The ant genus Pheidole Westwood, 1839 (Hymenoptera: Formicidae) in Madagascar-taxonomic revision of the bessonii species-group Author Salata, Sebastian Author Fisher, Brian L. 0000-0002-4653-3270 bfisher@calacademy.org text Zootaxa 2020 2020-08-31 4843 1 1 64 journal article 8556 10.11646/zootaxa.4843.1.1 a2aef684-44f1-429d-9dd5-dd394539fa36 1175-5326 4405729 57044928-1950-4525-850E-E381A52599A1 Pheidole maro sp. nov. Figs 19 A–F, 27K, 28K HOLOTYPE : 1s., Madagascar , Toamasina , Corridor Forestier Analamay-Mantadia , Tsaravoniana , -18.76465 48.41938 , 1039 m , 4-Dec-2012 , rainforest, ex rotten log, B. L. Fisher et al . leg. BLF30097, CASENT0923256 ( CASC ) . PARATYPES : 1w. , the same data as holotype, CASENT0299307 ( CASC ) . Other material. Antananarivo : 1w. , Forêt de galerie, Telomirahavavy , 23.4 km NNE Ankazobe , -18.12167 47.20627 , 1520 m , B. L. Fisher et al . leg. ( CASC ) . Antsiranana : 1w. , Ampasindava , Forêt d’Ambilanivy , 3.9 km 181° S Ambaliha , -13.79861 48.16167 , 600 m , B. L. Fisher et al . leg. ( CASC ) ; 1w. , Forêt Ambanitaza , 26.1 km 347° Antalaha , -14.67933 50.18367 , 240 m , B. L. Fisher et al . leg. ( CASC ) ; 1w. , Forêt d’ Andavakoera , 21.4 km 75° ENE Ambilobe ; 4.6 km 356° N Betsiaka , -13.11833 49.23, 425 m , B. L. Fisher et al . leg. ( CASC ) ; 1w. , Forêt de Binara , 9.1 km 233° SW Daraina , -13.26333 49.60333 , 800 m , B. L. Fisher et al . leg. ( CASC ) ; 1w. , Forêt de Binara , 9.1 km 233° SW Daraina , -13.26333 49.60333 , 650-800 m , B. L. Fisher et al . leg. ( CASC ) ; 1w. , Makirovana forest , -14.16506 49.9477 , 900 m , B. L. Fisher et al . leg. ( CASC ) ; 1w. , Montagne des Français , 7.2 km 142° SE Antsiranana (= Diego Suarez ), -12.32278 49.33817 , 180 m , B. L. Fisher et al . leg. ( CASC ) ; 1w. , Parc National de Marojejy , Manantenina River , 27.6 km 35° NE Andapa , 9.6 km 327° NNW Manantenina , -14.435 49.76, 775 m , B. L. Fisher et al . leg. ( CASC ) ; 1w. , 1s., R.S. Manongarivo , 14.5 km 220° SW Antanambao , -13.99833 48.42833 , 1175 m , B. L. Fisher et al . leg. ( CASC ) ; 2w. , Réserve Spéciale d’Ambre , 3.5 km 235° SW Sakaramy , -12.46889 49.24217 , 325 m , B. L. Fisher et al . leg. ( CASC ) ; 1w. , Réserve Spéciale de l’Ankarana , 13.6 km 192° SSW Anivorano Nord , -12.86361 49.22583 , 210 m , B. L. Fisher et al . leg. ( CASC ) ; 2w. , Réserve Spéciale de l’Ankarana , 22.9 km 224° SW Anivorano Nord , -12.90889 49.10983 , 80 m , B. L. Fisher et al . leg. ( CASC ) ; 3w. , Sakalava Beach [vegetated beach dunes], -12.26972 49.39167 , 10 m , R. Harin’Hala leg. ( CASC ) ; 2w. , SAVA Region , District of Sambava , Marojejy National Park , 5 km W of Manantenina village , 1st Camp site ( Mantella ), -14.43817 49.774 , 487 m, M. Rin’Ha leg. ( CASC ) . Fianarantsoa : 2w. , 43 km S Ambalavao , Rés. Andringitra , -22.23333 47, 825 m , B. L. Fisher et al . leg. ( CASC ) ; 4w. , 45 km S. Ambalavao , -22.21667 47.01667 , 785 m , B. L. Fisher et al . leg. ( CASC ) ; 2w. , 9.0 km NE Ivohibe , -22.42667 46.93833 , 900 m , B. L. Fisher et al . leg. ( CASC ) ; 21w. , Belle Vue trail, Ranomafana National Park , Fianarantsoa Prov. , -21.2665 47.42017 , 1020 m , M.E. Irwin et. al. leg. ( CASC ) ; 2w. , Fitovavy Fitovinany Region , District of Ifanadiana , 12 km W of Ranomafana , -21.25083 47.40717 , 1127 m , M. Rin’Ha leg. ( CASC ) ; 2w. , 2s., Parc National de Ranomafana , Vatoharanana River , 4.1 km 231° SW Ranomafana , -21.29 47.43333, 1100 m , B. L. Fisher et al . leg. ( CASC ) ; 1w. , R.S. Ivohibe 8.0 km E Ivohibe , -22.48333 46.96833 , 1200 m , B. L. Fisher et al . leg. ( CASC ) ; 1w. , 1s., R.S. Ivohibe , 7.5 km ENE Ivohibe , -22.47 46.96, 900 m , B. L. Fisher et al . leg. ( CASC ) ; 18w. , radio tower, Ranomafana National Park , Fianarantsoa Prov. , -21.25833 47.40717 , 1130 m , M.E. Irwin et. al. leg. ( CASC ) ; 10w. , Vohiparara broken bridge, Fianarantsoa Prov. , -21.22617 47.36983 , 1110 m , R. Harin’Hala leg. MA-02-09A-06 ( CASC ) . Mahajanga : 1w. , Réserve Spéciale Marotandrano , Marotandrano 48.3 km S Mandritsara , -16.28322 48.81443 , 865 m , B. L. Fisher et al . leg. ( CASC ) . Toamasina : 2w. , 7 km SE Andasibe National Park Headquarters , -18.969856 48.465894 , 1050 m , M.E. Irwin et. al. leg. ( CASC ) ; 1s., Ambanizana , Parc National Masoala , -15.57167 50.00611 , 800-897 m , D. Andriamalala et. al. leg. ( CASC ) ; 1w. , 1s., Ambanizana , Parc National Masoala , -15.57167 50.00611 , 900-950 m , D. Andriamalala et. al. leg. ( CASC ) ; 2w. , 2s., Ambatovy , 12.4 km NE Moramanga , -18.84963 48.2947 , 1010 m , B. L. Fisher et al . leg. ( CASC ) ; 2w. , 2s., Ambatovy , 12.4 km NE Moramanga , -18.83937 48.30842 , 1080 m , B. L. Fisher et al . leg. ( CASC ) ; 3w. , Ambatovy , 12.4 km NE Moramanga , -18.84773 48.29568 , 1000 m , B. L. Fisher et al . leg. ( CASC ) ; 1w. , Ambatovy , 12.4 km NE Moramanga , -18.85813 48.28488 , 1040 m , B. L. Fisher et al . leg. ( CASC ) ; 1w. , 1s., Analamay , -18.80623 48.33707 , 1068 m , B. L. Fisher et al . leg. ( CASC ) ; 2w. , Andasibe National Park , botanic garden near entrance, West of ANGAP office, -18.925172 48.418651 , 1025 m , M.E. Irwin et. al. leg. ( CASC ) ; 2w. , Ankerana , -18.40636 48.80254 , 1108 m , B. L. Fisher et al . leg. ( CASC ) ; 1w. , Ankerana , -18.4017 48.80605 , 1035 m , B. L. Fisher et al . leg. ( CASC ) ; 1s., Bevolota 17.1 km N Andasibe , -18.77071 48.43164 , 995 m , B. L. Fisher et al . leg. ( CASC ) ; 1s., Corridor Forestier Analamay-Mantadia , Ambatoharanana , -18.80398 48.40358 , 1064 m , B. L. Fisher et al . leg. ( CASC ) ; 2w. , 1s., Corridor Forestier Analamay-Mantadia , Ambatoharanana , -18.79956 48.4028 , 1058 m , B. L. Fisher et al . leg. ( CASC ) ; 2w. , 2s., Corridor Forestier Analamay-Mantadia , Ambatoharanana , -18.80438 48.40735 , 960 m , B. L. Fisher et al . leg. ( CASC ) ; 2w. , Corridor Forestier Analamay-Mantadia , Ambohibolakely , -18.77898 48.36375 , 918 m , B. L. Fisher et al . leg. ( CASC ) ; 2w. , Corridor Forestier Analamay-Mantadia , Ambohibolakely , -18.76087 48.37128 , 1044 m , B. L. Fisher et al . leg. ( CASC ) ; 1w. , 1s., Corridor Forestier Analamay-Mantadia , Tsaravoniana , -18.76124 48.42134 , 939 m , B. L. Fisher et al . leg. ( CASC ) ; 4w. , 