hypostomal teeth distinct, moderately high, narrow, triangular, with rounded apex; outer hypostomal teeth distinct, triangular, and with rounded tops directed outward; outer hypostomal teeth are slightly lower than inner hypostomal teeth (Fig. 85X). Mesosoma. In lateral view, promesonotum short, angular, and moderately high, posterior mesonotum convex, with indistinct tubercle-like projection; promesonotal groove absent; metanotal groove absent or very indistinct; propodeal spines short, triangular, with acute top; humeral area with wide and flat tubercles (Fig. 20D). Pronotum smooth and shiny, sometimes on lateral sides with indistinct and sparse rugulae; mesonotum and propodeum shiny, with fine, sparse to moderately sparse rugoreticulation; katepisternum indistinctly foveolate; anepisternum with thick rugoreticulation. Pilosity moderately dense, long, and erect (Fig. 20D, F). Petiole. Shiny; peduncle finely foveolate, without horizontal lobes on its basal part; node smooth, moderately high, with convex apex, in rear view node dorsoventrally concave; pilosity moderately dense and erect (Fig. 20D, F). Postpetiole. Finely foveolate and shiny; in dorsal view sides with moderately long, acute, and triangular projections; pilosity moderately long, moderately dense, and erect (Fig. 20D, F). Gaster. Smooth and shiny; pilosity dense, short, and erect (Fig. 20D, F). Colour. Yellow, malar area and lower part of frons dark yellow to bright brown (Fig. 20D, F).

Figure 20. 

Pheidole typhlos sp. nov., full-face view (A), profile (C), and dorsal view (E) of paratype minor worker (CASENT0923217) and full-face view (B), profile (D), and dorsal view (F) of holotype major worker (CASENT0302953).

Minor workers. Measurements (N = 1): HL: 0.55; HW: 0.46; SL: 0.45; EL: 0.05; WL: 0.62; PSL: 0.09; MTL: 0.35; PNW: 0.32; PTW: 0.09; PPW: 0.14; CI: 83.9; SI: 97.4; PSLI: 16.4; PPI: 61.2; PNI: 69.8; MTI: 76.7. Head. In full-face view rectangular, posterior and anterior of eyes slightly convex, occipital margin straight or indistinctly concave; occipital carina absent (Fig. 20A). Pilosity moderately dense, short, subdecumbent to suberect. Whole head shiny and smooth, only lateral sides of frons with short, indistinct, longitudinal rugulae; antennal sockets with sparse carinae curved outward. Clypeus with median longitudinal carina absent; two lateral longitudinal carinae absent. Scape, when laid back, reaching posterior head margin; pilosity suberect to erect (Fig. 20A, C). Mesosoma. In lateral view, promesonotum box-like; promesonotal groove absent; metanotal groove distinct; propodeal spines short and triangular, apex acute (Fig. 20C). Sculpture smooth and shiny, sometimes lateral sides with indistinct, irregular, and sparse rugae. Pilosity moderately sparse, short, and erect (Fig. 20C, E). Petiole. Peduncle short with ventral face slightly convex; node globular; with few short, erect setae (Fig. 20C, 20E). Postpetiole. Short, low, and convex; with few short, erect setae (Fig. 20C, E). Gaster. Pilosity moderately dense, erect, and short (Fig. 20C, E). Colour. Unicolourous, yellow (Fig. 20C, E).

Greek for blind [τυφλός], in reference to the reduced eyes.

The species was collected at elevation between 520–980 m, in rainforest, in montane forest. Nesting preferences unknown.

This species is most similar to P. parviocula sp. nov. Major workers. Pheidole typhlos sp. nov. can be distinguished from P. parviocula sp. nov. by sparse and short pilosity on sides of the head, presence of distinct, triangular inner hypostomal teeth and presence of short, triangular propodeal spines. Minor workers. Pheidole typhlos sp. nov. can be distinguished from P. parviocula sp. nov. by box-like promesonotum, absence of promesonotal groove and presence of short and triangular propodeal spines.

Pheidole longispinosa Forel, 1891: 170, pl. 5, fig. 4 (s.w.q.). Lectotype [designated here]: major worker (top specimen, CASENT0101682): Madagascar, Antananarivo, Ambatomanjaka, Miarinarivo (MHNG) [examined]. Paralectotypes: 1 major worker (CASENT0810542, bottom specimen, the same pin as lectotype) (MHNG) [examined], 3 minor workers (1 pin, CASENT0101598, CASENT0923202, CASENT0923203) (MHNG) [examined], 1 dealate queen (CASENT0101657) (MHNG) [examined]: the same data as lectotype.

Madagascar. – Antananarivo: •12w., 2s.; 3 km 41°NE Andranomay, 11.5 km 147°SSE Anjozorobe; -18.47333, 47.96; alt. 1300 m; 5 Dec 2000; B.L. Fisher et al. leg.; CASENT0406929, CASENT0406930, CASENT0413533, CASENT0413537, CASENT0427722, CASENT0427724, CASENT0427725, CASENT0427733. •1 w.; Ambatolaona; -18.928, 47.88283; alt. 1382 m; 19 Feb 2007; B.L. Fisher et al. leg.; CASENT0120674 (CASC). •4w., 1s, 1q.; Mandraka; -18.91813, 47.91717; alt. 1312 m; 20 Mar 2014; B.L. Fisher et al. leg.; CASENT0377078, CASENT0378205, CASENT0378163, CASENT0378568 (CASC). •4w, 1m.; Mandraka Park; -18.9019, 47.90786; 1360 m; 11 Mar 2012; B.L. Fisher et al. leg.; CASENT0275803, CASENT0275804, CASENT0275863 (CASC). •11w., 5s., 1m., 1q; Reg. Analamanga, St. Forestière Mandraka; -18.9183, 47.91687; alt. 1285 m; 25 Mar 2015; B.L. Fisher et al. leg.; CASENT0390404-CASENT0390406, CASENT0390415, CASENT0390429, CASENT0390496, CASENT0390497, CASENT0390506, CASENT0390533, CASENT0720697, CASENT0720833 (CASC); •3w., 1s.; Tsinjoarivo forest, Ankadivory; -19.71572, 47.82084; alt. 1385 m; 23 Aug 2014; B.L. Fisher et al. leg.; CASENT0389650, CASENT0389655, CASENT0389682 (CASC). – Antsiranana: •1 w.; 9.2 km WSW Befingotra, Rés. Anjanaharibe-Sud; -14.75, 49.46667; alt. 1280 m; 5 Nov 1994; B.L. Fisher leg.; CASENT0198233 (CASC). •3w., 1s., 2q; Betaolana Forest, along Bekona River; -14.52996, 49.44039; alt. 880 m; 4 Mar 2009; B.L. Fisher et al. leg.; CASENT0152375, CASENT0152403, CASENT0152410 (CASC).•3w.; Parc National de Marojejy, Manantenina River, 27.6 km 35°NE Andapa, 9.6 km 327°NNW Manantenina; -14.435, 49.76; alt. 775 m; 17 Nov 2003; B.L. Fisher leg.; CASENT0487914 (CASC). – Fianarantsoa: •5w., 2s.; 2 km W Andrambovato, along river Tatamaly; -21.51167, 47.41; alt. 1075 m; 3 Jun 2005; B.L. Fisher et al. leg.; CASENT0060852, CASENT0060853, CASENT0060932, CASENT0060933, CASENT0061669 (CASC). •1w.; 38 km S Ambalavao, Rés. Andringitra; -22.2, 46.96667; alt. 1680 m; 23 Oct 1993; B.L. Fisher leg.; CASENT0198397 (CASC). •3w., 1q.; 40 km S Ambalavao, Rés. Andringitra; -22.21667, 46.96667; alt. 1275 m; 19 Oct 1993; B.L. Fisher leg.; CASENT0198376, CASENT0198377 (CASC). •7w, 1s., 1q.; 8.0 km NE Ivohibe; -22.42167, 46.89833; alt. 1200 m; 3 Nov 1997; B.L. Fisher leg.; CASENT0198369, CASENT0198371, CASENT0198803, CASENT0198868 (CASC). •4w., 3s., 1q.; Parc National Befotaka-Midongy, Papango 28.5 km S Midongy-Sud, Mount Papango; -23.84083, 46.9575; alt. 1250 m; 17 Nov 2006; B.L. Fisher et al. leg.; CASENT0118416, CASENT0119618, CASENT0119592, CASENT0118442 (CASC). •7w., 1s., 1q.; R.S. Ivohibe 8.0 km E Ivohibe; -22.48333, 46.96833; alt. 1200 m; 15 Oct 1997; B.L. Fisher leg.; CASENT0196901, CASENT0198003, CASENT0198004, CASENT0198372, CASENT0198805, CASENT0198807, CASENT0198867 (CASC). – Toamasina: •1w.; [Morano-Chrome forêt, 25 km W]; Morarano Chrome; Amparafaravola; -17.75, 47.98333; alt. 1276 m; 15 Jun 1991; A. Pauly leg.; CASENT0198378 (CASC). •1w., 1s., 1q.; 6.9 km NE Ambanizana, Ambohitsitondroina; -15.56667, 50; alt. 1000 m; 9 Dec 1993; B.L. Fisher leg.; CASENT0198379 (CASC). •1w.; Ambanizana, Parc National Masoala; -15.57167, 50.00611; alt. 800–897 m; 26 Feb 2003; Andriamalala D. et al. leg.; CASENT0047728 (CASC). •8w.; Ankerana; -18.40636, 48.80254; alt. 1108 m; 19 Jan 2012; B.L. Fisher et al. leg.; CASENT0273728, CASENT0274171, CASENT0274172, CASENT0274173, CASENT0274993, CASENT0274994, CASENT0274994 (CASC). •1w., 1s.; Corridor Forestier Analamay-Mantadia, Tsaravoniana; -18.76124, 48.42134; alt. 939 m; 2 Dec 2012; B.L. Fisher et al. leg.; CASENT0300341 (CASC). •3w., 2s.; F.C. Didy; -18.19833, 48.57833; alt. 960 m; 16 Dec 1998; Ratsirarson H.J. leg.; CASENT0198005, CASENT0198006, CASENT0198373 (CASC). •2w., 1s.; Montagne d’Anjanaharibe, 19.5 km 27°NNE Ambinanitelo; -15.17833, 49.635; alt. 1100 m; 12 Mar 2003; B.L. Fisher et al. leg.; CASENT0037455, CASENT0037462, CASENT0048698, CASENT0048727 (CASC). •1w.; P.N. Mantadia; -18.79167, 48.42667; alt. 895 m; 25 Nov 1998; Ratsirarson H.J. leg.; CASENT0198384 (CASC). •1w., 1s.; Parc National d´ Andasibe-Mantadia, Forêt de Mantadia, 25.7 km 248° Moramanga; -18.81402, 48.43028; alt. 1040 m; 13 Jul 2006; Raharimalala & Blaimer leg.; CASENT0117442 (CASC). •1w., 1s.; Parc National de Zahamena, Onibe River; -17.75908, 48.85468; alt. 780 m; 21 Feb 2009; B.L. Fisher et al. leg.; CASENT0152053 (CASC). – Toliara: •7w., 1s., 1q.; 13 km NW Enkara, Rés Andohahela; -24.55, 46.8; alt. 1300 m; 2 Dec 1992; B.L. Fisher leg.; CASENT0198374, CASENT0198375, CASENT0198381, CASENT0198382 (CASC). •31w., 14s., 9q.; Anosy Region, Anosyenne Mts, 29.33 km NW Manantenina; -24.13993, 47.07418; alt. 540 m; 21 Feb 2015; B.L. Fisher et al. leg.; CASENT0704727, CASENT0704727, CASENT0704749, CASENT0704780, CASENT0723465, CASENT0704473, CASENT0704200, CASENT0704201, CASENT0704205, CASENT0704206, CASENT0704217, CASENT0704218, CASENT0704226, CASENT0704227, CASENT0704228, CASENT0704229, CASENT0704242, CASENT0704245, CASENT0704273, CASENT0704353, CASENT0704354, CASENT0704355, CASENT0704393, CASENT0704394, CASENT0704867, CASENT0704874, CASENT0704886, CASENT0704887, CASENT0705875, CASENT0705877, CASENT0721001, CASENT0721005, CASENT0721007, CASENT0721009, CASENT0721013 (CASC). •20w., 2s.; Parc National d’Andohahela, Col du Sedro, 3.8 km 113°ESE Mahamavo, 37.6 km 341°NNW Tolagnaro; -24.76389, 46.75167; alt. 900 m; 21 Jan 2002; B.L. Fisher et al. leg.; CASENT0078380, CASENT0430695, CASENT0430759, CASENT0451281, CASENT0451303, CASENT0451309, CASENT0456249, CASENT0460156, CASENT0460158, CASENT0483964, CASENT0484129, CASENT0484130 (CASC). •3w.; Parc National de Kirindy Mite, 16.3 km 127°SE Belo sur Mer; -20.79528, 44.147; alt. 80 m; 6 Dec 2001; B.L. Fisher et al. leg.; CASENT0477205 (CASC). •5w. 2s.,1q.; Réserve Spéciale Kalambatritra, Ambinanitelo; -23.4502, 46.45658; alt. 1325 m; 11 Feb 2009; B.L. Fisher et al. leg.; CASENT0148863, CASENT0148866, CASENT0149937–CASENT0149939 (CASC). •18w., 7s., 3m., 11q.; Réserve Spéciale Kalambatritra, Ampanihy; -23.463, 46.47057; alt. 1269 m; 10 Feb 2009; B.L. Fisher et al. leg.; CASENT0148871, CASENT0148872, CASENT0148875, CASENT0148945, CASENT0148953, CASENT0148964, CASENT0148967, CASENT0148972, CASENT0148983, CASENT0148994, CASENT0148995, CASENT0148996, CASENT0149001, CASENT0149008, CASENT0149801, CASENT0149992, CASENT0150511, CASENT0150535, CASENT0151012, CASENT0151013, CASENT0151019, CASENT0151020, CASENT0152223, CASENT0153227, CASENT0235023 (CASC). •6w., 3s., 2q.; Réserve Spéciale Kalambatritra, Befarara; -23.4178, 46.4478; alt. 1390 m; 7 Feb 2009; B.L. Fisher et al. leg.; CASENT0149598, CASENT0149599, CASENT0149869, CASENT0149871, CASENT0150723 (CASC). •1w., 1s.; Réserve Spéciale Kalambatritra, Betanana; -23.4144, 46.459; alt. 1360 m; 8 Feb 2009; B.L. Fisher et al. leg.; CASENT0148654 (CASC).

