Holotype. Madagascar. • 1 major worker; Antsiranana; Parc National de Marojejy, 25.7 km 32°NNE Andapa, 10.3 km 314°NW Manantenina; -14.445, 49.74167; alt. 1575 m; 22 Nov 2003; B. L. Fisher leg.; montane rainforest, ex rotten log; BLF09287; CASENT0923257, top specimen on the pin (CASC). Paratypes. • 6w., 2s.; same data as for holotype, CASENT0487334, CASENT0487336, CASENT0487335 (CASC, MHNG, PBZT).

Figure 19. 

Pheidole alina sp. nov., full-face view (A), profile (C), and dorsal view (E) of paratype minor worker (CASENT0487334) and full-face view (B), profile (D), and dorsal view (F) of holotype major worker (CASENT0923257).

Madagascar. –Antsiranana: • 6w., 3s.; Parc National de Marojejy, 25.4 km 30°NNE Andapa, 10.9 km 311°NW Manantenina; -14.445, 49.735; alt. 2000 m; 23 Nov 2003; B. L. Fisher leg.; montane shrubland, under moss, on ground; BLF09330 (CASC). •1w.; Parc National de Marojejy, 25.7 km 32°NNE Andapa, 10.3 km 314°NW Manantenina; -14.445, 49.74167; alt. 1575 m; 21 Nov 2003; B. L. Fisher leg.; montane rainforest; sifted litter (leaf mold, rotten wood); BLF09242 (CASC). • 6w.; Parc National de Marojejy, 25.7 km 32°NNE Andapa, 10.3 km 314°NW Manantenina; -14.445, 49.74167; alt. 1575 m; 22 Nov 2003; B. L. Fisher leg.; montane rainforest, under moss, above ground; BLF09264 (CASC). • 6w., 3s.; Parc National de Marojejy, 25.7 km 32°NNE Andapa, 10.3 km 314°NW Manantenina; -14.445, 49.74167; alt. 1575 m; 22 Nov 2003; B. L. Fisher leg.; montane rainforest, ex rotten log; BLF09302 (CASC). • 3w.; Parc National de Marojejy, 25.7 km 32°NNE Andapa, 10.3 km 314°NW Manantenina; -14.445, 49.74167; alt. 1575 m; 22 Nov 2003; B. L. Fisher leg.; montane rainforest, ex Melastomataceae; BLF09318 (CASC). • 3w.; Parc National de Marojejy, Antranohofa, 26.6 km 31°NNE Andapa, 10.7 km 318°NW Manantenina; -14.44333, 49.74333; alt. 1325 m; 19 Nov 2003; B. L. Fisher leg.; montane rainforest, under moss, above ground; BLF09142 (CASC). • 1w.; Prov. Antsiranana R.S. Manongarivo 17.3 km 218°SW Antanambao; -14.02167, 48.41833; alt. 1580 m; 27 Oct 1998; B. L. Fisher leg.; montane rainforest, sifted litter (leaf mold, rotten wood); BLF01970 (CASC). • 3w.; Prov. Antsiranana R.S. Manongarivo 17.3 km 218°SW Antanambao; -14.02167, 48.41833; alt. 1580 m; 27 Oct 1998; B. L. Fisher leg.; montane rainforest; BLF01972 (CASC). • 1w.; R.S. Manongarivo, 20.4 km 219°SW Antanambao; -14.04667, 48.40167; alt. 1860 m; 3 Nov 1998; B. L. Fisher leg.; montane rainforest; BLF01989 (CASC). –Mahajanga: • 10w.; Region Sofia, Ampotsidia; -14.40775, 48.70201; alt. 1625 m; 10 Jan 2017; B. L. Fisher leg.; montane rainforest, sifted litter; BLF39733 (CASC).

Moderately large species. Major workers. Head in full-face view sub-oval and slightly widening posteriorly, with anterior and posterior sides slightly convex, in lateral view sub-oval; ventral and dorsal faces convex; sides of the head with moderately dense, long, suberect to erect pilosity; medial part of frons with thick, interrupted, dense, and longitudinal rugae with smooth to indistinctly rugulate interspaces; occipital lobes and area posterolateral from eyes without smooth notches; scape, when laid back, exceeding the midlength of head by one-fifth of its length; inner hypostomal teeth indistinct, low, and wide, closely spaced, bulge-like; outer hypostomal teeth lobe-like, wider and higher than inner teeth, apex directed upward; inner and outer hypostomal teeth closely spaced and not connected by concavity; mesosoma rugofoveolate; pronotum with additional thin, moderately dense, and transverse rugae; katepisternum with reduced sculpture and sometimes with smooth notch; gaster smooth; body black. Minor workers. Head foveolate; median frons with short, indistinct, and longitudinal rugulae; area posterolateral from eyes with slightly to distinctly weaker sculpture; scape, when laid back, exceeding the posterior head margin by one-third of its length; promesonotum low and moderately long; promesonotal groove present; propodeal spines small and triangular; mesosoma foveolate; katepisternum with reduced sculpture or with smooth notch; body black to dark brown.

Major workers. Measurements (N = 9): HL: 1.3–1.79 (1.61); HW: 1.28–1.73 (1.56); SL: 0.88–1.05 (0.97); EL: 0.18–0.22 (0.19); WL: 1.22–1.48 (1.36); PSL: 0.22–0.28 (0.26); MTL: 0.91–1.07 (1.01); PNW: 0.54–0.7 (0.61); PTW: 0.16–0.22 (0.2); PPW: 0.35–0.53 (0.44); CI: 100.7–104.7 (102.9); SI: 57.5–70.4 (62.9); PSLI: 14.9–17.7 (16.1); PPI: 41.6–49.1 (45.1); PNI: 36.3–43.7 (39.4); MTI: 60.3–73.4 (65.4).

Head. In full-face view sub-oval, slightly widening posteriorly, with anterior and posterior sides convex (Fig. 19B). In lateral view sub-oval; ventral and dorsal faces convex; inner hypostomal teeth visible. Sides of the head with moderately dense, long, suberect to erect pilosity; whole head with dense, long, decumbent to erect pilosity. Medial part of frons with thick, interrupted, dense, and longitudinal rugae with smooth to indistinctly rugulate interspaces; lateral sides of frons with denser, thick, and longitudinal rugae, interspaces with dense and moderately thick rugulae. Occipital lobes with longitudinal and thinner rugae and indistinctly rugulate to smooth interspaces. Area posterolateral from eyes with dense, thick, and longitudinal rugae with distinctly rugulate interspaces, sculpture fading posteriorly. Gena with relatively sparse, thick, and longitudinal rugae with indistinctly rugulate interspaces. Centre of clypeus smooth and shiny, lateral sides with indistinct rugulae; median notch present, moderately wide, and shallow; median longitudinal carina present; lateral longitudinal carinae absent. Scape, when laid back, exceeding the midlength of head by one-fifth of its length; pilosity subdecumbent to erect (Fig. 19B, D). Inner hypostomal teeth indistinct, low, and wide, closely spaced, bulge-like; outer hypostomal teeth lobe-like, wider, and higher than inner teeth, apex directed upward; inner and outer hypostomal teeth closely spaced and not connected by concavity (Fig. 63A). Mesosoma. In lateral view, promesonotum short, angular, and moderately low, posterior mesonotum moderately steep, mesonotal process indistinct, tubercle-like; promesonotal groove absent; metanotal groove absent; propodeal spines moderately long, narrow, and with acute apex; humeral area weakly produced (Fig. 19D). Surface shiny and rugofoveolate; pronotum with additional thin, moderately dense, and transverse rugae; katepisternum with reduced sculpture and sometimes with smooth notch. Pilosity moderately dense, long, and erect (Fig. 19D, F). Petiole. Shiny with sparse foveolae; node finely foveolate to smooth, triangular, with rounded and thick apex, in rear view node dorsoventrally slightly convex; pilosity moderately sparse and erect (Fig. 19D, F). Postpetiole. Shiny and foveolate; dorsum with reduced sculpture and smooth notch; in dorsal view oval, lateral margins medially with two wide and dentate projections; pilosity long, moderately sparse, and erect (Fig. 19D, F). Gaster. Shiny and smooth; pilosity moderately sparse, long, and erect (Fig. 19D, F). Colour. Black, gena and posterior parts of lateral sides of promesonotum reddish, gaster, antenna, and legs yellowish (Fig. 19D, F).

Minor workers. Measurements (N = 10): HL: 0.65–0.85 (0.78); HW: 0.5–0.65 (0.61); SL: 0.87–1.07 (1.04); EL: 0.12–0.16 (0.14); WL: 0.89–1.12 (1.07); PSL: 0.11–0.17 (0.15); MTL: 0.72–0.89 (0.86); PNW: 0.37–0.47 (0.44); PTW: 0.08–0.13 (0.11); PPW: 0.15–0.21 (0.18); CI: 123.6–131.1 (127.5); SI: 155.1–172.1 (163.4); PSLI: 16.8–21.0 (19.1); PPI: 45.9–69.9 (58.8); PNI: 69.0–75.0 (71.8); MTI: 136.3–143.7 (139.7).

Head. Cephalic margin slightly convex (Fig. 19A). Pilosity relatively sparse, short, and subdecumbent. Sculpture shiny and foveolate; median frons with short and indistinct longitudinal rugulae; area posterolateral from eyes with slightly to distinctly weaker sculpture; antennal sockets with few indistinct, curved outward rugae and foveolate interspaces. Clypeus with median longitudinal carina absent; two lateral longitudinal carinae absent. Scape, when laid back, exceeding the posterior head margin by one-third of its length; pilosity dense, subdecumbent to erect (Fig. 19A, C). Mesosoma. In lateral view, promesonotum low and moderately long, arched; promesonotal groove indistinct; metanotal groove distinct; propodeal spines small and triangular (Fig. 19C). Sculpture shiny and foveolate; katepisternum with reduced sculpture or with smooth notch. Pilosity sparse, moderately long, and erect (Fig. 19C, E). Gaster. With sparse, erect pilosity (Fig. 19C, E). Colour. Black to dark brown, legs, gaster, and antenna yellowish (Fig. 19C, E).

Malagasy for night in reference to dark body colouration.

The species was collected between 1270–2000 m in elevation, in montane rainforest and montane shrubland. Nests were located in rotten logs, under moss above and on the ground, and in a Melastomataceae twig.

Pheidole alina sp. nov. belongs to the group of taxa characterised by dark body colouration, in major workers from brownish black to black and in minor workers from black (head and mesosoma) to dark brown with body predominantly foveolate (in dark brown specimens entirely). The group consists of three species: P. alina sp. nov., P. trichotos sp. nov., and P. mainty sp. nov. All members of this group are sympatric and their distribution is limited to northernmost parts of the island, predominantly in Antsiranana prefecture. Major workers of Pheidole alina sp. nov. differ from P. trichotos sp. nov. by medial part of frons with thick, interrupted, dense, and longitudinal rugae with smooth to indistinctly rugulate interspaces, base of first gastral tergite smooth and indistinct, bulge-like inner hypostomal teeth; from P. mainty sp. nov. in sides of the head with moderately dense, long, suberect to erect pilosity, indistinct, bulge-like inner hypostomal teeth, and narrow and moderately long propodeal spines. Minor workers of P. alina sp. nov. differ from P. trichotos sp. nov. in larger body size and black body colouration; from P. mainty sp. nov. in never entirely smooth area posterolateral from eyes and distinctly foveolate mesosoma with reduced or no foveolae only on the katepisternum.

Holotype. Madagascar. • 1 major worker; Antananarivo; Ambohimanga; -18.76036, 47.56372; alt. 1294 m; 8 Sep 2015; B.L. Fisher et al. leg.; montane rainforest, ground nest; BLF37302; CASENT0720924 (CASC). Paratypes. • 2w., 1s.; same data as for holotype; CASENT0923279, CASENT0720925 (CASC, MHNG).

Figure 20. 

Pheidole ambohimanga sp. nov., full-face view (A), profile (C), and dorsal view (E) of paratype minor worker (CASENT0923279) and full-face view (B), profile (D), and dorsal view (F) of holotype major worker (CASENT0720924).

Madagascar. –Antananarivo: • 2w., 1s.; Ambohitrangano; -18.72801, 47.57878; alt. 1383 m; 3 Oct 2015; B.L. Fisher et al. leg.; disturbed montane rainforest, under root mat, litter on rock; BLF37326 (CASC). • 1w., 1s.; Station Forestière Angavokely; -18.92207, 47.74157; alt. 1460 m; 9 Mar 2013; B.L. Fisher et al. leg.; rainforest, under root mat, litter on rock; BLF31333 (CASC). • 1w., 1s.; Tsimbazaza; -18.928, 47.527; alt. 1300 m; 16 Dec 2006; B.L. Fisher et al. leg.; park/garden, ex rotten log; BLF16240 (CASC).

Moderately large species. Major workers. Head in full-face view oval, not widening posteriorly, with anterior and posterior sides convex, in lateral view sub-oval, ventral and dorsal faces convex, occipital cleft moderately deep; sides of the head with dense, very long, erect pilosity; medial part of frons with thick, moderately sparse, and irregular rugae, interspaces predominantly smooth and sometimes with indistinct rugulae; lateral sides of head, area posterolateral from eyes and occipital lobes with relatively dense to dense, thick, and irregular rugae; interspaces predominantly smooth or indistinctly rugulate; scape, when laid back, exceeding the midlength of head by two-fifths of its length; inner hypostomal teeth distinct, large, closely spaced, and triangular, with rounded apex directed inward; outer hypostomal teeth lobe-like, slightly wider than inner hypostomal teeth and approximately the same height, apex directed outward; inner and outer hypostomal teeth closely spaced and not connected by concavity; lateral sides of pronotum, mesonotum, katepisternum, anepisternum, and propodeum rugofoveolate, pronotal dorsum with thick to thin, irregular to transverse rugae with smooth to indistinctly rugofoveolate interspaces; katepisternum with smooth notch; body dark brown. Minor workers. Head with sparse foveolae; frons with foveolae reduced medially with moderately thick, longitudinal, and interrupted rugae; area posterolateral from eyes with reduced sculpture, predominantly smooth; scape, when laid back, surpassing the posterior head margin by two-fifths of its length; promesonotum moderately high and short; promesonotal groove absent; propodeal spines minute and triangular; propodeum predominantly smooth, its dorsum with sparse, short, and transverse rugae and indistinct foveolae; mesonotum, anepisternum, katepisternum, and propodeum predominantly smooth, with sparse and indistinct foveolae on anterolateral and anterior sides; body brown.

Major workers. Measurements (N = 4): HL: 0.95–1.03 (0.98); HW: 0.93–1.02 (0.96); SL: 0.61–0.63 (0.62); EL: 0.13–0.13 (0.13); WL: 0.84–0.92 (0.88); PSL: 0.14–0.16 (0.15); MTL: 0.55–0.59 (0.57); PNW: 0.45–0.5 (0.47); PTW: 0.14–0.16 (0.15); PPW: 0.3–0.35 (0.31); CI: 97.5–104.4 (101.5); SI: 62.0–67.0 (64.6); PSLI: 13.2–15.9 (15.0); PPI: 41.1–49.8 (47.2); PNI: 46.2–50.9 (48.8); MTI: 55.6–61.6 (58.9).

Head. In full-face view oval, not widening posteriorly, with anterior and posterior sides convex (Fig. 20B). In lateral view sub-oval; ventral and dorsal faces convex; inner hypostomal teeth visible. Sides of the head with dense, very long, erect pilosity; whole head with dense, long, decumbent to erect pilosity. Medial part of frons with thick, moderately sparse, irregular rugae, interspaces predominantly smooth and sometimes with indistinct rugulae; lateral sides of head, area posterolateral from eyes, and occipital lobes with relatively dense to dense, thick, and irregular rugae; interspaces predominantly smooth or indistinctly rugulate. Gena with relatively dense, thick, and longitudinal rugae and smooth interspaces. Centre of clypeus smooth and shiny, lateral sides with indistinct rugulae; median notch present, moderately wide, and shallow; median longitudinal carina present; lateral longitudinal carinae absent. Scape, when laid back, exceeding the midlength of head by two-fifths of its length; pilosity subdecumbent to erect (Fig. 20B, D). Inner hypostomal teeth distinct, large, closely spaced, triangular, with rounded apex directed inward; outer hypostomal teeth lobe-like, slightly wider than inner hypostomal teeth and approximately the same height, apex directed outward; inner and outer hypostomal teeth closely spaced and not connected by concavity (Fig. 63B). Mesosoma. In lateral view, promesonotum short, angular, and moderately low, posterior mesonotum moderately steep, mesonotal process distinct, tubercle-like; promesonotal groove absent; metanotal groove absent; propodeal spines moderately short, with wide base and acute apex; humeral area laterally weakly produced (Fig. 20D). Surface shiny; lateral sides of pronotum, mesonotum, katepisternum, anepisternum, and propodeum rugofoveolate, pronotal dorsum with thick to thin, irregular to transverse rugae with smooth to indistinctly rugofoveolate interspaces; katepisternum with smooth notch. Pilosity moderately dense, long, and erect (Fig. 20D, F). Petiole. Shiny with fine foveolae; node smooth to finely foveolate, low, triangular, with rounded and thin apex, in rear view node dorsoventrally straight to slightly convex; pilosity moderately sparse and erect (Fig. 20D, F). Postpetiole. Shiny and foveolate; dorsum with reduced sculpture and smooth notch; in dorsal view oval, lateral margins medially with two dentate projections; pilosity long, moderately sparse, and erect (Fig. 20D, F). Gaster. Shiny and smooth; pilosity dense, long, and erect (Fig. 20D, F). Colour. Dark brown; legs and antennae yellow (Fig. 20D, F).

Minor workers. Measurements (N = 7): HL: 0.55–0.6 (0.56); HW: 0.46–0.51 (0.47); SL: 0.58–0.62 (0.6); EL: 0.09–0.11 (0.1); WL: 0.67–0.76 (0.7); PSL: 0.08–0.1 (0.09); MTL: 0.44–0.48 (0.46); PNW: 0.32–0.38 (0.33); PTW: 0.09–0.1 (0.1); PPW: 0.13–0.14 (0.14); CI: 116.0–123.7 (119.4); SI: 119.5–130.0 (126.6); PSLI: 14.4–17.8 (15.7); PPI: 64.7–72.5 (69.0); PNI: 67.9–73.9 (70.0); MTI: 93.6–101.7 (96.6).

Head. Cephalic margin indistinctly concave or straight (Fig. 20A). Pilosity relatively sparse, long, decumbent to suberect. Sculpture shiny with sparse foveolae; frons with foveolae reduced medially with moderately thick, longitudinal, and interrupted rugae; area posterolateral from eyes with reduced sculpture, predominantly smooth; antennal sockets with few thick, curved outward rugae and indistinctly foveolate interspaces. Clypeus with median longitudinal carina absent; two lateral longitudinal carinae absent. Scape, when laid back, surpassing the posterior head margin by two-fifths of its length; pilosity dense, subdecumbent to erect (Fig. 20A, C). Mesosoma. In lateral view, promesonotum moderately high and short, arched; promesonotal groove absent; metanotal groove distinct; propodeal spines minute and triangular (Fig. 20C). Sculpture shiny; propodeum predominantly smooth, its dorsum with sparse, short, transverse rugae and indistinct foveolae; mesonotum, anepisternum, katepisternum, and propodeum predominantly smooth, with sparse and indistinct foveolae on anterolateral and anterior sides. Pilosity moderately dense, long, and erect (Fig. 20C, E). Gaster. With sparse, erect pilosity (Fig. 20C, E). Colour. Brown, legs slightly brighter (Fig. 20C, E).

From the type locality.

The species was collected between 1294–1460 m in elevation, in rainforest, montane rainforest and park. Nests were located under rootmat on rock, in the soil, and in rotten logs.

