taxonID	type	description	language	source
03938025FF8BA91DA7D8B7D2FEA1F98A.taxon	description	Pheidole antsahabe sp. nov. Pheidole atsirakambiaty sp. nov. Pheidole bessonii Forel, 1891 Pheidole clara sp. nov. Pheidole decollata Forel, 1892 stat. nov. Pheidole flammea sp. nov. Pheidole flavodepressa sp. nov. Pheidole grallatrix Emery, 1899 Pheidole madecassa Forel, 1892 Pheidole mantadioflava sp. nov. Pheidole maro sp. nov. Pheidole oswaldi Forel, 1891 Pheidole ovalinoda sp. nov. Pheidole similis sp. nov. Pheidole tenebrovulgaris sp. nov. Pheidole uranus sp. nov. Pheidole voreios sp. nov. Pheidole zirafy sp. nov.	en	Salata, Sebastian, Fisher, Brian L. (2020): The ant genus Pheidole Westwood, 1839 (Hymenoptera: Formicidae) in Madagascar-taxonomic revision of the bessonii species-group. Zootaxa 4843 (1): 1-64, DOI: 10.11646/zootaxa.4843.1.1
03938025FF8BA91DA7D8B7D2FEA1F98A.taxon	description	The following repetitive characters occurring in the majority of species have been omitted from the individual species accounts. Unless stated otherwise, the following descriptions apply to all species treated here: Major workers. Dorsal face of head in lateral view not depressed posteriorly; antennal socket shallow; frontal lobe absent; head in full-face view with distinct posteromedian concavity; antenna 12 - segmented with 3 - segmented club; masticatory margin of mandible with large, stout apical tooth and preapical tooth, followed by long diastema and then short and crenulate tooth just before rounded basal angle; outer surface of mandible mostly smooth and shining, sometimes with weak and sparse puncta; antennal scrobe absent; promesonotum strongly convex, well above level of propodeum; postpetiole shiny and microreticulate, short with slightly convex dorsum, in dorsal view oval with lateromedian dentate projection, pilosity erect; petiole shiny and microrugulate, peduncle with small horizontal lobes on its basal part, node triangular with rounded top, in rear view node dorsoventrally concave, pilosity erect. Minor workers. Antennal socket shallow, surrounded with few indistinct wrinkles that curve posterolaterally, most often with sculptured interspaces; frontal lobe absent; head in full-face view oval, posterior and anterior of compound eye convex; antenna 12 - segmented with 3 - segmented club; humeral tubercle not developed into projection; clypeus smooth and shiny; anterior margin regularly convex; promesonotum well above level of propodeum; petiole smooth with ventral face slightly convex, node triangular with few erect setae; postpetiole smooth, convex with few erect setae; gaster smooth and shiny.	en	Salata, Sebastian, Fisher, Brian L. (2020): The ant genus Pheidole Westwood, 1839 (Hymenoptera: Formicidae) in Madagascar-taxonomic revision of the bessonii species-group. Zootaxa 4843 (1): 1-64, DOI: 10.11646/zootaxa.4843.1.1
03938025FF8BA91DA7D8B7D2FEA1F98A.taxon	diagnosis	Diagnosis. Major worker. Postpetiole in profile without conspicuous ventral convexity; antennal socket shallow; frontal lobe absent or indistinct; propodeal spine moderate to large, wide to narrow; head in full-face view sub-rectangular, with lateral margins varying from relatively straight to slightly convex, not or slightly widening posteriorly; head in lateral view sub-oval with distinctly convex dorsal and ventral cephalic margins; occipital lobe never with transverse costulae and usually entirely sculptured (except P. madecassa, P. maro and P. uranus); frons entirely sculptured, medially costulate to rugocostulate, lateral sides with usually weaker sculpture; antennal scrobe absent or very indistinct; costulae and rugulae on the rest of head thick to very thin; promesonotum short, angular, and moderately low to very low; mesonotal process usually distinct; postpetiole in dorsal view with lateromedian dentate projection; gaster usually at least partially shagreened. Minor worker. Postpetiole in profile without conspicuous ventral convexity; antennal socket shallow; frontal lobe absent or indistinct; propodeal spine minute to moderately large, wide to narrow; promesonotum in lateral view never box-like and posterior mesonotum never steep; promesonotal groove present and distinct (except P. tenebrovulgaris); posterior region of head usually elongated, forming short to long neck; nuchal collar present and weakly developed to distinct (except P. madecassa and P. maro); promesonotum moderately low to very low and moderately long to very long, and arched.	en	Salata, Sebastian, Fisher, Brian L. (2020): The ant genus Pheidole Westwood, 1839 (Hymenoptera: Formicidae) in Madagascar-taxonomic revision of the bessonii species-group. Zootaxa 4843 (1): 1-64, DOI: 10.11646/zootaxa.4843.1.1
03938025FF8BA91DA7D8B7D2FEA1F98A.taxon	discussion	Comments. The P. bessonii group consist of species widely distributed on the island. Six species — P. bessonii, P. decollata, P. grallatrix, P. maro, P. oswaldi and P. tenebrovulgaris can be considered common. The remaining 12 species have more restricted distributions primarily limited to the northern and northeastern parts of the island. The majority of the major and minor workers of the bessonii group can be easily separated from other taxa based on the combination of the above-mentioned characters. However, majors of P. madecassa and P. maro can be confused with major workers of the longispinosa group. They can be separated based on distinctly shorter propodeal spines, denser frons sculpture, and brighter body coloration. Additionally, minors of P. tenebrovulgaris can be confused with two members of the sikorae group: P. trichotos Salata & Fisher and P. veteratrix Forel. Pheidole tenebrovulgaris can be separated based on larger body size, longer propodeal spines, higher promesonotum, and dark brown body coloration. There is also one member of the bessonii group known from Mayotte: P. ragnax. Because its original description is comprehensive and the species is not recorded from Madagascar, we decided not to include it in our revision. However, we included it in comparative diagnoses within the descriptions of most similar species.	en	Salata, Sebastian, Fisher, Brian L. (2020): The ant genus Pheidole Westwood, 1839 (Hymenoptera: Formicidae) in Madagascar-taxonomic revision of the bessonii species-group. Zootaxa 4843 (1): 1-64, DOI: 10.11646/zootaxa.4843.1.1
03938025FF85A918A7D8B0ACFD95FDE4.taxon	description	Figs 9 A – F, 27 A, 28 A	en	Salata, Sebastian, Fisher, Brian L. (2020): The ant genus Pheidole Westwood, 1839 (Hymenoptera: Formicidae) in Madagascar-taxonomic revision of the bessonii species-group. Zootaxa 4843 (1): 1-64, DOI: 10.11646/zootaxa.4843.1.1
03938025FF85A918A7D8B0ACFD95FDE4.taxon	materials_examined	HOLOTYPE: 1 s., Madagascar, Antsiranana, Forêt d’ Antsahabe, 11.4 km 275 ° W Daraina, - 13.21167 49.55667, 550 m, 16 - Nov- 2004, tropical dry forest, ex rotten log, B. L. Fisher et al. leg. BLF 10469, CASENT 0107506 (CASC). PARATYPES: 1 w., the same data as holotype, CASENT 0923247 (CASC). Other material. Madagascar. Antsiranana: 4 w., Ampasindava, Forêt d’Ambilanivy, 3.9 km 181 ° S Ambaliha, - 13.79861, 48.16167, 600 m, B. L. Fisher et al. leg. (CASC); 1 s., Binara Forest, - 13.26206, 49.60672, 559 m, B. L. Fisher et al. leg. (CASC); 10 w., 7 s., 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); 4 w., Forêt d’ Antsahabe, 11.4 km 275 ° W Daraina, - 13.21167, 49.55667, 550 m, B. L. Fisher et al. leg. (CASC); 2 w., Forêt de Binara, 7.5 km 230 ° SW Daraina, - 13.255, 49.61667, 375 m, B. L. Fisher et al. leg. (CASC); 1 w., 1 s., Galoko chain, Mont Galoko, - 13.58745, 48.71419, 380 m, B. L. Fisher et al. leg. (CASC); 1 w., Masoala National Park, - 15.3014, 50.22776, 280 m, B. L. Fisher et al. leg. (CASC); 2 w., Montagne des Français, 7.2 km 142 ° SE Antsiranana (= Diego Suarez), - 12.32278, 49.33817, 180 m, B. L. Fisher et al. leg. (CASC). Toamasina: 1 w., Corridor Forestier Analamay-Mantadia, Ambatoharanana, - 18.80219, 48.40585, 995 m, B. L. Fisher et al. leg. (CASC). Geographic range. Madagascar, northern part of Antsiranana.	en	Salata, Sebastian, Fisher, Brian L. (2020): The ant genus Pheidole Westwood, 1839 (Hymenoptera: Formicidae) in Madagascar-taxonomic revision of the bessonii species-group. Zootaxa 4843 (1): 1-64, DOI: 10.11646/zootaxa.4843.1.1
03938025FF85A918A7D8B0ACFD95FDE4.taxon	diagnosis	Diagnosis. Major workers. Head in full-face view sub-rectangular, not widening posteriorly, lateral margins relatively straight; side of head with moderately dense, moderately long, subdecumbent to suberect pilosity; medial frons with dense and thick costulae; frons laterally rugose-punctate; occipital lobe obliquely rugulose; interspaces between rugulae indistinctly punctate to smooth; scape, when laid back, exceeding midlength of head by two-fifths of its length; inner hypostomal tooth indistinct, small, and bulge-like with tip directed posteriorly; outer hypostomal tooth lobe-like, high, and wide with tip directed posteriorly; median tooth absent; promesonotum short, angular, and moderately low; posterior mesonotum moderately steep; mesonotal process distinct and tubercle-like; mesosoma with sparse microreticulae; promesonotum with transverse rugulae; katepisternum and side of propodeum rugose; gaster shagreened; body brown. Minor workers. Occiput forming short and moderately narrow neck; nuchal collar distinct; head mostly smooth; frons with sparse puncta; scape, when laid back, exceeding posterior head margin by three-fifths of its length; promesonotum in lateral view low and moderately long, arched; promesonotal groove present; metanotal groove distinct; propodeal spines with narrow base; promesonotum weakly microreticulate; katepisternum and propodeum punctate; body orange.	en	Salata, Sebastian, Fisher, Brian L. (2020): The ant genus Pheidole Westwood, 1839 (Hymenoptera: Formicidae) in Madagascar-taxonomic revision of the bessonii species-group. Zootaxa 4843 (1): 1-64, DOI: 10.11646/zootaxa.4843.1.1