Major workers. Large species: HL: 2.62–3.0 (2.77), HW: 2.64–2.92 (2.75), WL: 1.88–2.14 (2.04); propodeal spines very long (PSL: 0.67–0.87 (0.73)); head in full-face view trapezoid, widened posteriorly; sides of the head without pilosity; frons with longitudinal rugae never reaching further than midlength of head; gaster smooth and shiny; body brown to dark brown. Minor workers. Large species: HL: 0.99–1.15 (1.1), HW: 0.98–1.06 (1.0), WL: 1.62–1.76 (1.66); propodeal spines very long (PSL: 0.62–0.76 (0.69)); scape, when laid back, surpassing posterior head margin by more than half its length; mesosoma smooth and shiny, only anepisternum, katepisternum, and propodeum sometimes with indistinct and sparse superficial rugulae; promesonotal groove absent; metanotal groove shallow and wide.

Major workers. Measurements (N = 10): HL: 2.62–3.0 (2.77); HW: 2.64–2.92 (2.75); SL: 1.24–1.35 (1.3); EL: 0.24–0.28 (0.26); WL: 1.88–2.14 (2.04); PSL: 0.67–0.87 (0.73); MTL: 1.39–1.55 (1.46); PNW: 0.8–0.97 (0.89); PTW: 0.23–0.31 (0.27); PPW: 0.68–0.82 (0.75); CI: 94.3–106.4 (99.3); SI: 43.9–50.9 (47.4); PSLI: 24.1–29.0 (26.5); PPI: 30.1–42.3 (35.4); PNI: 30.45–33.38 (32.4); MTI: 49.6–57.6 (53.0). Head. In full-face view trapezoid, widened posteriorly (Fig. 23B). In lateral view sub-oval; ventral and dorsal faces convex; inner hypostomal teeth visible. Sides of the head without pilosity; frons and vertex with few, long, erect setae. Antennal scrobes absent or very indistinct. Occipital lobes, genae, posterior part of frons smooth and shiny; lateral sides of head and genae with sparse, partially fading superficial rugulae; centre of frons shiny with fine, thin, longitudinal rugae reaching at most to midlength of head; malar area with fine, thin, longitudinal rugae. Centre of clypeus smooth and shiny, lateral sides with fine and sparse rugae; median notch present, shallow, and wide; median longitudinal carina absent; lateral longitudinal carinae absent. Scape, when laid back, reaching midlength of head; pilosity suberect to erect (Fig. 23B, D). Inner hypostomal teeth distinct, low, triangular and thick, with rounded apex, closely spaced; outer hypostomal teeth distinct, slightly smaller and thinner than inner hypostomal teeth, lobe-like (Fig. 84Y). Mesosoma. In lateral view, promesonotum low, short, and relatively flat; pronotum slightly convex; dorsal mesonotum concave; posterior mesonotum relatively steep; promesonotal groove absent; metanotal groove absent; propodeal spines very long, massive basally, with acute apex; humeral area laterally either weakly or not produced (Fig. 23D). Surface shiny and smooth with weak, sparse, and irregular rugae on pronotum and lateral sides of propodeum. Pilosity very sparse, short, and decumbent; dorsum with few additional, long, erect setae (Fig. 23D, F). Petiole. Smooth and shiny; peduncle long and thin, superficially rugulose ventrally; node triangular, with rounded apex, in rear view node dorsoventrally depressed; pilosity sparse, short, and erect (Fig. 23D, F). Postpetiole. Smooth and shiny; in dorsal view sides with acute, narrow, moderately short triangular projections; pilosity short to long, sparse, and erect (Fig. 23D, F). Gaster. Smooth and shiny; pilosity sparse, erect, and long (Fig. 23D, F). Colour. Unicolourous, brown to dark brown (Fig. 23D, F).

Figure 23. 

Pheidole longispinosa Forel, full-face view (A), profile (C), and dorsal view (E) of minor worker (CASENT0451282) and full-face view (B), profile (D), and dorsal view (F) of major worker (CASENT0451284).

Minor workers. Measurements (N = 10): HL: 0.99–1.15 (1.1); HW: 0.98–1.06 (1.0); SL: 1.35–1.47 (1.4); EL: 0.14–0.21 (0.18); WL: 1.62–1.76 (1.66); PSL: 0.62–0.76 (0.69); MTL: 1.21–1.29 (1.24); PNW: 0.65–0.7 (0.67); PTW: 0.13–0.17 (0.16); PPW: 0.22–0.27 (0.25); CI: 88.3–99.1 (91.3); SI: 136.7–144.4 (139.7); PSLI: 56.8–67.8 (62.5); PPI: 56.3–66.9 (62.9); PNI: 65.6–68.3 (66.9); MTI: 121.3–128.1 (124.4). Head. Occipital margin convex; occipital carina narrow, weakly developed (Fig. 23A). Pilosity sparse, long, and erect on frons and median part of occiput. Sculpture smooth and shiny on the whole surface, only antennal sockets with sparse carinae curved outward. Clypeus with median longitudinal carina absent; two lateral longitudinal carinae present. Scape, when laid back, surpassing posterior head margin by more than half of its length; pilosity suberect to erect (Fig. 23A, C). Mesosoma. In lateral view, promesonotum low, long, and slightly convex; promesonotal groove absent; metanotal groove indistinct and wide; propodeal spines very long, massive basally, with acute apex (Fig. 23C). Surface smooth and shiny, only anepisternum, katepisternum, and propodeum sometimes with indistinct and sparse superficial rugulae. Pilosity absent or short, very sparse, suberect (Fig. 23C, E). Petiole. Smooth and shiny; peduncle long and thin; node bulge-like with rounded apex; pilosity absent (Fig. 23C, E). Postpetiole. Moderately long, low, and with slightly convex dorsum; with two long, erect setae at the anterior edge (Fig. 23C, E). Gaster. With few long, erect setae (Fig. 23C, E). Colour. Unicolourous, brown to dark brown (Fig. 23C, E).