Pheidole ambohimanga sp. nov. is a member of the group of species characterised by major workers with head in full-face view oval and not widening posteriorly and medial part of frons with thick, moderately sparse, irregular rugae or medial frons with moderately dense, thin, longitudinal anteriorly to irregular posteriorly, interrupted rugae and very shallow occipital cleft. Minor workers of this group have short and moderately high promesonotum and dark body colouration ranging from orange to brown. The group consists of three species: P. vadum sp. nov., P. analavelona sp. nov., and P. ambohimanga sp. nov. Pheidole ambohimanga sp. nov. is sympatric with P. vadum sp. nov. but morphologically is most similar to P. analavelona sp. nov. It can be easily separated from P. vadum sp. nov. based on thicker and exclusively irregular rugae on frons, deeper occipital cleft in majors, and presence of additional rugae on head in minors. Majors of P. ambohimanga sp. nov. can be distinguished from P. analavelona sp. nov. based on smooth to very indistinctly rugoreticulate interspaces on head, while minor workers have head sculpture foveolae strongly reduced to absent on frons, predominantly smooth area posterolateral from eyes, and mesonotum, anepisternum, katepisternum, and propodeum predominantly smooth, with sparse and indistinct foveolae on anterolateral and anterior sides. Minors of P. ambohimanga sp. nov. can be easily separated from other members of the sikorae group based on short and moderately high promesonotum and additional, moderately thick to thick, longitudinal, and interrupted rugae on head.

Holotype. Madagascar. • 1 major worker; Toliara; Forêt Classée d’Analavelona, 33.2 km 344°NNW Mahaboboka; -22.64333, 44.17167; alt. 1300 m; 22 Feb 2003; Fisher et al. leg.; montane rainforest, ex rotten log; BLF07970; CASENT0491849, top specimen on the pin (CASC). Paratypes. • 9w., 6s., 2m.; same data as for holotype; CASENT0491847, CASENT0491846, CASENT0872248, CASENT0491850, CASENT0491845, CASENT0491844, CASENT0491848 (CASC, MHNG, PBZT).

Figure 21. 

Pheidole analavelona sp. nov., full-face view (A), profile (C), and dorsal view (E) of paratype minor worker (CASENT0491846) and full-face view (B), profile (D), and dorsal view (F) of holotype major worker (CASENT0491849).

Madagascar. –Toliara: • 3w.; Forêt Classée d’Analavelona, 29.2 km 343°NNW Mahaboboka; -22.675, 44.19; alt. 1100 m; 18 Feb 2003; Fisher et al. leg.; montane rainforest, ex rotten log; BLF07833 (CASC). • 3w., 3s.; ibid.; BLF07835 (CASC). • 3w.; ibid.; BLF07844 (CASC). • 3w., 3s.; ibid.; BLF07857 (CASC). •3w.; ibid.; BLF07887 (CASC). • 3s.; ibid.; under stone; BLF07853 (CASC). • 9w., 3s.; Forêt Classée d’Analavelona, 33.2 km 344°NNW Mahaboboka; -22.64333, 44.17167; alt. 1300 m; 22 Feb 2003; Fisher et al. leg.; montane rainforest, ex rotten log; BLF07953 (CASC). • 1w., 1s.; ibid.; BLF07966 (CASC). • 3., 3s.; ibid.; BLF07971 (CASC). • 6w., 1s., 1q., 6m.; ibid.; BLF07989 (CASC).

Moderately large species. Major workers. Head in full-face view oval, not widening posteriorly, with anterior and posterior sides convex, in lateral view sub-oval, ventral and dorsal faces convex, occipital cleft moderately deep; sides of the head with dense, very long, suberect to erect pilosity; frons with thick, irregular rugae, moderately dense on lateral sides and sparser on medial part; interspaces shiny with distinct and sparse rugoreticulae; occipital lobes, area posterolateral from eyes never smooth; scape, when laid back, exceeding the midlength of head by one-fifth of its length; inner hypostomal teeth distinct, large, closely spaced, triangular, with rounded apex directed inward; outer hypostomal teeth lobe-like, wider than inner hypostomal teeth and approximately the same height, apex directed outward; inner and outer hypostomal teeth not closely spaced and not connected by concavity; mesosoma with dense rugofoveolae; pronotum, mesonotum, katepisternum, and anepisternum with additional thick to thin, irregular to transverse rugae; body dark brown. Minor workers. Head foveolate; medial frons with foveolae sparser and with moderately thick longitudinal and interrupted rugae; lateral sides of frons and vertex with additional moderately thick rugae; area posterolateral from eyes with reduced sculpture and smooth notches; scape, when laid back, surpassing the posterior head margin by two-fifths of its length; promesonotum moderately high and short; promesonotal groove absent; propodeal spines minute and triangular; propodeum predominantly smooth with indistinct rugofoveolae on lateral sides; mesonotum, anepisternum, katepisternum, and propodeum with sparse and indistinct rugofoveolae; body dark orange.

Major workers. Measurements (N = 10): HL: 0.97–1.02 (1.0); HW: 0.95–1.0 (0.98); SL: 0.64–0.67 (0.66); EL: 0.11–0.13 (0.12); WL: 0.88–0.95 (0.92); PSL: 0.13–0.15 (0.14); MTL: 0.57–0.6 (0.58); PNW: 0.44–0.48 (0.46); PTW: 0.13–0.17 (0.14); PPW: 0.26–0.29 (0.28); CI: 101.0–104.4 (102.0); SI: 65.3–69.3 (67.1); PSLI: 13.1–15.4 (14.1); PPI: 44.8–57.2 (52.2); PNI: 46.2–48.8 (47.4); MTI: 57.0–61.8 (59.3).

Head. In full-face view oval, not widening posteriorly, with anterior and posterior sides convex (Fig. 21B). In lateral view sub-oval; ventral and dorsal faces convex; inner hypostomal teeth visible. Sides of the head with dense, very long, suberect to erect pilosity; whole head with dense, long, decumbent to erect pilosity. Frons with thick, irregular rugae, moderately dense on lateral sides and sparser on medial part; interspaces shiny with distinct and sparse rugoreticulae. Occipital lobes with relatively dense, thick, and irregular rugae; interspaces rugoreticulate. Gena with relatively dense, thick, and longitudinal rugae and rugoreticulate interspaces. Area posterolateral from eyes with moderately dense, thick, and irregular rugae; interspaces shiny with distinct and sparse rugoreticulae. Centre of clypeus smooth and shiny, lateral sides with indistinct rugulae; median notch present, moderately wide, and shallow; median longitudinal carina present; lateral longitudinal carinae absent. Scape, when laid back, exceeding the midlength of head by one-fifth of its length; pilosity subdecumbent to erect (Fig. 21B, D). Inner hypostomal teeth distinct, large, closely spaced, triangular, with rounded apex directed inward; outer hypostomal teeth lobe-like, wider than inner hypostomal teeth and approximately the same height, apex directed outward; inner and outer hypostomal teeth not closely spaced and not connected by concavity (Fig. 63C). Mesosoma. In lateral view, promesonotum short, angular, and moderately low, posterior mesonotum moderately steep, mesonotal process indistinct, tubercle-like; promesonotal groove absent; metanotal groove indistinct; propodeal spines moderately short, with wide base and acute apex; humeral area laterally weakly produced (Fig. 21D). Surface shiny with dense rugofoveolae; pronotum, mesonotum, katepisternum, and anepisternum with additional thick to thin, irregular to transverse rugae. Pilosity moderately dense, long, and erect (Fig. 21D, F). Petiole. Shiny with fine foveolae; node smooth to finely foveolate, low, triangular, with rounded and thin apex, in rear view node dorsoventrally straight to slightly convex; pilosity moderately sparse and erect (Fig. 21D, F). Postpetiole. Shiny and foveolate; dorsum with reduced sculpture and smooth notch; in dorsal view oval, lateral margins medially with two dentate projections; pilosity long, moderately sparse and erect (Fig. 21D, F). Gaster. Shiny and smooth; pilosity dense, long, and erect (Fig. 21D, F). Colour. Dark brown; legs and antennae yellow (Fig. 21D, F).

Minor workers. Measurements (N = 10): HL: 0.55–0.59 (0.57); HW: 0.46–0.48 (0.47); SL: 0.59–0.63 (0.62); EL: 0.09–0.11 (0.1); WL: 0.69–0.73 (0.71); PSL: 0.06–0.09 (0.08); MTL: 0.46–0.49 (0.47); PNW: 0.32–0.35 (0.33); PTW: 0.07–0.1 (0.08); PPW: 0.12–0.14 (0.13); CI: 116.6–124.7 (120.7); SI: 125.9–133.5 (130.5); PSLI: 10.6–15.1 (14.0); PPI: 55.0–81.5 (66.0); PNI: 67.4–72.0 (69.8); MTI: 97.3–102.3 (99.4).

Head. Cephalic margin indistinctly concave or straight (Fig. 21A). Pilosity relatively dense, long, decumbent to suberect. Sculpture foveolate; medial frons with foveolae sparser and with moderately thick, longitudinal, and interrupted rugae; lateral sides of frons and vertex with additional moderately thick rugae; area posterolateral from eyes with reduced sculpture and smooth notches; antennal sockets with few thick, curved outward rugae and foveolate interspaces. Clypeus with median longitudinal carina absent; two lateral longitudinal carinae absent. Scape, when laid back, surpassing the posterior head margin by two-fifths of its length; pilosity dense, subdecumbent to erect (Fig. 21A, C). Mesosoma. In lateral view, promesonotum moderately high and short, arched; promesonotal groove absent; metanotal groove distinct; propodeal spines minute, triangular (Fig. 21C). Sculpture shiny; propodeum predominantly smooth with indistinct rugofoveolae on lateral sides; mesonotum, anepisternum, katepisternum, and propodeum with sparse and indistinct rugofoveolae. Pilosity moderately dense, long, and erect (Fig. 21C, E). Gaster. With sparse, erect pilosity (Fig. 21C, E). Colour. Dark orange, gaster slightly darker (Fig. 21C, E).

From the type locality.

The species was collected between 1100–1300 m in elevation, in montane rainforest. Nests were located in rotten logs and under stones.

Pheidole analavelona sp. nov. is a member of the group of species characterised by major workers with head in full-face view oval and not widening posteriorly and medial part of frons with thick, moderately sparse, and irregular rugae or medial frons with moderately dense, thin, longitudinal anteriorly to irregular posteriorly, and interrupted rugae and very shallow occipital cleft. Minor workers of this group have short and moderately high promesonotum and dark body colouration ranging from orange to brown. The group consists of three species: P. vadum sp. nov., P. analavelona sp. nov., and P. ambohimanga sp. nov. Pheidole analavelona sp. nov. is known from Forêt Classée d’Analavelona in Toliara and its distribution does not overlap with that of the two remaining members of the group known from the vicinity of Antananarivo. Morphologically both minor and major workers of P. analavelona sp. nov. are most similar to P. ambohimanga sp. nov. Majors of P. analavelona sp. nov. can be distinguished based on distinctly rugoreticulate interspaces on head while minor workers have head sculpture foveolate with foveolae sparser on frons and area posterolateral from eyes, and sparse but never absent rugofoveolae on mesonotum, anepisternum, katepisternum, and propodeum. Minors of P. analavelona sp. nov. can be easily separated from other members of the sikorae group based on short and moderately high promesonotum and additional, moderately thick to thick, longitudinal, and interrupted rugae on head.

Holotype. Madagascar. • 1 major worker; Toliara; Anosy Region, Parc National Andohahela, Col de Tanatana; -24.74969, 46.84949; alt. 400 m; 9 Mar 2015; B. L. Fisher et al. leg.; rainforest, ex rotten log; BLF36798; CASENT0700917 (CASC). Paratype. • 1w.; same data as for holotype, CASENT0923281 (CASC).

Figure 22. 

Pheidole andohahela sp. nov., full-face view (A), profile (C), and dorsal view (E) of paratype minor worker (CASENT0923281) and full-face view (B), profile (D), and dorsal view (F) of holotype major worker (CASENT0700917).

Madagascar. –Toliara: • 1w., 1s.; Anosy Region, Parc National Andohahela, Col de Tanatana; -24.74969, 46.84949; alt. 400 m; 9 Mar 2015; B. L. Fisher et al. leg.; rainforest, ex rotten log; BLF36803 (CASC).

Minute species. Major workers. HL < 0.95 mm and WL < 0.8 mm; head in full-face view sub-oval; body dark brown; sides of head with very dense, relatively short, suberect pilosity; entire head distinctly sculptured; scape, when laid back, exceeding the midlength of head by two-fifths of its length; inner hypostomal teeth distinct, large, with sharp apex; outer hypostomal teeth lobe-like with apex directed distinctly outward; inner and outer hypostomal teeth not closely spaced and not connected by concavity. Minor workers. HL < 0.5 mm and WL < 0.6 mm, scape, when laid back, surpassing the posterior head margin by two-fifths of its length, propodeal spines reduced to small lobes, head elongate and oval and body dark brown; head sculpture shiny and predominantly smooth, vertex and frons with sparse, short, and transverse rugulae; anterolateral sides of propodeum and anepisternum with indistinct and sparse rugoreticulae.

Major workers. Measurements (N = 2): HL: 0.87, 0.92; HW: 0.87, 0.92; SL: 0.64, 0.63; EL: 0.12, 0.11; WL: 0.77, 0.79; PSL: 0.1, 0.11; MTL: 0.53, 0.56; PNW: 0.36, 0.39; PTW: 0.11, 0.12; PPW: 0.28, 0.29; CI: 100.0, 99.9; SI: 72.9, 67.8; PSLI: 11.9, 12.4; PPI: 40.1, 40.7; PNI: 41.3, 42.0; MTI: 60.8, 60.5.

Head. In full-face view sub-oval, slightly widening posteriorly, with anterior and posterior sides convex (Fig. 22B). In lateral view sub-oval; ventral and dorsal faces convex; inner hypostomal teeth visible. Sides of the head with very dense, relatively short, suberect pilosity; whole head with dense, long, decumbent to erect pilosity. Medial part of frons with thick, dense, longitudinal, and interrupted rugae, interspaces smooth; lateral sides with thick, dense, and irregular rugae, interspaces shiny with dense and distinct rugoreticulae. Occipital lobes with thick, irregular rugae and smooth to rugoreticulate interspaces. Area posterolateral from eyes rugoreticulate to rugofoveolate with additional longitudinal and thick rugae. Gena with relatively dense, thick, and longitudinal rugae and rugoreticulate interspaces. Centre of clypeus smooth and shiny, lateral sides with indistinct rugulae; median notch present, moderately wide, and shallow; median longitudinal carina present; lateral longitudinal carinae absent. Scape, when laid back, exceeding the midlength of head by two-fifths of its length; pilosity subdecumbent to erect (Fig. B, D). Inner hypostomal teeth distinct, large, closely spaced, triangular, with sharp apex directed upward; outer hypostomal teeth lobe-like, more narrow and lower than inner hypostomal teeth, apex directed outward; inner and outer hypostomal teeth not closely spaced and not connected by concavity (Fig. 63D). Mesosoma. In lateral view, promesonotum short, angular, and moderately low, posterior mesonotum moderately steep, mesonotal process indistinct, tubercle-like; promesonotal groove absent; metanotal groove absent; propodeal spines short, with moderately wide base and acute apex; humeral area laterally weakly produced (Fig. 22D). Surface shiny with fine rugofoveolae; anterolateral sides of pronotum with smooth notches. Pilosity moderately dense, long, and erect (Fig. 22D, F). Petiole. Shiny with fine foveolae;node smooth to finely foveolate, low, triangular, with rounded and thin apex, in rear view node dorsoventrally straight to slightly convex; pilosity moderately sparse and erect (Fig. 22D, F). Postpetiole. Shiny and foveolate; dorsum with reduced sculpture and smooth notch; in dorsal view oval, lateral margins medially with two dentate projections; pilosity long, moderately sparse, and erect (Fig. 22D, F). Gaster. Shiny and smooth; pilosity dense, long, and erect (Fig. D, F). Colour. Dark brown; legs, gaster and antennae yellowish brown (Fig. 22D, F).

Minor workers. Measurements (N = 2): HL: 0.48, 0.49; HW: 0.39, 0.43; SL: 0.55, 0.57; EL: 0.09, 0.09; WL: 0.58, 0.58; PSL: 0.05, 0.05; MTL: 0.4, 0.42; PNW: 0.28, 0.28; PTW: 0.07, 0.08; PPW: 0.12, 0.14; CI: 122.4, 115.0; SI: 140.3, 132.8; PSLI: 10.8, 9.4; PPI: 55.6, 55.6; PNI: 70.9, 66.0; MTI: 102.0, 98.4.

Head. Cephalic margin indistinctly convex or straight (Fig. 22A). Pilosity relatively dense, long, decumbent to suberect. Sculpture shiny and smooth; vertex and frons with sparse, short, and transverse rugulae; antennal sockets with few thick, curved outward rugae and smooth interspaces. Clypeus with median longitudinal carina absent; two lateral longitudinal carinae absent. Scape, when laid back, surpassing the posterior head margin by two-fifths of its length; pilosity dense, subdecumbent to erect (Fig. 22A, C). Mesosoma. In lateral view, promesonotum moderately high and short, arched; promesonotal groove absent; metanotal groove distinct; propodeal spines reduce to small tubercles (Fig. 22C). Sculpture shiny and smooth; anterolateral sides of propodeum and anepisternum with indistinct and sparse rugoreticulae. Pilosity moderately sparse, long, and erect (Fig. 22C, E). Gaster. With sparse, erect pilosity (Fig. 22C, E). Colour. Dark brown, legs and antenna yellowish brown (Fig. 22C, E).

From the type locality.

The species was collected between 400–775 m in elevation, in rainforest. Nests were located in rotten logs and rotten sticks on the ground.

Pheidole andohahela sp. nov. is known only from its terra typica: Col de Tanatana in Parc National Andohahela, Toliara and is most similar to parapatric P. lavasoa sp. nov. recorded so far only from Grand Lavasoa, Toliara. Major workers of both taxa are extremely similar and can be distinguished based on slight differences in the setosity of lateral sides of head (which is sparser and longer in P. andohahela sp. nov.) and shape of the hypostomal teeth (P. andohahela sp. nov. has inner hypostomal teeth with sharp apices and outer hypostomal teeth that are lobe-like and distinctly directed outward). Minor workers are a better resource for species separation. Minor workers of P. andohahela sp. nov. have transverse rugulae on vertex and frons, while anterolateral sides of their propodeum, and anepisternum have indistinct and sparse rugoreticulae; body colour is dark brown.

Holotype. Madagascar. • 1 major worker; Antsiranana; Parc National Montagne d’Ambre, Mahasarika; -12.53176, 49.17662; alt. 1135 m; 17 Nov 2007; B. L. Fisher et al. leg.; montane rainforest, ex rotten log; BLF18461; CASENT0134498 (CASC). Paratype. • 1w.; same data as for holotype, CASENT0235124 (CASC).

Figure 23. 

Pheidole anomala sp. nov., full-face view (A), profile (C), and dorsal view (E) of paratype minor worker (CASENT0235124) and full-face view (B), profile (D), and dorsal view (F) of holotype major worker (CASENT0134498).

Moderately large species. Major workers. Head in full-face view sub-oval and slightly widening posteriorly, with anterior and posterior sides slightly convex, in lateral view sub-oval; ventral and dorsal faces convex; sides of the head with dense, long, suberect to erect pilosity; medial part of frons with irregular and dense rugae, interspaces distinctly rugulate; occipital lobes, and area posterolateral from eyes without smooth notches; scape, when laid back, exceeding the midlength of head by one-fifth of its length; inner hypostomal teeth distinct, large, closely spaced, triangular, with rounded apex directed inward; outer hypostomal teeth lobe-like, wider than and approximately as high as inner teeth; inner and outer hypostomal teeth closely spaced and not connected by concavity; mesosoma rugofoveolate; gaster smooth; body ferruginous. Minor workers. Head foveolate; vertex with a few arcing, sparse, and moderately thick rugae; scape, when laid back, exceeding the posterior head margin by one-third of its length; promesonotum very low and moderately long; promesonotal groove absent; propodeal spines small, triangular; mesosoma foveolate; body yellow.

Major workers. Measurements (N = 1): HL: 1.42; HW: 1.37; SL: 0.87; EL: 0.18; WL: 1.25; PSL: 0.21; MTL: 0.87; PNW: 0.57; PTW: 0.16; PPW: 0.34; CI: 103.0; SI: 63.2; PSLI: 14.8; PPI: 47.8; PNI: 41.6; MTI: 63.2.