03938025FF85A918A7D8B0ACFD95FDE4.taxon	description	Description. Major workers. Measurements (n = 8): HL: 1.59 – 1.78 (1.7); HW: 1.49 – 1.61 (1.55); SL: 1.19 – 1.25 (1.21); EL: 0.21 – 0.28 (0.24); WL: 1.65 – 1.71 (1.68); PSL: 0.27 – 0.33 (0.3); MTL: 1.32 – 1.43 (1.38); PNW: 0.61 – 0.66 (0.64); PTW: 0.18 – 0.22 (0.21); PPW: 0.56 – 0.62 (0.59); CI: 106.9 – 110.8 (109.4); SI: 75.1 – 80.8 (78.2); PSLI: 15.9 – 19.3 (17.5); PPI: 30.0 – 37.5 (34.8); PNI: 38.7 – 42.4 (41.0); MTI: 85.9 – 91.5 (88.7). Head. In full-face view sub-rectangular, not widening posteriorly, side relatively straight (Fig. 9 B). In lateral view sub-oval, not depressed posteriorly; ventral and dorsal margins convex; inner hypostomal tooth not visible. Side of head with moderately dense, moderately long, subdecumbent to suberect pilosity; head dorsum with relatively dense, long, decumbent to erect pilosity. Medial frons with dense and thick costulae; interspaces between costulae smooth or indistinctly punctate. Frons laterally rugulate; interspaces between rugae punctate; rugae and costulae on posterior frons arch posterolaterally. Occipital lobes with thin rugulae that arch posterolaterally; interspaces between rugulae indistinctly punctate to smooth. Gena with dense and thick costulae; interspaces between costulae indistinctly punctate. Sides posterolateral from eyes with network of dense but very thin rugoreticulae. Center of clypeus shiny and with weakly developed puncta, lateral sides with distinct rugulae; median notch present, moderately wide and shallow; median longitudinal carina present; lateral longitudinal carinae present. Scape, when laid back, exceeding the midlength of head by two-fifths of its length; pilosity subdecumbent to erect (Fig. 9 B, D). Inner hypostomal tooth indistinct, small and bulge-like, with top directed posteriorly; outer hypostomal tooth lobe-like, high, and wide, with top directed posteriorly; median tooth absent (Fig. 27 A). 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 present; propodeal spines moderately long with moderately wide base and acute top; humeral tubercle weakly produced (Fig. 9 D). Surface shiny and with sparse microreticulae; promesonotum with additional transverse and thin rugae; katepisternum and lateral sides of propodeum with additional rugae. Pilosity moderately dense, moderately long, and erect (Fig. 9 D, F). Gaster. Shiny and shagreened; pilosity moderately sparse, long, and erect (Fig. 9 D, F). Color. Brown, legs and antennae yellowish-brown (Fig. 9 D, F). Description. Minor workers. Measurements (n = 10): HL: 0.72 – 0.89 (0.82); HW: 0.46 – 0.59 (0.52); SL: 0.99 – 1.27 (1.15); EL: 0.14 – 0.18 (0.16); WL: 0.95 – 1.21 (1.1); PSL: 0.11 – 0.16 (0.13); MTL: 0.82 – 1.13 (1.03); PNW: 0.33 – 0.41 (0.38); PTW: 0.08 – 0.12 (0.09); PPW: 0.13 – 0.18 (0.15); CI: 150.8 – 161.4 (155.8); SI: 206.7 – 227.3 (220.2); PSLI: 14.8 – 17.4 (16.2); PPI: 51.2 – 71.2 (62.9); PNI: 67.1 – 79.3 (73.6); MTI: 180.3 – 206.2 (196.6). Head. In full-face view oval, posterior region elongated forming short and moderately narrow neck; nuchal collar distinct (Fig. 9 A). Pilosity relatively sparse, long, subdecumbent to erect. Sculpture shiny and mostly smooth; frons with sparse puncta. Clypeus without median longitudinal carina; two lateral longitudinal carinae absent. Scape, when laid back, exceeding posterior head margin by three-fifths its length; pilosity dense, subdecumbent to erect (Fig. 9 A, C). Mesosoma. In lateral view, promesonotum low, moderately long, and arched; promesonotal groove present; metanotal groove distinct; propodeal spines moderate, triangular and with narrow base (Fig. 9 C). Promesonotum with weak microreticulae; katepisternum and propodeum punctate. Pilosity very sparse, long, and erect (Fig. 9 C, E). Gaster. With sparse and erect pilosity (Fig. 9 C, E). Color. Orange, legs and sometimes head brighter (Fig. 9 C, E).	en	Salata, Sebastian, Fisher, Brian L. (2020): The ant genus Pheidole Westwood, 1839 (Hymenoptera: Formicidae) in Madagascar-taxonomic revision of the bessonii species-group. Zootaxa 4843 (1): 1-64, DOI: 10.11646/zootaxa.4843.1.1
03938025FF85A918A7D8B0ACFD95FDE4.taxon	biology_ecology	Biology. The species was collected between 180 – 600 m in elevation, in rainforest and tropical dry forest. Nests were located in rotten logs, in soil, and in rotten sticks on the ground. Workers were collected from sifted litter.	en	Salata, Sebastian, Fisher, Brian L. (2020): The ant genus Pheidole Westwood, 1839 (Hymenoptera: Formicidae) in Madagascar-taxonomic revision of the bessonii species-group. Zootaxa 4843 (1): 1-64, DOI: 10.11646/zootaxa.4843.1.1
03938025FF85A918A7D8B0ACFD95FDE4.taxon	discussion	Comments. Pheidole antsahabe is most similar to the sympatric P. voreios. Major workers of P. antsahabe differ in the smooth to indistinctly punctate interspaces between costulae on the medial frons, and the presence of sparse microreticulae on the promesonotal dorsum. In contrast, P. voreios majors have distinctly punctate interspaces between costulae on the medial frons and mostly to entirely smooth promesonotum. Minors can be separated from P. voreios by the orange body coloration, longer and thinner neck, and sparser sculpture on the promesonotum. Minors of P. voreios have brown body, shorter and thicker neck, and distinctly stronger sculpture on the promesonotum. Additionally, minor workers of P. antsahabe can be confused with P. mantadioflava and P. clara but differ from them in having a longer, narrower neck and a promesonotum that is never entirely smooth nor entirely punctate.	en	Salata, Sebastian, Fisher, Brian L. (2020): The ant genus Pheidole Westwood, 1839 (Hymenoptera: Formicidae) in Madagascar-taxonomic revision of the bessonii species-group. Zootaxa 4843 (1): 1-64, DOI: 10.11646/zootaxa.4843.1.1
03938025FF85A918A7D8B0ACFD95FDE4.taxon	etymology	Etymology. From the type locality.	en	Salata, Sebastian, Fisher, Brian L. (2020): The ant genus Pheidole Westwood, 1839 (Hymenoptera: Formicidae) in Madagascar-taxonomic revision of the bessonii species-group. Zootaxa 4843 (1): 1-64, DOI: 10.11646/zootaxa.4843.1.1
03938025FF80A906A7D8B462FD95FC34.taxon	description	Figs 10 A – F, 27 B, 28 B	en	Salata, Sebastian, Fisher, Brian L. (2020): The ant genus Pheidole Westwood, 1839 (Hymenoptera: Formicidae) in Madagascar-taxonomic revision of the bessonii species-group. Zootaxa 4843 (1): 1-64, DOI: 10.11646/zootaxa.4843.1.1
03938025FF80A906A7D8B462FD95FC34.taxon	materials_examined	HOLOTYPE: 1 s., Madagascar, Fianarantsoa, Forêt d’Atsirakambiaty, 7.6 km 285 ° WNW Itremo, - 20.59333 46.56333, 1550 m, 22 - Jan- 2003, montane rainforest, ex rotten log / termite mound, Fisher et al. leg. BLF 07274, CASENT 0491660 (CASC). PARATYPES: 6 w., 2 s., 1 m., 1 q., the same data as holotype, CASENT 0491661, CASENT 0491664, CASENT 0491665, CASENT 0491666, CASENT 0872258, CASENT 0491662 (CASC, MHNG, PBZT). Other material. Madagascar. Fianarantsoa: 11 w., 3 s., Forêt d’Atsirakambiaty, 7.6 km 285 ° WNW Itremo, - 20.59333 46.56333, 1550 m, Fisher et al. leg. (CASC). Geographic range. Madagascar, Fianarantsoa, Forêt d’Atsirakambiaty.	en	Salata, Sebastian, Fisher, Brian L. (2020): The ant genus Pheidole Westwood, 1839 (Hymenoptera: Formicidae) in Madagascar-taxonomic revision of the bessonii species-group. Zootaxa 4843 (1): 1-64, DOI: 10.11646/zootaxa.4843.1.1
03938025FF80A906A7D8B462FD95FC34.taxon	diagnosis	Diagnosis. Major workers. Head in full-face view sub-rectangular, not widening posteriorly, lateral margin relatively straight; sides of head with moderately dense, moderately long, suberect pilosity; frons with moderately thick and moderately dense costulae that arch posterolaterally; interspaces between costulae smooth to indistinctly rugopunctate; occipital lobes with thin and sparse rugulae that arch posterolaterally; interspaces between rugulae sparsely rugopunctate; scape, when laid back, exceeding the midlength of head by two-fifths of its length; inner hypostomal tooth absent; outer hypostomal tooth lobe-like, high, and wide, with top arching posterolaterally; median tooth present; promesonotum short, angular, and moderately low; posterior mesonotum moderately steep; mesonotal process tubercle-like; promesonotum with thick, dense and transverse rugae; interspaces between rugae distinctly punctate; katepisternum and propodeum with sparse rugae; interspaces between rugae rugopunctate; gaster shagreened; body black. Minor workers. Occiput slightly elongated; nuchal collar distinct; head mostly smooth; frons rugopunctate; scape, when laid back, exceeding posterior head margin by half its length; promesonotum low, long, and arched; promesonotal groove present; metanotal groove distinct; propodeal spines moderately small with wide base; mesosoma punctate; promesonotal dorsum with sparser puncta and transverse, thin rugae; body black.	en	Salata, Sebastian, Fisher, Brian L. (2020): The ant genus Pheidole Westwood, 1839 (Hymenoptera: Formicidae) in Madagascar-taxonomic revision of the bessonii species-group. Zootaxa 4843 (1): 1-64, DOI: 10.11646/zootaxa.4843.1.1