The species was collected at between 80–1680 m in elevation, in montane rainforest, montane shrubland, thicket, tropical dry forest, and urban areas. Nests were located in rotten logs, tree stumps, soil, and rock crevasses.

This species is most similar to P. mahaboensis sp. nov. and P. praegrandis sp. nov. Major workers. Pheidole longispinosa can be distinguished from P. mahaboensis sp. nov. by absence of metanotal groove, lack of pilosity at the sides of head, and longitudinal rugae never reaching further than midlength of head; from P. praegrandis sp. nov. by brighter body colouration, reduced head sculpture, and not shagreened first gastral tergite and head. Minor workers. Pheidole longispinosa can be distinguished from P. mahaboensis sp. nov. by absence of promesonotal groove, shallow and wide metanotal groove and reduced sculpture on mesosoma; from P. praegrandis sp. nov. by mostly smooth and shiny sculpture of head and mesosoma, which is never rugoreticulate.

Holotype. Madagascar. •1 major worker; Fianarantsoa; Réserve Forestière d’Agnalazaha, Mahabo, 42.9 km 215° Farafangana; -23.19383, 47.723; alt. 20 m; 19 Apr 2006; B.L. Fisher et al. leg.; BLF13900, CASENT0070925 (CASC). Paratypes. Madagascar. •2w. 1m.; same data as for holotype; CASENT0872077, CASENT0070926, CASENT0872197 (CASC).