Head. In full-face sub-oval, widening posteriorly, with anterior and posterior sides convex (Fig. 23B). In lateral view sub-oval; ventral and dorsal faces convex; inner hypostomal teeth visible. Sides of the head with dense, long, suberect to erect pilosity; whole head with dense, long, decumbent to erect pilosity. Frons with thick, irregular, and dense rugae, interspaces distinctly rugulate. Occipital lobes with slightly thinner, dense, and irregular rugae and rugulate interspaces. Area posterolateral from eyes with dense rugoreticulae. Gena with relatively sparse and thick, longitudinal rugae with distinctly rugofoveolate interspaces. Centre of clypeus with sparse foveolae and shiny, lateral sides with indistinct rugulae; median notch present, moderately wide, and shallow; median longitudinal carina present; lateral longitudinal carinae absent. Scape, when laid back, exceeding the midlength of head by one-fifth of its length; pilosity subdecumbent to erect (Fig. 23B, D). Inner hypostomal teeth distinct, large, closely spaced, triangular, with rounded apex directed inward; outer hypostomal teeth lobe-like, wider than and approximately as high as inner teeth; inner and outer hypostomal teeth closely spaced and not connected by concavity (Fig. 63E). Mesosoma. In lateral view, promesonotum short, angular, and moderately low, posterior mesonotum moderately steep, mesonotal process indistinct, tubercle-like; promesonotal groove absent; metanotal groove absent; propodeal spines moderate, with wide base and acute apex; humeral area produced (Fig. 23D). Surface shiny and rugofoveolate. Pilosity moderately dense, long, and erect (Fig. 23D, F). Petiole. Shiny with dense foveolae; node finely foveolate, triangular, with rounded and thick apex, in rear view node dorsoventrally slightly convex; pilosity moderately sparse and erect (Fig. 23D, F). Postpetiole. Shiny and foveolate; dorsum with reduced sculpture and smooth notch; in dorsal view oval, lateral margins medially with two dentate projections; pilosity long, moderately sparse, and erect (Fig. 23D, F). Gaster. Shiny and smooth; pilosity moderately dense, long, and erect (Fig. D, F). Colour. Ferruginous with yellowish legs (Fig. 23D, F).

Minor workers. Measurements (N = 1): HL: 0.66; HW: 0.52; SL: 0.82; EL: 0.15; WL: 0.89; PSL: 0.12; MTL: 0.7; PNW: 0.36; PTW: 0.1; PPW: 0.16; CI: 128.2; SI: 158.8; PSLI: 17.4; PPI: 62.7; PNI: 69.5; MTI: 136.7.

Head. Cephalic margin slightly convex (Fig. 23A). Pilosity relatively sparse, moderately long, subdecumbent to erect. Sculpture shiny and foveolate; vertex with a few arcing, sparse, and moderately thick rugae; antennal sockets with few indistinct, curved outward rugae and foveolate interspaces. Clypeus with median longitudinal carina absent; two lateral longitudinal carinae absent. Scape, when laid back, exceeding the posterior head margin by one-third of its length; pilosity dense, subdecumbent to erect (Fig. 23A, C). Mesosoma. In lateral view, promesonotum very low and moderately long, arched; promesonotal groove absent; metanotal groove distinct; propodeal spines small, triangular (Fig. 23C). Sculpture shiny and foveolate. Pilosity very sparse, moderately long, and erect (Fig. 23C, E). Gaster. With sparse, erect pilosity (Fig. 23C, E). Colour. Yellow, legs brighter (Fig. 23C, E).

Latin for irregular in reference to strongly irregular rugae on frons of major workers.

The species was collected at 1135 m in elevation, in montane rainforest. Nest was located in a rotten log.

Pheidole anomala sp. nov. is most similar to P. veteratrix Forel, a widespread species known from Antananarivo north to Taolagnaro, and sympatric P. trichotos sp. nov. Major workers of P. anomala sp. nov. differ from both taxa in ferruginous body colouration and distinctly rugulate interspaces on frons; minor workers are easy to distinguish based on their bright yellow body colouration and entirely foveolate katepisternum.

Holotype. Madagascar. • 1 major worker; Toliara; Anosy Region, Anosyenne Mts, 31.2 km NW Manantenina; -24.13894, 47.06804; 1125 m; 26 Feb 2015; B. L. Fisher et al. leg.; rainforest, ex rotten log; BLF36544; CASENT0923297 (CASC). Paratype. • 1w.; same data as for holotype; CASENT0704255 (CASC).

Figure 24. 

Pheidole anosyenne sp. nov., full-face view (A), profile (C), and dorsal view (E) of paratype minor worker (CASENT0704255) and full-face view (B), profile (D), and dorsal view (F) of holotype major worker (CASENT0923297).

Moderately large species. Major workers. Head in full-face view sub-oval and slightly widening posteriorly, with anterior and posterior sides slightly convex, in lateral view sub-oval; ventral and dorsal faces convex; sides of the head with moderately dense, short, decumbent to subdecumbent pilosity; anteromedial part of frons with moderately dense and thick, longitudinal rugae, posteromedial part with irregular rugae, interspaces with sparse and distinct rugofoveolae; occipital lobes, and area posterolateral from eyes without smooth notches; scape, when laid back, exceeding the midlength of head by approximately one-fifth of its length; inner hypostomal teeth distinct, moderate, closely spaced, triangular, with rounded apex and relatively narrow base; outer hypostomal teeth lobe-like, wider, and higher than inner hypostomal teeth, apex directed forward; inner and outer hypostomal teeth closely spaced and connected by indistinct concavity; mesosoma with fine rugofoveolae, dorsal and lateral sides of pronotum with additional transverse and thin rugae; gaster smooth; body ochreous. Minor workers. Head foveolate with additional indistinct longitudinal rugulae on medial frons, area posterolateral from eyes with reduced sculpture; scape, when laid back, surpassing the posterior head margin by two-fifths of its length; promesonotum moderately high and long, arched; promesonotal groove present; propodeal spines very small, triangular; mesosomal dorsum rugofoveolate; lateral sides of pronotum foveolate, anepisternum and katepisternum smooth, lateral sides of propodeum with smooth notches; body dark orange.

Major workers. Measurements (N = 1): HL: 1.13; HW: 1.18; SL: 0.69; EL: 0.14; WL: 1.0; PSL: 0.17; MTL: 0.63; PNW: 0.52; PTW: 0.14; PPW: 0.34; CI: 96.0; SI: 58.3; PSLI: 14.8; PPI: 40.2; PNI: 44.0; MTI: 53.2.

Head. In full-face view sub-oval, widening posteriorly, with anterior and posterior sides convex (Fig. 24B). In lateral view sub-oval; ventral and dorsal faces convex; inner hypostomal teeth visible. Sides of the head with moderately dense, short, decumbent to subdecumbent pilosity; whole head with relatively dense, short, decumbent to erect pilosity. Anteromedial part of frons with moderately dense and thick, longitudinal rugae, posteromedial part with irregular rugae, interspaces with sparse and distinct rugofoveolae; lateral sides with irregular and thick rugae with distinctly rugofoveolate interspaces. Occipital lobes with thick, sparse, irregular rugae, interspaces smooth or with fine, indistinct rugulae. Gena with sparse and thick, longitudinal rugae and indistinctly rugofoveolate interspaces. Area posterolateral from eyes with dense and thin rugofoveolae. Centre of clypeus smooth and shiny, lateral sides with indistinct rugulae; median notch present, wide, and shallow; median longitudinal carina present; lateral longitudinal carinae absent. Scape, when laid back, exceeding the midlength of head by approximately one-fifth of its length; pilosity decumbent to suberect (Fig. 24B, D). Inner hypostomal teeth distinct, moderate, closely spaced, triangular, with rounded apex and relatively narrow base; outer hypostomal teeth lobe-like, wider and higher than inner hypostomal teeth, apex directed forward; inner and outer hypostomal teeth closely spaced and connected by indistinct concavity (Fig. 63F). Mesosoma. In lateral view, promesonotum short, angular, and moderately high, posterior mesonotum moderately steep, mesonotal process indistinct, tubercle-like; promesonotal groove absent; metanotal groove absent; propodeal spines moderate, relatively wide, with acute apex; humeral area laterally weakly produced (Fig. 24D). Surface shiny with fine rugofoveolae, dorsal and lateral sides of pronotum with additional transverse and thin rugae. Pilosity relatively dense, long, and erect (Fig. 24D, F). Petiole. Shiny with fine and dense foveolae; node foveolate, low, triangular, with rounded and thin apex, in rear view node dorsoventrally convex; pilosity moderately sparse and erect (Fig. 24D, F). Postpetiole. Shiny, with fine and sparse rugofoveolae; in dorsal view postpetiole oval, lateral margins medially with two dentate projections; pilosity long, moderately sparse, and erect (Fig. 24D, F). Gaster. Shiny and smooth; pilosity moderately dense, long, and erect (Fig. 24D, F). Colour. Ochreous; legs dark yellow, gaster and mandibles brown (Fig. 24D, F).

Minor workers. Measurements (N = 1): HL: 0.55; HW: 0.49; SL: 0.62; EL: 0.11; WL: 0.72; PSL: 0.07; MTL: 0.47; PNW: 0.34; PTW: 0.07; PPW: 0.13; CI: 113.8; SI: 126.7; PSLI: 12.8; PPI: 54.3; PNI: 70.4; MTI: 96.5.

Head. Cephalic margin indistinctly convex (Fig. 24A). Pilosity relatively sparse, long, decumbent to subdecumbent. Head foveolate with additional indistinct longitudinal rugulae on medial frons, area posterolateral from eyes with reduced sculpture; antennal sockets with few thick, curved outward rugae and foveolate interspaces. Clypeus with median longitudinal carina absent; two lateral longitudinal carinae absent. Scape, when laid back, surpassing the posterior head margin by two-fifths of its length; pilosity dense, subdecumbent to erect (Fig. 24A, C). Mesosoma. In lateral view, promesonotum moderately high and long, arched; promesonotal groove present but indistinct; metanotal groove present and distinct; propodeal spines very small, triangular (Fig. 24C). Mesosomal dorsum rugofoveolate; lateral sides of pronotum foveolate, anepisternum and katepisternum smooth, lateral sides of propodeum with smooth notches. Pilosity sparse, moderately long, and erect (Fig. 24C, E). Petiole. Peduncle with ventral face relatively straight (Fig. 24C, E). Gaster. With sparse, erect pilosity (Fig. 24C, E). Colour. Dark orange (Fig. 24C, E).

From the type locality.

The species was collected at 1125 m in elevation, in rainforest. Nest was located in rotten log.

Pheidole anosyenne sp. nov., described from Anosyenne Mts. in Toliara, has major workers with dense and thick rugae that are anteromedially longitudinal and posteromedially irregular with sparsely and distinctly rugofoveolate interspaces, and ferruginous body colouration. Most similar are P. joffreville sp. nov. and P. mivory sp. nov., known from remote localities in Antsiranana. Major workers of P. anosyenne sp. nov. can be easily separated from both of those taxa based on sparse to moderately sparse, moderately long to long, decumbent to erect pilosity on sides of head, brighter body colouration and presence of smooth notches on lateral sides of pronotum and katepisternum; minor workers differ from P. joffreville sp. nov. in brighter body colouration, presence of smooth notches on lateral sides of propodeum, and entirely smooth anepisternum and katepisternum, from P. mivory sp. nov. in foveolate mesosomal dorsum, brighter body colouration, and less developed propodeal spines.

Holotype. Madagascar. • 1 major worker; Antsiranana; Parc National de Marojejy, Antranohofa, 26.6 km 31°NNE Andapa, 10.7 km 318°NW Manantenina; -14.44333, 49.74333; alt. 1325 m; 20 Nov 2003; B. L. Fisher et al. leg.; montane rainforest, ex root mat, ground layer; BLF09196; CASENT0923295, top specimen on the pin (CASC). Paratypes. • 3w., 2s; same data as for holotype; CASENT0923294, CASENT0499827 CASENT0499829 (CASC, MHNG, PBZT).

Figure 25. 

Pheidole antranohofa sp. nov., full-face view (A), profile (C), and dorsal view (E) of paratype minor worker (CASENT0923294) and full-face view (B), profile (D), and dorsal view (F) of holotype major worker (CASENT0923295).

Madagascar. –Antsiranana: •1w., 1q.; Parc National de Marojejy, Antranohofa, 26.6 km 31°NNE Andapa, 10.7 km 318°NW Manantenina; -14.44333, 49.74333; alt. 1325 m; 20 Nov 2003; B. L. Fisher et al. leg.; montane rainforest, ex rotten tree stump; BLF09202 (CASC). •1w., 1s.; Parc National de Marojejy, Antranohofa, 26.6 km 31°NNE Andapa, 10.7 km 318°NW Manantenina; -14.44333, 49.74333; alt. 1325 m; 20 Nov 2003; B. L. Fisher et al. leg.; montane rainforest, ex rotten log; BLF09215 (CASC).

Moderately large species. Major workers. Head in full-face view sub-oval, widening posteriorly, with anterior and posterior sides convex, in lateral view sub-oval; ventral and dorsal faces convex; sides of the head with dense, short, suberect pilosity; anterior and medial parts of frons with moderately dense and thick, longitudinal and sometimes interrupted rugae with distinctly rugofoveolate interspaces; posterolateral sides with longitudinally irregular and thick rugae with distinctly rugofoveolate interspaces, area posterolateral from eyes without smooth notches; scape, when laid back, exceeding the midlength of head by approximately one-fifth of its length; inner hypostomal teeth distinct, moderately long, closely spaced, triangular, with rounded apex and wide base; outer hypostomal teeth lobe-like, wider and higher than inner hypostomal teeth, apex directed outward; inner and outer hypostomal teeth closely spaced and connected by indistinct concavity; promesonotum predominantly rugofoveolate, with additional sparse, thin, transverse rugae on pronotum; propodeum with fine rugofoveolae; gaster smooth; body yellowish orange. Minor workers. Head foveolate with additional indistinct rugulae on frons and vertex, area posterolateral from eyes with reduced sculpture, on posterior part predominantly smooth; scape, when laid back, surpassing the posterior head margin by two-fifths of its length; promesonotum moderately high and short; promesonotal groove present; propodeal spines indistinct, triangular; mesosoma smooth and shiny, only dorsum with few transverse, indistinct rugulae and propodeum with indistinct and sparse foveolae; body dark yellow.

Major workers. Measurements (N = 4): HL: 1.18–1.23 (1.21); HW: 1.21–1.27 (1.24); SL: 0.67–0.7 (0.69); EL: 0.14–0.16 (0.15); WL: 1.1–1.13 (1.12); PSL: 0.2–0.21 (0.2); MTL: 0.7–0.73 (0.72); PNW: 0.54–0.59 (0.57); PTW: 0.15–0.17 (0.16); PPW: 0.33–0.35 (0.34); CI: 96.7–97.8 (97.3); SI: 54.1–57.8 (55.4); PSLI: 16.3–17.7 (16.9); PPI: 43.4–50.5 (46.6); PNI: 43.9–46.9 (45.6); MTI: 56.7–58.8 (57.6).

Head. In full-face view sub-oval, widening posteriorly, with anterior and posterior sides convex (Fig. 25B). In lateral view sub-oval; ventral and dorsal faces convex; inner hypostomal teeth invisible. Sides of the head with dense, short, suberect pilosity; whole head with relatively dense, short, decumbent to erect pilosity. Anterior and medial parts of frons with moderately dense and thick, longitudinal and sometimes interrupted rugae with distinctly rugofoveolate interspaces; posterolateral sides with longitudinally irregular and thick rugae with distinctly rugofoveolate interspaces. Occipital lobes with thick, sparse, irregular rugae, interspaces smooth or with fine, indistinct rugofoveolae. Gena with sparse and thick, longitudinal rugae and smooth interspaces. Area posterolateral from eyes with thin, longitudinal rugae with distinctly rugofoveolate interspaces. Centre of clypeus smooth and shiny, lateral sides with indistinct rugulae; median notch present, wide and shallow; median longitudinal carina absent; lateral longitudinal carinae absent. Scape, when laid back, exceeding the midlength of head by approximately one-fifth of its length; pilosity decumbent to suberect (Fig. 25B, D). Inner hypostomal teeth distinct, moderate, closely spaced, triangular, with rounded apex and wide base; outer hypostomal teeth lobe-like, wider and higher than inner hypostomal teeth, apex directed outward; inner and outer hypostomal teeth closely spaced and connected by indistinct concavity (Fig. 63G). Mesosoma. In lateral view, promesonotum short, angular, and moderately high, posterior mesonotum moderately steep, mesonotal process indistinct, tubercle-like; promesonotal groove absent; metanotal groove absent; propodeal spines moderate, thin, with acute apex; humeral area laterally weakly produced (Fig. 25D). Surface shiny, promesonotum predominantly rugofoveolate, with additional sparse, thin, transverse rugae on pronotum; propodeum with fine rugofoveolae. Pilosity sparse, moderately long, and erect (Fig. 25D, F). Petiole. Shiny with fine and sparse rugofoveolae; node smooth to indistinctly rugulose, low, triangular, with rounded apex, in rear view node dorsoventrally convex; pilosity moderately sparse and erect (Fig. 25D, F). Postpetiole. Shiny, with fine and sparse rugulae; in dorsal view postpetiole oval, lateral margins medially with two dentate projections; pilosity long, moderately sparse, and erect (Fig. 25D, F). Gaster. Shiny and smooth; pilosity sparse, long, and erect (Fig. 25D, F). Colour. Yellowish orange; mandibles and anterior part of head slightly darker (Fig. 25D, F).

Minor workers. Measurements (N = 9): HL: 0.57–0.64 (0.59); HW: 0.47–0.53 (0.5); SL: 0.59–0.66 (0.62); EL: 0.09–0.11 (0.1); WL: 0.7–0.79 (0.74); PSL: 0.06–0.08 (0.07); MTL: 0.48–0.56 (0.52); PNW: 0.3–0.38 (0.33); PTW: 0.07–0.1 (0.09); PPW: 0.13–0.16 (0.14); CI: 112.2–122.3 (119.5); SI: 121.6–130.3 (125.5); PSLI: 10.8–12.6 (11.8); PPI: 51.1–70.3 (61.2); PNI: 64.0–70.2 (66.2); MTI: 101.9–108.0 (104.8).

Head. Cephalic margin indistinctly convex (Fig. 25A). Pilosity relatively sparse, long, and subdecumbent. Head foveolate with additional indistinct rugulae on frons and vertex, area posterolateral from eyes with reduced sculpture and partially smooth; antennal sockets with few thick, curved outward rugae and foveolate interspaces. Clypeus with median longitudinal carina absent; two lateral longitudinal carinae absent. Scape, when laid back, surpassing the posterior head margin by two-fifths of its length; pilosity dense, subdecumbent to erect (Fig. 25A, C). Mesosoma. In lateral view, promesonotum moderately high, short, arched; promesonotal groove present, distinct; metanotal groove present and distinct; propodeal spines indistinct, triangular, apex acute (Fig. 25C). Sculpture smooth and shiny, only dorsum with few transverse, indistinct rugulae and propodeum with indistinct and sparse foveolae. Pilosity sparse, moderately long, and erect (Fig. 25C, E). Petiole. Peduncle with ventral face relatively straight; node low, knobbed, and small; with few short, erect setae (Fig. 25C, E). Gaster. With sparse, erect pilosity (Fig. 25C, E). Colour. Dark yellow (Fig. 25C, E).

From the type locality.

The species was collected at 1325 m in elevation, in montane rainforest. Nests were located in rotten logs, root mat and rotten tree stump.

Pheidole antranohofa sp. nov. is known only from its type locality: Parc National de Marojejy in Antsiranana. Morphologically it is most similar to P. sikorae Forel which is widely distributed from Forêt Ambohidena in Toamasina to Andrambovato in Fianarantsoa. Major workers of P. antranohofa sp. nov. can be distinguished based on developed sculpture of mesosoma, absence of impressed and smooth concavity placed lateral to antennal socket and tentorial pit and distinctly rugofoveolate interspaces on frons. However, minors of P. antranohofa sp. nov. and P. sikorae are indistinguishable.

Holotype. Madagascar. • 1 major worker; Mahajanga; Réserve forestière Beanka, 53.6 km E Maintirano; -18.04014, 44.53394; alt. 272 m; 25 Oct 2009; B. L. Fisher et al. leg.; tropical dry forest on tsingy, ex rotten log; BLF22970; CASENT0156684 (CASC). Paratype. • 1w.; same data as for holotype, CASENT0923276 (CASC).

Figure 26. 

Pheidole beanka sp. nov., full-face view (A), profile (C), and dorsal view (E) of paratype minor worker (CASENT0923276) and full-face view (B), profile (D), and dorsal view (F) of holotype major worker (CASENT0156684).