03938025FF80A906A7D8B462FD95FC34.taxon	description	Description. Major workers. Measurements (n = 7): HL: 1.88 – 2.1 (1.99); HW: 1.75 – 1.97 (1.86); SL: 1.21 – 1.25 (1.24); EL: 0.23 – 0.29 (0.26); WL: 1.62 – 1.74 (1.69); PSL: 0.23 – 0.32 (0.28); MTL: 1.36 – 1.49 (1.42); PNW: 0.59 – 0.74 (0.64); PTW: 0.19 – 0.31 (0.23); PPW: 0.52 – 0.73 (0.6); CI: 103.4 – 110.8 (107.0); SI: 61.5 – 71.5 (66.7); PSLI: 11.2 – 15.9 (14.0); PPI: 32.2 – 42.7 (39.3); PNI: 32.6 – 37.6 (34.5); MTI: 72.1 – 79.2 (76.1). Head. In full-face view sub-rectangular, not widening posteriorly, lateral margin relatively straight (Fig. 10 B). In lateral view sub-oval, not depressed posteriorly; ventral and dorsal margins convex; inner hypostomal tooth not visible. Side of the head with moderately dense, moderately long, suberect pilosity; head dorsum with relatively dense, long, decumbent to erect pilosity. Frons with moderately thick and moderately dense costulae that arch posterolaterally; interspaces between costulae smooth to indistinctly rugopunctate. Occipital lobe with thin and sparse rugulae that arch posterolaterally; interspaces between rugulae sparsely rugopunctate. Gena with dense and thick costulae; interspaces between costulae indistinctly rugopunctate. Sides posterolateral from eyes with dense but thin rugocostulae; interspaces between rugocostulae rugopunctate. Center of clypeus shiny and indistinctly punctate, lateral sides with distinct rugulae; median notch present, moderately wide and shallow; median and lateral longitudinal carinae present. Scape, when laid back, exceeding the midlength of head by two-fifths of its length; pilosity subdecumbent to erect (Fig. 10 B, D). Inner hypostomal tooth absent; outer hypostomal tooth lobe-like, high and wide, with top arching posterolaterally; median tooth present (Fig. 27 B). Mesosoma. In lateral view, promesonotum short, angular, and moderately low; posterior mesonotum moderately steep; mesonotal process distinct and tuberclelike; promesonotal groove absent; metanotal groove present; propodeal spines moderate with wide base and acute top; humeral tubercle weakly produced (Fig. 10 D). Surface shiny; promesonotum with thick, dense, and transverse rugae; interspaces between rugae distinctly punctate; katepisternum and propodeum with sparse rugae; interspaces between rugae rugopunctate. Pilosity moderately dense, long, and erect (Fig. 10 D, F). Petiole. In rear view node dorsoventrally convex (Fig. 10 D, F). Gaster. Dull and shagreened; pilosity moderately sparse, long, and erect (Fig. 10 D, F). Color. Black, antennae and legs brown, mandibles ochraceous (Fig. 10 D, F). Description. Minor workers. Measurements (n = 10): HL: 0.85 – 0.95 (0.91); HW: 0.59 – 0.71 (0.65); SL: 1.13 – 1.29 (1.23); EL: 0.16 – 0.2 (0.18); WL: 1.16 – 1.29 (1.23); PSL: 0.12 – 0.15 (0.13); MTL: 1.07 – 1.21 (1.13); PNW: 0.41 – 0.47 (0.44); PTW: 0.1 – 0.14 (0.12); PPW: 0.17 – 0.22 (0.2); CI: 133.4 – 143.6 (139.1); SI: 180.3 – 196.8 (189.1); PSLI: 13.0 – 16.5 (14.8); PPI: 47.7 – 68.3 (59.2); PNI: 64.1 – 72.1 (67.9); MTI: 163.4 – 183.0 (173.8). Head. In full-face view oval, occiput slightly elongated; nuchal collar distinct (Fig. 10 A). Pilosity relatively dense, moderately long, subdecumbent to erect. Sculpture shining and smooth; frons rugopunctate. Clypeus with median longitudinal carina present; two lateral longitudinal carinae absent. Scape, when laid back, exceeding posterior head margin by half its length; pilosity dense, subdecumbent to erect (Fig. 10 A, C). Mesosoma. In lateral view, promesonotum low, long and arched; promesonotal groove present; metanotal groove distinct; propodeal spines moderately small, triangular and with wide base (Fig. 10 C). Surface punctate; promesonotal dorsum with sparser puncta and thin, transverse rugae. Pilosity very sparse, long, and erect (Fig. 10 C, E). Gaster. With sparse and erect pilosity (Fig. 10 C, E). Color. Black, with legs, mandibles, and antennae brown (Fig. 10 C, E).	en	Salata, Sebastian, Fisher, Brian L. (2020): The ant genus Pheidole Westwood, 1839 (Hymenoptera: Formicidae) in Madagascar-taxonomic revision of the bessonii species-group. Zootaxa 4843 (1): 1-64, DOI: 10.11646/zootaxa.4843.1.1
03938025FF80A906A7D8B462FD95FC34.taxon	biology_ecology	Biology. The specimens were collected at 1550 m elevation, in montane rainforest and grassland. Nests were located in rotten logs on termite mounds and under stones.	en	Salata, Sebastian, Fisher, Brian L. (2020): The ant genus Pheidole Westwood, 1839 (Hymenoptera: Formicidae) in Madagascar-taxonomic revision of the bessonii species-group. Zootaxa 4843 (1): 1-64, DOI: 10.11646/zootaxa.4843.1.1
03938025FF80A906A7D8B462FD95FC34.taxon	discussion	Comments. Considering the ants known from northern Madagascar, Pheidole atsirakambiaty is most similar to P. voreios and parapatric P. bessonii. Its majors can be separated based on the black body and the presence of moderately thick, moderately dense costulae that arch posterolaterally, and rugulate frons and occipital lobes with smooth to indistinctly rugopunctate interspaces between rugulae. Majors of P. voreios and P. bessonii have a body that is brown to blackish brown, their head sculpture is never arches uniformly posterolaterally, and interspaces between rugocostulae are smooth to punctate. Minors can be distinguished based on their black body and distinctly punctate mesosoma. In contrast, minors of P. voreios and P. bessonii have a body that is bright brown to brown and their mesosoma is never distinctly punctate.	en	Salata, Sebastian, Fisher, Brian L. (2020): The ant genus Pheidole Westwood, 1839 (Hymenoptera: Formicidae) in Madagascar-taxonomic revision of the bessonii species-group. Zootaxa 4843 (1): 1-64, DOI: 10.11646/zootaxa.4843.1.1
03938025FF80A906A7D8B462FD95FC34.taxon	etymology	Etymology. From the type locality.	en	Salata, Sebastian, Fisher, Brian L. (2020): The ant genus Pheidole Westwood, 1839 (Hymenoptera: Formicidae) in Madagascar-taxonomic revision of the bessonii species-group. Zootaxa 4843 (1): 1-64, DOI: 10.11646/zootaxa.4843.1.1
03938025FF9EA902A7D8B212FD19FBEE.taxon	description	Figs 11 A – F, 27 C, 28 C	en	Salata, Sebastian, Fisher, Brian L. (2020): The ant genus Pheidole Westwood, 1839 (Hymenoptera: Formicidae) in Madagascar-taxonomic revision of the bessonii species-group. Zootaxa 4843 (1): 1-64, DOI: 10.11646/zootaxa.4843.1.1
03938025FF9EA902A7D8B212FD19FBEE.taxon	description	Status as species: Dalla Torre, 1893: 88. LECTOTYPE [designated here]: 1 w., top specimen of the pin, Madagascar, Fianarantsoa, coll. Besson, ANTC 3392, CASENT 0101838 (MHNG). PARALECTOTYPES: 1 w., bottom specimen, the same pin as lectotype; 3 w., the same data as lectotype (CASENT 0923201).	en	Salata, Sebastian, Fisher, Brian L. (2020): The ant genus Pheidole Westwood, 1839 (Hymenoptera: Formicidae) in Madagascar-taxonomic revision of the bessonii species-group. Zootaxa 4843 (1): 1-64, DOI: 10.11646/zootaxa.4843.1.1
03938025FF9EA902A7D8B212FD19FBEE.taxon	materials_examined	Other material. Madagascar. Antananarivo: 11 w., Ambohitantely 1410, - 18.2253 47.28683, 1410 m, Fisher et al. leg. (CASC). Fianarantsoa: 1 w., 9.0 km NE Ivohibe, - 22.42667 46.93833, 900 m, Fisher et al. leg. (CASC); 12 w., Belle Vue trail, Ranomafana National Park, Fianarantsoa Prov., - 21.2665 47.42017, 1020 m, R. Harin’Hala leg. (CASC); 4 w., dry wash, 1 km E of Isalo National Park Interpretive Center, Fianarantsoa Prov., - 22.62667 45.35817, 885 m, Irwin et al. leg (CASC); 5 w., Fitovavy Fitovinany Region, District of Ifanadiana Belle vue area 1200 m S of Ranomafana National Park entrance, - 21.2665 47.42017, 1018 m, M. Rin’ha leg. (CASC); 9 w., 2 s., Forêt d’Analalava, 29.6 km 280 ° W Ranohira, - 22.59167 45.12833, 700 m, Fisher et al. leg. (CASC); 4 w., JIRAMA water works near river, Ranomafana National Park, Fianarantsoa Prov., - 21.2485 47.45217, 690 m, R. Harin’Hala leg. (CASC); 1 w., Mampiarika IV Non Protected Area, 27.98 km SW Ambositra, - 20.73528 47.08382, 1486 m, A. Ravelomanana leg. (CASC); 8 w., 4 s., Parc National d’Isalo, 9.1 km 354 ° N Ranohira, - 22.48167 45.46167, 725 m, Fisher et al. leg. (CASC); 1 w., Parc National d’Isalo, Ambovo Springs, 29.3 km 4 ° N Ranohira, - 22.29833 45.35167, 990 m, Fisher et al. leg. (CASC); 3 w., 1 s., Parc National d’Isalo, Sahanafa River, 29.2 km 351 ° N Ranohira, - 22.31333 45.29167, 500 m, Fisher et al. leg. (CASC); 1 w., Parc National de Ranomafana, Vatoharanana River, 4.1 km 231 ° SW Ranomafana, - 21.29 47.43333, 1100 m, Fisher et al. leg. (CASC); 4 w., R. S. Ivohibe 8.0 km E Ivohibe, - 22.48333 46.96833, 1200 m, Fisher et al. leg. (CASC); 2 w., R. S. Ivohibe, 7.5 km ENE Ivohibe, - 22.47 46.96, 900 