Madagascar. – Antsiranana: •7w., 4s, 2q., 1m; Masoala National Park; -15.3014, 50.22776; alt. 280 m; 7 Mar 2014; B.L. Fisher et al. leg.; CASENT0376080, CASENT0376081, CASENT0377018, CASENT0377035, CASENT0377036, CASENT0377038, CASENT0377568 (CASC). – Fianarantsoa: •1w., 1s.; 2 km W Andrambovato, along river Tatamaly; -21.51167, 47.41; alt. 1075 m; 3 Jun 2005; B.L. Fisher et al. leg.; CASENT0060947 (CASC). •4w., 2q; 40 km S Ambalavao, Rés. Andringitra; -22.21667, 46.96667; alt. 1275 m; 19 Oct 1993; B.L. Fisher leg.; CASENT0198822, CASENT0198823, CASENT0198815, CASENT0198831 (CASC). •5w., 3s., 2q.; 45 km S Ambalavao; -22.21667, 47.01667; alt. 720 m; 26 Sep 1993; B.L. Fisher leg.; CASENT0198830, CASENT0198832, CASENT0198824, CASENT0198829 (CASC). •14w., 7s., 1q.; 7.6 km 122° Kianjavato, Forêt Classée Vatovavy; -21.4, 47.94; alt. 175 m; 6 Jun 2005; B.L. Fisher et al. leg.; CASENT0059894, CASENT0059895, CASENT0059940, CASENT0059941, CASENT0059944, CASENT0060019, CASENT0060020, CASENT0060166, CASENT0060167, CASENT0060298, CASENT0060495, CASENT0061087, CASENT0061148, CASENT0061233 (CASC). •10w., 1s.; 8.0 km NE Ivohibe; -22.42167, 46.89833; alt. 1200 m; 3 Nov 1997; B.L. Fisher et al. leg.; CASENT0198821, CASENT0009788, CASENT0198231, CASENT0198806, CASENT0198825, CASENT0198871, CASENT0198874 (CASC). •2w., 1s.; Belle Vue trail, Ranomafana National Park, Fianarantsoa Prov.; -21.2665, 47.42017; alt. 1020 m; 23 May 2002; Harin’Hala leg.; CASENT0079551, CASENT0080544, CASENT0080545, CASENT0080546, CASENT0112227, CASENT0112304, CASENT0112699, CASENT0112768, CASENT0112778, CASENT0113028, CASENT0113154, CASENT0113191, CASENT0113192, CASENT0113227, CASENT0113325, CASENT0113341, CASENT0113369, CASENT0113399, CASENT0113408, CASENT0113411, CASENT0113816, CASENT0113817, CASENT0114099, CASENT0114211, CASENT0114327, CASENT0114751, CASENT0114767 (CASC). •4w.; Fitovavy Fitovinany Region, District of Ifanadiana Belle vue area1200 m S of Ranomafana National Park entrance; -21.2665, 47.42017; alt. 1018 m; 28 May 2003; Rin’ha leg.; CASENT0114066, CASENT0114710, CASENT0112925, CASENT0113635 (CASC). •8w., 6s.; Forêt de Vevembe, 66.6 km 293° Farafangana; -22.791, 47.18183; alt. 600 m; 24 Apr 2006; B.L. Fisher et al. leg.; CASENT0070669, CASENT0070706, CASENT0070718, CASENT0070740, CASENT0070753, CASENT0071145, CASENT0108002, CASENT0108034 (CASC). •1w.; Ifanadiana Pref: Ranomafana S.-Pref: Ranomafana, Talatakely site near R. Namorona; -21.26806, 47.4247; alt. 967 m; 5 Dec 2004; Lees et al. leg.; CASENT0056933 (CASC). •3w.; JIRAMA water works near river, Ranomafana National Park; -21.2485, 47.45217; alt. 690 m, 28 Jan 2002; Harin’Hala leg.; CASENT0114140, CASENT0114293, CASENT0114295 (CASC). •12w., 6s., 1q.; Parc National Befotaka-Midongy, Papango 27.7 km S Midongy-Sud, Mount Papango; -23.83517, 46.96367, alt. 940 m, 14 Nov 2006; B.L. Fisher et al. leg.; CASENT0119097, CASENT0119108, CASENT0119110, CASENT0119111, CASENT0119113, CASENT0119121, CASENT0119424, CASENT0119441, CASENT0125642, CASENT0128545, CASENT0128694, CASENT0128699 (CASC). •1w.; Parc National Befotaka-Midongy, Papango 28.5 km S Midongy-Sud, Mount Papango; -23.84083, 46.9575; alt. 1250 m; 17 Nov 2006; B.L. Fisher et al. leg.; CASENT0128569 (CASC). •129w., 19s., 1q.; Parc National de Ranomafana, Vatoharanana River, 4.1 km 231°SW Ranomafana; -21.29, 47.43333; alt. 1100 m; 27 Mar 2003; Fisher et al. leg.; CASENT0039679, CASENT0039915, CASENT0039916, CASENT0039920, CASENT0039923, CASENT0039930, CASENT0039941, CASENT0039951, CASENT0039954, CASENT0039959, CASENT0039963, CASENT0039967, CASENT0039967, CASENT0039970, CASENT0039978, CASENT0040051, CASENT0040056, CASENT0040058, CASENT0040075, CASENT0040079, CASENT0040084, CASENT0040181, CASENT0040200, CASENT0040215, CASENT0040232, CASENT0040235, CASENT0040238, CASENT0040240, CASENT0040244, CASENT0040251, CASENT0040253, CASENT0040264, CASENT0040270, CASENT0040326, CASENT0040331, CASENT0040336, CASENT0040337, CASENT0040344, CASENT0040350, CASENT0040360, CASENT0040415, CASENT0040421, CASENT0049613, CASENT0050290, CASENT0073552, CASENT0488608, CASENT0488609, CASENT0488627, CASENT0488640, CASENT0488641, CASENT0488642, CASENT0488643, CASENT0488659, CASENT0488662, CASENT0488677, CASENT0488678, CASENT0488679, CASENT0488680, CASENT0497159, CASENT0497161, CASENT0497211–CASENT0497216, CASENT0497327–CASENT0497333, CASENT0497372–CASENT0497377, CASENT0497412–CASENT0497416, CASENT0497428, CASENT0497429, CASENT0497595–CASENT0497598, CASENT0497643–CASENT0497646, CASENT0497685 (CASC). •11w., 1s.; Parc Nationale Ranomafana: Talatakely; -21.24833, 47.42667; 9 Apr 1998; Griswold et al. leg.; CASENT0096608, CASENT0096609, CASENT0096712, CASENT0096713, CASENT0096714, CASENT0096765, CASENT0096770, CASENT0096771, CASENT0096772, CASENT0096774, CASENT0096775, CASENT0097967 (CASC). •3w., 4s.; R.S. Ivohibe 8.0 km E Ivohibe; -22.48333, 46.96833; alt. 1200 m; 15 Oct 1997; B.L. Fisher et al. leg.; CASENT0198804, CASENT0198872 (CASC). •2w., 1s.; R.S. Ivohibe, 7.5 km ENE Ivohibe; -22.47,46.96; alt. 900 m; 7 Oct 1997; B.L. Fisher et al. leg.; CASENT0198232, CASENT0198802, CASENT0198869 (CASC). •21w., 1s.; radio tower, Ranomafana National Park; -21.25833, 47.40717; alt. 1130 m; 14 Jun 2002; Irwin & Harin’Hala leg.; CASENT0052853, CASENT0053113, CASENT0053232, CASENT0053238, CASENT0078772, CASENT0079894, CASENT0080504, CASENT0111901, CASENT0112824, CASENT0112862, CASENT0113134, CASENT0113293, CASENT0113643, CASENT0113751, CASENT0114008, CASENT0114062, CASENT0114517, CASENT0114536, CASENT0114537, CASENT0114642, CASENT0114661 (CASC). •3w.; Ranomafana; -21.25, 47.36667; 1 Mar 1994; A. Pauly leg.; CASENT0096206-CASENT0096208 (CASC). •2w.; Ranomafana National Park, Talatakely area, 0.4 km WSW of Park Entrance; -21.41667, 47.68333; alt. 900 m; 2 Jan 2001; Kavanaugh leg.; CASENT0007643, CASENT0007644 (CASC). •3w.; Ranomafana National Park, Talatakely; Sahambavy; Fianarantsoa Rural; -21.451179, 47.3023894; alt. 1139 m; 30 Oct 1998; Lee & Ribardo leg.; CASENT0198826 (CASC). •1w.; research cabin at Talatakely, Ranomafana National Park; -21.25041, 47.41945; alt. 900 m; 11 Apr 1998; Irwin & Schlinger leg.; CASENT0198817 (CASC). •9w., 4s., 2q.; Réserve Forestière d’Agnalazaha, Mahabo, 42.9 km 215° Farafangana; -23.19383, 47.723; alt. 20 m; 19 Apr 2006; B.L. Fisher et al. leg.; CASENT0070855–CASENT0070858, CASENT0070976, CASENT0070977, CASENT0071584, CASENT0072846, CASENT0072853 (CASC). •17w., 8s., 1q.; Réserve Spéciale Manombo 24.5 km 228° Farafangana; -23.01583, 47.719; alt. 30 m; 21 Apr 2006; B.L. Fisher et al. leg.; CASENT0071522–CASENT0071525, CASENT0071528, CASENT0071547, CASENT0071605, CASENT0071606, CASENT0072089, CASENT0072093, CASENT0072098, CASENT0073134, CASENT0073136, CASENT0073142, CASENT0073153, CASENT0073157, CASENT0108512, CASENT0108514, CASENT0108519, CASENT0108527 (CASC). •13w., 1s.; Vohiparara broken bridge; -21.22617, 47.36983; alt. 1110 m; 25 Jul 2002; Harin’Hala leg.; CASENT0052215, CASENT0078742, CASENT0079658, CASENT0079845, CASENT0111141, CASENT0111855, CASENT0111955, CASENT0111993, CASENT0112046, CASENT0112070, CASENT0112157, CASENT0112176, CASENT0112606, CASENT0113065 (CASC). – Toamasina: •2w., 1s.; 6.2 km SSE Ambanizana, Be Dinta; -15.66667, 49.99806; alt. 600 m; 20 Nov 1993; B.L. Fisher leg.; CASENT0198816 (CASC). •2w., 1s., 1q.; Analalava, 7.0 km 255° Mahavelona; -17.7095, 49.454; alt. 50 m; 27 Nov 2005; B.L. Fisher et al. leg.; CASENT0067391, CASENT0067392 (CASC). •1w.; F.C. Andriantantely; -18.695, 48.81333; alt. 530 m; 4 Dec 1998; Ratsirarson leg.; CASENT0198230 (CASC). •2w.; F.C. Sandranantitra; -18.04833, 49.09167; alt. 450 m; 21 Jan 1999; Ratsirarson leg.; CASENT0198808 (CASC). •2w., 2s.; Mahavelona (Foulpointe); -17.66667, 49.5; 25 Dec 1993; A. Pauly leg.; CASENT0096124, CASENT0096181, CASENT0096365, CASENT0096366 (CASC). •1w., 1s., 1 q.; Manakambahiny, near Vavatenina Forest; -17.46667, 49.35; 9 Feb 1995; A. Pauly leg.; CASENT0094829, CASENT0095095, CASENT0095102 (CASC). •3q.; Montagne d’Anjanaharibe, 18.0 km 21°NNE Ambinanitelo; -15.18833, 49.615; alt. 470 m; 8 Mar 2003; Fisher et al. leg.; CASENT0495177 (CASC). •1w.; Parc National de Zahamena, Onibe River; -17.75908, 48.85468; alt. 780 m; 21 Feb 2009; B.L. Fisher et al. leg.; CASENT0152016 (CASC). •6w., 4s.; Parc National Mananara-Nord, 7.1 km 261° Antanambe; -16.455, 49.7875; alt. 225 m; 15 Nov 2005; B.L. Fisher et al. leg.; CASENT0067356, CASENT0067462, CASENT0067464, CASENT0067473, CASENT0067601, CASENT0071327 (CASC). •4w., 2s.; Reserve Betampona, Camp Rendrirendry 34.1 km 332° Toamasina; -17.924, 49.19967; alt. 390 m; 30 Nov 2005; B.L. Fisher et al. leg.; CASENT0067806, CASENT0068019, CASENT0071898, CASENT0072529 (CASC). •40w., 7s., 5q.; Réserve Spéciale Ambatovaky, Sandrangato River; -16.81745, 49.2925; alt. 400 m; 26 Feb 2010 B.L. Fisher et al. leg.; CASENT0160439, CASENT0160440, CASENT0161071, CASENT0161072, CASENT0161084, CASENT0161085, CASENT0161474, CASENT0161475, CASENT0161806-CASENT0161808; CASENT0161881, CASENT0161953, CASENT0161954, CASENT0162120, CASENT0162121, CASENT0162670, CASENT0163018, CASENT0163071, CASENT0163072, CASENT0163114, CASENT0163138, CASENT0163160, CASENT0163552, CASENT0163614, CASENT0163667, CASENT0163871, CASENT0163941, CASENT0163985, CASENT0164288, CASENT0164294, CASENT0164299, CASENT0164304, CASENT0164319, CASENT0164433, CASENT0164441, CASENT0164456, CASENT0164469, CASENT0164487, CASENT0164496 (CASC). – Toliara: •1w., 1s.; 10 km NW Enakara, Rés. Andohahela; -24.56667, 46.81667; alt. 430 m; 22 Nov 1992; B.L. Fisher leg.; CASENT0198809 (CASC). •4w., 1s.; 10 km NW Enakara, Rés. Andohahela; -24.56667, 46.81667; alt. 425 m; 24 Nov 1992; B.L. Fisher leg.; CASENT0198811, CASENT0709102 (CASC). •4w., 1s.; 11 km NW Enakara, Rés. Andohahela; -24.56667, 46.83333; alt. 800 m; 20 Nov 1992; B.L. Fisher leg.; CASENT0198812, CASENT0198813 (CASC). •2w.; 13 km NW Enakara, Rés. Andohahela; -24.55, 46.8; alt. 1250 m; 30 Nov 1992; B.L. Fisher leg.; CASENT0198380, CASENT0198814 (CASC). •6w., 2s., 1q.; 2.7 km WNW 302° Ste. Luce; -24.77167, 47.17167; alt. 20 m; 9 Dec 1998; B.L. Fisher et al. leg.; CASENT0198818–CASENT0198820, CASENT0198870 (CASC). •12w., 4s., 2m.; Anosy Region, Anosyenne Mts, 29.33 km NW Manantenina; -24.13993, 47.07418; alt. 540 m; 21 Feb 2015; B.L. Fisher et al. leg.; CASENT0704710, CASENT0705814, CASENT0723419, CASENT0723782, CASENT0723788, CASENT0723798, CASENT0723801, CASENT0723811, CASENT0723833, CASENT0704455, CASENT0704456, CASENT0704499, CASENT0704520 (CASC). •2w., 1s., 1m.; Anosy Region, Parc National Andohahela, Col de Tanatana; -24.7585, 46.85367; alt. 275 m; 7 Mar 2015; B.L. Fisher et al. leg.; CASENT0724259, CASENT0724260 (CASC). •5w., 1s.; Cul du Marosohy; -24.55, 46.83333; alt. 600 m; 14 Nov 1992; B.L. Fisher leg.; CASENT0198810, CASENT0198827 (CASC). •8w., 6s., 1q.; Forêt Ivohibe 55.0 km N Tolagnaro; -24.569, 47.204; alt. 200 m; 3 Dec 2006; B.L. Fisher et al. leg.; CASENT0122618, CASENT0121993, CASENT0122577, CASENT0122635, CASENT0122790, CASENT0122792, CASENT0122793, CASENT0122800 (CASC). •4w., 4s.; Grand Lavasoa, 25.9 km W Tolagnaro; -25.08767, 46.749; alt. 450 m; 30 Nov 2006; B.L. Fisher et al. leg.; CASENT0122875, CASENT0122896, CASENT0122907, CASENT0122908 (CASC). •1w., 1q.; Manatantely, 8.9 km NW Tolagnaro; -24.9815, 46.92567; alt. 100 m; 27 Nov 2006; B.L. Fisher et al. leg.; CASENT0125791, CASENT0125793 (CASC). •3w., 2s.; Parc National Andohahela, Col de Tanatana, 33.3 km NW Tolagnaro; -24.7585, 46.85367; alt. 275 m; 22 Nov 2006; B.L. Fisher et al. leg. CASENT0121898, CASENT0121900, CASENT0121917 (CASC).

Major workers. Large species: HL: 2.38–2.71 (2.56), HW: 2.34–2.61 (2.48), WL: 1.78–1.96 (1.83); propodeal spines very long (PSL: 0.64–0.75 (0.7)); head in full-face view trapezoid, widened posteriorly; sides of the head with very sparse, short, decumbent pilosity; frons with longitudinal rugae reaching further than midlength of head; gaster smooth and shiny; body reddish brown to dark brown. Minor workers. Large species: 0.93–1.08 (1.0), HW: 0.79–0.93 (0.88), WL: 1.2–1.5 (1.4); propodeal spines very long (PSL: 0.53–0.66 (0.61)); scape, when laid back, surpassing posterior head margin by more than half its length; pronotum and mesonotal dorsum shiny, smooth or with indistinct, sparse, superficial rugulae; anepisternum, katepisternum and propodeum shiny, with slightly denser superficial rugulae; promesonotal groove present; metanotal groove narrow and relatively deep.