4s., Corridor Forestier Analamay-Mantadia , Tsaravoniana , -18.76465 48.41938 , 1039 m , B. L. Fisher et al . leg. ( CASC ) ; 1w. , Corridor Forestier Analamay-Mantadia , Tsaravoniana , -18.75641 48.42195 , 1036 m , B. L. Fisher et al . leg. ( CASC ) ; 1w. , Corridor Forestier Analamay-Mantadia , Tsaravoniana , -18.75737 48.42302 , 1018 m , B. L. Fisher et al . leg. ( CASC ) ; 3w. , F.C. Andriantantely , -18.695 48.81333 , 530 m , H. J. Ratsirarson leg. ( CASC ) ; 2w. , 1s., F.C. Didy , -18.19833 48.57833 , 960 m , H. J. Ratsirarson leg. ( CASC ) ; 1w. , F.C. Sandranantitra , -18.04833 49.09167 , 450 m , H. J. Ratsirarson leg. ( CASC ) ; 2w. , Forêt Ambatovy , 14.3 km 57° Moramanga , -18.85083 48.32, 1075 m , B. L. Fisher et al . leg. ( CASC ) ; 2w. , Montagne d’Akirindro 7.6 km 341° NNW Ambinanitelo , -15.28833 49.54833 , 600 m , B. L. Fisher et al . leg. ( CASC ) ; 2w. , 2s., Montagne d’Anjanaharibe , 18.0 km 21° NNE Ambinanitelo , -15.18833 49.615 , 470 m, B. L. Fisher et al . leg. ( CASC ) ; 5w. , 2s., Montagne d’Anjanaharibe , 19.5 km 27° NNE Ambinanitelo , -15.17833 49.635 , 1100 m , B. L. Fisher et al . leg. ( CASC ) ; 3w. , P.N. Mantadia , -18.79167 48.42667 , 895 m , H. J. Ratsirarson leg. ( CASC ) ; 1w. , Parc National de Zahamena , -17.73359 48.72625 , 950 m , B. L. Fisher et al . leg. ( CASC ) ; 1w. , 1s., Parc National de Zahamena , Onibe River , -17.75908 48.85468 , 780 m , B. L. Fisher et al . leg. ( CASC ) ; 1s., Parc National Mananara-Nord , 7.1 km 261° Antanambe , -16.455 49.7875 , 225 m , B. L. Fisher et al . leg. ( CASC ) ; 1w. , Parcelle K 9 Tampolo , -17.175 49.268 , 10 m, B. L. Fisher et al . leg. ( CASC ) ; 4w. , Réserve Spéciale Ambatovaky , Sandrangato river , -16.77274 49.26551 , 450 m , B. L. Fisher et al . leg. ( CASC ) ; 3w. , Réserve Spéciale Ambatovaky , Sandrangato river , -16.81739 49.29402 , 360 m , B. L. Fisher et al . leg. ( CASC ) ; 2w. , 1s., Sahafina forest 11.4 km W Brickaville , -18.81445 48.96205 , 140 m , B. L. Fisher et al . leg. ( CASC ) ; 1w. , 1s., Station forestière Analamazaotra , Analamazaotra 1.3 km S Andasibe , -18.38466 48.41271 , 980 m , B. L. Fisher et al . leg. ( CASC ) ; 4w. , 1s., Torotorofotsy , -18.87082 48.34737 , 1070 m , B. L. Fisher et al . leg. ( CASC ) ; 1w. , Torotorofotsy , -18.77048 48.43043 , 1005 m , B. L. Fisher et al . leg. ( CASC ) ; 1w. , Toliara , 13 km NW Enakara , Rés. Andohahela , -24.55 46.8, 1250 m , B. L. Fisher et al . leg. ( CASC ) ; 1w. , Parc National Andohahela , Col de Tanatana , 33.3 km NW Tolagnaro , -24.7585 46.85367 , 275 m , B. L. Fisher et al . leg. ( CASC ) . Geographic range. Madagascar , widely distributed across the eastern part of the island. Diagnosis. Major workers. Head in full-face view sub-rectangular, slightly widening posteriorly, with lateral margins slightly convex; side of head with moderately dense, long, suberect to erect pilosity; antennal scrobes very indistinct, with sparse and very thin rugocostulae; interspaces between rugocostulae indistinctly punctate; frons with sparse and thin costulae; interspaces between costulae mostly smooth and