m, Fisher et al. leg. (CASC); 19 w., radio tower, Ranomafana National Park, Fianarantsoa Prov., - 21.25833 47.40717, 1130 m, Irwin et al. leg. (CASC); 1 s., Ranomafana National Park, Talatakely; Sahambavy; Fianarantsoa Rural, - 21.451179 47.3023894, 1139 m, Lee & Ribardo leg. (CASC); 2 w., stream area, 900 m E of Isalo National Park Interpretive Center, Fianarantsoa Prov., - 22.62667 45.35817, 750 m, R. Harin’Hala leg. (CASC); 12 w., Vohiparara broken bridge, Fianarantsoa Prov., - 21.22617 47.36983, 1110 m, R. Harin’Hala leg. (CASC); 1 w., Ampangabe I Non Protected Area, 21.4 km W Itremo, - 20.61111 46.60688, 1414 m, A. Ravelomanana leg. (CASC); 1 w., Ampangabe II Non Protected Area, 21.29 km W Itremo, - 20.61139 46.60809, 1402 m, A. Ravelomanana leg. (CASC); 2 w., Antapia III Non Protected Area, 26.43 km SW Ambositra, - 20.72 47.08785, 1494 m, A. Ravelomanana leg. (CASC); 1 w., Antohatsahomby II Non Protected Area, 23.38 km NW Itremo, - 20.55444 46.58438, 1640 m, A. Ravelomanana leg. (CASC); 3 w., Forêt d’Analalava, 29.6 km 280 ° W Ranohira, - 22.59167 45.12833, 700 m, Fisher et al. leg. (CASC); 1 w., Mampiarika II Non Protected Area, 28.08 km SW Ambositra, - 20.73556 47.08384, 1464 m, A. Ravelomanana leg. (CASC); 3 w., Mampiarika III Non Protected Area, 28.93 km SW Ambositra, - 20.73583 47.08399, 1487 m, A. Ravelomanana leg. (CASC); 1 w., 2 s., Mampiarika IV Non Protected Area, 27.98 km SW Ambositra, - 20.73528 47.08382, 1486 m, A. Ravelomanana leg. (CASC); 1 w., Manandriana II Non Protected Area, 27.12 km SW Ambositra, - 20.7325 47.09461, 1589 m, A. Ravelomanana leg. (CASC); 1 s., Parc National d’Isalo, 9.1 km 354 ° N Ranohira, - 22.48167 45.46167, 725 m, Fisher et al. leg. (CASC); 1 w., Parc National d’Isalo, Sahanafa River, 29.2 km 351 ° N Ranohira, - 22.31333 45.29167, 500 m, Fisher et al. leg. (CASC); 1 w., Soanierenana II Non Protected Area, 25.61 km SW Ambositra, - 20.72194 47.10896, 1732 m, A. Ravelomanana leg. (CASC). Mahajanga: 9 w., 5 s., Forêt Ambohimanga, 26.1 km 314 ° Mampikony, - 15.96267 47.43817, 250 m, Fisher et al. leg. (CASC); 1 w., Mahavavy River, 6.2 km 145 ° SE Mitsinjo, - 16.05167 45.90833, 20 m, Fisher et al. leg. (CASC); 4 w., 8 s., 1 q., Parc National d’Ankarafantsika, Ampijoroa Station Forestière, 5.4 km 331 ° NW Andranofasika, - 16.29889 46.813, 70 m, Fisher et al. leg. (CASC); 2 w., Parc National d’Ankarafantsika, Ampijoroa Station Forestière, 40 km 306 ° NW Andranofasika, - 16.32083 46.81067, 130 m, Fisher et al. leg. (CASC); 8 w., Parc National d’Ankarafantsika, Forêt de Tsimaloto, 18.3 km 46 ° NE de Tsaramandroso, - 16.22806, 47.14361, 135 m, Fisher et al. leg. (CASC); 7 w., 5 s., Parc National de Baie de Baly, 12.4 km 337 ° NNW Soalala, - 16.01 45.265, 10 m, Fisher et al. leg. (CASC); 12 w., 3 s., Parc National de Namoroka, 16.9 km 317 ° NW Vilanandro, - 16.40667 45.31, 100 m, Fisher et al. leg. (CASC); 2 w., 1 s., Parc National de Namoroka, 9.8 km 300 ° WNW Vilanandro, - 16.46667 45.35, 140 m, Fisher et al. leg. (CASC); 1 w., Parc National Tsingy de Bemaraha, 10.6 km ESE 123 ° Antsalova, - 18.70944 44.71817, 150 m, Fisher et al. leg. (CASC); 4 w., 1 s., Parc National Tsingy de Bemaraha, 2.5 km 62 ° ENE Bekopaka, Ankidrodroa River, - 19.13222 44.81467, 100 m, Fisher et al. leg. (CASC); 6 w., 1 s., Parc National Tsingy de Bemaraha, 3.4 km 93 ° E Bekopaka, Tombeau Vazimba, - 19.14194 44.828, 50 m, Fisher et al. leg. (CASC); 1 w, 1 s., Réserve d’Ankoririka, 10.6 km 13 ° NE de Tsaramandroso, - 16.26722 47.04861, 210 m, Fisher et al. leg. (CASC); 2 w., 1 s., Réserve Spéciale de Bemarivo, 23.8 km 223 ° SW Besalampy, - 16.925 44.36833, 30 m, Fisher et al. leg. (CASC); 1 w., Réserve forestière Beanka, 50.2 km E Maintirano, - 18.02649 44.05051, 250 m, Fisher et al. leg. (CASC). Toamasina: 1 w., 1 s., 6.3 km S Ambanizana, Andranobe, - 15.6813 49.958, 25 m, Fisher et al. leg. (CASC). Toliara: 1 w., 50 km N Morondava, - 20.06667, 44.58333, A. Pauly leg. (CASC); 1 w., 1 s., Anosy Region, Parc National d’Andohahela, Forêt de Manatalinjo, - 24.82505 46.57811, 90 m, Fisher et al. leg. (CASC); 1 w., Antafoky, - 23.48778 44.0775, 60 m, Fisher et al. leg. (CASC); 1 w., 1 s., Beza-Mahafaly, 27 km E Betioky, - 23.65 44.63333, 135 m, Fisher et al. leg. (CASC); 1 s., Fiherenana, - 23.17694 43.96083, 100 m, Fisher et al. leg. (CASC); 1 w., Fiherenana, - 23.23528 43.87083, 50 m, Fisher et al. leg. (CASC); 1 w., Forêt Classée d’Analavelona, 29.2 km 343 ° NNW Mahaboboka, - 22.675 44.19, 1100 m, Fisher et al. leg. (CASC); 3 w., 3 s., Forêt de Beroboka, 5.9 km 131 ° SE Ankidranoka, - 22.23306 43.36633, 80 m, Fisher et al. leg. (CASC); 4 w., Forêt de Kirindy, 15.5 km 64 ° ENE Marofandilia, - 20.045 44.66222, 100 m, Fisher et al. leg. (CASC); 7 w., Forêt de Kirindy, 15.5 km 64 ° ENE Marofandilia, - 20.06855 44.6595667, 30 m, Fisher et al. leg. (CASC); 4 w., Forêt de Mahavelo, Isantoria River, - 24.75833 46.15717, 110 m, Fisher et al. leg. (CASC); 1 w., Forêt de Mahavelo, Isantoria River, 5.5 km 37 ° NE Ifotaka, - 24.75361 46.1515, 115 m, Fisher et al. leg. (CASC); 3 w., 3 s., Forêt de Mite, 20.7 km 29 ° WNW Tongobory, - 23.52417 44.12133, 75 m, Fisher et al. leg. (CASC); 1 w., Forêt Ivohibe 55.0 km N Tolagnaro, - 24.569 47.204, 200 m, Fisher et al. leg. (CASC); 1 w., Forêt Vohidava 88.9 km N Amboasary, - 24.24067 46.28783, 500 m, Fisher et al. leg. (CASC); 1 w., 1 s., Forêt Vohidava 89.2 km N Amboasary, - 24.239 46.28233, 850 m, Fisher et al. leg. (CASC); 2 w., Ivahona, - 23.45591 46.17376, 820 m, Fisher et al. leg. (CASC); 1 w., Makay Mts., - 21.21836 45.3106, 510 m, Fisher et al. leg. (CASC); 2 w., Makay Mts., - 21.30997, 45.12946, 590 m, Fisher et al. leg. (CASC); 3 w., Manderano, - 23.52722 44.0875, 70 m, Fisher et al. leg. (CASC); 1 w., Manombo, - 22.81092 43.7344, 177 m, Fisher et al. leg. (CASC); 1 w., Manombo, - 22.8123 43.73932, 165 m, Fisher et al. leg. (CASC); 1 w., Parc National d’Andohahela, Col du Sedro, 3.8 km 113 ° ESE Mahamavo, 37.6 km 341 ° NNW Tolagnaro, - 24.76389 46.75167, 900 m, Fisher et al. leg. (CASC); 6 w., Parc National d’Andohahela, Forêt d’Ambohibory, 1.7 km 61 ° ENE Tsimelahy, 36.1 km 308 ° NW Tolagnaro, - 24.93 46.6455, 300 m, Fisher et al. leg. (CASC); 3 w., Parc National d’Andohahela, Forêt de Manatalinjo, 33.6 km 63 ° ENE Amboasary, 7.6 km 99 ° E Hazofotsy, - 24.81694 46.61, 150 m, Fisher et al. leg. (CASC); 1 w., Parc National de Kirindy Mite, 16.3 km 127 ° SE Belo sur Mer, - 20.79528 44.147, 80 m, Fisher et al. leg. (CASC); 4 w., Parc National de Zombitse, 19.8 km 84 ° E Sakaraha, - 22.84333 44.71, 770 m, Fisher et al. leg. (CASC); 11 w., 5 s., 3 m., Réserve Privé Berenty, Forêt de Bealoka, Mandraré River, 14.6 km 329 ° NNW Amboasary, - 24.95694 46.2715, 35 m, Fisher et al. leg. (CASC); 5 w., Réserve Spéciale d’Ambohijanahary, Forêt d’Ankazotsihitafototra, 34.6 km 314 ° NW Ambaravaranala, - 18.26 45.41833, 1100 m, Fisher et al. leg. (CASC); 2 w., Réserve Spéciale d’Ambohijanahary, Forêt d’Ankazotsihitafototra, 35.2 km 312 ° NW Ambaravaranala, - 18.26667 45.40667, 1050 m, Fisher et al. leg. (CASC); 1 w., Sakaraha, - 22.91233 44.53283, 470 m, Fisher et al. leg. (CASC); 1 w., Sept Lacs, - 23.52833, 44.15556, 80 m, Fisher et al. leg. (CASC); 1 w., southern Isoky-Vohimena Forest, 59 km NE Sakaraha, - 22.46667 44.85, 730 m, Fisher et al. leg. (CASC); 14 w., 9 s., Forêt Classée d’Analavelona, 29.2 km 343 ° NNW Mahaboboka, - 22.675 44.19, 1100 m, Fisher et al. leg. (CASC); 1 w., Forêt Classée d’Analavelona, 29.4 km 343 ° NNW Mahaboboka, - 22.675 44.18667, 1050 m, Fisher et al. leg. (CASC); 1 s., Forêt Vohidava 89.2 km N Amboasary, - 24.239 46.28233, 850 m, Fisher et al. leg. (CASC); 2 w., Makay Mts., - 21.21761, 45.33917, 500 m, Fisher et al. leg. (CASC); 2 w., Makay Mts., - 21.22336 45.32628, 480 m, Fisher et al. leg. (CASC); 4 w., Makay Mts., - 21.30997 45.12946, 590 m, Fisher et al. leg. (CASC); 2 w., 2 m., Makay Mts., - 21.31664 45.1296, 620 m, Fisher et al. leg. (CASC); 3 w., 2 s., Parc National de Zombitse, 19.8 km 84 ° E Sakaraha, - 22.84333 44.71, 770 m, Fisher et al. leg. (CASC); 1 w., Réserve Privé Berenty, Forêt de Malaza, Mandraré River, 8.6 km 314 ° NW Amboasary, - 25.00778 46.306, 40 m, Fisher et al. leg. (CASC). Geographic range. Madagascar, widely distributed on the island except Antsiranana.	en	Salata, Sebastian, Fisher, Brian L. (2020): The ant genus Pheidole Westwood, 1839 (Hymenoptera: Formicidae) in Madagascar-taxonomic revision of the bessonii species-group. Zootaxa 4843 (1): 1-64, DOI: 10.11646/zootaxa.4843.1.1