Major workers. Measurements (N = 10): HL: 2.38–2.71 (2.56); HW: 2.34–2.61 (2.48); SL: 1.1–1.24 (1.16); EL: 0.22–0.28 (0.25); WL: 1.78–1.96 (1.83); PSL: 0.64–0.75 (0.7); MTL: 1.21–1.32 (1.27); PNW: 0.74–0.86 (0.8); PTW: 0.21–0.26 (0.24); PPW: 0.73–0.83 (0.77); CI: 93.0–101.0 (96.9); SI: 44.3–49.7 (46.6); PSLI: 25.5–29.6 (27.3); PPI: 27.8–33.7 (30.9); PNI: 30.1–35.6 (32.4); MTI: 49.8–53.5 (51.3). Head. In full-face view trapezoid, widened posteriorly (Fig. 24B). In lateral view sub-oval; ventral and dorsal faces convex; inner hypostomal teeth visible. Sides of the head with very sparse, short, decumbent pilosity; frons and vertex with few additional, long, erect setae. Antennal scrobes absent or very indistinct. Occipital lobes smooth and shiny; genae smooth and shiny or dull, superficially rugulose with median shiny patch widening posteriorly; centre of frons shiny with longitudinal rugae reaching at least midlength of head, sometimes surface between rugae with superficial rugulae; malar area and lateral sides of head superficially rugulose, sometimes with several short, longitudinal rugae; posterior part of frons smooth and shiny or superficially rugulose; head sculpture weakens posteriorly. Centre of clypeus smooth and shiny, lateral sides superficially rugulose; median notch present, shallow, and wide; median longitudinal carina absent; lateral longitudinal carinae absent. Scape, when laid back, reaching midlength of head; pilosity decumbent to subdecumbent (Fig. 24B, D). Inner hypostomal teeth distinct, low, triangular, and thick, with rounded apex, closely spaced; outer hypostomal teeth distinct, slightly smaller and thinner than inner hypostomal teeth, lobe-like (Fig. 85B). Mesosoma. In lateral view, promesonotum low, short, and relatively flat; pronotum slightly convex; mesonotum slightly concave; posterior mesonotum relatively steep; promesonotal groove absent; metanotal groove narrow and relatively deep; propodeal spines very long, massive basally, with acute apex; humeral area laterally weakly or not produced (Fig. 24D). Surface shiny and smooth with weak, sparse, and irregular rugae on pronotum, lateral sides of propodeum, katepisternum, and anepisternum. Pilosity very sparse, short, and decumbent; dorsum with few additional, long erect setae (Fig. 24D, F). Petiole. Smooth and shiny; peduncle long and thin, superficially rugulose ventrally; node triangular with rounded apex, in rear view node dorsoventrally depressed; pilosity sparse, short, and erect (Fig. 24D, F). Postpetiole. Smooth and shiny; in dorsal view sides with acute, narrow, triangular projections; pilosity short to long, sparse, and erect (Fig. 24D, F). Gaster. Pilosity sparse, long, and erect (Fig. 24D, F). Colour. Unicolourous, reddish brown to dark brown (Fig. 24D, F).

Figure 24. 

Pheidole mahaboensis sp. nov., full-face view (A), profile (C), and dorsal view (E) of paratype minor worker (CASENT0070926) and full-face view (B), profile (D), and dorsal view (F) of holotype major worker (CASENT0070925).

Minor workers. Measurements (N = 10): HL: 0.93–1.08 (1.0); HW: 0.79–0.93 (0.88); SL: 1.16–1.32 (1.26); EL: 0.13–0.19 (0.16); WL: 1.2–1.5 (1.4); PSL: 0.53–0.66 (0.61); MTL: 0.96–1.11 (1.06); PNW: 0.53–0.68 (0.6); PTW: 0.12–0.13 (0.13); PPW: 0.19–0.22 (0.21); CI: 85.0–90.1 (87.4); SI: 141.5–146.5 (144.1); PSLI: 57.3–63.2 (60.5); PPI: 55.2–66.8 (61.8); PNI: 62.7–73.1 (68.3); MTI: 116.8–126.5 (120.8). Head. Occipital margin convex; occipital carina narrow, weakly developed (Fig. 24A). Pilosity sparse, short, and appressed on the whole surface and additionally long and erect on frons and median part of occiput. Sculpture smooth and shiny on the whole surface, only antennal sockets with sparse carinae curved outward. Clypeus smooth and shiny; its anterior margin regularly convex, with weakly developed teeth; median longitudinal carina absent; two lateral longitudinal carinae present. Scape, when laid back, surpassing posterior head margin by more than half of its length; pilosity suberect to erect (Fig. 24A, C). Mesosoma. In lateral view, promesonotum low, long, and slightly convex; promesonotal groove indistinct; metanotal groove narrow and relatively deep; propodeal spines very long, massive basally, with acute apex (Fig. 24C). Pronotum and mesonotal dorsum shiny, smooth or with indistinct, sparse, superficial rugulae; anepisternum, katepisternum, and propodeum shiny, with slightly denser superficial rugulae. Pilosity short, very sparse, suberect (Fig. 24C, E). Petiole. Peduncle long and thin; node bulge-like with rounded apex; pilosity absent (Fig. 24C, E). Postpetiole. Moderately long, low, and slightly convex; with two long, erect setae at the anterior edge (Fig. 24C, E). Gaster. With few long, erect setae (Fig. 24C, E). Colour. Unicolourous, brown to dark brown (Fig. 24C, E).

From the type locality.

The species was collected at between 20–1275 m in elevation, in rainforest and once in open area near stream. Nests were located in rotten logs and tree stumps.

This species is most similar to P. longispinosa and P. praegrandis sp. nov. Major workers. Pheidole mahaboensis sp. nov. can be distinguished from P. longispinosa by presence of metanotal groove, presence of very sparse, short, decumbent pilosity at the sides of head, and longitudinal rugae reaching at least midlength of head; from P. praegrandis sp. nov. by presence of metanotal groove, presence of very sparse, short, decumbent pilosity at the sides of head, and not shagreened first gastral tergite and head. Minor workers. Pheidole mahaboensis sp. nov. can be distinguished from P. longispinosa by presence of promesonotal groove, narrow and relatively deep metanotal groove, and presence of fine superficial rugulae on most of mesosoma; from P. praegrandis sp. nov. by mostly smooth and shiny sculpture of head and mesosoma, which is never rugoreticulate.

Holotype. Madagascar. •1 major worker; Antsiranana; Parc National de Marojejy, Manantenina River, 27.6 km 35°NE Andapa, 9.6 km 327°NNW Manantenina; -14.435, 49.76; alt. 775 m; 15 Nov 2003; B.L. Fisher et al. leg.; BLF08889, CASENT0494942, top specimen on the pin (CASC). Paratypes. Madagascar. •9w., 2s.; same data as for holotype; CASENT0494952, CASENT0872086, CASENT0494943-CASENT0494945, CASENT0872223–CASENT0872229 (CASC).

Madagascar. –Antsiranana: •1w., 1s.; Makirovana Forest; -14.16044, 49.95216; alt. 550 m; 1 May 2011; B.L. Fisher et al. leg. CASENT0212471 (CASC). •7w.; Makirovana Forest; -14.17066, 49.95409; alt. 415 m; 29 Apr 2011; B.L. Fisher et al. leg. CASENT0212806, CASENT0231274, CASENT0236088, CASENT0236098 (CASC). •6w., 2s., 1q.; Makirovana Forest; -14.16506, 49.9477; alt. 900 m; 30 Apr 2011; B.L. Fisher et al. leg.; CASENT0231001, CASENT0231002, CASENT0231012, CASENT0231014, CASENT0231043 (CASC). •4w., 2s., 2q.; Makirovana Forest; -14.16666, 49.95, alt. 715 m; 2 May 2011; B.L. Fisher et al. leg.; CASENT0231097, CASENT0231098, CASENT0231113, CASENT0231167 (CASC). •2w., 2s.; Masoala National Park; -15.33058, 50.30279, alt. 250 m; 13 Mar 2014; B.L. Fisher et al. leg.; CASENT0374505, CASENT0377081 (CASC). •21w., 11s., 1q.; Parc National de Marojejy, Manantenina River, 27.6 km 35°NE Andapa, 9.6 km 327°NNW Manantenina; -14.435, 49.76; alt. 775 m; 15 Nov 2003; B.L. Fisher et al. leg.; CASENT0045333, CASENT0045341, CASENT0045357, CASENT0045368, CASENT0045372, CASENT0048887, CASENT0048895, CASENT0235122, CASENT0487704, CASENT0487706, CASENT0487825, CASENT0487903, CASENT0494780, CASENT0494781, CASENT0077076, CASENT0077079, CASENT0077119, CASENT0077122 (CASC). –Fianarantsoa: •4w.; Forêt d’Ambalagoavy Nord, Ikongo, Ambatombe; -21.857068, 47.37849; alt. 625 m; 1 Dec 2000; Harin’Hala & Irwin leg.; CASENT0009561, CASENT0009562, CASENT0009566, CASENT0009571 (CASC). –Toamasina: •10w., 8s., 3q.; Montagne d’Anjanaharibe, 18.0 km 21°NNE Ambinanitelo; -15.18833, 49.615; alt. 470 m; 8 Mar 2003; B.L. Fisher et al. leg.; CASENT0495134, CASENT0495141, CASENT0495145, CASENT0495193, CASENT0495220, CASENT0495221, CASENT0495222, CASENT0495393, CASENT0495395, CASENT0495396 (CASC). •4w., 3s.; Reserve Betampona, Camp Vohitsivalana, 37.1 km 338° Toamasina; -17.88667, 49.2025; alt. 520 m; 2 Dec 2005; B.L. Fisher et al. leg.; CASENT0067653, CASENT0067655, CASENT0067915, CASENT0069180 (CASC).