Minute species. Major workers. HL < 0.9 mm and WL < 0.7 mm; head in full-face view elongate, not widening posteriorly with posterior sides slightly convex, in lateral view sub-oval and short with convex dorsal and ventral sides; body ocherous; sides of head with moderately dense, moderately short, suberect to erect pilosity; entire head distinctly sculptured, medial part of frons with thick and longitudinal rugae and distinctly rugulate and never smooth interspaces, occipital lobes with dense and irregular rugae with distinctly rugoreticulate interspaces; scape, when laid back, exceeding the midlength of head by one-fifth of its length; lateral sides of promesonotum distinctly foveolate without additional sparse and indistinct rugulae; inner hypostomal teeth distinct, large, closely spaced, triangular, with rounded apex directed upward; outer hypostomal teeth lobe-like, approximately the same height and width as inner teeth; inner and outer hypostomal teeth closely spaced and not connected by concavity; base of first gastral tergite shagreened. Minor workers. HL < 0.45 mm and WL < 0.5 mm, scape, when laid back, surpassing the posterior head margin by one-fifth of its length; propodeal spines reduced to small tubercles; head relatively rectangular; body yellowish brown; lateral sides of frons smooth; vertex with transverse, sparse, and moderately thick, short rugulae; mesosoma smooth.

Major workers. Measurements (N = 1): HL: 0.86; HW: 0.74; SL: 0.45; EL: 0.1; WL: 0.7; PSL: 0.09; MTL: 0.38; PNW: 0.39; PTW: 0.12; PPW: 0.3; CI: 115.3; SI: 60.3; PSLI: 10.5; PPI: 40.4; PNI: 51.9; MTI: 51.1.

Head. In full-face view elongate, not widening posteriorly, with anterior and posterior sides slightly convex (Fig. 26B). In lateral view sub-oval; ventral and dorsal faces convex; inner hypostomal teeth visible. Sides of the head with moderately dense, moderately short, suberect to erect pilosity; whole head with dense, long, decumbent to erect pilosity. Medial part of frons with thick, longitudinal, and dense rugae and distinctly rugulate interspaces, rugae more irregular and with denser sculpture of interspaces on posteromedial part; lateral sides with thick, dense and longitudinal rugae with distinctly rugofoveolate interspaces, rugae more irregular on posterolateral parts. Occipital lobes with dense, irregular rugae and distinctly rugoreticulate interspaces. Area posterolateral from eyes with dense rugofoveolae. Gena with relatively sparse and thick, longitudinal rugae and indistinctly rugoreticulate interspaces. Centre of clypeus smooth and shiny, lateral sides with indistinct rugulae; median notch present, moderately wide, and shallow; median longitudinal carina absent; lateral longitudinal carinae absent. Scape, when laid back, exceeding the midlength of head by one-fifth of its length; pilosity subdecumbent to erect (Fig. 26B, D). Inner hypostomal teeth distinct, large, closely spaced, triangular, with rounded apex directed upward; outer hypostomal teeth lobe-like, approximately the same height and width as inner teeth; inner and outer hypostomal teeth closely spaced and not connected by concavity (Fig. 63H). Mesosoma. In lateral view, promesonotum short, angular, and moderately high, posterior mesonotum moderately steep, mesonotal process very indistinct, tubercle-like; promesonotal groove absent; metanotal groove absent; propodeal spines very small, with wide base and acute apex; humeral area produced (Fig. 26D). Surface shiny and distinctly foveolate; katepisternum with sparser foveolae. Pilosity sparse, moderately long, and erect (Fig. 26D, F). Petiole. Shiny with dense foveolae; node finely foveolate, triangular, with rounded and thin apex, in rear view node dorsoventrally concave; pilosity moderately sparse and erect (Fig. 26D, F). Postpetiole. Shiny and foveolate; dorsum with reduced sculpture; in dorsal view oval, lateral margins medially with two distinct dentate projections; pilosity long, moderately sparse, and erect (Fig. 26D, F). Gaster. Shiny and smooth with shagreened base; pilosity moderately dense, moderately short, and erect (Fig. 26D, F). Colour. Ochreous with yellow legs and antenna (Fig. 26D, F).

Minor workers. Measurements (N = 1): HL: 0.44; HW: 0.39; SL: 0.39; EL: 0.1; WL: 0.48; PSL: 0.05; MTL: 0.3; PNW: 0.25; PTW: 0.06; PPW: 0.11; CI: 112.8; SI: 98.7; PSLI: 10.9; PPI: 48.7; PNI: 65.0; MTI: 77.0.

Head. Cephalic margin slightly concave (Fig. 26A). Pilosity relatively sparse, moderately long, decumbent to subdecumbent. Sculpture shiny and smooth; vertex with transverse, sparse, and moderately thick, short rugulae; antennal sockets with few thick, curved outward rugae and smooth interspaces. Clypeus with median longitudinal carina absent; two lateral longitudinal carinae absent. Scape, when laid back, exceeding the posterior head margin by one-fifth of its length; pilosity dense, subdecumbent to erect (Fig. 26A, C). Mesosoma. In lateral view, promesonotum moderately high and short, box-like; promesonotal groove absent; metanotal groove distinct; propodeal spines reduced to small tubercles (Fig. 26C). Sculpture shiny and smooth. Pilosity sparse, moderately long, and erect (Fig. 26C, E). Gaster. With sparse, erect pilosity (Fig. 26C, E). Colour. Yellow (Fig. 26C, E).

From the type locality.

The species was collected at 272 m in elevation, in tropical dry forest. Nest was located in rotten log.

Pheidole beanka sp. nov. belongs to the group of species characterised by small body size (major workers: HL < 1.05 mm, WL < 0.9 mm and minor workers HL < 0.5 mm, WL < 0.6 mm), elongate and not widening posteriorly head in major workers, and minor workers with predominantly smooth and relatively rectangular head and yellow to brown body colouration. The group includes five species: P. flavominuta sp. nov., P. nitidobruna sp. nov., P. mikros sp. nov., and P. beanka sp. nov. Pheidole beanka sp. nov., described from Réserve forestière Beanka, is the only member of this group known from the Mahajanga and its distribution doesn’t overlap with the remaining four taxa. However, morphology of P. beanka sp. nov. is most similar to P. mikros sp. nov., which is distributed on the southernmost part of the Antsiranana prefecture. Major workers of Pheidole beanka sp. nov. differ from P. mikros sp. nov. in the combination of the following characters: medial part of frons with distinctly rugulate interspaces and very small propodeal spines; minor workers can be separated based on smooth lateral sides of frons and mesosoma, yellow body, and propodeal spines reduced to small tubercles.

Holotype. Madagascar. • 1 major worker; Fianarantsoa; Parc National Befotaka-Midongy, Papango 28.5km S Midongy-Sud, Mount Papango; -23.84083, 46.9575; alt. 1250 m; 17 Nov 2006; B. L. Fisher et al. leg.; montane rainforest, ex rotten log; BLF14974; CASENT0119590 (CASC). Paratype. • 1w.; same data as for holotype; CASENT0235125 (CASC).

Figure 27. 

Pheidole befotaka sp. nov., full-face view (A), profile (C), and dorsal view (E) of paratype minor worker (CASENT0235125) and full-face view (B), profile (D), and dorsal view (F) of holotype major worker (CASENT0119590).

Madagascar. –Fianarantsoa: •1w., 1s.; Parc National Befotaka-Midongy, Papango 27.7km S Midongy-Sud, Mount Papango; -23.83517, 46.96367; alt. 940 m; 16 Nov 2006; B. L. Fisher et al. leg.; rainforest, ex dead twig above ground; BLF14931 (CASC).

Moderately large species. Major workers. Head in full-face view sub-oval, slightly widening posteriorly, with anterior and posterior sides convex, in lateral view sub-oval, ventral and dorsal faces convex; sides of the head with dense, moderately long, erect pilosity; medial part of frons with moderately dense and thick rugae, anteriorly rugae longitudinal and interrupted, posteromedially rugae irregular, interspaces shiny with sparse and indistinct irregular rugulae; lateral sides with irregular, thick, and relatively sparse rugae with indistinctly rugoreticulate interspaces; occipital lobes, and area posterolateral from eyes never smooth; scape, when laid back, exceeding the midlength of head by two-fifths of its length; inner hypostomal teeth distinct, large, closely spaced, triangular, with rounded apex directed upward; outer hypostomal teeth lobe-like, wider than inner hypostomal teeth and approximately the same height, apex directed outward; inner and outer hypostomal teeth closely spaced and connected by indistinct concavity; pronotum and mesonotum with fine rugoreticulae and additional transverse to irregular, thick rugae; anepisternum, katepisternum and propodeum with fine rugoreticulae; body orange. Minor workers. Head foveolate; medial part of frons with smooth notch; vertex and lateral sides of frons with additional, irregular to arcing rugae, area posterolateral from eyes smooth; scape, when laid back, surpassing the posterior head margin by two-fifths of its length; promesonotum moderately low and moderately short; promesonotal groove indistinct; propodeal spines small and triangular; mesosoma with very indistinct and sparse rugofoveolae; dorsal pronotum and medial parts of lateral sides of pronotum, katepisternum, and anepisternum smooth; body yellow.

Major workers. Measurements (N = 2): HL: 1.18, 1.12; HW: 1.18, 1.1; SL: 0.71, 0.73; EL: 0.16, 0.16; WL: 1.06, 1.05; PSL: 0.19, 0.17; MTL: 0.73, 0.72; PNW: 0.51, 0.48; PTW: 0.17, 0.14; PPW: 0.31, 0.31; CI: 100.5, 101.8; SI: 60.6, 66.3; PSLI: 16.0, 15.2; PPI: 56.9, 45.4; PNI: 43.0, 44.0; MTI: 62.2, 65.7.

Head. In full-face view sub-oval, slightly widening posteriorly, with anterior and posterior sides convex (Fig. 27B). In lateral view sub-oval; ventral and dorsal faces convex; inner hypostomal teeth visible. Sides of the head with dense, moderately long, erect pilosity; whole head with dense, long, decumbent to erect pilosity. Medial part of frons with moderately dense and thick rugae, anteriorly rugae longitudinal and interrupted, posteromedially rugae irregular, interspaces shiny with sparse and indistinct irregular rugulae; lateral sides with irregular, thick, and relatively sparse rugae with indistinctly rugoreticulate interspaces. Occipital lobes with sparse, thick, and irregular rugae; interspaces smooth. Gena with relatively dense and thick longitudinal rugae and smooth interspaces. Area posterolateral from eyes with thin and dense rugofoveolae, sculpture weakening posteriorly. Centre of clypeus smooth and shiny, lateral sides with indistinct rugulae; median notch present, moderately wide, and shallow; median longitudinal carina present; lateral longitudinal carinae absent. Scape, when laid back, exceeding the midlength of head by two-fifths of its length; pilosity subdecumbent to erect (Fig. 27B, D). Inner hypostomal teeth distinct, large, closely spaced, triangular, with rounded apex directed upward; outer hypostomal teeth lobe-like, wider than inner hypostomal teeth and approximately the same height, apex directed outward; inner and outer hypostomal teeth closely spaced and connected by indistinct concavity (Fig. 63I). Mesosoma. In lateral view, promesonotum short, angular, and moderately low, posterior mesonotum moderately steep, mesonotal process indistinct, tubercle-like; promesonotal groove absent; metanotal groove indistinct; propodeal spines moderately long, moderately wide, with acute apex; humeral area laterally weakly produced (Fig. 27D). Surface shiny; pronotum and mesonotum with fine rugoreticulae and additional transverse to irregular, thick rugae; anepisternum, katepisternum, and propodeum with fine rugoreticulae. Pilosity relatively dense, long, and erect (Fig. 27D, F). Petiole. Shiny with fine and dense rugofoveolae; node smooth, low, triangular, with rounded and thin apex, in rear view node dorsoventrally slightly convex; pilosity moderately sparse and erect (Fig. 27D, F). Postpetiole. Shiny and smooth; in dorsal view oval, lateral margins medially with two dentate projections; pilosity long, moderately sparse, and erect (Fig. 27D, F). Gaster. Shiny and smooth; pilosity moderately dense, long, and erect (Fig. 27D, F). Colour. Orange; mandibles and gaster slightly darker; legs yellowish (Fig. 27D, F).

Minor workers. Measurements (N = 2): HL: 0.66, 0.64; HW: 0.54, 0.52; SL: 0.76, 0.74; EL: 0.13, 0.13; WL: 0.85, 0.86; PSL: 0.09, 0.1; MTL: 0.61, 0.61; PNW: 0.36, 0.36; PTW: 0.1, 0.08; PPW: 0.17, 0.15; CI: 121.2, 123.2; SI: 140.0, 141.0; PSLI: 13.5, 14.9; PPI: 58.2, 55.0; PNI: 66.4, 69.2; MTI: 112.0, 117.0.

Head. Cephalic margin indistinctly convex or straight (Fig. 27A). Pilosity relatively sparse, long, decumbent to suberect. Sculpture foveolate; medial part of frons with smooth notch; vertex and lateral sides of frons with additional, irregular to arcing rugae; area posterolateral from eyes smooth. Clypeus with median longitudinal carina absent; two lateral longitudinal carinae absent. Scape, when laid back, surpassing the posterior head margin by two-fifths of its length; pilosity dense, subdecumbent to erect (Fig. 27A, C). Mesosoma. In lateral view, promesonotum moderately low and moderately short, arched; promesonotal groove absent; metanotal groove indistinct; propodeal spines small and triangular (Fig. 27C). Sculpture with very indistinct and sparse rugofoveolae; dorsal pronotum and medial parts of lateral sides of pronotum, katepisternum, and anepisternum smooth. Pilosity very sparse, moderately long, and erect (Fig. 27C, E). Postpetiole. Short, low, and relatively flat; with few short, erect setae (Fig. 27C, E). Gaster. With sparse, erect pilosity (Fig. 27C, E). Colour. Yellow, vertex slightly darker (Fig. 27C, E).

From the type locality.

The species was collected between 940–1250 m in elevation, in rainforest and montane rainforest. Nests were located in rotten logs and dead twigs above ground.

Pheidole befotaka sp. nov. is a member of a group of species characterised by body coloration that is bright, yellow to orange in majors and yellow in minors, head sub-oval, not widening posteriorly with sides not convex or convex indistinctly, and medial part of frons with longitudinal and interrupted rugae anteriorly and distinctly irregular rugae posteriorly. The group includes three taxa: P. vony sp. nov., P. befotaka sp. nov., and P. mamiratra sp. nov. Pheidole befotaka sp. nov. is the only member of the group known from Parc National Befotaka-Midongy in Fianarantsoa and can be easily separated based on orange body of major workers with slightly convex lateral sides of head and lateral sides of frons with distinctly irregular rugae. Minor workers of P. befotaka sp. nov. can be separated from P. vony sp. nov. and P. mamiratra sp. nov. based on presence of additional, irregular to arcing rugae on vertex and lateral sides of frons. Additionally, minors of P. befotaka sp. nov. can be confused with minors of sympatric P. veteratrix from which they differ in reduced sculpture of mesosoma and higher promesonotum.

Holotype. Madagascar. • 1 major worker; Antsiranana; Makirovana forest; -14.104, 50.03574; alt. 225 m; 4 May 2011; B. L. Fisher et al. leg.; rainforest, ex rotten log; BLF27046; CASENT0212441 (CASC). Paratype. • 1w.; same data as for holotype, CASENT0923284 (CASC).

Figure 28. 

Pheidole dasos sp. nov., full-face view (A), profile (C), and dorsal view (E) of paratype minor worker (CASENT0923284) and full-face view (B), profile (D), and dorsal view (F) of holotype major worker (CASENT0212441).

Madagascar. –Antsiranana: •1w., 1s.; Makirovana forest; -14.17066, 49.95409; alt. 415 m; 29 Apr 2011; B. L. Fisher et al. leg.; rainforest, ex rotten log; BLF26619 (CASC).

Moderately large species. Major workers. Head in full-face view sub-oval, not widening posteriorly, with anterior and posterior sides relatively straight, in lateral view sub-oval; ventral and dorsal faces convex; sides of the head with moderately dense, moderately long, subdecumbent to suberect pilosity; medial part of frons with thick, interrupted, sparse, and longitudinal rugae with indistinctly foveolate interspaces; anterolateral sides of frons with sparse, thick, and longitudinal rugae, posterolateral sides with rugae more irregular, interspaces with sparse foveolae; occipital lobes, and area posterolateral from eyes without smooth notches; scape, when laid back, exceeding the midlength of head by two-fifths of its length; inner hypostomal teeth distinct, large, closely spaced, triangular, with rounded apex directed upward; outer hypostomal teeth lobe-like, narrower, and approximately as high as inner teeth; inner and outer hypostomal teeth closely spaced and not connected by concavity; mesosoma rugofoveolate; dorsal promesonotum with additional thick and transverse rugae; lateral sides of pronotum with smooth notches; gaster smooth; body orange. Minor workers. Head foveolate; vertex and median frons with fading sculpture; area posterolateral from eyes smooth; antennal sockets with few indistinct, curved outward rugae and foveolate interspaces; scape, when laid back, exceeding the posterior head margin by one-third of its length; promesonotum low and moderately long; promesonotal groove absent; propodeal spines small and triangular; mesosoma predominantly smooth; lateral sides with very sparse and indistinct, thin rugulae; body brown.

Major workers. Measurements (N = 2): HL: 1.1, 1.14; HW: 1.04, 1.12; SL: 0.79, 0.81; EL: 0.16, 0.14; WL: 0.98, 1.05; PSL: 0.14, 0.15; MTL: 0.72, 0.77; PNW: 0.4, 0.45; PTW: 0.13, 0.13; PPW: 0.29, 0.31; CI: 105.9, 101.4; SI: 76.2, 72.6; PSLI: 12.4, 13.5; PPI: 45.8, 41.7; PNI: 38.8, 39.9; MTI: 68.8, 68.6.

Head. In full-face view sub-oval, not widening posteriorly, with anterior and posterior sides relatively straight (Fig. 28B). In lateral view sub-oval; ventral and dorsal faces convex; inner hypostomal teeth visible. Sides of the head with moderately dense, moderately long, subdecumbent to suberect pilosity; whole head with dense, long, decumbent to erect pilosity. Medial part of frons with thick, interrupted, sparse, and longitudinal rugae with indistinctly foveolate interspaces; anterolateral sides of frons with sparse, thick, and longitudinal rugae, posterolateral sides with rugae more irregular, interspaces with sparse foveolae. Occipital lobes with slightly thinner, sparse, and irregular rugae and smooth to indistinctly foveolate interspaces. Area posterolateral from eyes with denser and thinner longitudinal rugae with rugofoveolate interspaces. Gena with relatively sparse and thick, longitudinal rugae with distinctly rugofoveolate interspaces. Centre of clypeus smooth and shiny, lateral sides with indistinct rugulae; median notch present, moderately wide, and shallow; median longitudinal carina absent; lateral longitudinal carinae absent. Scape, when laid back, exceeding the midlength of head by two-fifths of its length; pilosity subdecumbent to erect (Fig. 28B, D). Inner hypostomal teeth distinct, large, closely spaced, triangular, with rounded apex directed upward; outer hypostomal teeth lobe-like, narrower than and approximately as high as inner teeth; inner and outer hypostomal teeth closely spaced and not connected by concavity (Fig. 63J). Mesosoma. In lateral view, promesonotum short, angular and moderately low, posterior mesonotum moderately steep, mesonotal process indistinct, tubercle-like; promesonotal groove absent; metanotal groove absent; propodeal spines short, with wide base and acute apex; humeral area produced (Fig. 28D). Surface shiny and rugofoveolate; dorsal promesonotum with additional thick and transverse rugae; lateral sides of pronotum with smooth notches. Pilosity moderately dense, long, and erect (Fig. 28D, F). Petiole. Shiny with dense foveolae; node finely foveolate, triangular, with rounded and thick apex, in rear view node dorsoventrally slightly convex; pilosity moderately sparse and erect (Fig. 28D, F). Postpetiole. Shiny and foveolate; dorsum with reduced sculpture and smooth notch; in dorsal view oval, lateral margins medially with two dentate projections; pilosity long, moderately sparse and erect (Fig. 28D, F). Gaster. Shiny and smooth; pilosity moderately dense, long and erect (Fig. 28D, F). Colour. Orange with yellowish legs (Fig. 28D, F).