03938025FF9EA902A7D8B212FD19FBEE.taxon	diagnosis	Diagnosis. Major workers. Head in full-face view sub-rectangular, not or slightly widening posteriorly, lateral margin slightly convex with sparse, long, decumbent to subdecumbent pilosity; medial frons with moderately thick to thin and dense costulae that are on the posterior part; interspaces between costulae smooth to indistinctly punctate; frons laterally with sparser and thinner rugocostulae; interspaces between rugocostulae distinctly punctate; occipital lobes punctate, sometimes with additional network of sparse and thin rugae, sculpture weakening posteriorly; scape, when laid back, exceeding the midlength of head by two-fifths of its length; inner hypostomal tooth absent or indistinct, bulge-like; outer hypostomal tooth lobe-like, high and wide, directed posteriorly; median tooth absent; promesonotum short, angular, and moderately low; posterior mesonotum moderately steep; mesonotal process distinct and tubercle-like; mesosoma microreticulate; promesonotum with additional thin, sparse, and transverse rugae; gaster shagreened; body brown to blackish brown. Minor workers. Occiput elongated; nuchal collar distinct; head sculpture smooth; anterior frons with sparse puncta; scape, when laid back, exceeding the posterior head margin by two-fifths of its length; promesonotum low, long, and arched; promesonotal groove present; metanotal groove distinct; propodeal spines very small with relatively wide base; promesonotum mostly smooth with sparse microreticulae laterally; mesopleuron and propodeum with sparse microreticulae; body bright brown to brown.	en	Salata, Sebastian, Fisher, Brian L. (2020): The ant genus Pheidole Westwood, 1839 (Hymenoptera: Formicidae) in Madagascar-taxonomic revision of the bessonii species-group. Zootaxa 4843 (1): 1-64, DOI: 10.11646/zootaxa.4843.1.1
03938025FF9EA902A7D8B212FD19FBEE.taxon	description	Description. Major workers. Measurements (n = 10): HL: 1.78 – 2.13 (1.97); HW: 1.74 – 2.01 (1.89); SL: 1.13 – 1.2 (1.17); EL: 0.22 – 0.27 (0.25); WL: 1.61 – 1.78 (1.68); PSL: 0.26 – 0.34 (0.3); MTL: 1.24 – 1.38 (1.3); PNW: 0.66 – 0.75 (0.7); PTW: 0.22 – 0.31 (0.28); PPW: 0.62 – 0.81 (0.7); CI: 99.7 – 106.5 (104.3); SI: 57.8 – 65.4 (61.7); PSLI: 13.9 – 16.6 (15.3); PPI: 32.1 – 47.0 (40.4); PNI: 34.2 – 39.1 (37.0); MTI: 64.8 – 74.8 (69.1). Head. In full-face view sub-rectangular, slightly widening posteriorly with sides slightly convex (Fig. 11 B); in lateral view sub-oval, not depressed posteriorly; ventral and dorsal margins convex; inner hypostomal tooth not visible. Sides of head with sparse, long, decumbent to subdecumbent pilosity; head dorsum with relatively dense, long, decumbent to erect pilosity. Medial frons with moderately thick to thin and dense costulae on the posterior part; interspaces between costulae smooth to indistinctly punctate. Frons laterally with sparser and thinner rugocostulae; interspaces between rugocostulae distinctly punctate. Occipital lobes punctate, sometimes with additional network of sparse and thin rugae; sculpture weakening posteriorly. Gena with dense, moderately thick to thin costulae; interspaces between costulae distinctly punctate. Sides posterolateral from eyes densely punctate to microreticulate, sometimes with additional network of thin and sparse rugulae. Center of clypeus shiny and with a few rugulae, lateral sides with distinct rugulae; median notch present, moderately wide, and shallow; median longitudinal carina present; lateral longitudinal carinae present. Scape, when laid back, exceeding the midlength of head by two-fifths of its length; pilosity subdecumbent to erect (Fig. 11 B, D). Inner hypostomal tooth absent or indistinct, bulge-like; outer hypostomal tooth lobe-like, high, and wide ,, with top directed posteriorly; median tooth absent (Fig. 27 C). Mesosoma. In lateral view, promesonotum short, angular, and moderately low; posterior mesonotum moderately steep; mesonotal process distinct and tubercle-like; promesonotal groove weakly developed; metanotal groove present; propodeal spines moderately long with wide base and acute top; humeral tubercle weakly produced (Fig. 11 D). Surface shiny and microreticulate; promesonotum with additional thin, sparse, and transverse rugae. Pilosity moderately dense, long, and erect (Fig. 11 D, F). Gaster. Dull and shagreened; pilosity moderately sparse, long, and erect (Fig. 11 D, F). Color. Brown to blackish brown, antennae and legs yellowish brown, mandibles ochraceous (Fig. 11 D, F). Description. Minor workers. Measurements (n = 10): HL: 0.69 – 0.82 (0.77); HW: 0.52 – 0.61 (0.57); SL: 0.9 – 1.09 (1.0); EL: 0.13 – 0.19 (0.16); WL: 0.96 – 1.12 (1.04); PSL: 0.07 – 0.12 (0.1); MTL: 0.78 – 0.95 (0.88); PNW: 0.33 – 0.41 (0.37); PTW: 0.09 – 0.12 (0.1); PPW: 0.17 – 0.21 (0.18); CI: 131.4 – 137.6 (133.5); SI: 169.3 – 183.2 (175.0); PSLI: 10.2 – 15.3 (13.3); PPI: 48.7 – 60.0 (55.3); PNI: 61.0 – 68.7 (65.2); MTI: 144.7 – 158.1 (152.8). Head. In full-face view oval, posterior region elongated; nuchal collar distinct (Fig. 11 A). Pilosity relatively sparse, moderately long, subdecumbent to erect. Sculpture shiny and smooth; anterolateral frons with sparse puncta. Clypeus with median longitudinal carina absent; two lateral longitudinal carinae absent. Scape, when laid back, exceeding the posterior head margin by two-fifths of its length; pilosity dense, subdecumbent to erect (Fig. 11 A, C). Mesosoma. In lateral view, promesonotum low, long, and arched; promesonotal groove present; metanotal groove distinct; propodeal spines very small, triangular, and with relatively wide base (Fig. 11 C). Promesonotum mostly smooth with sparse microreticulae on lateral sides; katepisternum, anepisternum, and propodeum with sparse microreticulae. Pilosity very sparse, moderately long, and erect (Fig. 11 C, E). Gaster. With sparse and erect pilosity (Fig. 11 C, E). Color. Bright brown to brown, legs, mandibles, and antennae yellowish (Fig. 11 C, E).	en	Salata, Sebastian, Fisher, Brian L. (2020): The ant genus Pheidole Westwood, 1839 (Hymenoptera: Formicidae) in Madagascar-taxonomic revision of the bessonii species-group. Zootaxa 4843 (1): 1-64, DOI: 10.11646/zootaxa.4843.1.1
03938025FF9EA902A7D8B212FD19FBEE.taxon	biology_ecology	Biology. The species was collected between 30 – 1732 m elevation, in Uapaca woodland, montane rainforest, tropical dry forest, savanna and woodland, gallery forest and rainforest. Nests were located in rotten logs, in the soil, and under stones. Workers were collected from sifted litter.	en	Salata, Sebastian, Fisher, Brian L. (2020): The ant genus Pheidole Westwood, 1839 (Hymenoptera: Formicidae) in Madagascar-taxonomic revision of the bessonii species-group. Zootaxa 4843 (1): 1-64, DOI: 10.11646/zootaxa.4843.1.1
03938025FF9EA902A7D8B212FD19FBEE.taxon	discussion	Comments. Pheidole bessonii is probably one of the most widespread Pheidole species known from Madagascar. Morphologically it is most similar to P. voreios and P. atsirakambiaty, and its major workers can be separated from both species by punctate occipital lobes (which sometimes have additional thin and very sparse rugulae that arch posterolaterally), and sparsely rugocostulate lateral frons with distinctly punctate interspaces between rugocostulae. In contrast, majors of P. voreios and P. atsirakambiaty always have distinctly rugulate to costulate occipital lobes and frons. Minors of P. bessonii can be separated from P. voreios and P. atsirakambiaty by the less elongated posterior part of the neck, a mostly smooth promesonotum with sparse microreticulae on its sides, and very small propodeal spines. Pheidole voreios and P. atsirakambiaty have a more elongated posterior part of the neck, mostly or entirely sculptured promesonotum, and larger propodeal spines. In addition, majors and minors of P. bessonii can be distinguished from P. voreios by the brown body.	en	Salata, Sebastian, Fisher, Brian L. (2020): The ant genus Pheidole Westwood, 1839 (Hymenoptera: Formicidae) in Madagascar-taxonomic revision of the bessonii species-group. Zootaxa 4843 (1): 1-64, DOI: 10.11646/zootaxa.4843.1.1
03938025FF9AA900A7D8B26AFACEF932.taxon	description	Figs 12 A – F, 27 D, 28 D	en	Salata, Sebastian, Fisher, Brian L. (2020): The ant genus Pheidole Westwood, 1839 (Hymenoptera: Formicidae) in Madagascar-taxonomic revision of the bessonii species-group. Zootaxa 4843 (1): 1-64, DOI: 10.11646/zootaxa.4843.1.1
03938025FF9AA900A7D8B26AFACEF932.taxon	materials_examined	HOLOTYPE: 1 s., Madagascar, Antsiranana, Parc National de Marojejy, Manantenina River, 27.6 km 35 ° NE Andapa, 9.6 km 327 ° NNW Manantenina, - 14.435 49.76, 775 m, 17 - Nov- 2003, rainforest, ex dead twig above ground, B. L. Fisher et al. leg. BLF 09023, CASENT 0494851 (CASC). PARATYPES: 2 w., 1 s., 1 q the same data as holotype, CASENT 0494852, CASENT 0494850, CASENT 0494849 (CASC, MHNG, PBZT). Other material. Madagascar. Toamasina: 3 w., 2 s., 1 q., 5.3 km SSE Ambanizana, Andranobe, - 15.67133 49.97395, 425 m, B. L. Fisher et al. leg. (CASC). Geographic range. Madagascar, from Parc National de Marojejy, Antsiranana south to Andranobe, Toamasina.	en	Salata, Sebastian, Fisher, Brian L. (2020): The ant genus Pheidole Westwood, 1839 (Hymenoptera: Formicidae) in Madagascar-taxonomic revision of the bessonii species-group. Zootaxa 4843 (1): 1-64, DOI: 10.11646/zootaxa.4843.1.1