Major workers. Large species: HL: 2.94–3.2 (3.0), HW: 2.68–3.1 (2.8), WL: 2.01–2.24 (2.11); propodeal spines very long (PSL: 0.81–0.96 (0.89)); head in full-face view trapezoid, widened posteriorly; sides of the head without pilosity; frons with fine, dense, longitudinal rugae reaching at most midlength of head; first gastral tergite shagreened, at least on its basal half; body dark brown to black. Minor workers. Large species: HL: 1.15–1.29 (1.2), HW: 1.04–1.14 (1.1), WL: 1.86–2.01 (1.92); propodeal spines very long (PSL: 0.91–1.0 (0.97)); scape, when laid back, surpassing posterior head margin by more than half its length; mesosoma finely rugoreticulate; promesonotal groove absent or very indistinct; metanotal groove shallow and wide.

Major workers. Measurements (N = 10): HL: 2.94–3.2 (3.0); HW: 2.68–3.1 (2.8); SL: 1.36–1.45 (1.39); EL: 0.24–0.3 (0.27); WL: 2.01–2.24 (2.11); PSL: 0.81–0.96 (0.89); MTL: 1.53–1.7 (1.6); PNW: 0.84–1.0 (0.92); PTW: 0.28–0.32 (0.3); PPW: 0.76–0.9 (0.82); CI: 90.1–96.6 (93.4); SI: 47.4–51.0 (49.3); PSLI: 27.4–31.7 (29.5); PPI: 33.1–38.4 (36.5); PNI: 30.1–35.9 (32.6); MTI: 54.9–59.9 (56.5). Head. In full-face view trapezoid, widened posteriorly (Fig. 25B). In lateral view sub-oval; ventral and dorsal faces convex; inner hypostomal teeth visible. Sides of the head without pilosity; frons and vertex with few, long, erect setae. Antennal scrobes absent. Occipital lobes shiny and smooth or partially shagreened; genae smooth and shiny, sometimes partially shagreened; centre of frons shiny with fine, dense, longitudinal rugae reaching at most midlength of head; malar area shagreened with several short, longitudinal rugae; lateral sides of head and posterior part of frons shagreened; head sculpture weakens posteriorly. Centre of clypeus smooth and shiny, lateral sides with longitudinal rugae; median notch present, shallow and wide; median longitudinal carina absent; lateral longitudinal carinae absent. Scape, when laid back, reaching midlength of head; pilosity suberect to erect (Fig. 25B, D). Inner hypostomal teeth distinct, triangular and thick, with rounded apex, closely spaced; outer hypostomal teeth distinct, slightly smaller and thinner than inner hypostomal teeth, lobe-like (Fig. 85S). Mesosoma. In lateral view, promesonotum low, short, and relatively flat; dorsal pronotum slightly convex; dorsal mesonotum slightly concave; posterior mesonotum relatively convex; promesonotal groove absent; metanotal groove absent; propodeal spines very long, massive basally, with acute apex; humeral area laterally weakly or not produced (Fig. 25D). Surface shiny, rugoreticulate, propodeal dorsum with weaker sculpture. Pilosity very sparse, short, and decumbent; dorsum with few additional, long, erect setae (Fig. 25D, F). Petiole. Weakly shagreened; peduncle relatively long and thin; node triangular with rounded apex, in rear view node dorsoventrally depressed; pilosity sparse, short, and erect (Fig. 25D, F). Postpetiole. Weakly shagreened; in dorsal view sides with acute, moderately wide, short, and triangular projections; pilosity short to long, sparse and erect (Fig. 25D, F). Gaster. First gastral tergite shagreened, at least on its basal half; pilosity sparse, long, and erect (Fig. 25D, F). Colour. Unicolourous, dark brown to black (Fig. 25D, F).

Figure 25. 

Pheidole praegrandis sp. nov., full-face view (A), profile (C), and dorsal view (E) of paratype minor worker (CASENT0494945) and full-face view (B), profile (D), and dorsal view (F) of holotype major worker (CASENT0494942).

Minor workers. Measurements (N = 10): HL: 1.15–1.29 (1.2); HW: 1.04–1.14 (1.1); SL: 1.52–1.69 (1.62); EL: 0.17–0.2 (0.19); WL: 1.86–2.01 (1.92); PSL: 0.91–1.0 (0.97); MTL: 1.42–1.54 (1.49); PNW: 0.75–0.81 (0.77); PTW: 0.15–0.2 (0.17); PPW: 0.27–0.35 (0.32); CI: 87.5–92.7 (89.7); SI: 140.8–156.6 (147.6); PSLI: 75.8–84.0 (78.5); PPI: 45.6–60.8 (53.1); PNI: 67.6–72.1 (70.1); MTI: 126.2–143.4 (134.8). Head. Occipital margin convex; occipital carina narrow, weakly developed (Fig. 25A). Pilosity sparse, short, and appressed on the whole surface and additionally long and erect on frons and median part of occiput. Sculpture variable, shiny, smooth to finely rugulose on vertex, centre, and posterior part of frons and genae; lateral sides of head finely rugulose; antennal sockets with sparse carinae curved outward. Clypeus with median longitudinal carina absent; two lateral longitudinal carinae present. Scape, when laid back, surpassing posterior head margin by more than half of its length; pilosity suberect to erect (Fig. 25A, C). Mesosoma. In lateral view, promesonotum low, long, and slightly convex; promesonotal groove absent or very indistinct; metanotal groove shallow and wide; propodeal spines very long, massive basally, with acute apex (Fig. 25C). Surface finely rugoreticulate. Pilosity short, very sparse, suberect (Fig. 25C, E). Petiole. Peduncle long and thin; node triangular with rounded apex; pilosity absent (Fig. 25C, E). Postpetiole. Moderately short, low, and slightly convex; with two long, erect setae at the anterior edge (Fig. 25C, E). Gaster. With few long, erect setae (Fig. 25C, E). Colour. Unicolourous, dark brown to black (Fig. 25C, E).

Latin for huge, in reference to large body size and very long propodeal spines.

The species was collected between 250–900 m in elevation, in rainforest and montane rainforest. Nests were located in rotten logs and tree stumps.

This species is most similar to P. longispinosa and P. mahaboensis sp. nov. Major workers. Pheidole praegrandis sp. nov. can be distinguished from P. longispinosa by shagreened surface of head and first gastral tergite, and darker body colouration; from P. mahaboensis sp. nov. by absence of metanotal groove, absence of very sparse, short, decumbent pilosity at the sides of head, and shagreened first gastral tergite and head. Minor workers. Pheidole praegrandis sp. nov. can be distinguished from P. longispinosa by rugoreticulate mesosoma sculpture; from P. mahaboensis sp. nov. by rugoreticulate mesosoma sculpture, and shallow and wide metanotal groove.

Pheidole longispinosa scabrata Forel, 1895a: 249 (s.w.). Lectotype [designated here]: major worker (CASENT0101695): Madagascar, Est Imerina, coll. Sikora (MHNG) [examined]. Paralectotypes: 3 minor workers (1 pin, CASENT0101869) (MHNG) [examined], 1 major worker (CASENT0923189) (MHNG) [examined]: the same data as lectotype.

Madagascar. –Antsiranana: •4w., 4q.; 9.2 km WSW Befingotra, Rés. Anjanaharibe-Sud; -14.75, 49.46667; alt. 1200 m; 9 Nov 1994; B.L. Fisher leg.; CASENT0198388, CASENT0198392, CASENT0198395, CASENT0198396 (CASC). •32w., 7s.; Forêt de Binara, 9.4 km 235°SW Daraina; -13.26333, 49.6; alt. 1100 m; 5 Dec 2003; B.L. Fisher leg.; CASENT0043382, CASENT0043383, CASENT0494064–CASENT0494066, CASENT0494103–CASENT0494105, CASENT0494126–CASENT0494128, CASENT0494134–CASENT0494136, CASENT0494152 (CASC). •4w.; R.S. Manongarivo, 14.5 km 220°SW Antanambao; -13.99833, 48.42833; alt. 1175 m; 20 Oct 1998; B.L. Fisher leg.; CASENT0198007, CASENT0198386 (CASC). •3w.; R.S. Manongarivo, 14.5 km 220°SW Antanambao; -14, 48.43167; alt. 1220 m; 18 Oct 1998; B.L. Fisher leg.; CASENT0198387 (CASC). •1w.; R.S. Manongarivo, 14.5 km 220°SW Antanambao; -13.99833, 48.42833; alt. 1175 m; 20 Oct 1998; B.L. Fisher leg.; CASENT0198767 (CASC). –Mahajanga: •2w.; Réserve Spéciale Marotandrano, Marotandrano 48.3km S Mandritsara; -16.28322, 48.81443; alt. 865 m; 6 Dec 2007; B.L. Fisher et al. leg.; CASENT0140632, CASENT0140653 (CASC). –Toamasina: •1w., 1s.; 23 km E Moramanga; -18.98028, 48.45306; alt. 900 m; 10 Nov 1996; B.L. Fisher leg.; CASENT0198385 (CASC). •1w.; Ambatovy, 12.4 km NE Moramanga; -18.85813, 48.28488; alt. 1040 m; 5 Mar 2007; B.L. Fisher et al. leg.; CASENT0124242 (CASC). •2w., 2s.; Bevolota 17.1 km N Andasibe; -18.77071, 48.43164; alt. 995 m; 12 Dec 2007; B.L. Fisher et al. leg.; CASENT0135100, CASENT0235126 (CASC). •2w., 1s.; Forêt Ambatovy, 14.3 km 57° Moramanga; -18.85083, 48.32; alt. 1075 m; 21 Mar 2004; B.L. Fisher et al. leg.; CASENT0047076, CASENT0052866 (CASC). –Toliara: •4w., 3s.; Réserve Spéciale d’Ambohijanahary, Forêt d’Ankazotsihitafototra, 34.6 km 314°NW Ambaravaranala; -18.26, 45.41833; alt. 1100 m; 16 Jan 2003; B.L. Fisher et al. leg.; CASENT0029499, CASENT0029619, CASENT0029620, CASENT0029719–CASENT0029722 (CASC). •22w., 5s.; Réserve Spéciale d’Ambohijanahary, Forêt d’Ankazotsihitafototra, 35.2 km 312°NW Ambaravaranala; -18.26667, 45.40667; alt. 1050 m; 13 Jan 2003; B.L. Fisher et al. leg.; CASENT0028299, CASENT0028313, CASENT0028333, CASENT0485910, CASENT0485911, CASENT0485912, CASENT0485913, CASENT0496690, CASENT0496691, CASENT0496743, CASENT0496744, CASENT0496745, CASENT0496747 (CASC).