Minor workers. Measurements (N = 2): HL: 0.56, 0.61; HW: 0.46, 0.49; SL: 0.71, 0.74; EL: 0.1, 0.1; WL: 0.74, 0.79; PSL: 0.08, 0.09; MTL: 0.57, 0.59; PNW: 0.31, 0.32; PTW: 0.09, 0.07; PPW: 0.12, 0.14; CI: 122.0, 122.7; SI: 153.6, 150.9; PSLI: 13.8, 15.4; PPI: 74.3, 52.9; PNI: 67.5, 65.7; MTI: 123.3, 120.5.

Head. Cephalic margin slightly convex (Fig. 28A). Pilosity relatively sparse, moderately long, subdecumbent to erect. Sculpture shiny and foveolate; vertex and median frons with fading sculpture; area posterolateral from eyes smooth; antennal sockets with few indistinct, curved outward rugae and foveolate interspaces. Clypeus with median longitudinal carina absent; two lateral longitudinal carinae absent. Scape, when laid back, exceeding the posterior head margin by one-third of its length; pilosity dense, subdecumbent to erect (Fig. 28A, C). Mesosoma. In lateral view, promesonotum low and moderately long, arched; promesonotal groove absent; metanotal groove distinct; propodeal spines small, triangular (Fig. 28C). Sculpture shiny and predominantly smooth; lateral sides with very sparse and indistinct, thin rugulae. Pilosity very sparse, moderately long, and erect (Fig. 28C, E). Gaster. With sparse, erect pilosity (Fig. 28C, E). Colour. Brown, legs and gaster brighter (Fig. 28C, E).

Greek for forest in reference to habitat preferences of the species.

The species was collected between 225–415 m in elevation, in rainforest. Nests were located in rotten logs.

Pheidole dasos sp. nov., described from Makirovana forest in Antsiranana, is most similar to P. hazo sp. nov., known from the vicinity of Andranoma in Antananarivo, and P. sparsa sp. nov., recorded so far only from Bemanevika in Mahajanga. Majors of P. dasos sp. nov. can be distinguished from both taxa by medial part of frons with moderately dense rugae with interspaces with indistinct foveolae, and pronotum predominantly sculptured with smooth notches on lateral sides; minors of P. dasos sp. nov. can be distinguished from P. sparsa sp. nov. and P. hazo sp. nov. by brown body colouration, longer scape exceeding the posterior head margin by one-third of its length, and predominantly smooth mesosoma with very sparse and indistinct, thin rugulae on lateral sides.

Holotype. Madagascar. • 1 major worker; Toamasina; Réserve Spéciale Ambatovaky, Sandrangato river; -16.81745, 49.2925; alt. 400 m; 26 Feb 2010; B. L. Fisher et al. leg.; rainforest, under root mat on ground; BLF24911; CASENT0923272 (CASC). Paratype. • 1w.; same data as for holotype, CASENT0162186 (CASC).

Figure 29. 

Pheidole flavominuta sp. nov., full-face view (A), profile (C), and dorsal view (E) of paratype minor worker (CASENT0162186) and full-face view (B), profile (D), and dorsal view (F) of holotype major worker (CASENT0923272).

Madagascar. –Toamasina: • 1w., 1s.; Reserve Betampona, Camp Vohitsivalana, 37.1 km 338° Toamasina; -17.88667, 49.2025; alt. 520 m; 1 Dec 2005; B. L. Fisher et al. leg.; rainforest, ex rotten log; BLF13262 (CASC). • 1w., 1s.; Réserve Spéciale Ambatovaky, Sandrangato river; -16.77468, 49.26551; alt. 355 m; 21 Feb 2010; B. L. Fisher et al. leg.; rainforest along river, under rootmat, litter on rock; BLF24505 (CASC). • 1w., 1s.; Réserve Spéciale Ambatovaky, Sandrangato river; -16.81745, 49.2925; alt. 400 m; 26 Feb 2010; B. L. Fisher et al. leg.; rainforest, ex rotten log; BLF24913 (CASC). • 2w.; Réserve Spéciale Ambatovaky, Sandrangato river; -16.81745, 49.2925; alt. 400 m; 26 Feb 2010; B. L. Fisher et al. leg.; rainforest, ex rotten log; BLF24943 (CASC). • 1w., 1s.; Réserve Spéciale Ambatovaky, Sandrangato river; -16.81745, 49.2925; alt. 400 m; 26 Feb 2010; B. L. Fisher et al. leg.; rainforest, ex rotten log; BLF24958 (CASC).

Minute species. Major workers. HL < 0.85 mm and WL < 0.65 mm; head in full-face view elongate, not widening posteriorly with posterior sides slightly convex, in lateral view sub-oval and elongate with slightly convex dorsal and ventral sides; body yellow; sides of head with moderately dense, short, suberect to erect pilosity; entire head distinctly sculptured but with smooth interspaces; scape, when laid back, reaching the midlength of head; inner hypostomal teeth distinct, moderate, closely spaced, triangular, with rounded apex directed upward; outer hypostomal teeth lobe-like, approximately as high and wide as inner teeth; inner and outer hypostomal teeth moderately closely spaced and not connected by concavity; base of first gastral tergite smooth. Minor workers. HL < 0.45 mm and WL < 0.5 mm, scape, when laid back, reaching the posterior head margin, propodeal spines small and triangular; head relatively rectangular; body yellow; head predominantly smooth; mesosoma smooth.

Major workers. Measurements (N = 5): HL: 0.76–0.83 (0.8); HW: 0.68–0.73 (0.71); SL: 0.39–0.41 (0.4); EL: 0.08–0.08 (0.08); WL: 0.61–0.66 (0.64); PSL: 0.1–0.13 (0.12); MTL: 0.33–0.36 (0.35); PNW: 0.36–0.4 (0.37); PTW: 0.09–0.1 (0.1); PPW: 0.17–0.19 (0.18); CI: 111.3–115.3 (113.0); SI: 53.7–58.7 (56.2); PSLI: 12.8–15.5 (14.5); PPI: 47.4–60.8 (54.6); PNI: 51.4–53.9 (52.6); MTI: 48.2–49.6 (49.0).

Head. In full-face view elongate, not widening posteriorly, with anterior and posterior sides slightly convex (Fig. 29B). In lateral view sub-oval and elongate; ventral and dorsal faces convex; inner hypostomal teeth visible. Sides of the head with moderately dense, short, suberect to erect pilosity; whole head with dense, long, decumbent to erect pilosity. Medial part of frons with thick, longitudinal, and dense rugae and smooth to indistinctly rugulate interspaces, rugae directed slightly outward on posteromedial part; lateral sides with thick, dense, and irregular rugae, interspaces shiny and distinctly rugoreticulate. Occipital lobes with dense, irregular rugae and distinctly rugoreticulate interspaces. Area posterolateral from eyes with weaker sculpture, rugoreticulate. Gena with relatively sparse and thick longitudinal rugae and smooth interspaces. Centre of clypeus smooth and shiny, lateral sides with indistinct rugulae; median notch present, moderately wide, and shallow; median longitudinal carina absent; lateral longitudinal carinae absent. Scape, when laid back, reaching the midlength of head; pilosity subdecumbent to erect (Fig. 29B, D). Inner hypostomal teeth distinct, moderate, closely spaced, triangular, with rounded apex directed upward; outer hypostomal teeth lobe-like, approximately as high and wide as inner teeth; inner and outer hypostomal teeth moderately closely spaced and not connected by concavity (Fig. 63K). Mesosoma. In lateral view, promesonotum short, angular, and moderately low, posterior mesonotum steep, mesonotal process very indistinct, bulge-like; promesonotal groove absent; metanotal groove absent; propodeal spines moderately long, with wide base and acute apex; humeral area laterally weakly produced (Fig. 29D). Surface shiny; promesonotum predominantly smooth with very fine network of rugoreticulae; anepisternum, katepisternum, and propodeum with rugoreticulae denser and thicker. Pilosity moderately dense, moderately long, and erect (Fig. 29D, F). Petiole. Shiny with dense foveolae; node smooth to finely foveolate, triangular, with rounded and thick apex, in rear view node dorsoventrally convex; pilosity moderately sparse and erect (Fig. 29D, F). Postpetiole. Shiny and foveolate; dorsum with reduced sculpture and smooth notch; in dorsal view oval, lateral margins medially with two dentate projections; pilosity long, moderately sparse, and erect (Fig. 29D, F). Gaster. Shiny and smooth; pilosity moderately dense, moderately short, and erect (Fig. 29D, F). Colour. Yellow (Fig. 29D, F).

Minor workers. Measurements (N = 5): HL: 0.42–0.44 (0.43); HW: 0.37–0.39 (0.38); SL: 0.34–0.36 (0.35); EL: 0.06–0.07 (0.06); WL: 0.46–0.49 (0.47); PSL: 0.05–0.07 (0.06); MTL: 0.25–0.28 (0.26); PNW: 0.23–0.26 (0.25); PTW: 0.05–0.07 (0.06); PPW: 0.09–0.11 (0.1); CI: 108.4–117.2 (114.8); SI: 88.2–95.7 (93.2); PSLI: 12.2–16.9 (14.6); PPI: 47.7–70.7 (63.8); PNI: 59.6–69.4 (66.4); MTI: 67.7–70.3 (69.0).

Head. Cephalic margin indistinctly concave to straight (Fig. 29A). Pilosity relatively dense, moderately long, decumbent to suberect. Sculpture shiny and smooth; lateral sides of frons with fine, sparse, longitudinal, and interrupted rugulae; vertex with indistinct, sparse, and transverse rugulae; antennal sockets with few thick, curved outward rugae and smooth interspaces. Clypeus with median longitudinal carina absent; two lateral longitudinal carinae absent. Scape, when laid back, reaching the posterior head margin; pilosity dense, subdecumbent to erect (Fig. 29A, C). Mesosoma. In lateral view, promesonotum moderately high and short, arched; promesonotal groove absent; metanotal groove distinct; propodeal spines small and triangular (Fig. 29C). Sculpture shiny and smooth. Pilosity moderately sparse, moderately long, and erect (Fig. 29C, E). Gaster. With sparse, erect pilosity (Fig. 29C, E). Colour. Yellow, head and gaster slightly darker (Fig. 29C, E).

Latin for bright and small in reference to small body size and bright colouration.

The species was collected between 355–520 m in elevation, in rainforest. Nests were located in rotten logs, and under rootmats on the ground.

Pheidole flavominuta sp. nov. belongs to the group of species characterised by small body size (major workers: HL < 1.05 mm, WL < 0.9 mm and minor workers HL < 0.5 mm, WL < 0.6 mm), head in major workers elongate and not widening posteriorly, minor workers with predominantly smooth and relatively rectangular head, and yellow to brown body colouration. The group includes four species: P. flavominuta sp. nov., P. nitidobruna sp. nov., P. mikros sp. nov., and P. beanka sp. nov. Distribution of P. flavominuta sp. nov. covers area north from Toamasina up to Antenina and doesn’t overlap with other members of this group. Pheidole flavominuta sp. nov. strongly differs from other members of the group by having distinctly yellow body colouration in major and minor workers. Additionally, major workers have head, in lateral view, sub-oval and elongate with slightly convex dorsal and ventral sides, and first gastral tergite is entirely smooth. Minor workers of this species can be also separated by a short antennal scape which doesn’t exceed the posterior head margin.

Holotype. Madagascar. • 1 major worker; Antsiranana; Parc National Montagne d’Ambre, 12.2 km 211°SSW Joffreville; -12.59639, 49.1595; alt. 1300 m; 2 Feb 2001; Fisher et al. leg.; montane rainforest, ex dead branch above ground; BLF02825; CASENT0403308, middle specimen on the pin (CASC). Paratypes. • 6w., 4s.; same data as for holotype, CASENT0403307, CASENT0872252, CASENT0403309, CASENT0403310 (CASC, MHNG, PBZT).

Figure 30. 

Pheidole gracilis sp. nov., full-face view (A), profile (C), and dorsal view (E) of paratype minor worker (CASENT0403309) and full-face view (B), profile (D), and dorsal view (F) of holotype major worker (CASENT0403308).

Madagascar. –Antsiranana: •2w., 3s.; Parc National Montagne d’Ambre, 12.2 km 211°SSW Joffreville; -12.59639, 49.1595; alt. 1300 m; 2 Feb 2001; Fisher et al. leg.; montane rainforest, ex dead branch above ground; BLF02844 (CASC).

Moderately large species. Major workers. Head in full-face view sub-oval and not widening posteriorly, with anterior and posterior sides slightly convex, in lateral view sub-oval; ventral and dorsal faces convex; sides of the head with dense, short, decumbent to suberect pilosity; medial part of frons with moderately thick, longitudinal, interrupted, and moderately dense rugae, interspaces rugulate, rugae directed slightly outward on posteromedial part; occipital lobes predominantly smooth, only anterior part with indistinct, dense, thin, and longitudinal rugae; area posterolateral from eyes with indistinct, dense, thin, and longitudinal rugae, posteriormost part smooth; scape, when laid back, exceeding the midlength of head by approximately two-fifths of its length; inner hypostomal teeth distinct, low, closely spaced, triangular, with rounded apex directed upward; outer hypostomal teeth dentate, wider and higher then inner teeth; inner and outer hypostomal teeth closely spaced and connected by indistinct concavity; mesosoma distinctly foveolate; promesonotum with additional sparse and moderately thick, irregular to transverse rugae on dorsum; katepisternum with smooth notch; body yellowish orange. Minor workers. Head foveolate; vertex with additional, short, and transverse rugulae; area posterolateral from eyes with weaker sculpture that appears smooth; scape, when laid back, exceeding the posterior head margin by two-fifths of its length; promesonotum low and moderately long; promesonotal groove present; propodeal spines moderately large with narrow base; mesosoma foveolate; katepisternum with smooth notch; body yellow to orange.

Major workers. Measurements (N = 8): HL: 1.22–1.32 (1.27); HW: 1.22–1.33 (1.27); SL: 0.72–0.82 (0.77); EL: 0.16–0.19 (0.17); WL: 1.08–1.22 (1.15); PSL: 0.16–0.21 (0.18); MTL: 0.76–0.83 (0.79); PNW: 0.48–0.53 (0.51); PTW: 0.14–0.17 (0.16); PPW: 0.3–0.37 (0.33); CI: 97.3–103.5 (99.8); SI: 56.7–63.3 (60.7); PSLI: 12.8–16.9 (14.6); PPI: 43.1–55.8 (49.0); PNI: 38.8–41.4 (39.8); MTI: 59.3–65.7 (62.1).

Head. In full-face sub-oval, not widening posteriorly, with anterior and posterior sides slightly convex (Fig. 30B). In lateral view sub-oval; ventral and dorsal faces convex; inner hypostomal teeth visible. Sides of the head with dense, short, decumbent to suberect pilosity; whole head with dense, long, decumbent to erect pilosity. Medial part of frons with moderately thick, longitudinal, interrupted, and moderately dense rugae, interspaces rugulate, rugae directed slightly outward on posteromedial part; lateral sides with thick, dense, and longitudinal rugae with distinctly rugulate interspaces. Occipital lobes predominantly smooth, only anterior part with indistinct, dense, thin, and longitudinal rugae. Area posterolateral from eyes with indistinct, dense, thin, and longitudinal rugae, posteriormost part smooth and shiny. Gena with relatively sparse and thick longitudinal rugae and smooth to indistinctly rugulate interspaces. Centre of clypeus smooth and shiny, lateral sides with indistinct rugulae; median notch present, moderately wide, and shallow; median longitudinal carina absent; lateral longitudinal carinae absent. Scape, when laid back, exceeding the midlength of head by two-fifths of its length; pilosity subdecumbent to erect (Fig. 30B, D). Inner hypostomal teeth distinct, low, closely spaced, triangular, with rounded apex directed upward; outer hypostomal teeth dentate, wider and higher than inner teeth; inner and outer hypostomal teeth closely spaced and connected by indistinct concavity (Fig. 63L). Mesosoma. In lateral view, promesonotum short, angular, and moderately low, posterior mesonotum moderately steep, mesonotal process indistinct, tubercle-like; promesonotal groove absent; metanotal groove absent; propodeal spines moderately long, with narrow base and acute apex; humeral area weakly produced (Fig. 30D). Surface shiny and distinctly foveolate; promesonotum with additional sparse and moderately thick, irregular to transverse rugae on dorsum; katepisternum with smooth notch. Pilosity sparse, long, and erect (Fig. 30D, F). Petiole. Shiny with sparse foveolae; node finely foveolate, triangular, with rounded and thick apex, in rear view node dorsoventrally slightly concave; pilosity moderately sparse and erect (Fig. 30D, F). Postpetiole. Shiny and foveolate; dorsum with reduced sculpture and smooth notch; in dorsal view oval, lateral margins medially with two wide and dentate projections; pilosity long, moderately sparse, and erect (Fig. 30D, F). Gaster. Shiny and smooth; pilosity moderately sparse, moderately long, and erect (Fig. 30D, F). Colour. Yellowish orange with yellow legs (Fig. 30D, F).

Minor workers. Measurements (N = 10): HL: 0.65–0.71 (0.69); HW: 0.55–0.58 (0.56); SL: 0.69–0.74 (0.71); EL: 0.12–0.14 (0.13); WL: 0.84–0.89 (0.86); PSL: 0.1–0.12 (0.11); MTL: 0.58–0.66 (0.61); PNW: 0.36–0.4 (0.38); PTW: 0.07–0.1 (0.09); PPW: 0.15–0.19 (0.16); CI: 118.3–123.9 (121.5); SI: 123.7–132.8 (126.8); PSLI: 15.0–17.8 (16.3); PPI: 49.7–66.0 (57.0); PNI: 65.1–69.6 (67.4); MTI: 103.8–112.7 (108.9).

Head. Cephalic margin slightly convex (Fig. 30A). Pilosity relatively sparse, moderately long, decumbent to subdecumbent. Sculpture shiny and foveolate; vertex with additional, short, and transverse rugulae; area posterolateral from eyes with weaker sculpture that appears smooth; antennal sockets with few thick, curved outward rugae and foveolate interspaces. Clypeus with median longitudinal carina absent; two lateral longitudinal carinae absent. Scape, when laid back, exceeding the posterior head margin by two-fifths of its length; pilosity dense, suberect to erect (Fig. 30A, C). Mesosoma. In lateral view, promesonotum low and moderately long, arched; promesonotal groove present; metanotal groove distinct; propodeal spines moderately large with narrow base, triangular (Fig. 30C). Sculpture shiny and foveolate; katepisternum with smooth notch. Pilosity very sparse, moderately long, and erect (Fig. 30C, E). Gaster. With sparse, erect pilosity (Fig. 30C, E). Colour. Orange to yellow (Fig. 30C, E).

Latin for slender in reference to body sculpture.

The species was collected at 1300 m in elevation, in montane rainforest. Nests were located in dead branches above ground.

Pheidole gracilis sp. nov. is a member of a group of species characterised by distinctly reduced head sculpture in major workers with occipital lobes entirely or predominantly smooth, area posterolateral from eyes partially or entirely smooth and shiny or with reduced sculpture and smooth notches. The group consists of four species: P. litigiosa, P. masoandro sp. nov., P. gracilis sp. nov., and P. tampony sp. nov. Pheidole gracilis sp. nov. is known only from Parc National Montagne d’Ambre in Antsiranana and its distribution doesn’t overlap with other members of this group. Morphologically the species is most similar to P. litigiosa, known from the vicinity of Antananarivo, and P. masoandro sp. nov., described from Anosyenne Mts. in Toliara. Its major workers can be easily separated from both of those taxa based on more developed head sculpture with occipital lobes anteriorly covered by indistinct, dense, thin, and longitudinal rugae and distinctly foveolate promesonotum; minor workers can be separated based on foveolate mesosoma and low and long promesonotum. Minor workers of P. gracilis sp. nov. are similar to minors of P. tampony sp. nov. and P. befotaka sp. nov. but can be separated based on more developed head sculpture with additional, short, and transverse rugulae on vertex and area posterolateral from eyes with weak but distinct sculpture and lacking smooth notches.

Holotype. Madagascar. • 1 major worker; Fianarantsoa; Forêt d’Atsirakambiaty, 7.6 km 285°WNW Itremo; -20.59333, 46.56333; alt. 1550 m; 22 Jan 2003; Fisher et al. leg.; montane rainforest, ex rotten log; BLF07262; CASENT0491605, bottom specimen on the pin (CASC). Paratypes. • 3w., 7s.; same data as for holotype, CASENT0491603, CASENT0491604, CASENT0872253, CASENT0491601 (CASC, MHNG, PBZT).

Figure 31. 