03938025FF9AA900A7D8B26AFACEF932.taxon	diagnosis	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; medial frons with dense costulae; interspaces between costulae smooth to indistinctly punctate; frons laterally with thick and dense rugae; interspaces between rugae distinctly punctate; occipital lobes with weakly developed and sparse rugae; interspaces between rugae punctate; sculpture weakening posteriorly; scape, when laid back, exceeding the midlength of head by two-fifths of its length; inner hypostomal tooth distinct, large, triangular, and directed posteriorly; outer hypostomal tooth lobe-like, high, and wide, with top arching posterolaterally; median tooth absent; promesonotum short, angular, and moderately low; posterior mesonotum moderately steep; mesonotal process indistinct and tubercle-like; mesosoma punctate; promesonotal dorsum with sculpture reduced and additional transverse rugae; gaster smooth; body orange. Minor workers. Occiput elongated forming short and moderately narrow neck; nuchal collar distinct; head sculpture smooth; anterolateral frons with sparse puncta; scape, when laid back, exceeding the posterior head margin by three-fifths of its length; promesonotum low, long, and arched; promesonotal groove present; metanotal groove distinct; propodeal spines small and thin; mesosoma punctate; katepisternum with a smooth notch; body yellow.	en	Salata, Sebastian, Fisher, Brian L. (2020): The ant genus Pheidole Westwood, 1839 (Hymenoptera: Formicidae) in Madagascar-taxonomic revision of the bessonii species-group. Zootaxa 4843 (1): 1-64, DOI: 10.11646/zootaxa.4843.1.1
03938025FF9AA900A7D8B26AFACEF932.taxon	description	Description. Major workers. Measurements (n = 3): HL: 1.77 – 1.9 (1.81); HW: 1.67 – 1.79 (1.73); SL: 1.01 – 1.14 (1.05); EL: 0.22 – 0.23 (0.23); WL: 1.55 – 1.59 (1.57); PSL: 0.24 – 0.27 (0.26); MTL: 1.16 – 1.24 (1.19); PNW: 0.62 – 0.66 (0.64); PTW: 0.18 – 0.2 (0.18); PPW: 0.4 – 0.44 (0.42); CI: 103.0 – 105.7 (104.3); SI: 58.1 – 63.6 (60.7); PSLI: 13.1 – 15.2 (14.4); PPI: 42.5 – 46.0 (44.5); PNI: 36.4 – 37.1 (36.8); MTI: 68.3 – 69.2 (68.9). Head. In full-face view sub-rectangular, not widening posteriorly, with lateral sides slightly convex (Fig. 12 B). 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. Medial frons with dense costulae; interspaces between costulae smooth to indistinctly punctate. Frons laterally with thick and dense rugae; interspaces between rugae distinctly punctate. Occipital lobes with weakly developed and sparse rugae; interspaces between rugae punctate; sculpture weakening posteriorly. Gena with dense and moderately thick costulae; interspaces between costulae smooth. Sides posterolateral from eyes with dense but very thin network of rugoreticulae; sculpture weakening posteriorly. 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, exceeding the midlength of head by two-fifths of its length; pilosity subdecumbent to erect (Fig. 12 B, D). Inner hypostomal tooth distinct, large, triangular, and directed posteriorly; outer hypostomal tooth lobe-like, high, and wide ,, with top arching posterolaterally; median tooth absent (Fig. 27 D). Mesosoma. In lateral view, promesonotum short, angular, and moderately low; posterior mesonotum moderately steep; mesonotal process indistinct and tubercle-like; promesonotal groove absent; metanotal groove indistinct; propodeal spines moderately long with narrow base and acute top; humeral tubercle weakly produced (Fig. 12 D). Surface punctate; promesonotal dorsum with reduced puncta and additional transverse rugae. Pilosity dense, long, and erect (Fig. 12 D, F). Gaster. Shiny and smooth; pilosity moderately sparse, long, and erect (Fig. 12 D, F). Color. Orange, antennae and legs yellow (Fig. 12 D, F). Description. Minor workers. Measurements (n = 5): HL: 0.75 – 0.82 (0.78); HW: 0.52 – 0.59 (0.55); SL: 1.04 – 1.1 (1.06); EL: 0.14 – 0.17 (0.16); WL: 1.03 – 1.12 (1.07); PSL: 0.1 – 0.13 (0.11); MTL: 0.92 – 1.03 (0.96); PNW: 0.36 – 0.4 (0.38); PTW: 0.08 – 0.11 (0.09); PPW: 0.14 – 0.17 (0.14); CI: 140.3 – 145.9 (143.2); SI: 188.4 – 206.6 (195.5); PSLI: 12.9 – 15.8 (14.7); PPI: 52.1 – 77.5 (62.3); PNI: 65.5 – 72.1 (69.1); MTI: 171.1 – 177.3 (175.2). Head. In full-face view oval, posterior region elongated forming short and moderately narrow neck; nuchal collar distinct (Fig. 12 A). Pilosity relatively sparse, long, subdecumbent to erect. Sculpture shiny and smooth; anterolateral frons with sparse puncta; antennal sockets with few indistinct rugulae that are curved posterolaterally; interspaces between rugulae smooth. Clypeus with median longitudinal carina absent; two lateral longitudinal carinae absent. Scape, when laid back, exceeding the posterior head margin by three-fifths of its length; pilosity dense, subdecumbent to erect (Fig. 12 A, C). Mesosoma. In lateral view, promesonotum low, long, and arched; promesonotal groove present; metanotal groove distinct; propodeal spines small, thin, and triangular (Fig. 12 C). Sculpture punctate, katepisternum with a smooth notch. Pilosity very dense, long, and erect (Fig. 12 C, E). Gaster. With sparse and erect pilosity (Fig. 12 C, E). Color. Yellow (Fig. 12 C, E).	en	Salata, Sebastian, Fisher, Brian L. (2020): The ant genus Pheidole Westwood, 1839 (Hymenoptera: Formicidae) in Madagascar-taxonomic revision of the bessonii species-group. Zootaxa 4843 (1): 1-64, DOI: 10.11646/zootaxa.4843.1.1
03938025FF9AA900A7D8B26AFACEF932.taxon	biology_ecology	Biology. The species was collected between 425 – 775 m elevation, in rainforest. Nests were located in rotten logs and dead twigs above the ground.	en	Salata, Sebastian, Fisher, Brian L. (2020): The ant genus Pheidole Westwood, 1839 (Hymenoptera: Formicidae) in Madagascar-taxonomic revision of the bessonii species-group. Zootaxa 4843 (1): 1-64, DOI: 10.11646/zootaxa.4843.1.1
03938025FF9AA900A7D8B26AFACEF932.taxon	discussion	Comments. Pheidole clara, a species recorded from southeastern Antsiranana and the northernmost part of Toamasina, is most similar to P. mantadioflava, known from the northeastern part of Toamasina. Majors of P. clara differ in the absence of antennal scrobes and the anterior mesonotum being at the same level or lower than the pronotum. In contrast, P. mantadioflava has weakly developed antennal scrobes and its anterior mesonotum is placed slightly higher than the pronotum. Minors of P. clara can be separated by sparse puncta restricted only to anterolateral frons and by their small and thin propodeal spines. Minors of P. mantadioflava have entire frons sparsely punctate and their propodeal spines are small with a wide base.	en	Salata, Sebastian, Fisher, Brian L. (2020): The ant genus Pheidole Westwood, 1839 (Hymenoptera: Formicidae) in Madagascar-taxonomic revision of the bessonii species-group. Zootaxa 4843 (1): 1-64, DOI: 10.11646/zootaxa.4843.1.1
03938025FF9AA900A7D8B26AFACEF932.taxon	etymology	Etymology. Latin for clear or bright, in reference to the bright body coloration of minor workers.	en	Salata, Sebastian, Fisher, Brian L. (2020): The ant genus Pheidole Westwood, 1839 (Hymenoptera: Formicidae) in Madagascar-taxonomic revision of the bessonii species-group. Zootaxa 4843 (1): 1-64, DOI: 10.11646/zootaxa.4843.1.1
03938025FF98A90DA7D8B11EFC50FC34.taxon	description	Figs 13 A – F, 27 E, 28 E	en	Salata, Sebastian, Fisher, Brian L. (2020): The ant genus Pheidole Westwood, 1839 (Hymenoptera: Formicidae) in Madagascar-taxonomic revision of the bessonii species-group. Zootaxa 4843 (1): 1-64, DOI: 10.11646/zootaxa.4843.1.1
03938025FF98A90DA7D8B11EFC50FC34.taxon	description	pin as lectotype, CASENT 0876547, ANTC 3357 (MHNG); 2 w., the same data as lectotype, CASENT 0101720, ANTC 3358 (MHNG).	en	Salata, Sebastian, Fisher, Brian L. (2020): The ant genus Pheidole Westwood, 1839 (Hymenoptera: Formicidae) in Madagascar-taxonomic revision of the bessonii species-group. Zootaxa 4843 (1): 1-64, DOI: 10.11646/zootaxa.4843.1.1
03938025FF98A90DA7D8B11EFC50FC34.taxon	materials_examined	Other material. Madagascar. Antananarivo: 29 w., 9 s., 3 km 41 ° NE Andranomay, 11.5 km 147 ° SSE Anjozorobe, - 18.47333 47.96, 1300 m, Fisher et al. leg. (CASC); 1 w., Analamanga Region, District of Ankazobe, Ambohitantely, 46 km NE of Ankazobe, - 18.198 47.2815, 701 m, M. Rinha leg. (CASC); 3 w., Forêt de galerie, Andranorovitra, 24.0 km NNE Ankazobe, - 18.11243 47.19757, 1491 m, Fisher et al. leg. (CASC); 5 w., 4 s., Forêt de galerie, Telomirahavavy, 23.4 km NNE Ankazobe, - 18.12167 47.20627, 1520 m, Fisher et al. leg. (CASC); 5 w., 2 s., 1 m., Mandraka, - 18.91813 47.91717, 1312 m, Fisher et al. leg. (CASC); 3 w., 3 s., Réserve Naturelle Sohisika, Sohisika 24.6 km NNE Ankazobe, - 18.10322 47.18692, 1464 m, Fisher et al. leg. (CASC); 4 w., 3 s., Réserve Spéciale d’Ambohitantely, Forêt d Ambohitantely, 20.9 km 72 ° NE d Ankazobe, - 18.22528 47.28683, 1410 m, Fisher et al. leg. (CASC); Antsiranana: 2 w., 1 s., 2.0 km S Andrakata, - 14.65 49.71667, 520 m, Fisher et al. leg. (CASC); 5 w., 5 s., 1 q., 6.5 km SSW Befingotra, Rés. Anjanaharibe-Sud, - 14.75 49.5, 875 m, Fisher et al. leg. (CASC); 1 w., 7 km N Joffreville [camp 2 of Fisher], - 12.33333 49.25, 360 m, R. Harin’Hala leg. (CASC); 1 w., 9.2 km WSW Befingotra, Rés. Anjanaharibe-Sud, - 14.75 49.46667, 1280 m, Fisher et al. leg. (CASC); 3 w., 9.2 km WSW Befingotra, Rés. Anjanaharibe-Sud, - 14.75 49.46667, 1200 m, Fisher et al. leg. (CASC); 2 w., 1 s., Ambondrobe, 41.1 km 175 ° Vohemar, - 13.71533 50.10167, 10 m, Fisher et al. leg. (CASC); 1 s., Ampasindava, Forêt d’Ambilanivy, 3.9 km 181 ° S Ambaliha, - 13.79861 48.16167, 600 m, Fisher et al. leg. (CASC); 17 w., 13 s., 3 m., Betaolana Forest, along Bekona River, - 14.52996 49.44039, 880 m, Fisher et al. leg. (CASC); 4 w., 4 s., Binara Forest, - 13.26392 49.59919, 1065 m, Fisher et al. leg. (CASC); 2 w., 2 s., Binara Forest, - 13.26388 49.60141, 