Major workers. Large species: HL: 2.44–2.72 (2.56), HW: 2.3–2.58 (2.43), WL: 1.69–1.85 (1.77); propodeal spines very long (PSL: 0.5–0.63 (0.55)); head in full-face view trapezoid, widened posteriorly; sides of the head with sparse, relatively long, erect pilosity; frons with rugoreticulation and additional longitudinal rugae on the whole surface, sculpture weakening posteriorly; first gastral tergite smooth to slightly shagreened; body brown to dark brown; inner hypostomal teeth distinct, triangular and thick, with rounded apex, closely spaced; outer hypostomal teeth distinct, low, lobe-like, with wide base, smaller and thinner than inner hypostomal teeth. Minor workers. Large species: HL: 0.93–1.06 (0.99); HW: 0.9–1.06 (0.96), 1.41–1.65 (1.48); propodeal spines very long (PSL: 0.41–0.48 (0.43)); scape, when laid back, surpassing posterior head margin by one third of its length; head sculpture never rugoreticulate; mesosoma with dense rugoreticulation, sometimes dorsum with weaker sculpture, but never smooth.

Major workers. Measurements (N = 10): HL: 2.44–2.72 (2.56); HW: 2.3–2.58 (2.43); SL: 1.0–1.12 (1.06); EL: 0.23–0.3 (0.27); WL: 1.69–1.85 (1.77); PSL: 0.5–0.63 (0.55); MTL: 1.14–1.29 (1.22); PNW: 0.77–0.94 (0.88); PTW: 0.25–0.29 (0.27); PPW: 0.71–0.89 (0.8); CI: 93.1–96.5 (94.8); SI: 41.3–45.7 (43.5); PSLI: 19.2–24.9 (21.4); PPI: 31.0–38.3 (34.0); PNI: 33.5–38.4 (36.1); MTI: 48.2–52.5 (50.3). Head. In full-face view trapezoid, widened posteriorly (Fig. 26B). In lateral view sub-oval; ventral and dorsal faces convex; inner hypostomal teeth visible. Sides of the head with sparse, relatively long, erect pilosity; whole head with moderately sparse, long erect setae. Antennal scrobes absent. Occipital lobes shiny and smooth or partially rugulose; genae smooth and shiny or with indistinct rugulae; centre of frons shiny, rugoreticulate, longitudinal rugae directed outward posteriorly; malar area rugoreticulate; lateral sides of head shiny, with sparse rugoreticulation; head sculpture weakens posteriorly. Centre of clypeus smooth and shiny, lateral sides with longitudinal rugae; median notch present, shallow and wide; median longitudinal carina absent; lateral longitudinal carinae absent. Scape, when laid back, reaching midlength of head; pilosity suberect to erect (Fig. 26B, D). Inner hypostomal teeth distinct, triangular and thick, with rounded apex, closely spaced; outer hypostomal teeth distinct, low, lobe-like, with wide base, smaller and thinner than inner hypostomal teeth; inner and outer teeth closely spaced and connected by indistinct concavity (Fig. 85W). Mesosoma. In lateral view, promesonotum short, low, and evenly convex, with relatively steep posterior declivity; promesonotal groove absent; metanotal groove present; propodeal spines very long, massive basally, with acute apex; humeral area laterally weakly produced (Fig. 26D). Surface shiny, with sparse to relatively dense rugoreticulation, mesosomal dorsum and propodeum with weaker sculpture, sometimes with smooth patches; katepisternum with smooth area. Pilosity moderately sparse, long, and erect (Fig. 26D, F). Petiole. Smooth to weakly shagreened; peduncle relatively long and thin; node triangular with rounded apex, in rear view node dorsoventrally slightly depressed; pilosity moderately sparse and erect (Fig. 26D, F). Postpetiole. Smooth to weakly shagreened; in dorsal view sides with acute, and short to moderately short angular projections; pilosity long, moderately sparse, and erect (Fig. 26D, F). Gaster. First gastral tergite smooth to slightly shagreened; pilosity moderately sparse, long, and erect (Fig. 26D, F). Colour. Unicolourous, brown to dark brown, sometimes malar area and lower part of frons with brighter colouration than the rest of body (Fig. 26D, F).

Figure 26. 

Pheidole scabrata Forel, full-face view (A), profile (C), and dorsal view (E) of minor worker (CASENT0496745) and full-face view (B), profile (D), and dorsal view (F) of major worker (CASENT0496747).

Minor workers. Measurements (N = 10): HL: 0.93–1.06 (0.99); HW: 0.9–1.06 (0.96); SL: 1.14–1.35 (1.23); EL: 0.17–0.19 (0.18); WL: 1.41–1.65 (1.48); PSL: 0.41–0.48 (0.43); MTL: 1.05–1.21 (1.12); PNW: 0.64–0.72 (0.67); PTW: 0.11–0.16 (0.13); PPW: 0.24–0.37 (0.3); CI: 95.4–100.5 (97.4); SI: 123.8–132.7 (127.5); PSLI: 41.1–45.6 (43.7); PPI: 41.8–51.3 (45.7); PNI: 67.6–73.2 (69.8); MTI: 111.8–119.5 (116.3). Head. Occipital margin convex; occipital carina narrow, weakly developed (Fig. 26A). Pilosity moderately sparse, long, and erect. Sculpture variable, shiny, smooth to finely rugoreticulate on vertex, centre and posterior part of frons; genae smooth; lateral sides of head with few short, longitudinal rugulae, and sometimes finely shagreened; antennal sockets with sparse carinae curved outward. Clypeus with median longitudinal carina absent; two lateral longitudinal carinae present. Scape, when laid back, surpassing posterior head margin by one third of its length; pilosity suberect to erect (Fig. 26A, C). Mesosoma. In lateral view, promesonotum low, long, and slightly convex; promesonotal groove absent or very indistinct; metanotal groove very indistinct; propodeal spines very long, massive basally, with acute apex (Fig. 26C). Surface shiny, with dense rugoreticulation, sometimes dorsum with weaker sculpture, but never smooth. Pilosity sparse, long, and erect (Fig. 26C, E). Petiole. Peduncle long and thin; node triangular with rounded apex; with few long, erect setae (Fig. 26C, E). Postpetiole. Moderately short, low, and indistinctly convex; with few long, erect setae at the anterior edge (Fig. 26C, E). Petiole. With few long, erect setae (Fig. 26C, E). Colour. Unicolourous, brown to dark brown (Fig. 26C, E).

The species was collected at between 825–1220 m in elevation, in montane rainforest, rainforest, transitional humid forest, and tropical dry forest. Nests were located in rotten logs, and in rotten sticks on the ground.

This species is most similar to P. maizina sp. nov. Major workers. Pheidole scabrata can be distinguished from P. maizina sp. nov. by smooth and shiny genae or genae with fine sculpture; smooth area on katepisternum; tips of outer hypostomal teeth not directed outward, never densely shagreened surface of first gastral tergite, and brighter body colouration. Minor workers. Pheidole scabrata can be distinguished from P. maizina sp. nov. by weak head sculpture, and always smooth genae.

Holotype. Madagascar. •1 major worker; Toamasina; Réserve Spéciale Ambatovaky, Sandrangato River; -16.80561, 49.29507; alt. 480 m; 27 Feb 2010; B.L. Fisher et al. leg.; BLF25039, CASENT0162231 (CASC). Paratypes. Madagascar. •2w., 1q.; same data as for holotype; CASENT0162230, CASENT0923223, CASENT0872222 (CASC).