Pheidole haboka sp. nov., full-face view (A), profile (C), and dorsal view (E) of paratype minor worker (CASENT0491601) and full-face view (B), profile (D), and dorsal view (F) of holotype major worker (CASENT0491605).

Madagascar. –Fianarantsoa: • 3w., 3s.; Forêt d’Atsirakambiaty, 7.6 km 285°WNW Itremo; -20.59333, 46.56333; alt. 1550 m; 22 Jan 2003; Fisher et al. leg.; montane rainforest, ex rotten log; BLF07170 (CASC). • 1w.; Forêt d’Atsirakambiaty, 7.6 km 285°WNW Itremo; -20.59333, 46.56333; alt. 1550 m; 22 Jan 2003; Fisher et al. leg.; montane rainforest, ex rotten log; BLF07182 (CASC). • 3w.; Forêt d’Atsirakambiaty, 7.6 km 285°WNW Itremo; -20.59333, 46.56333; alt. 1550 m; 22 Jan 2003; Fisher et al. leg.; montane rainforest, under stone; BLF07184 (CASC). • 1w.; Forêt d’Atsirakambiaty, 7.6 km 285°WNW Itremo; -20.59333, 46.56333; alt. 1550 m; 22 Jan 2003; Fisher et al. leg.; montane rainforest, ex rotten log; BLF07249 (CASC). • 3w.; Forêt d’Atsirakambiaty, 7.6 km 285°WNW Itremo; -20.59333, 46.56333; alt. 1550 m; 22 Jan 2003; Fisher et al. leg.; montane rainforest, ex rotten log; BLF07250 (CASC).

Moderately large species. Major workers. Head in full-face view sub-oval, not widening posteriorly, with anterior and posterior sides relatively straight, in lateral view sub-oval; ventral and dorsal faces convex; sides of the head with dense, moderately long, suberect to erect pilosity; medial part of frons with thick, longitudinal, and dense rugae, interspaces predominantly smooth to indistinctly rugulate, rugae more irregular and directed slightly outward on posteromedial part; lateral sides with thick, dense, and longitudinal rugae with sparsely rugulate interspaces, rugae more irregular on posterolateral parts; occipital lobes without smooth notches; area posterolateral from eyes with sculpture weakening posteriorly and smooth notches; scape, when laid back, exceeding the midlength of head by two-fifths of its length; inner hypostomal teeth distinct, small, closely spaced, triangular, with rounded apex directed upward; outer hypostomal teeth lobe-like, wider than inner teeth, and approximately the same height; inner and outer hypostomal teeth closely spaced and not connected by concavity; mesosoma foveolate; promesonotum with additional thin, moderately sparse, transverse to irregular rugae; katepisternum with smooth notch; gaster smooth; body orange. Minor workers. Head smooth; lateral sides of frons with longitudinal, short, and thick rugae; vertex with very short, sparse, and transverse to arcing rugae; scape, when laid back, exceeding the posterior head margin by two-fifths of its length; promesonotum moderately low and short, arched; promesonotal groove absent; propodeal spines very small, triangular; mesosoma smooth; dorsal promesonotum with very sparse and irregular rugae; propodeum with sparse, irregular, and indistinct rugulae; body yellow.

Major workers. Measurements (N = 8): HL: 0.94–1.13 (1.06); HW: 0.97–1.13 (1.06); SL: 0.66–0.72 (0.68); EL: 0.12–0.14 (0.13); WL: 1.05–1.13 (1.08); PSL: 0.15–0.19 (0.17); MTL: 0.65–0.7 (0.67); PNW: 0.52–0.58 (0.55); PTW: 0.14–0.17 (0.15); PPW: 0.29–0.34 (0.31); CI: 97.0–101.9 (99.6); SI: 59.5–69.1 (64.5); PSLI: 13.7–19.3 (16.0); PPI: 46.5–53.6 (50.1); PNI: 48.2–58.5 (52.2); MTI: 59.4–70.8 (63.7).

Head. In full-face view elongate, not widening posteriorly, with anterior and posterior sides relatively straight (Fig. 31B). In lateral view sub-oval; ventral and dorsal faces convex; inner hypostomal teeth visible. Sides of the head with dense, moderately long, suberect to erect pilosity; whole head with dense, long, decumbent to erect pilosity. Medial part of frons with thick, longitudinal, and dense rugae, interspaces predominantly smooth to indistinctly rugulate, rugae more irregular and directed slightly outward on posteromedial part; lateral sides with thick, dense, and longitudinal rugae with sparsely rugulate interspaces, rugae more irregular on posterolateral parts. Occipital lobes with dense, irregular rugae and sparsely rugulate interspaces. Area posterolateral from eyes with weaker and more irregular sculpture, sculpture weakening posteriorly and smooth on the posteriormost part. Gena with relatively sparse and thick longitudinal rugae and distinctly rugulate interspaces. Centre of clypeus smooth and shiny, lateral sides with indistinct rugulae; median notch present, moderately wide, and shallow; median longitudinal carina absent; lateral longitudinal carinae absent. Scape, when laid back, exceeding the midlength of head by two-fifths of its length; pilosity subdecumbent to erect (Fig. 31B, D). Inner hypostomal teeth distinct, small, closely spaced, triangular, with rounded apex directed upward; outer hypostomal teeth lobe-like, wider than inner teeth and approximately the same height; inner and outer hypostomal teeth closely spaced and not connected by concavity (Fig. 63M). Mesosoma. In lateral view, promesonotum short, angular, and moderately low, posterior mesonotum moderately steep, mesonotal process indistinct, tubercle-like; promesonotal groove absent; metanotal groove indistinct; propodeal spines short, with wide base and acute apex; humeral area weakly produced (Fig. 31D). Surface shiny and foveolate; promesonotum with additional thin, moderately sparse, transverse to irregular rugae; katepisternum with smooth notch. Pilosity dense, long, and erect (Fig. 31D, F). Petiole. Shiny with sparse foveolae; node finely foveolate, triangular, with rounded and thick apex, in rear view node dorsoventrally convex; pilosity moderately sparse and erect (Fig. 31D, F). Postpetiole. Shiny and foveolate; dorsum smooth; in dorsal view oval, lateral margins medially with two dentate projections; pilosity long, moderately sparse, and erect (Fig. 31D, F). Gaster. Shiny and smooth; pilosity very dense, moderately long, and erect (Fig. 31D, F). Colour. Orange with yellow legs (Fig. 31D, F).

Minor workers. Measurements (N = 3): HL: 0.61–0.64 (0.62); HW: 0.53–0.55 (0.54); SL: 0.59–0.63 (0.61); EL: 0.08–0.09 (0.09); WL: 0.78–0.81 (0.79); PSL: 0.08–0.1 (0.08); MTL: 0.52–0.56 (0.54); PNW: 0.37–0.39 (0.38); PTW: 0.09–0.1 (0.09); PPW: 0.16–0.16 (0.16); CI: 113.3–117.9 (115.4); SI: 111.9–115.5 (113.1); PSLI: 12.2–15.4 (13.6); PPI: 56.1–61.0 (58.5); PNI: 70.1–70.7 (70.4); MTI: 98.5–101.4 (99.6).

Head. Cephalic margin slightly concave (Fig. 31A). Pilosity relatively dense, moderately long, decumbent to subdecumbent. Sculpture shiny and smooth; lateral sides of frons with longitudinal, short and thick rugae; vertex with very short, sparse, and transverse to arcing rugae; antennal sockets with few thick, curved outward rugae and smooth interspaces. Clypeus with median longitudinal carina absent; two lateral longitudinal carinae absent. Scape, when laid back, exceeding the posterior head margin by two-fifths of its length; pilosity dense, suberect to erect (Fig. 31A, C). Mesosoma. In lateral view, promesonotum moderately low and short, arched; promesonotal groove absent; metanotal groove distinct; propodeal spines very small, triangular (Fig. 31C). Sculpture shiny and smooth; dorsal promesonotum with very sparse and irregular rugae; propodeum with sparse, irregular, and indistinct rugulae. Pilosity sparse, moderately long, and erect (Fig. 31C, E). Gaster. With moderately dense, erect pilosity (Fig. 31C, E). Colour. Yellow (Fig. 31C, E).

Malagasy for orange in reference to body colouration.

The species was collected at 1550 m in elevation, in montane rainforest. Nests were located in rotten logs and under stones.

Pheidole haboka sp. nov., described from Forêt d’Atsirakambiaty in Fianarantsoa, is most similar to P. sparsa sp. nov. and P. dasos sp. nov., both known from the northern part of the island, and P. hazo sp. nov., recorded so far only from the vicinity of Andranoma in Antananarivo. Majors of P. haboka sp. nov. can be easily distinguished from all taxa mentioned above by area posterolateral from eyes with sculpture weakening posteriorly and smooth notches and very dense pilosity on gaster; minors can be separated based on body sculpture predominantly smooth and lacking foveolae.

Holotype. Madagascar. • 1 major worker; Toliara; Anosy Region, Anosyenne Mts, 31.2 km NW Manantenina; -24.13894, 47.06804; alt. 1125 m; 26 Feb 2015; B. L. Fisher et al. leg.; rainforest, ex root mat; BLF36492; CASENT0704280 (CASC). Paratypes. • 2w., 1m.; same data as for holotype; CASENT0923290, CASENT0704279 (CASC, MHNG).

Figure 32. 

Pheidole havoana sp. nov., full-face view (A), profile (C), and dorsal view (E) of paratype minor worker (CASENT0923290) and full-face view (B), profile (D), and dorsal view (F) of holotype major worker (CASENT0704280).

Madagascar. –Toliara: • 2w., 1s., 1m.; Anosy Region, Anosyenne Mts, 31.2 km NW Manantenina; -24.13894, 47.06804; alt. 1125 m; 26 Feb 2015; B. L. Fisher et al. leg.; rainforest, under stone; BLF36463 (CASC). • 2w.,1s., 1q.; Anosy Region, Anosyenne Mts, 31.2 km NW Manantenina; -24.13894, 47.06804; alt. 1125 m; 26 Feb 2015; B. L. Fisher et al. leg.; rainforest, under root mat on rock; BLF36477 (CASC). •2 w., 1s., 1m.; Anosy Region, Anosyenne Mts, 31.2 km NW Manantenina; -24.13894, 47.06804; alt. 1125 m; 26 Feb 2015; B. L. Fisher et al. leg.; rainforest, ex root mat; BLF36496 (CASC). • 1w., 1s.; Anosy Region, Anosyenne Mts, 31.2 km NW Manantenina; -24.13894, 47.06804; alt. 1125 m; 26 Feb 2015; B. L. Fisher et al. leg.; rainforest, ex rotten log; BLF36561 (CASC).

Minute species. Major workers. HL < 1.1 mm and WL < 0.9 mm; head, in full-face view, sub-oval, slightly widening posteriorly, with anterior and posterior sides convex, in lateral view sub-oval; ventral and dorsal faces convex; body dark orange; sides of head with moderately dense, moderately long, subdecumbent to suberect pilosity; entire head distinctly sculptured, medial part of frons with longitudinal rugae, rugae in posteromedial part more irregular, interspaces shiny with dense, distinct, and irregular rugulae; scape, when laid back, slightly exceeding the midlength of head; mesosoma predominantly rugofoveolate, pronotal dorsum and lateral sides of propodeum with reduced sculpture; inner hypostomal teeth distinct, moderately high, closely spaced, triangular, with rounded apex directed upward; outer hypostomal teeth lobe-like, wider and higher than inner hypostomal teeth, apex directed outward; inner and outer hypostomal teeth closely spaced, connected by indistinct concavity; base of first gastral tergite smooth. Minor workers. HL < 0.5 mm and WL < 0.6 mm, scape, when laid back, surpassing the posterior head margin by one-fifth of its length; propodeal spines very small, triangular; head relatively rectangular; body yellow; head foveolate, frons with additional indistinct longitudinal and interrupted rugae in medial part, area posterolateral from eyes smooth; mesosoma smooth.

Major workers. Measurements (N = 4): HL: 0.99–1.06 (1.02); HW: 1.0–1.05 (1.0); SL: 0.55–0.56 (0.56); EL: 0.12–0.12 (0.12); WL: 0.85–0.91 (0.88); PSL: 0.18–0.2 (0.19); MTL: 0.53–0.55 (0.54); PNW: 0.46–0.48 (0.47); PTW: 0.11–0.16 (0.14); PPW: 0.24–0.31 (0.27); CI: 98.7–103.1 (101.2); SI: 53.4–56.7 (55.2); PSLI: 17.1–20.5 (18.6); PPI: 40.6–60.4 (52.2); PNI: 45.3–48.9 (46.3); MTI: 52.3–56.0 (53.7).

Head. In full-face view sub-oval, slightly widening posteriorly, with anterior and posterior sides convex (Fig. 32B). In lateral view sub-oval; ventral and dorsal faces convex; inner hypostomal teeth visible. Sides of the head with moderately dense, moderately long, subdecumbent to suberect pilosity; whole head with dense, long, decumbent to erect pilosity. Medial part of frons with moderately dense, thick, longitudinal rugae, rugae in posteromedial part more irregular and directed outward, interspaces shiny with dense and distinct irregular rugulae; lateral sides with longitudinal to irregular and thick rugae with distinctly rugofoveolate interspaces. Occipital lobes with sparse, thick, and irregular rugae; interspaces smooth to indistinctly rugulate. Gena with relatively dense and thick longitudinal rugae and distinctly rugofoveolate interspaces. Area posterolateral from eyes with dense and thick rugofoveolae. Centre of clypeus smooth and shiny, lateral sides with indistinct rugulae; median notch present, moderately wide, and shallow; median longitudinal carina present; lateral longitudinal carinae absent. Scape, when laid back, slightly exceeding the midlength of head; pilosity decumbent to erect (Fig. 32B, D). Inner hypostomal teeth distinct, moderately high, closely spaced, triangular, with rounded apex directed upward; outer hypostomal teeth lobe-like, wider and higher than inner hypostomal teeth, apex directed outward; inner and outer hypostomal teeth closely spaced, connected by indistinct concavity (Fig. 63N). Mesosoma. In lateral view, promesonotum short, angular, and moderately high, posterior mesonotum moderately steep, mesonotal process indistinct, tubercle-like; promesonotal groove absent; metanotal groove absent; propodeal spines moderately long, moderately wide, with acute apex; humeral area laterally weakly produced (Fig. 32D). Surface shiny and predominantly rugofoveolate, pronotal dorsum and lateral sides of propodeum with reduced sculpture. Pilosity relatively dense, long, and erect (Fig. 32D, F). Petiole. Shiny with fine and dense rugofoveolae; node partially smooth, low, triangular, with rounded and thin apex, in rear view node dorsoventrally slightly concave; pilosity moderately sparse and erect (Fig. 32D, F). Postpetiole. Shiny and smooth; in dorsal view oval, lateral margins medially with two dentate projections; pilosity long, moderately sparse, and erect (Fig. 32D, F). Gaster. Shiny and smooth; pilosity moderately dense, long, and erect (Fig. 32D, F). Colour. Dark orange; legs yellow (Fig. 32D, F).

Minor workers. Measurements (N = 5): HL: 0.52–0.53 (0.52); HW: 0.44–0.47 (0.45); SL: 0.51–0.53 (0.52); EL: 0.09–0.11 (0.09); WL: 0.6–0.62 (0.61); PSL: 0.05–0.08 (0.07); MTL: 0.39–0.41 (0.4); PNW: 0.28–0.33 (0.3); PTW: 0.06–0.09 (0.07); PPW: 0.12–0.13 (0.12); CI: 112.2–118.6 (115.6); SI: 112.2–115.8 (114.1); PSLI: 10.3–14.3 (12.9); PPI: 50.4–70.5 (58.2); PNI: 63.4–69.3 (66.7); MTI: 87.2–90.2 (88.4).

Head. Cephalic margin indistinctly concave or straight (Fig. 32A). Pilosity relatively sparse, long, decumbent to subdecumbent. Sculpture foveolate; frons with additional indistinct longitudinal and interrupted rugae in medial part; area posterolateral from eyes smooth. 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 dense, subdecumbent to erect (Fig. 32A, C). Mesosoma. In lateral view, promesonotum moderately low and short, arched; promesonotal groove absent; metanotal groove present and distinct; propodeal spines very small and triangular (Fig. 32C). Sculpture smooth. Pilosity very sparse, moderately long, and erect (Fig. 32C, E). Postpetiole. Short, low, and relatively flat; with few short, erect setae (Fig. 32C, E). Gaster. With sparse, erect pilosity (Fig. 32C, E). Colour. Yellow, vertex slightly darker (Fig. 32C, E).

Malagasy for hill or mountain in reference to type locality.

The species was collected at 1125 m in elevation, in rainforest. Nests were located in rotten logs under stone and in root mats.

Pheidole havoana sp. nov. belongs to the group of species characterised by small body size (major workers: HL < 1.05 mm, WL < 0.9 mm and minor workers HL < 0.5 mm, WL < 0.6 mm), head sub-oval, slightly widening posteriorly with anterior and posterior sides convex in major workers, and minor workers with yellow to brown body colouration and head foveolate or predominantly smooth and relatively oval. The group includes six species: P. havoana sp. nov., P. kely sp. nov., P. parvula sp. nov., P. parvulogibba sp. nov., P. volontany sp. nov., and P. midongy sp. nov. Within this group, P. havoana sp. nov., described form Anosyenne Mts. in Toliara, is most similar to P. kely sp. nov., distributed in the northern part of the island, P. parvula sp. nov., known from two localities in the vicinity of Antananarivo, and sympatric P. parvulogibba sp. nov. Minor workers of P. havoana sp. nov. distinctly differ from those of P. parvula sp. nov. and P. parvulogibba sp. nov. by predominantly foveolate head and presence of additional indistinct longitudinal and interrupted rugae in medial part of frons. Minor workers of P. havoana sp. nov. and P. kely sp. nov. are indistinguishable. Major workers of P. havoana sp. nov. differ from P. kely sp. nov. and P. parvulogibba sp. nov. by medial part of frons with longitudinal rugae, rugae in posteromedial part more irregular, with densely and distinctly irregular rugulae in interspaces; and from P. parvula sp. nov. in reduced sculpture on pronotal dorsum and lateral sides of propodeum.

Holotype. Madagascar. • 1 major worker; Antananarivo; 3 km 41°NE Andranomay, 11.5 km 147°SSE Anjozorobe; -18.47333, 47.96; alt. 1300 m; 5 Dec 2000; Fisher et al. leg.; montane rainforest, ex dead branch above ground; BLF02441; CASENT0427790, bottom specimen on the pin (CASC). Paratypes. • 5w., 8s.; same data as for holotype; CASENT0427762, CASENT00872251, CASENT0413711, CASENT0413712, CASENT0427765 (CASC, MHNG, PBZT).

Figure 33. 

Pheidole hazo sp. nov., full-face view (A), profile (C), and dorsal view (E) of paratype minor worker (CASENT0427762) and full-face view (B), profile (D), and dorsal view (F) of holotype major worker (CASENT0427790).

Moderately large species. Major workers. Head in full-face view sub-oval, not widening posteriorly, with anterior and posterior sides relatively straight, in lateral view sub-oval; ventral and dorsal faces convex; sides of the head with dense, long, suberect to erect pilosity; medial part of frons with moderately dense, thin, longitudinal rugae, rugae in posteromedial portion directed slightly outward, interspaces shiny with sparse and distinct rugofoveolae; lateral sides with longitudinal, thicker, and relatively sparser rugae; interspaces shiny with relatively dense and distinct rugofoveolae; occipital lobes, and area posterolateral from eyes without smooth notches; scape, when laid back, exceeding the midlength of head by two-fifths of its length; inner hypostomal teeth distinct, large, closely spaced, triangular, with rounded apex directed slightly outward; outer hypostomal teeth lobe-like, wider than inner hypostomal teeth and approximately the same height, apex directed outward; inner and outer hypostomal teeth closely spaced and connected by concavity; mesosoma with thick and sparse foveolae; pronotum with additional transverse to irregular, thin rugae; katepisternum with smooth notch; gaster smooth; body orange. Minor workers. Head foveolate with additional longitudinal to irregular, interrupted, and moderately thick rugae; area posterolateral from eyes with weaker foveolae and smooth notches; scape, when laid back, surpassing the posterior head margin by one-fifth of its length; promesonotum moderately low and moderately long, arched; promesonotal groove absent; propodeal spines minute and triangular; mesosoma with thick foveolae; lateral sides of propodeum, katepisternum, and anepisternum with smooth notches; body yellow.