sometimes indistinctly punctate; occipital lobes mostly smooth, sometimes with weakly developed rugopuncta; scape, when laid back, not reaching the midlength of head; inner hypostomal tooth distinct, bulge-like, closely spaced, and arching posteroventrally; outer hypostomal tooth lobe-like, high, and moderately wide; median tooth absent; promesonotum short, angular, and moderately low; posterior mesonotum moderately steep; mesonotal process distinct and tubercle-like; mesosoma indistinctly microreticulate; gaster shagreened; body reddish brown. Minor workers. Occiput not elongated; nuchal collar absent; head sculpture mostly smooth; frons and sometimes vertex with very sparse puncta; scape, when laid back, exceeding the posterior head margin by two-fifths of its length; promesonotum moderately low, moderately short, and arched; promesonotal groove present; metanotal groove distinct; propodeal spines small and thin; mesosoma with very sparse and weakly developed puncta; body yellowish brown. Description. Major workers. Measurements (n=10): HL: 2.01–2.31 (2.17); HW: 1.8–2.13 (2.04); SL: 0.86– 1.00 (0.94); EL: 0.23–0.27 (0.25); WL: 1.35–1.61 (1.49); PSL: 0.24–0.31 (0.27); MTL: 1.05–1.17 (1.11); PNW: 0.69–0.88 (0.76); PTW: 0.23–0.31 (0.25); PPW: 0.59–0.88 (0.72); CI: 99.9–111.4 (106.7); SI: 42.7–49.3 (46.5); PSLI: 10.8–13.6 (12.4); PPI: 32.9–41.7 (35.6); PNI: 35.0–41.9 (37.3); MTI: 50.2–58.4 (54.4). Head. In full-face view sub-rectangular, slightly widening posteriorly, with lateral sides slightly convex ( Fig. 19B ). In lateral view sub-oval, not depressed posteriorly; ventral and dorsal margins convex; inner hypostomal tooth visible. Side of head with moderately dense, long, suberect to erect pilosity; head dorsum with relatively dense, long, suberect to erect pilosity. Antennal scrobes very indistinct; with sparse and very thin rugocostulae; interspaces between rugocostulae indistinctly punctate. Frons with sparse and thin costulae; interspaces between costulae mostly smooth and sometimes indistinctly punctate. Occipital lobes mostly smooth, sometimes with weakly developed rugopuncta. Gena with moderately dense and thin costulae; interspaces between costulae mostly punctate. Sides posterolateral from eyes smooth; sometimes anterior parts with dense and very thin network of rugoreticulae. Center of clypeus shiny and smooth, lateral sides with distinct rugulae; median notch present, moderately wide and shallow; median longitudinal carina present; lateral longitudinal carinae present. Scape, when laid back, not reaching the midlength of head; pilosity subdecumbent to erect ( Fig. 19B, D ). Inner hypostomal tooth distinct, bulge-like, closely spaced, and arching posteroventrally; outer hypostomal tooth lobe-like, high, and moderately wide; median tooth absent ( Fig. 27K ). Mesosoma. In lateral view, promesonotum short, angular, and moderately low; posterior mesonotum moderately steep; mesonotal process distinct and tubercle-like; promesonotal groove absent; metanotal groove absent; propodeal spines moderately long with wide base and acute top; humeral