Madagascar. –Antsiranana: •3w., 1s.; 6.5 km SSW Befingotra, Rés. Anjanaharibe-Sud; -14.75, 49.5; alt. 875 m; 19 Oct 1994; B.L. Fisher leg.; CASENT0198389, CASENT0198390 (CASC). •7w., 4s., 1q.; 9.2 km WSW Befingotra, Rés. Anjanaharibe-Sud; -14.75, 49.46667; alt. 1200 m; 8 Nov 1994; B.L. Fisher leg.; CASENT0198391, CASENT0198393, CASENT0198394, CASENT0235113, CASENT0235114 (CASC). –Fianarantsoa: •1w.; Parc National de Ranomafana, Vatoharanana River, 4.1 km 231°SW Ranomafana; -21.29, 47.43333; alt. 1100 m; 27 Mar 2003; B.L. Fisher et al. leg.; CASENT0497619 (CASC). –Mahajanga: •3w., 2s.; Réserve Spéciale Marotandrano, Marotandrano 48.3 km S Mandritsara; -16.28322, 48.81443; alt. 865 m; 7 Dec 2007; B.L. Fisher et al. leg.; CASENT0134242, CASENT0134279, CASENT0235121 (CASC). –Toamasina: •1w.; Ambanizana, Parc National Masoala; -15.57222, 50.00694; alt. 930–1110 m; 2 Mar 2003; Andriamalala et al. leg.; CASENT0073482 (CASC). •1w.; F.C. Andriantantely; -18.695, 48.81333; alt. 530 m; 4 Dec 1998; Ratsirarson leg.; CASENT0198237 (CASC). •19w., 1s.; Montagne d’Akirindro 7.6 km 341°NNW Ambinanitelo; -15.28833, 49.54833; alt. 600 m; 17 Mar 2003; B.L. Fisher et al. leg.; CASENT0039054, CASENT0039064, CASENT0039103, CASENT0039188, CASENT0039200, CASENT0039209, CASENT0039219, CASENT0235138, CASENT0496323, CASENT0496345, CASENT0496346, CASENT0496347 (CASC). •15w., 4s.; Montagne d’Anjanaharibe, 18.0 km 21°NNE Ambinanitelo; -15.18833, 49.615; alt. 470 m; 8 Mar 2003; B.L. Fisher et al. leg.; CASENT0037623, CASENT0037626, CASENT0048951, CASENT0495382, CASENT0495383, CASENT0495384, CASENT0495480, CASENT0495481, CASENT0495482 (CASC). •1w.; Réserve Spéciale Ambatovaky, Sandrangato River; -16.7633, 49.26692; alt. 520 m; 22 Feb 2010; B.L. Fisher et al. leg.; CASENT0160480 (CASC). •2w.; Réserve Spéciale Ambatovaky, Sandrangato River; -16.7702, 49.26638; alt. 470 m; 23 Feb 2010; B.L. Fisher et al. leg.; CASENT0161455 (CASC). •1w.; Réserve Spéciale Ambatovaky, Sandrangato River; -16.8162, 49.29202; alt. 425 m; 25 Feb 2010; B.L. Fisher et al. leg.; CASENT0161854 (CASC). •2w.; Réserve Spéciale Ambatovaky, Sandrangato River; -16.81745, 49.2925; alt. 400 m; 26 Feb 2010; B.L. Fisher et al. leg.; CASENT0162160 (CASC). •4w., 2s., 1q.; Réserve Spéciale Ambatovaky, Sandrangato River; -16.77274, 49.26551; alt. 450 m; 20 Feb 2010; B.L. Fisher et al. leg.; CASENT0162675, CASENT0162686, CASENT0162688, CASENT0162689 (CASC). •4w., 4s., 3q.; Réserve Spéciale Ambatovaky, Sandrangato River; -16.80561, 49.29507; alt. 480 m; 27 Feb 2010; B.L. Fisher et al. leg.; CASENT0163009, CASENT0163019, CASENT0163021, CASENT0163027, CASENT0163032, CASENT0163040, CASENT0163073 (CASC). •2w.; Réserve Spéciale Ambatovaky, Sandrangato River; -16.7633, 49.26692; alt. 520 m; 22 Feb 2010; B.L. Fisher et al. leg.; CASENT0163856, CASENT0163945 (CASC).

Major workers. Large species: HL: 2.38–2.71 (2.55), HW: 2.17–2.54 (2.4), WL: 1.6–1.95 (1.76); propodeal spines very long (PSL: 0.48–0.59 (0.54)); head in full-face view trapezoid, widened posteriorly; sides of the head with sparse, relatively long, erect pilosity; frons with longitudinal, sparse, and directed outward rugae, interspaces with dense and fine rugulae, sculpture weakening posteriorly; first gastral tergite shagreened, at least on its basal part; body dark brown to black; inner hypostomal teeth distinct, triangular and thick, with rounded apex, closely spaced; outer hypostomal teeth distinct, low, lobe-like, with wide base and tops directed outward, smaller and thinner than inner hypostomal teeth. Minor workers. Large species: HL: 0.85–1.03 (0.93), HW: 0.8–1.0 (0.89), WL: 1.21–1.47 (1.36); propodeal spines very long (PSL: 0.34–0.44 (0.39)); scape, when laid back, surpassing posterior head margin by one-third of its length; head sculpture rugoreticulate; mesosoma with dense rugoreticulation, sometimes dorsum with weaker sculpture, but never smooth.

Major workers. Measurements (N = 10): HL: 2.38–2.71 (2.55); HW: 2.17–2.54 (2.4); SL: 0.95–1.17 (1.08); EL: 0.24–0.3 (0.27); WL: 1.6–1.95 (1.76); PSL: 0.48–0.59 (0.54); MTL: 1.16–1.28 (1.23); PNW: 0.8–0.95 (0.9); PTW: 0.24–0.31 (0.28); PPW: 0.7–0.92 (0.83); CI: 91.3–98.5 (94.0); SI: 43.6–46.3 (45.2); PSLI: 19.8–22.6 (21.2); PPI: 31.0–36.0 (34.1); PNI: 35.5–38.8 (37.4); MTI: 49.5–53.8 (51.2). Head. In full-face view trapezoid, widened posteriorly (Fig. 27B). In lateral view sub-oval; ventral and dorsal faces convex; inner hypostomal teeth visible. Sides of the head with sparse, relatively long, erect pilosity; whole head with moderately sparse, long, erect setae. Antennal scrobes absent. Occipital lobes shiny, and smooth at posterior part, anterior part with fine and dense rugulae; genae shiny, with fine and dense rugulae, sometimes posterior part smooth or with reduced sculpture; malar area and frons with longitudinal, sparse, and directed outward rugae, interspaces with dense and fine rugulae; lateral sides of head shiny, with dense and fine rugulae and thin, indistinct, longitudinal rugae. Centre of clypeus smooth and shiny, lateral sides with longitudinal rugae; median notch present, shallow, and wide; median longitudinal carina absent; lateral longitudinal carinae absent. Scape, when laid back, reaching mid-length of head; pilosity suberect to erect (Fig. 27B, D). Inner hypostomal teeth distinct, triangular, and thick, with rounded apex, closely spaced; outer hypostomal teeth distinct, low, lobe-like, with wide base and tops directed outward, smaller and thinner than inner hypostomal teeth (Fig. 85C). Mesosoma. In lateral view, promesonotum short, low, and evenly convex, with relatively steep posterior declivity; promesonotal groove absent; metanotal groove absent; propodeal spines very long, massive basally, with acute apex; humeral area laterally weakly produced (Fig. 27D). Surface shiny, with dense rugoreticulation, mesosomal dorsum and propodeum with weaker sculpture, but never smooth. Pilosity relatively dense, long, and erect (Fig. 27D, F). Petiole. Shagreened; peduncle relatively long and thin; node triangular with rounded apex, in rear view node dorsoventrally slightly depressed; pilosity moderately sparse and erect (Fig. 27D, F). Postpetiole. Shagreened; in dorsal view sides with acute, and short to moderately short angular projections; pilosity long, moderately sparse, and erect (Fig. 27D, F). Petiole. First gastral tergite shagreened, at least on its basal part; pilosity sparse, long and erect (Fig. 27D, F). Colour. Unicolourous, dark brown to black (Fig. 27D, F).

Figure 27. 

Pheidole maizina sp. nov., full-face view (A), profile (C), and dorsal view (E) of paratype minor worker (CASENT0923223) and full-face view (B), profile (D), and dorsal view (F) of holotype major worker (CASENT0162231).

Minor workers. Measurements (N = 10): HL: 0.85–1.03 (0.93); HW: 0.8–1.0 (0.89); SL: 1.07–1.2 (1.16); EL: 0.16–0.21 (0.18); WL: 1.21–1.47 (1.36); PSL: 0.34–0.44 (0.39); MTL: 0.91–1.1 (1.01); PNW: 0.55–0.66 (0.61); PTW: 0.1–0.16 (0.13); PPW: 0.19–0.28 (0.23); CI: 93.3–101.0 (95.5); SI: 122.0–135.6 (130.7); PSLI: 40.0–44.9 (42.3); PPI: 50.5–60.2 (55.3); PNI: 66.3–70.4 (68.7); MTI: 111.2–117.0 (114.1). Head. Occipital margin convex; occipital carina narrow, weakly developed (Fig. 27A). Pilosity sparse, long, and erect. Sculpture variable, finely to strongly rugoreticulate on the whole surface; only centre of frons sometimes with sculpture reduced or smooth; antennal sockets with sparse carinae curved outward. Clypeus with median longitudinal carina absent; two lateral longitudinal carinae present. Scape, when laid back, surpassing posterior head margin by one-third of its length; pilosity suberect to erect (Fig. 27A, C). Mesosoma. In lateral view, promesonotum low, long, and slightly convex; promesonotal groove absent; metanotal groove very indistinct; propodeal spines very long, massive basally, with acute apex (Fig. 27C). Surface shiny, with dense rugoreticulation, sometimes dorsum with weaker sculpture, but never smooth. Pilosity sparse, long, and erect (Fig. 27C, E). Petiole. Finely shagreened; peduncle long and thin; node triangular with rounded apex; with few long, erect setae (Fig. 27C, E). Postpetiole. Moderately shagreened; moderately short, low, and convex; with few long, erect setae at the anterior edge (Fig. 27C, E). Petiole. With few long, erect setae (Fig. 27C, E). Colour. Unicolourous, dark brown to black (Fig. 27C, E).

Malagasy for dark, in reference to dark body colouration.

The species was collected at between 400–1200 m in elevation, in rainforest, montane rainforest, and transitional humid forest. Nests were located in in rotten logs and stumps, and in rotten sticks on the ground.

This species is most similar to P. scabrata. Major workers. Pheidole maizina sp. nov. can be distinguished from P. scabrata by having distinct rugulae on at least half of the genae, tips of outer hypostomal teeth directed outward, never smooth katepisternum, densely shagreened surface of first gastral tergite, and darker body colouration. Minor workers. Pheidole maizina sp. nov. can be distinguished from P. scabrata by rugoreticulate head sculpture with never smooth genae.