Major workers. Measurements (N = 9): HL: 1.06–1.14 (1.09); HW: 1.02–1.13 (1.08); SL: 0.63–0.66 (0.64); EL: 0.14–0.17 (0.15); WL: 0.98–1.08 (1.03); PSL: 0.16–0.18 (0.17); MTL: 0.63–0.65 (0.64); PNW: 0.41–0.52 (0.47); PTW: 0.13–0.15 (0.14); PPW: 0.25–0.3 (0.27); CI: 97.6–104.3 (101.0); SI: 56.4–63.3 (59.7); PSLI: 15.1–17.1 (16.0); PPI: 47.3–55.8 (51.5); PNI: 39.2–47.0 (43.3); MTI: 57.1–63.9 (59.9).

Head. In full-face view sub-oval, not widening posteriorly, with anterior and posterior sides relatively straight (Fig. 33B). In lateral view sub-oval; ventral and dorsal faces convex; inner hypostomal teeth visible. Sides of the head with dense, long, suberect to erect pilosity; whole head with dense, long, decumbent to erect pilosity. Medial part of frons with moderately dense, thin, longitudinal rugae, rugae in posteromedial directed slightly outward, interspaces shiny with sparse and distinct rugofoveolae; lateral sides with longitudinal, thicker, and relatively sparser rugae; interspaces shiny with relatively dense and distinct rugofoveolae. Occipital lobes with sparse, thick, and longitudinal to irregular rugae; interspaces rugofoveolate. Gena with relatively dense and thick, longitudinal rugae and rugofoveolate interspaces. Area posterolateral from eyes shiny, with dense, longitudinal rugofoveolae. Centre of clypeus smooth and shiny, lateral sides with indistinct rugulae; median notch present, moderately wide, and shallow; median longitudinal carina absent; lateral longitudinal carinae absent. Scape, when laid back, exceeding the midlength of head by two-fifths of its length; pilosity subdecumbent to erect (Fig. 33B, D). Inner hypostomal teeth distinct, large, closely spaced, triangular, with rounded apex directed slightly outward; outer hypostomal teeth lobe-like, wider than inner hypostomal teeth and approximately the same height, apex directed outward; inner and outer hypostomal teeth closely spaced and connected by concavity (Fig. 63O). Mesosoma. In lateral view, promesonotum short, angular, and moderately low, posterior mesonotum moderately steep, mesonotal process indistinct, tubercle-like; promesonotal groove absent; metanotal groove absent; propodeal spines moderately long, moderately narrow, with acute apex; humeral area laterally weakly produced (Fig. 33D). Surface shiny with thick and sparse foveolae; pronotum with additional transverse to irregular, thin rugae; katepisternum with smooth notch. Pilosity sparse, long, and erect (Fig. 33D, F). Petiole. Shiny with fine and sparse foveolae; node smooth to finely foveolate, low, triangular, with rounded and thin apex, in rear view node dorsoventrally straight to slightly convex; pilosity moderately sparse and erect (Fig. 33D, F). Postpetiole. Shiny and smooth; in dorsal view oval, lateral margins medially with two dentate projections; pilosity long, moderately sparse, and erect (Fig. 33D, F). Gaster. Shiny and smooth; pilosity moderately dense, long, and erect (Fig. 33D, F). Colour. Dark orange; mandibles slightly darker; legs yellow (Fig. 33D, F).

Minor workers. Measurements (N = 5): HL: 0.6–0.63 (0.61); HW: 0.5–0.53 (0.52); SL: 0.61–0.64 (0.62); EL: 0.11–0.12 (0.12); WL: 0.76–0.82 (0.79); PSL: 0.08–0.09 (0.09); MTL: 0.52–0.55 (0.54); PNW: 0.33–0.35 (0.34); PTW: 0.09–0.1 (0.09); PPW: 0.14–0.15 (0.15); CI: 114.8–121.9 (118.8); SI: 115.4–123.9 (120.9); PSLI: 12.9–14.6 (13.9); PPI: 57.3–70.4 (62.7); PNI: 63.4–67.4 (66.0); MTI: 101.5–106.2 (103.8).

Head. Cephalic margin indistinctly convex or straight (Fig. 33A). Pilosity relatively sparse, long, decumbent to suberect. Sculpture foveolate with additional longitudinal to irregular, interrupted, and moderately thick rugae; area posterolateral from eyes with weaker foveolae and smooth notches. 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 dense, subdecumbent to erect (Fig. 33A, C). Mesosoma. In lateral view, promesonotum moderately low and moderately long, arched; promesonotal groove absent; metanotal groove distinct; propodeal spines minute and triangular (Fig. 33C). Sculpture shiny and with thick foveolae; lateral sides of propodeum, katepisternum, and anepisternum with smooth notches. Pilosity very sparse, moderately long, and erect (Fig. 33C, E). Postpetiole. Short, low, and relatively flat; with few short, erect setae (Fig. 33C, E). Gaster. With sparse, erect pilosity (Fig. 33C, E). Colour. Yellow, vertex slightly darker (Fig. 33C, E).

Malagasy for tree in reference to arboreal nesting preferences of the species.

The species was collected at 1300 m in elevation, in montane rainforest. Nest was located in dead branch above ground.

Pheidole hazo sp. nov., described from the vicinity of Andranoma in Antananarivo, is most similar to P. dasos sp. nov., known from Makirovana forest in Antsiranana, and P. sparsa sp. nov., recorded so far only from Bemanevika in Mahajanga. Majors of P. hazo sp. nov. can be distinguished from both taxa by medial part of frons with moderately dense rugae with interspaces with distinct rugofoveolae, and pronotum lacking smooth notches; minors of P. hazo sp. nov. can be distinguished from P. sparsa sp. nov. by yellow body colouration, and mesosoma predominantly with dense foveolae, and from P. dasos sp. nov. by yellow body colouration, shorter scape surpassing the posterior head margin by one-fifth of its length, and pronotum with distinct and dense foveolae.

Holotype. Madagascar. • 1 major worker; Fianarantsoa; Forêt d’Atsirakambiaty, 7.6 km 285°WNW Itremo; -20.59333, 46.56333; alt. 1550 m; 22 Jan 2003; Fisher et al. leg.; montane rainforest, ex rotten log; BLF07251; CASENT0491584, top specimen on the pin (CASC). Paratypes. • 6w., 1s.; same data as for holotype, CASENT0872249, CASENT0491586, CASENT0491585 (CASC, MHNG, PBZT).

Figure 34. 

Pheidole itremo sp. nov., full-face view (A), profile (C), and dorsal view (E) of paratype minor worker (CASENT0491586) and full-face view (B), profile (D), and dorsal view (F) of holotype major worker (CASENT0491584).

Madagascar. –Antananarivo: • 1w., 2s.; Réserve Spéciale d’Ambohitantely, Forêt d Ambohitantely, 20.9 km 72°NE d Ankazobe; -18.22528, 47.28683; alt. 1410 m; 17 Apr 2001; Fisher et al. leg.; montane rainforest, ex root mat, ground layer; BLF03700 (CASC). • 3s.; Réserve Spéciale d’Ambohitantely, Forêt d Ambohitantely, Jardin Botanique, 24.1km 59°NE d Ankazobe; -18.17139, 47.28182; alt. 1620 m; 17 Apr 2001; Fisher et al. leg.; montane rainforest, sifted litter; BLF03720 (CASC). • 2w.; Réserve Speciale d’Ambohitantely; -18.18762, 47.28576; alt. 1580 m; 8 Mar 2012; Fisher et al. leg.; montane forest, ex rotten log; BLF28200 (CASC). • 3w., 1q.; Réserve Speciale d’Ambohitantely; -18.18762, 47.28576; alt. 1580 m; 8 Mar 2012; Fisher et al. leg.; montane forest, ex rotten log; BLF28206 (CASC). • 2w.; Réserve Speciale d’Ambohitantely; -18.18762, 47.28576; alt. 1580 m; 8 Mar 2012; Fisher et al. leg.; montane forest, ex rotten log; BLF28218 (CASC). • 3w., 1q.; Réserve Speciale d’Ambohitantely; -18.18762, 47.28576; alt. 1580 m; 8 Mar 2012; Fisher et al. leg.; montane forest, ex rotten log; BLF28227 (CASC).

Moderately large species. Major workers. Head in full-face view sub-oval, widening posteriorly, with anterior and posterior sides convex, in lateral view sub-oval; ventral and dorsal faces convex; sides of the head with moderately dense, moderately long, suberect to erect pilosity; medial part of frons with thick, dense, longitudinal, and interrupted rugae and smooth interspaces, posteromedial frons with more irregular rugae directed slightly outward; lateral sides with thick and dense rugae, interspaces shiny and smooth or indistinctly rugulate; rugae longitudinal on anterolateral sides and more irregular on posterolateral sides; area posterolateral from eyes without smooth notches; scape, when laid back, exceeding the midlength of head by one-fifth of its length; inner hypostomal teeth distinct, large, closely spaced, triangular, with rounded apex directed upward; outer hypostomal teeth lobe-like, wider than inner teeth and approximately the same height; inner and outer hypostomal teeth closely spaced and connected by indistinct concavity; mesosoma with fine rugofoveolae; pronotum with rugofoveolae sparser and additional thin and transverse rugae; gaster smooth; body brown. Minor workers. Head smooth; posterolateral sides of frons with sparse and indistinct foveolae; scape, when laid back, surpassing the posterior head margin by two-fifths of its length; promesonotum moderately high and short, arched; promesonotal groove absent; propodeal spines minute and triangular; mesosoma smooth and shiny; body yellowish brown.

Major workers. Measurements (N = 6): HL: 1.14–1.24 (1.18); HW: 1.16–1.21 (1.17); SL: 0.64–0.69 (0.66); EL: 0.12–0.14 (0.13); WL: 0.99–1.06 (1.03); PSL: 0.18–0.2 (0.19); MTL: 0.65–0.68 (0.66); PNW: 0.51–0.54 (0.52); PTW: 0.15–0.17 (0.16); PPW: 0.28–0.34 (0.31); CI: 98.3–103.3 (100.8); SI: 53.7–58.8 (56.0); PSLI: 15.5–16.9 (16.1); PPI: 43.0–56.6 (51.5); PNI: 42.7–46.2 (44.3); MTI: 54.6–57.7 (56.6).

Head. In full-face view sub-oval, widening posteriorly, with anterior and posterior sides convex (Fig. 34B). In lateral view sub-oval; ventral and dorsal faces convex; inner hypostomal teeth visible. Sides of the head with moderately dense, moderately long, suberect to erect pilosity; whole head with dense, long, decumbent to erect pilosity. Medial part of frons with thick, dense, longitudinal, and interrupted rugae and smooth interspaces, posteromedial frons with rugae more irregular and directed slightly outward; lateral sides with thick and dense rugae, interspaces shiny and smooth or indistinctly rugulate; rugae longitudinal on anterolateral sides and more irregular on posterolateral sides. Occipital lobes with thinner, irregular rugae and smooth to indistinctly rugoreticulate interspaces. Area posterolateral from eyes with irregular to longitudinal, relatively thick to thin rugulae and rugoreticulate interspaces. Gena with relatively dense, thick, longitudinal rugae and smooth to indistinctly rugoreticulate interspaces. Centre of clypeus smooth and shiny, lateral sides with indistinct rugulae; median notch present, moderately wide, and shallow; median longitudinal carina present; lateral longitudinal carinae absent. Scape, when laid back, exceeding the midlength of head by one-fifth of its length; pilosity subdecumbent to erect (Fig. 34B, D). Inner hypostomal teeth distinct, large, closely spaced, triangular, with rounded apex directed upward; outer hypostomal teeth lobe-like, wider than inner teeth and approximately the same height; inner and outer hypostomal teeth closely spaced and connected by indistinct concavity (Fig. 63P). Mesosoma. In lateral view, promesonotum short, angular, and moderately low, posterior mesonotum moderately steep, mesonotal process very indistinct, tubercle-like; promesonotal groove absent; metanotal groove absent; propodeal spines moderately long, with moderately narrow base and acute apex; humeral area laterally weakly produced (Fig. 34D). Surface shiny with fine rugofoveolae; pronotum with rugofoveolae sparser and additional thin and transverse rugae. Pilosity moderately sparse, long, and erect (Fig. 34D, F). Petiole. Shiny with fine foveolae; node smooth to finely foveolate, triangular, with rounded and thick apex, in rear view node dorsoventrally straight to slightly convex; pilosity moderately sparse and erect (Fig. 34D, F). Postpetiole. Shiny and foveolate; dorsum with reduced sculpture and smooth notch; in dorsal view oval, lateral margins medially with two dentate projections; pilosity long, moderately sparse, and erect (Fig. 34D, F). Gaster. Shiny and smooth; pilosity dense, long, and erect (Fig. 34D, F). Colour. Brown; legs yellow (Fig. 34D, F).

Minor workers. Measurements (N = 10): HL: 0.54–0.59 (0.56); HW: 0.46–0.5 (0.48); SL: 0.59–0.63 (0.6); EL: 0.1–0.13 (0.11); WL: 0.68–0.74 (0.71); PSL: 0.06–0.1 (0.08); MTL: 0.45–0.5 (0.47); PNW: 0.3–0.35 (0.32); PTW: 0.08–0.1 (0.08); PPW: 0.13–0.16 (0.14); CI: 115.7–121.8 (117.9); SI: 120.4–131.4 (125.6); PSLI: 11.8–17.0 (14.0); PPI: 49.4–67.0 (58.5); PNI: 65.8–71.0 (68.2); MTI: 94.5–101.7 (99.2).

Head. Cephalic margin indistinctly convex (Fig. 34A). Pilosity relatively sparse, long, decumbent to suberect. Sculpture shiny and smooth; posterolateral sides of frons with sparse and indistinct foveolae; antennal sockets with few thick, curved outward rugae and smooth interspaces. Clypeus with median longitudinal carina absent; two lateral longitudinal carinae absent. Scape, when laid back, surpassing the posterior head margin by two-fifths of its length; pilosity dense, subdecumbent to erect (Fig. 34A, C). Mesosoma. In lateral view, promesonotum moderately high and short, arched; promesonotal groove absent; metanotal groove distinct; propodeal spines minute and triangular (Fig. 34C). Sculpture shiny and smooth. Pilosity moderately sparse, long, and erect (Fig. 34C, E). Gaster. With sparse, erect pilosity (Fig. 34C, E). Colour. Yellowish brown (Fig. 34C, E).

From the type locality.

The species was collected between 1410–1620 m in elevation, in montane forest and montane rainforest. Nests were located in rotten logs and root mats.

Pheidole itremo sp. nov., known from Forêt d’Atsirakambiaty in Fianarantsoa and Forêt d’Ambohitantely in Antananarivo, is most similar to P. sava sp. nov., described from Parc National de Marojejy in Antsiranana. Major workers of P. itremo sp. nov. can be separated based on darker body colouration, presence of moderately dense, moderately long, suberect to erect pilosity on sides of the head, medial frons with smooth interspaces, finely rugofoveolae mesosoma lacking smooth notches; minor workers can be separated based on predominantly smooth head with posterolateral sides of frons with sparse and indistinct foveolae, and entirely smooth mesosoma.

Holotype. Madagascar. • 1 major worker; Antsiranana; Parc National Montagne d’Ambre, 3.6 km 235°SW Joffreville; -12.53444, 49.1795; alt. 925 m; 20 Jan 2001; Fisher et al. leg.; montane rainforest, ex rotten log; BLF02568; CASENT0406355 (CASC). Paratypes. • 4w., 2s.; same data as for holotype, CASENT0406358, CASENT0406356, CASENT0406352, CASENT0406354, CASENT0406350 (CASC, MHNG).

Figure 35. 

Pheidole joffreville sp. nov., full-face view (A), profile (C), and dorsal view (E) of paratype minor worker (CASENT0406352) and full-face view (B), profile (D), and dorsal view (F) of holotype major worker (CASENT0406355).

Moderately large species. Major workers. Head in full-face view sub-oval and widening slightly posteriorly, with anterior and posterior sides slightly convex, in lateral view sub-oval; ventral and dorsal faces convex; sides of the head with sparse, moderately long, suberect pilosity; anteromedial part of frons with thick, longitudinal, interrupted, and sparse rugae, posteromedial frons with rugae longitudinally irregular, interspaces distinctly rugofoveolate, rugae on posteromedial part directed outward; lateral sides with thick, sparse, and irregular rugae with distinctly rugofoveolate interspaces; occipital lobes, and area posterolateral from eyes without smooth notches; scape, when laid back, exceeding the midlength of head by one-fifth of its length; inner hypostomal teeth distinct, large, closely spaced, triangular, with rounded apex directed upward; outer hypostomal teeth lobe-like, narrower, and approximately as high as inner teeth; inner and outer hypostomal teeth closely spaced and not connected by concavity; mesosoma foveolate; promesonotum with additional sparse, thick, and transverse rugulae on dorsum; gaster smooth; body brown. Minor workers. Head foveolate; vertex with a few arcing, sparse, and moderately thick rugae; scape, when laid back, exceeding the posterior head margin by one-third of its length; promesonotum low and moderately long, arched; promesonotal groove present; propodeal spines moderate, triangular; mesosoma foveolate; katepisternum with big smooth notch; body brown.

Major workers. Measurements (N = 3): HL: 1.04–1.1 (1.06); HW: 1.03–1.16 (1.06); SL: 0.73–0.75 (0.74); EL: 0.14–0.15 (0.14); WL: 0.9–0.93 (0.92); PSL: 0.17–0.19 (0.17); MTL: 0.62–0.67 (0.64); PNW: 0.44–0.47 (0.45); PTW: 0.15–0.15 (0.15); PPW: 0.31–0.34 (0.32); CI: 98.6–101.0 (100.1); SI: 67.3–70.9 (69.3); PSLI: 15.4–17.9 (16.3); PPI: 44.4–47.3 (46.3); PNI: 42.2–43.0 (42.5); MTI: 59.7–60.1 (59.9).

Head. In full-face view sub-oval, widening posteriorly, with anterior and posterior sides convex (Fig. 35B). In lateral view sub-oval; ventral and dorsal faces convex; inner hypostomal teeth visible. Sides of the head with sparse, moderately long, suberect pilosity; whole head with dense, long, decumbent to erect pilosity. Anteromedial part of frons with thick, longitudinal, interrupted and sparse rugae, posteromedial frons with rugae longitudinally irregular, interspaces distinctly rugofoveolate, rugae on posteromedial part directed outward; lateral sides with thick, sparse, and irregular rugae with distinctly rugofoveolate interspaces. Occipital lobes with thinner and denser irregular rugae and rugofoveolate interspaces. Area posterolateral from eyes with dense rugofoveolae and additional sparse rugae. Gena with relatively sparse, thick, longitudinal rugae and distinctly rugofoveolate interspaces. Centre of clypeus smooth and shiny, lateral sides with indistinct rugulae; median notch present, moderately wide, and shallow; median longitudinal carina present; lateral longitudinal carinae absent. Scape, when laid back, exceeding the midlength of head by one-fifth of its length; pilosity subdecumbent to erect (Fig. 35B, D). Inner hypostomal teeth distinct, large, closely spaced, triangular, with rounded apex directed upward; outer hypostomal teeth lobe-like, narrower, and approximately as high as inner teeth; inner and outer hypostomal teeth closely spaced and not connected by concavity (Fig. 63Q). Mesosoma. In lateral view, promesonotum short, angular, and moderately low, posterior mesonotum moderately steep, mesonotal process indistinct, tubercle-like; promesonotal groove absent; metanotal groove absent; propodeal spines moderate, with wide base and acute apex; humeral area produced (Fig. 35D). Surface shiny and foveolate; promesonotum with additional sparse, thick, and transverse rugulae on dorsum. Pilosity moderately sparse, long, and erect (Fig. 35D, F). Petiole. Shiny with dense foveolae; node finely foveolate, triangular, with rounded and thick apex, in rear view node dorsoventrally slightly convex; pilosity moderately sparse and erect (Fig. 35D, F). Postpetiole. Shiny and foveolate; dorsum with reduced sculpture and smooth notch; in dorsal view oval, lateral margins medially with two dentate projections; pilosity long, moderately sparse, and erect (Fig. 35D, F). Gaster. Shiny and smooth; pilosity moderately sparse, long, and erect (Fig. 35D, F). Colour. Brown with yellowish legs (Fig. 35D, F).