900 m, Fisher et al. leg. (CASC); 2 w., 2 s., Binara Forest, - 13.26206 49.60672, 559 m, Fisher et al. leg. (CASC); 8 w., 4 s., 1 m., 1 q., Forêt Ambanitaza, 26.1 km 347 ° Antalaha, - 14.67933 50.18367, 240 m, Fisher et al. leg. (CASC); 2 w., Forêt de Binara, 9.1 km 233 ° SW Daraina, - 13.26333 49.60333, 650 - 800 m, Fisher et al. leg. (CASC); 3 w., 7 s., Forêt de Binara, 9.4 km 235 ° SW Daraina, - 13.26333 49.6, 1100 m, Fisher et al. leg. (CASC); 3 w., Makirovana forest, - 14.17066 49.95409, 415 m, Fisher et al. leg. (CASC); 2 w., Makirovana forest, - 14.16666 49.95, 715 m, Fisher et al. leg. (CASC); 4 w., Makirovana forest, - 14.16044 49.95216, 550 m, Fisher et al. leg. (CASC); 1 w., Marojejy, tributary Manantenina R., - 14.43333 49.75, 750 m, Quinter & Nguyen leg. (CASC); 1 w., Masoala National Park, - 15.3014 50.22776, 280 m, Fisher et al. leg. (CASC); 2 w., 1 s., Nosy Be, Réserve Naturelle Intégrale de Lokobe, 6.3 km 112 ° ESE Hellville, - 13.41933 48.33117, 30 m, Fisher et al. leg. (CASC); 13 w., 2 s., Parc National de Marojejy, Antranohofa, 26.6 km 31 ° NNE Andapa, 10.7 km 318 ° NW Manantenina, - 14.44333 49.74333, 1325 m, Fisher et al. leg. (CASC); 3 w., 1 s., Parc National de Marojejy, Manantenina River, 27.6 km 35 ° NE Andapa, 9.6 km 327 ° NNW Manantenina, - 14.435 49.76, 775 m, Fisher et al. leg. (CASC); 27 w., 10 s., Parc National de Marojejy, Manantenina River, 28.0 km 38 ° NE Andapa, 8.2 km 333 ° NNW Manantenina, - 14.43667 49.775, 450 m, Fisher et al. leg. (CASC); 6 w., Parc National Montagne d’Ambre [1 st campsite], 960 m, Irwin et al. leg. (CASC); 1 w., Parc National Montagne d’Ambre [lemur trail], 975 m, R. Harin’Hala leg. (CASC); 4 w., Parc National Montagne d’Ambre [Petit Lac road], - 12.533333, 49.166667, 1125 m, R. Harin’Hala leg. (CASC); 1 s., Parc National Montagne d’Ambre, 12.2 km 211 ° SSW Joffreville, - 12.59639 49.1595, 1300 m, Fisher et al. leg. (CASC); 12 w., 9 s., 1 q. Parc National Montagne d’Ambre, 3.6 km 235 ° SW Joffreville, - 12.53444 49.1795, 925 m, J. Boutin leg. (CASC); 2 w., 1 s., Parc National Montagne d’Ambre, Antomboka, - 12.51269 49.17807, 970 m, Fisher et al. leg. (CASC); 2 w., 1 s., Parc National Montagne d’Ambre, Lac maudit, - 12.58502 49.15147, 1250 m, Fisher et al. leg. (CASC); 2 w., 1 s., 1 q., Parc National Montagne d’Ambre, Mahasarika, - 12.53176 49.17662, 1135 m, Fisher et al. leg. (CASC); 1 w., 1 s., Parc National Montagne d’Ambre, Roussettes, - 12.52574 49.17238, 1025 m, Fisher et al. leg. (CASC); 3 w., 3 s., Prov. Antsiranana R. S. Manongarivo 17.3 km 218 ° SW Antanambao, - 14.02167 48.41833, 1580 m, Fisher et al. leg. (CASC); 1 s., R. S. Manongarivo, 12.8 km 228 ° SW Antanambao, - 13.97667 48.42333, 780 m, Fisher et al. leg. (CASC); 3 w., 3 s., R. S. Manongarivo, 14.5 km 220 ° SW Antanambao, - 14 48.43167, 1220 m, Fisher et al. leg. (CASC); 1 w., 1 s., R. S. Manongarivo, 14.5 km 220 ° SW Antanambao, - 13.99833 48.42833, 1175 m, Fisher et al. leg. (CASC); 3 w., 3 s., R. S. Manongarivo, 14.5 km 220 ° SW Antanambao, - 13.99833 48.42833, 1175 m, Fisher et al. leg. (CASC); 4 w., 1 s., Réserve Spéciale d’Ambre, 3.5 km 235 ° SW Sakaramy, - 12.46889 49.24217, 325 m, Fisher et al. leg. (CASC); 1 w., RNI Marojejy, 10 km NW Manantenina, - 14.43333, 49.76667, 750 m, E. L. Quinter leg. (CASC); 1 w., RNI Marojejy, 8 km NW Manantenina, - 14.43333 49.78333, 450 m, E. L. Quinter leg. (CASC); 2 w., Sakalava Beach [vegetated beach dunes], - 12.26972 49.39167, 10 m, R. Harin’Hala leg. (CASC); 4 w., 1 s., SAVA Region, District of Sambava, Marojejy National Park, 5 km W of Manantenina village, 1 st Camp site (Mantella), - 14.43817 49.774, 487 m, M. Rin’Ha leg. (CASC); 2 w., 1 s., 1 q., Sava Region: Parc National de Marojejy, Manantenina River, 21.3 km 27.0 ° NE Andapa, - 14.43686 49.74291, 1350 m, Fisher et al. leg. (CASC); 3 w., 1 s., 1 m., Sava Region: Parc National de Marojejy, Manantenina River, 28.0 km 24.8 ° NE Andapa, - 14.43461 49.76074, 780 m, Fisher et al. leg. (CASC); 4 w., 2 s., 1 m., Sava Region: Parc National de Marojejy, Manantenina River, 28.0 km 24.8 ° NE Andapa, - 14.43461 49.76074, 780 m, Fisher et al. leg. (CASC); 2 w., 1 s., 1 m., Sava Region: Parc National de Marojejy, Manantenina River, 28.1 km 25.7 ° NE Andapa, - 14.43553 49.76463, 680 m, Fisher et al. leg. (CASC); 5 w., 3 s., 2 q., Sava Region: Parc National de Marojejy, near Manantenina River, - 14.43677 49.77541, 475 m, Fisher et al. leg. (CASC). Fianarantsoa: 6 w., 2 s., 2 km W Andrambovato, along river Tatamaly, - 21.51167 47.41, 1075 m, Fisher et al. leg. (CASC); 2 w., 40 km S Ambalavao, Rés. Andringitra, - 22.21667 46.96667, 1275 m, Fisher et al. leg. (CASC); 1 w., 1 q., 43 km S Ambalavao, Rés. Andringitra, - 22.23333 47, 825 m, Fisher et al. leg. (CASC); 2 w., 1 s., 45 km S Ambalavao, - 22.21667 47.01667, 720 m, Fisher et al. leg. (CASC); 7 w., 4 s., 1 q., 45 km S. Ambalavao, - 22.21667 47.01667, 785 m, Fisher et al. leg. (CASC); 5 w., 3 s., 7.6 km 122 º Kianjavato, Forêt Classée Vatovavy, - 21.4 47.94, 175 m, Fisher et al. leg. (CASC); 2 w., 1 s., 8.0 km NE Ivohibe, - 22.42167 46.89833, 1200 m, Fisher et al. leg. (CASC); 2 w., 9.0 km NE Ivohibe, - 22.42667 46.93833, 900 m, Fisher et al. leg. (CASC); 1 s., dry wash, 1 km E of Isalo National Park Interpretive Center, Fianarantsoa Prov., - 22.62667 45.35817, 885 m, R. Harin’Hala leg. (CASC); 1 s., Forêt d’Ambalagoavy Nord, Ikongo, Ambatombe, - 21.857068 47.37849, 625 m, R. Harin’Hala leg. (CASC); 1 w., Forêt d’Atsirakambiaty, 7.6 km 285 ° WNW Itremo, - 20.59333 46.56333, 1550 m, Fisher et al. leg. (CASC); 3 w., Forêt d’Atsirakambiaty, 7.6 km 285 ° WNW Itremo, - 20.59333 46.56333, 1550 m, Fisher et al. leg. (CASC); 4 w., 2 s., Forêt de Vevembe, 66.6 km 293 ° Farafangana, - 22.791 47.18183, 600 m, 23 - Apr- 2006, rainforest, transition to montane forest, Fisher et al. leg. (CASC); 3 w., 4 s., Parc National Befotaka-Midongy, Papango 27.7 km S Midongy-Sud, Mount Papango, - 23.83517 46.96367, 940 m, Fisher et al. leg. (CASC); 2 w., 2 s., Parc National Befotaka-Midongy, Papango 28.5 km S Midongy-Sud, Mount Papango, - 23.84083 46.9575, 1250 m, Fisher et al. leg. (CASC); 1 w., Parc National d’Isalo, 9.1 km 354 ° N Ranohira, - 22.48167 45.46167, 725 m, Fisher et al. leg. (CASC); 10 w., 2 s., Parc National de Ranomafana, Vatoharanana River, 4.1 km 231 ° SW Ranomafana, - 21.29 47.43333, 1100 m, Fisher et al. leg. (CASC); 2 w., 1 s., Parc Nationale Ranomafana: Talatakely, - 21.24833 47.42667, Griswold et al. leg. (CASC); 8 w., 1 s., R. S. Ivohibe 8.0 km E Ivohibe, - 22.48333 46.96833, 1200 m, Fisher et al. leg. (CASC); 2 w., 1 s., R. S. Ivohibe, 7.5 km ENE Ivohibe, - 22.47 46.96, 900 m, Fisher et al. leg. (CASC); 1 w., radio tower, Ranomafana National Park, Fianarantsoa Prov., - 21.25833 47.40717, 1130 m, Irwin et al. leg. (CASC); 1 s., Ranomafana, - 21.25 47.36667, A. Pauly leg. (CASC); 1 w., Ranomafana National Park, Talatakely area, 0.4 km WSW of Park Entrance, - 21.41667 47.68333, 900 m, D. H. & K. M. Kavanaugh leg. (CASC); 1 w., Vohiparara broken bridge, Fianarantsoa Prov., - 21.22617 47.36983, 1110 m, R. Harin’Hala leg. (CASC); 1 s., Vohiparara Kidonavo 1, - 21.22632 47.37007, 1100 m, V. C. Clark leg. (CASC). Mahajanga: 28 w., 3 m., 3 q., Region Sofia, Ampotsidia, - 14.42935 48.69863, 1193 m, Fisher et al. leg. (CASC); 11 w., Region Sofia, Ampotsidia, - 14.40775 48.70201, 1625 m, Fisher et al. leg. (CASC); 6 w., 2 m., Region Sofia, Ampotsidia, - 14.41592 48.71118, 1364 m, Fisher et al. leg. (CASC); 4 w., 1 m., 1 q., Region Sofia, Ampotsidia, - 14.42188 48.72083, 1400 m, Fisher et al. leg. (CASC); 2 w., 3 s., Réserve Spéciale Marotandrano, Marotandrano 48.3 km S Mandritsara, - 16.28322 48.81443, 865 m, Fisher et al. leg. (CASC); 1 w., 1 s., 1 q., Toamasina, 5.3 km SSE Ambanizana, Andranobe, - 15.67133 49.97395, 425 m, Fisher et al. leg. (CASC); 1 w., 2 s., 6.8 km S Ambanizana, - 15.68333, 49.8, 25 m, Fisher et al. leg. (CASC); 1 w., 2 s., 6.9 km NE Ambanizana, Ambohitsitondroina, - 15.58506 50.00952, 825 m, Fisher et al. leg. (CASC); 1 w., 2 s., 6.9 km NE Ambanizana, Ambohitsitondroina, - 15.56667 50, 1000 m, Fisher et al. leg. (CASC); 1 w., Ambanizana, Parc National Masoala, - 15.57167 50.00611, 800 - 897 m, Andriamalala et al. leg. (CASC); 1 w., 1 s., Ambanizana, Parc National Masoala, - 15.57222 50.00694, 930 - 1110 m, Andriamalala et al. leg. (CASC); 1 s., Ambatovy, 12.4 km NE Moramanga, - 18.83937 48.30842, 1080 m, Fisher et al. leg. (CASC); 1 w., 1 s., Ambatovy, 12.4 km NE Moramanga, - 18.84773 48.29568, 1000 m, Fisher et al. leg. (CASC); 1 w., Ambatovy, 12.4 km NE Moramanga, - 18.85813 48.28488, 1040 m, Fisher et al. leg. (CASC); 1 w., Analamay, - 18.80623 48.33707, 1068 m, Fisher et al. leg. (CASC); 3 w., Andasibe National Park, botanic garden near entrance, West of ANGAP office, - 18.925172 48.418651, 1025 m, Irwin et al. leg. (CASC); 1 w., F. C. Didy, - 18.19833 48.57833, 960 m, H. J. Ratsirarson leg. (CASC); 2 w., F. C. Sandranantitra, - 18.04833 49.09167, 450 m, H. J. Ratsirarson leg. (CASC); 4 w., 1 s., Forêt Ambatovy, 14.3 km 57 ° Moramanga, - 18.85083 48.32, 1075 m, Fisher et al. leg. (CASC); 1 w., Manakambahiny, near Vavatenina Forest, - 17.46667 49.35, A. Pauly leg. (CASC); 1 w., Montagne d’Akirindro 7.6 km 341 ° NNW Ambinanitelo, - 15.28833 49.54833, 600 m, Fisher et al. leg. (CASC); 5 w., 1 s., Montagne d’Anjanaharibe, 18.0 km 21 ° NNE Ambinanitelo, - 15.18833 49.615, 470 m, Fisher et al. leg. (CASC); 3 w., 2 s., Montagne d’Anjanaharibe, 19.5 km 27 ° NNE Ambinanitelo, - 15.17833 49.635, 1100 m, Fisher et al. leg. (CASC); 1 w., P. N. Mantadia, - 18.79167 48.42667, 895 m, H. J. Ratsirarson leg. (CASC); 2 w., 2 s., Parc National de Zahamena, Besaky River, - 17.75244 48.85321, 760 m, Fisher et al. leg. (CASC); 2 w., Parc National Mananara-Nord, 7.1 km 261 ° Antanambe, - 16.455 49.7875, 225 m, Fisher et al. leg. (CASC); 1 w., Reserve Betampona, Camp Vohitsivalana, 37.1 km 338 ° Toamasina, - 17.88667, 49.2025 520 m, Fisher et al. leg. (CASC); 3 w., 1 s., Réserve Nationale Intégrale Betampona, Betampona 35.1 km NW Toamasina, - 17.91801 49.20074, 500 m, Fisher et al. leg. (CASC); 1 w., 1 s., Sahafina forest 11.4 km W Brickaville, - 18.81445 48.96205, 140 m, Fisher et al. leg. (CASC); 2 w., Toliara, 10 km NW Enakara, Rés. Andohahela, - 24.56667 46.81667, 430 m, Fisher et al. leg. (CASC); 4 w., 1 s., 11 km NW Enakara, Rés. Andohahela, - 24.56667 46.83333, 800 m, Fisher et al. leg. (CASC); 1 w., 2 s., 11 km NW Enkara, Rés Andohahela, - 24.56667 46.81667, 950 m, Fisher et al. leg. (CASC); 1 w., 13 km NW Enakara, Rés. Andohahela, - 24.55. 