tubercle weakly produced ( Fig. 19D ). Surface indistinctly microreticulate. Pilosity moderately dense, long, and erect ( Fig. 19D, F ). Postpetiole. In dorsal view trapezoidal and distinctly wider than long ( Fig. 19D, F ). Gaster. Shiny and shagreened; pilosity moderately sparse, long, and erect ( Fig. 19D, F ). Color. Reddish brown, antenna, legs and propodeum yellowish, gaster usually darker than mesosoma ( Fig. 19D, F ). FIGURE 19. Pheidole maro , full-face view (A), profile (C), and dorsal view (E) of paratype minor worker (CASENT0299307) and full-face view (B), profile (D), and dorsal view (F) of holotype major worker (CASENT0923256). Description. Minor workers. Measurements (n=9): HL: 0.67–0.75 (0.71); HW: 0.58–0.68 (0.62); SL: 0.8– 0.91 (0.82); EL: 0.13–0.15 (0.14); WL: 0.83–0.98 (0.92); PSL: 0.07–0.11 (0.1); MTL: 0.68–0.82 (0.72); PNW: 0.38–0.43 (0.4); PTW: 0.09–0.12 (0.1); PPW: 0.17–0.22 (0.2); CI: 107.7–123.5 (115.0); SI: 123.9–150.7 (134.2); PSLI: 10.0–15.5 (13.6); PPI: 44.0–60.0 (51.4); PNI: 62.8–68.8 (64.9); MTI: 106.8–135.8 (116.3). Head. In full-face view oval, posterior region not elongated; nuchal collar absent ( Fig. 19A ). Pilosity relatively sparse, moderately short, subdecumbent to erect. Sculpture mostly smooth; frons and sometimes vertex with very sparse puncta. Clypeus with median longitudinal carina absent; two lateral longitudinal carinae absent. Scape, when laid back, exceeding the posterior head margin by two-fifths of its length; pilosity dense, subdecumbent to erect ( Fig. 19A, C ). Mesosoma. In lateral view, promesonotum moderately low, moderately short, and arched; promesonotal groove present; metanotal groove distinct; propodeal spines small, thin, and triangular ( Fig. 19C ). Sculpture with very sparse and weakly developed puncta. Pilosity sparse, moderately long, and erect ( Fig. 19C, E ). Gaster. With sparse and erect pilosity ( Fig. 19C, E ). Color. Yellowish brown, mandibles, antenna, and gaster usually brighter than the rest of the body ( Fig. 19C, E ). Biology. The species was collected between 10–1520 m elevation, in rainforest, montane rainforest, mixed tropical forest, tropical dry forest, and montane forest. Nests were located in rotten logs and under moss. Workers were collected from sifted litter and on low vegetation. Comments. Pheidole maro is most similar to sympatric P. madecassa . Its major workers can be separated based on indistinctly punctate interspaces between rugocostulae on antennal scrobes, sculptured mesosoma, large inner hypostomal tooth, and reddish-brown body. Majors of P. madecassa have smooth interspaces between rugocostulae on antennal scrobes, mostly smooth mesosoma with very indistinct and thin rugulae, small inner hypostomal tooth, and yellow to orange body. Minor workers can be distinguished based on presence of indistinct sculpture on frons and mesosoma. In contrast, minors of P. madecassa have smooth mesosoma, and mostly smooth frons with weakly developed puncta on anterolateral areas. Etymology. Malagasy for many or plentiful, in reference to the wide distribution of the species.