Minor workers. Measurements (N = 4): HL: 0.6–0.63 (0.62); HW: 0.52–0.54 (0.53); SL: 0.7–0.74 (0.72); EL: 0.13–0.13 (0.13); WL: 0.74–0.8 (0.76); PSL: 0.11–0.12 (0.11); MTL: 0.53–0.55 (0.54); PNW: 0.35–0.37 (0.36); PTW: 0.09–0.1 (0.09); PPW: 0.14–0.15 (0.15); CI: 116.0–120.5 (117.6); SI: 133.0–138.5 (136.2); PSLI: 17.6–18.9 (18.3); PPI: 58.6–67.8 (63.7); PNI: 66.9–70.2 (68.3); MTI: 99.3–105.6 (102.4).

Head. Cephalic margin slightly convex (Fig. 35A). Pilosity relatively sparse, moderately long, subdecumbent to erect. Sculpture shiny and foveolate; vertex with a few arcing, sparse, and moderately thick rugae; antennal sockets with few thick, curved outward rugae and foveolate interspaces. Clypeus with median longitudinal carina absent; two lateral longitudinal carinae absent. Scape, when laid back, exceeding the posterior head margin by one-third of its length; pilosity dense, subdecumbent to erect (Fig. 35A, C). Mesosoma. In lateral view, promesonotum low and moderately long, arched; promesonotal groove present; metanotal groove distinct; propodeal spines moderate, triangular (Fig. 35C). Sculpture shiny and foveolate; katepisternum with big smooth notch. Pilosity very sparse, moderately long, and erect (Fig. 35C, E). Gaster. With sparse, erect pilosity (Fig. 35C, E). Colour. Brown, legs, antenna and gaster yellowish (Fig. 35C, E).

From the type locality.

The species was collected at 925 m in elevation, in montane rainforest. Nest was located in rotten log.

Pheidole joffreville sp. nov., described from Parc National Montagne d’Ambre in Antsiranana, has major workers with dense and thick rugae, anteromedially longitudinal and posteromedially irregular, with distinctly rugofoveolate interspaces, and brown body colouration. Morphologically is most similar to allopatric P. mivory sp. nov., known from Parc National de Marojejy in Antsiranana. Majors of both taxa are extremely similar and species separation should be supported or based exclusively on minors. Majors of P. joffreville sp. nov. differ from P. mivory sp. nov. in distinctly rugofoveolate interspaces on frons and lack of longitudinal rugae on lateral sides of frons and lack of shagreened sculpture on first gastral tergite; minor workers differ in distinctly foveolate mesosoma with only katepisternum with smooth notch.

Holotype. Madagascar. • 1 major worker; Toamasina; Parc National de Zahamena; -17.73359, 48.72625; alt. 950 m; 19 Feb 2009; B. L. Fisher et al. leg.; rainforest, ex rotten log; BLF22067; CASENT0235022 (CASC). Paratype. • 1w.; same data as for holotype, CASENT0149839 (CASC).

Figure 36. 

Pheidole kely sp. nov., full-face view (A), profile (C), and dorsal view (E) of paratype minor worker (CASENT0149839) and full-face view (B), profile (D), and dorsal view (F) of holotype major worker (CASENT0235022).

Madagascar. –Antsiranana: • 2w., 2s.; Betaolana Forest, along Bekona River; -14.52996, 49.44039; alt. 880 m; 4 Mar 2009; B. L. Fisher et al. leg.; rainforest, ex rotten log; BLF22452 (CASC). • 2w., 2q.; Betaolana Forest, along Bekona River; -14.52996, 49.44039; alt. 880 m; 4 Mar 2009; B. L. Fisher et al. leg.; rainforest, ex rotten log; BLF22480 (CASC). • 2w., 2s.; Betaolana Forest, along Bekona River; -14.52996, 49.44039; alt. 880 m; 4 Mar 2009; B. L. Fisher et al. leg.; rainforest, ex rotten log; BLF22559 (CASC). • 1w., 1s.; Betaolana Forest, along Bekona River; -14.52996, 49.44039; alt. 880 m; 5 Mar 2009; B. L. Fisher et al. leg.; rainforest, ex rotten log; BLF22591 (CASC). • 1w., 1s.; Betaolana Forest, along Bekona River; -14.52996, 49.44039; alt. 880 m; 5 Mar 2009; B. L. Fisher et al. leg.; rainforest, ex rotten log; BLF22628 (CASC). • 2w., 2q.; Makirovana forest; -14.17066, 49.95409; alt. 415 m; 29 Apr 2011; B. L. Fisher et al. leg.; rainforest, ex rotten log; BLF26641 (CASC). • 2w.; Makirovana forest; -14.17066, 49.95409; alt. 415 m; 29 Apr 2011; B. L. Fisher et al. leg.; rainforest, ex root mat, ground layer; BLF26670 (CASC). • 2w.; Makirovana forest; -14.16044, 49.95216; alt. 550 m; 1 May 2011; B. L. Fisher et al. leg.; rainforest, under rootmat, litter on rock; BLF26911 (CASC). • 3w., 2s.; 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.; rainforest, ex rotten log; BLF08904 (CASC). • 3w., 6s.; 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.; rainforest, ex rotten log; BLF08928 (CASC). • 2w.; 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; 16 Nov 2003; B. L. Fisher et al. leg.; rainforest, ex rotten log; BLF08943 (CASC). • 2w., 3s.; Parc National de Marojejy, Manantenina River, 28.0 km 38°NE Andapa, 8.2 km 333°NNW Manantenina; -14.43667, 49.775; alt. 450 m; 12 Nov 2003; B. L. Fisher et al. leg.; rainforest, ex rotten log; BLF08749 (CASC). • 3w., 1s., 1q.; Sava Region: Parc National de Marojejy, Manantenina River, 28.0 km 24.8°NE Andapa; -14.43461, 49.76074; alt. 780 m; 8 Feb 2018; B. L. Fisher et al. leg.; rainforest, ex rotten log; BLF40819 (CASC). • 3w., 1s., 1m.; Sava Region: Parc National de Marojejy, Manantenina River, 28.0 km 24.8°NE Andapa; -14.43461, 49.76074; alt. 780 m; 13 Feb 2018; B. L. Fisher et al. leg.; rainforest, ex rotten log; BLF41152 (CASC). • 2w., 1s., 1q.; Sava Region: Parc National de Marojejy, near Manantenina River; -14.43677, 49.77541; alt. 475 m; 7 Feb 2018; B. L. Fisher et al. leg.; rainforest, ex rotten log; BLF40761 (CASC).–Fianarantsoa: • 2w.; Parc National d’Isalo, Ambovo Springs, 29.3 km 4°N Ranohira; -22.29833, 45.35167; alt. 990 m; 9 Feb 2003; B. L. Fisher et al. leg.; Uapaca woodland, under stone; BLF07796 (CASC).–Toamasina: • 3w.; 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.; rainforest, ex rotten log; BLF08293 (CASC). • 1w., 1s.; Parc National de Zahamena; -17.73359, 48.72625; alt. 950 m; 19 Feb 2009; B. L. Fisher et al. leg.; rainforest, ex rotten log; BLF22093 (CASC).

Minute species. Major workers. HL < 1.0 mm and WL < 0.8 mm; head in full-face view sub-oval, widening slightly posteriorly, with anterior and posterior sides convex, in lateral view sub-oval; ventral and dorsal faces convex; body yellowish orange; sides of head with moderately dense, moderately long, suberect to erect pilosity; entire head distinctly sculptured, medial part of frons with thick, dense, longitudinal, and interrupted rugae and smooth to indistinctly rugulate interspaces; scape, when laid back, slightly exceeding the midlength of head; mesosoma with fine and sparse rugofoveolae, pronotum with rugofoveolae reduced and smooth notches on medial parts of dorsum and lateral sides; katepisternum with smooth notch; inner hypostomal teeth distinct, large, closely spaced, triangular, with rounded apex directed upward; outer hypostomal teeth lobe-like, lower than inner teeth and approximately the same weight; inner and outer hypostomal teeth moderately closely spaced and not connected by concavity; base of first gastral tergite smooth. Minor workers. HL < 0.5 mm and WL < 0.6 mm, scape, when laid back, surpassing the posterior head margin by one-fifth of its length; propodeal spines small and triangular; head relatively rectangular; body dark yellow; head foveolate; medial side of frons with sparse, fine, and interrupted rugulae; area posterolateral from eyes smooth; mesosoma smooth.

Major workers. Measurements (N = 10): HL: 0.9–0.97 (0.93); HW: 0.87–0.95 (0.91); SL: 0.48–0.53 (0.5); EL: 0.1–0.13 (0.11); WL: 0.77–0.84 (0.8); PSL: 0.12–0.15 (0.14); MTL: 0.44–0.5 (0.47); PNW: 0.39–0.46 (0.43); PTW: 0.1–0.14 (0.12); PPW: 0.25–0.31 (0.28); CI: 100.2–104.6 (102.4); SI: 53.1–58.3 (55.0); PSLI: 13.2–15.7 (14.5); PPI: 38.7–48.6 (41.8); PNI: 43.0–49.2 (47.3); MTI: 49.3–54.1 (51.5).

Head. In full-face view sub-oval, widening slightly posteriorly, with anterior and posterior sides convex (Fig. 36B). In lateral view sub-oval; ventral and dorsal faces convex; inner hypostomal teeth visible. Sides of the head with moderately dense, moderately long, suberect to erect pilosity; whole head with dense, long, decumbent to erect pilosity. Medial part of frons with thick, dense, longitudinal, and interrupted rugae and smooth to indistinctly rugulate interspaces; lateral sides with thick, dense, and irregular rugae, interspaces shiny and distinctly rugofoveolate; rugae more longitudinal on anterolateral sides. Occipital lobes with thinner, irregular rugae and smooth to indistinctly rugofoveolate interspaces. Area posterolateral from eyes with sculpture weaker and rugoreticulate; sculpture weakening posteriorly. Gena with relatively sparse, thick, longitudinal rugae and smooth to indistinctly rugoreticulate interspaces. Centre of clypeus smooth and shiny, lateral sides with indistinct rugulae; median notch present, moderately wide, and shallow; median longitudinal carina present; lateral longitudinal carinae absent. Scape, when laid back, slightly exceeding the midlength of head; pilosity subdecumbent to erect (Fig. 36B, D). Inner hypostomal teeth distinct, large, closely spaced, triangular, with rounded apex directed upward; outer hypostomal teeth lobe-like, lower than inner teeth and approximately the same weight; inner and outer hypostomal teeth moderately closely spaced and not connected by concavity (Fig. 63R). Mesosoma. In lateral view, promesonotum short, angular, and moderately low, posterior mesonotum moderately steep, mesonotal process very indistinct, tubercle-like or absent; promesonotal groove absent; metanotal groove absent; propodeal spines moderately long, with moderately wide base and acute apex; humeral area laterally weakly produced (Fig. 36D). Surface shiny with fine and sparse rugofoveolae; pronotum with rugofoveolae reduced and smooth notches on medial parts of dorsum and lateral sides; katepisternum with smooth notch. Pilosity moderately sparse, long, and erect (Fig. 36D, F). Petiole. Shiny with fine foveolae; node smooth to finely foveolate, triangular, with rounded and thick apex, in rear view node dorsoventrally straight to slightly convex; pilosity moderately sparse and erect (Fig. 36D, F). Postpetiole. Shiny and foveolate; dorsum with reduced sculpture and smooth notch; in dorsal view oval, lateral margins medially with two dentate projections; pilosity long, moderately sparse, and erect (Fig. 36D, F). Gaster. Shiny and smooth; pilosity dense, moderately long, and erect (Fig. 36D, F). Colour. Yellowish orange; mandibles and gaster brown (Fig. 36D, F).

Minor workers. Measurements (N = 10): HL: 0.47–0.52 (0.49); HW: 0.4–0.45 (0.42); SL: 0.43–0.48 (0.45); EL: 0.09–0.11 (0.1); WL: 0.53–0.61 (0.57); PSL: 0.04–0.06 (0.05); MTL: 0.33–0.41 (0.37); PNW: 0.27–0.3 (0.28); PTW: 0.05–0.08 (0.06); PPW: 0.1–0.14 (0.12); CI: 111.9–121.1 (117.0); SI: 103.9–108.0 (106.1); PSLI: 7.9–11.7 (10.3); PPI: 48.3–64.2 (55.5); PNI: 64.9–69.7 (67.0); MTI: 80.2–98.8 (85.4).

Head. Cephalic margin indistinctly concave to straight (Fig. 36A). Pilosity relatively dense, moderately long, decumbent to suberect. Sculpture shiny and foveolate; medial side of frons with sparse, fine, and interrupted rugulae; area posterolateral from eyes smooth; antennal sockets with few thick, curved outward rugae and indistinctly foveolate interspaces. 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 dense, subdecumbent to erect (Fig. 36A, C). Mesosoma. In lateral view, promesonotum moderately high and short, arched; promesonotal groove absent; metanotal groove distinct; propodeal spines small and triangular (Fig. 36C). Sculpture shiny and smooth. Pilosity moderately sparse, moderately long, and erect (Fig. 36C, E). Gaster. With sparse, erect pilosity (Fig. 36C, E). Colour. Dark yellow, gaster and vertex brown (Fig. 36C, E).

Malagasy for small in reference to body size.

The species was collected between 325–990 m in elevation, in rainforest and Uapaca woodland. Nests were located in rotten logs, under rootmats, and under stones.

Pheidole kely sp. nov. belongs to the group of species characterised by small body size (major workers: HL < 1.05 mm, WL < 0.9 mm and minor workers HL < 0.5 mm, WL < 0.6 mm), sub-oval, slightly widening posteriorly head with anterior and posterior sides convex in major workers, and minor workers with yellow to brown body colouration and foveolate head or head predominantly smooth and relatively oval. The group includes six species: P. havoana sp. nov., P. kely sp. nov., P. parvula sp. nov., P. parvulogibba sp. nov., P. volontany sp. nov., and P. midongy sp. nov. Pheidole kely sp. nov. is distributed from Toamasina north to Sambava and has a single record from Parc National d’Isalo in Fianarantsoa. Morphologically it is most similar to P. havoana sp. nov. and P. parvulogibba sp. nov., described from the Anosyenne Mts. in Toliara, and parapatric P. parvula sp. nov., known from two localities in the vicinity of Antananarivo. Minor workers of P. kely sp. nov. distinctly differ from those of P. parvula sp. nov. and P. parvulogibba sp. nov. in having a predominantly foveolate head. Minor workers of P. havoana sp. nov. and P. kely sp. nov. are indistinguishable. Major workers of P. kely sp. nov. differ from P. havoana sp. nov. and P. parvula sp. nov. in medial part of frons with thick, dense, longitudinal, and interrupted rugae and smooth to indistinctly rugulate interspaces; from P. parvulogibba sp. nov. in mesosoma with fine and sparse rugofoveolae, pronotum with rugofoveolae reduced, and smooth notches on medial parts of dorsum and its lateral sides.

Holotype. Madagascar. •1 major worker; Toliara; Grand Lavasoa, 25.9km W Tolagnaro; -25.08767, 46.749; alt. 450 m; 30 Nov 2006; B. L. Fisher et al. leg.; rainforest, ex rotten log; BLF15392; CASENT0122898 (CASC). Paratype. •1w.; same data as for holotype, CASENT0923282 (CASC).

Figure 37. 

Pheidole lavasoa sp. nov., full-face view (A), profile (C), and dorsal view (E) of paratype minor worker (CASENT0923282) and full-face view (B), profile (D), and dorsal view (F) of holotype major worker (CASENT0122898).

Minute species. Major workers. HL < 0.95 mm and WL < 0.8 mm; head in full-face view sub-oval; body dark brown; sides of head with very dense, short, suberect pilosity; entire head distinctly sculptured; scape, when laid back, exceeding the midlength of head by two-fifths of its length; inner hypostomal teeth distinct, large, with rounded apex; outer hypostomal teeth dentate with apex directed indistinctly outward; inner and outer hypostomal teeth not closely spaced and not connected by concavity. Minor workers. HL < 0.5 mm and WL < 0.6 mm, scape, when laid back, surpassing the posterior head margin by two-fifths of its length, propodeal spines reduced to small lobes, head elongate, and oval and body yellowish brown; head sculpture shiny and predominantly smooth, frons with sparse, short, and longitudinal rugulae; mesosoma shiny and smooth.

Major workers. Measurements (N = 1): HL: 0.94; HW: 0.98; SL: 0.66; EL: 0.12; WL: 0.83; PSL: 0.13; MTL: 0.57; PNW: 0.36; PTW: 0.13; PPW: 0.28; CI: 95.6; SI: 67.0; PSLI: 13.9; PPI: 44.0; PNI: 36.3; MTI: 57.9.

Head. In full-face view sub-oval, slightly widening posteriorly, with anterior and posterior sides convex (Fig. 37B). In lateral view sub-oval; ventral and dorsal faces convex; inner hypostomal teeth visible. Sides of the head with very dense, short, suberect to erect pilosity; whole head with dense, long, decumbent to erect pilosity. Medial part of frons with thick, dense, longitudinal and interrupted rugae and smooth interspaces, posteromedial frons with rugae more irregular and directed slightly outward; lateral sides with thick, dense, and irregular rugae, interspaces shiny and smooth or indistinctly rugulate. Occipital lobes with thick, irregular rugae and smooth to rugoreticulate interspaces. Area posterolateral from eyes rugoreticulate to rugofoveolate with additional longitudinal thick rugae. Gena with relatively dense, thick, and longitudinal rugae and smooth to indistinctly rugoreticulate interspaces. Centre of clypeus smooth and shiny, lateral sides with indistinct rugulae; median notch present, moderately wide, and shallow; median longitudinal carina present; lateral longitudinal carinae absent. Scape, when laid back, exceeding the midlength of head by three-fifths of its length; pilosity subdecumbent to erect (Fig. 37B, D). Inner hypostomal teeth distinct, large, closely spaced, triangular, with rounded apex directed upward; outer hypostomal teeth dentate and lower than inner teeth with sharp apex directed slightly outward; inner and outer hypostomal teeth not closely spaced and not connected by concavity (Fig. 63S). Mesosoma. In lateral view, promesonotum short, angular, and moderately low, posterior mesonotum moderately steep, mesonotal process very indistinct, tubercle-like; promesonotal groove absent; metanotal groove absent; propodeal spines moderately short, with moderately wide base and acute apex; humeral area laterally weakly produced (Fig. 37D). Surface shiny with fine rugofoveolae; medial parts of lateral sides of pronotum with reduced sculpture. Pilosity moderately dense, long, and erect (Fig. 37D, F). Petiole. Shiny with fine foveolae; node smooth to finely foveolate, triangular, with rounded and thick apex, in rear view node dorsoventrally straight to slightly convex; pilosity moderately sparse and erect (Fig. 37D, F). Postpetiole. Shiny and foveolate; dorsum with reduced sculpture and smooth notch; in dorsal view oval, lateral margins medially with two dentate projections; pilosity long, moderately sparse, and erect (Fig. 37D, F). Gaster. Shiny and smooth; pilosity dense, long, and erect (Fig. 37D, F). Colour. Dark brown; legs, gaster and antennae yellowish brown (Fig. 37D, F).

Minor workers. Measurements (N = 1): HL: 0.48; HW: 0.42; SL: 0.54; EL: 0.1; WL: 0.58; PSL: 0.05; MTL: 0.4; PNW: 0.27; PTW: 0.07; PPW: 0.11; CI: 116.4; SI: 130.8; PSLI: 9.7; PPI: 65.5; PNI: 65.1; MTI: 95.4.

Head. Cephalic margin indistinctly convex (Fig. 37A). Pilosity relatively dense, long, decumbent to suberect. Sculpture shiny and smooth; frons with sparse, short, and longitudinal rugulae; antennal sockets with few thick, curved outward rugae and smooth interspaces. Clypeus with median longitudinal carina absent; two lateral longitudinal carinae absent. Scape, when laid back, surpassing the posterior head margin by two-fifths of its length; pilosity dense, subdecumbent to erect (Fig. 37A, C). Mesosoma. In lateral view, promesonotum moderately high and short, arched; promesonotal groove absent; metanotal groove distinct; propodeal spines reduced to small lobes (Fig. 37C). Sculpture shiny and smooth. Pilosity moderately sparse, long, and erect (Fig. 37C, E). Gaster. With sparse, erect pilosity (Fig. 37C, E). Colour. Yellowish brown (Fig. 37C, E).

From the type locality.