46.8, 1250 m, Fisher et al. leg. (CASC); 2 w., 1 s., 13 km NW Enkara, Rés Andohahela, - 24.56667 46.81667, 1000 m, Fisher et al. leg. (CASC); 6 w., 4 s., 1 m., Anosy Region, Anosyenne Mts, 29.33 km NW Manantenina, - 24.13993 47.07418, 540 m, Fisher et al. leg. (CASC); 3 w., 3 s., Forêt Ivohibe 55.0 km N Tolagnaro, - 24.569 47.204, 200 m, Fisher et al. leg. (CASC); 4 w., 2 s., 1 q., Forêt Ivohibe 55.6 km N Tolagnaro, - 24.56167 47.20017, 650 m, Fisher et al. leg. (CASC); 2 w., 3 s., Grand Lavasoa, 25.9 km W Tolagnaro, - 25.08767 46.749, 450 m, Fisher et al. leg. (CASC); 4 w., 3 s., Parc National Andohahela, Col de Tanatana, 33.3 km NW Tolagnaro, - 24.7585 46.85367, 275 m, Fisher et al. leg. (CASC); 1 w., 1 s., Parc National Andohahela, Manangotry, 33.8 km NW Tolagnaro, - 24.75117 46.85783, 575 m, Fisher et al. leg. (CASC); 13 w., 7 s., Parc National d’Andohahela, Col du Sedro, 3.8 km 113 ° ESE Mahamavo, 37.6 km 341 ° NNW Tolagnaro, - 24.76389 46.75167, 900 m, Fisher et al. leg. (CASC); 7 w., 3 s., Parc National d’Andohahela, Manampanihy River, 5.4 km 113 ° ESE Mahamavo, 36.7 km 343 ° NNW Tolagnaro, - 24.76389 46.76683, 650 m, Fisher et al. leg. (CASC); 1 w., 1 s., Réserve Spéciale d’Ambohijanahary, Forêt d’Ankazotsihitafototra, 34.6 km 314 ° NW Ambaravaranala, - 18.26 45.41833, 1100 m, Fisher et al. leg. (CASC); 19 w., 11 s., Réserve Spéciale d’Ambohijanahary, Forêt d’Ankazotsihitafototra, 35.2 km 312 ° NW Ambaravaranala, - 18.26667 45.40667, 1050 m, Fisher et al. leg. (CASC). Geographic range. Madagascar, widely distributed mostly across the central and eastern parts of the island.	en	Salata, Sebastian, Fisher, Brian L. (2020): The ant genus Pheidole Westwood, 1839 (Hymenoptera: Formicidae) in Madagascar-taxonomic revision of the bessonii species-group. Zootaxa 4843 (1): 1-64, DOI: 10.11646/zootaxa.4843.1.1
03938025FF98A90DA7D8B11EFC50FC34.taxon	diagnosis	Diagnosis. Major workers. Head in full-face view sub-rectangular, slightly widening posteriorly, with lateral margins relatively straight; side of head with dense, long, suberect to erect pilosity; medial frons with sparse and thick costulae; interspaces between costulae indistinctly punctate; frons laterally with thick and sparse rugae; interspaces between rugae distinctly punctate; occipital lobes punctate with sparse and weakly developed rugae; scape, when laid back, exceeding the midlength of head by one-fifth of its length; inner hypostomal tooth distinct, bulge-like; outer hypostomal tooth lobe-like, moderately high and wide, with top arching posterolaterally; median tooth absent; promesonotum short, angular, and moderately low; posterior mesonotum smoothly declining towards propodeum; mesonotal process indistinct and tubercle-like; mesosoma microreticulate; lateral sides of pronotum, propodeum, and katepisternum with additional, thin and transverse rugae; gaster smooth; body yellowish orange to brown. Minor workers. Occiput not elongated; nuchal collar indistinct; head sculpture distinctly to indistinctly punctate and fading posteriorly to the vertex, gena mostly or entirely smooth; scape, when laid back, exceeding the posterior head margin by two-fifths of its length; promesonotum moderately low, moderately long, and arched; promesonotal groove present; metanotal groove distinct; propodeal spines thin and very small to small; mesosoma punctate; body yellow to brown.	en	Salata, Sebastian, Fisher, Brian L. (2020): The ant genus Pheidole Westwood, 1839 (Hymenoptera: Formicidae) in Madagascar-taxonomic revision of the bessonii species-group. Zootaxa 4843 (1): 1-64, DOI: 10.11646/zootaxa.4843.1.1
03938025FF98A90DA7D8B11EFC50FC34.taxon	description	Description. Major workers. Measurements (n = 10): HL: 1.54 – 1.7 (1.6); HW: 1.49 – 1.61 (1.53); SL: 1.04 – 1.09 (1.06); EL: 0.15 – 0.18 (0.17); WL: 1.49 – 1.6 (1.54); PSL: 0.22 – 0.27 (0.24); MTL: 1.12 – 1.21 (1.17); PNW: 0.59 – 0.64 (0.62); PTW: 0.17 – 0.21 (0.18); PPW: 0.35 – 0.43 (0.39); CI: 101.9 – 107.8 (104.3); SI: 64.8 – 71.4 (68.9); PSLI: 12.8 – 16.6 (15.1); PPI: 39.2 – 42.0 (40.2); PNI: 38.5 – 42.0 (40.2); MTI: 69.4 – 80.5 (76.5). Head. In full-face view sub-rectangular, slightly widening posteriorly, with lateral sides slightly convex (Fig. 13 B). In lateral view sub-oval, not depressed posteriorly; ventral and dorsal margins convex; inner hypostomal tooth not visible. Side of head with dense, long, suberect to erect pilosity; head dorsum with relatively dense, long, suberect to erect pilosity. Medial frons with sparse and thick costulae; interspaces between costulae indistinctly punctate. Frons laterally with thick and sparse rugae; interspaces between rugae distinctly punctate. Occipital lobes punctate with sparse and indistinct rugae. Gena with dense and moderately thick costulae; interspaces between costulae mostly punctate. Sides posterolateral from eyes microreticulate with additional network of thin and sparse rugulae. Center of clypeus shiny with weakly developed rugulae, lateral sides with distinct rugulae; median notch present, moderately wide and shallow; median longitudinal carina present; lateral longitudinal carinae present. Scape, when laid back, exceeding the midlength of head by one-fifth of its length; pilosity subdecumbent to erect (Fig. 13 B, D). Inner hypostomal tooth distinct, bulge-like; outer hypostomal tooth lobe-like, moderately high and wide ,, with top arching posterolaterally; median tooth absent (Fig. 27 E). Mesosoma. In lateral view, promesonotum short, angular, and moderately low; posterior mesonotum smoothly declining towards propodeum; mesonotal process indistinct and tubercle-like; promesonotal groove absent; metanotal groove indistinct; propodeal spines moderately long with moderately wide base and acute top; humeral tubercle weakly produced (Fig. 13 D). Surface microreticulate; lateral sides of pronotum, propodeum, and katepisternum with additional thin and transverse rugae. Pilosity dense, long, and erect (Fig. 13 D, F). Gaster. Shiny and smooth; pilosity moderately sparse, long, and erect (Fig. 13 D, F). Color. Yellowish orange to brown, antennae and legs yellow (Fig. 13 D, F). Description. Minor workers. Measurements (n = 9): HL: 0.72 – 0.91 (0.85); HW: 0.54 – 0.66 (0.62); SL: 0.99 – 1.13 (1.09); EL: 0.11 – 0.15 (0.13); WL: 0.99 – 1.19 (1.12); PSL: 0.08 – 0.14 (0.12); MTL: 0.93 – 1.04 (0.99); PNW: 0.34 – 0.47 (0.42); PTW: 0.09 – 0.13 (0.11); PPW: 0.16 – 0.23 (0.19); CI: 131.0 – 142.5 (138.0); SI: 171.1 – 183.7 (176.6); PSLI: 10.3 – 15.6 (13.8); PPI: 51.1 – 60.9 (56.7); PNI: 63.1 – 72.7 (68.8); MTI: 152.0 – 173.5 (160.4). Head. In full-face view oval, posterior region not elongated; nuchal collar weakly developed (Fig. 13 A). Pilosity relatively sparse, moderately long, subdecumbent to erect. Sculpture distinctly to indistinctly punctate, fading posteriorly; vertex and gena mostly or entirely smooth. 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. 13 A, C). Mesosoma. In lateral view, promesonotum moderately low and moderately long and arched; promesonotal groove present; metanotal groove distinct; propodeal spines very small to small, thin, and triangular (Fig. 13 C). Sculpture punctate. Pilosity sparse, long, and erect (Fig. 13 C, E). Gaster. With sparse and erect pilosity (Fig. 13 C, E). Color. Yellow to brown, legs and antennae usually brighter than the rest of the body (Fig. 13 C, E).	en	Salata, Sebastian, Fisher, Brian L. (2020): The ant genus Pheidole Westwood, 1839 (Hymenoptera: Formicidae) in Madagascar-taxonomic revision of the bessonii species-group. Zootaxa 4843 (1): 1-64, DOI: 10.11646/zootaxa.4843.1.1
03938025FF98A90DA7D8B11EFC50FC34.taxon	biology_ecology	Biology. The species was collected between 10 – 1625 m elevation, in montane rainforest, rainforest, montane ravine forest, montane shrubland, disturbed gallery montane forest, tropical forest, and montane forest. Nest were located in rotten logs, branches, and sticks on the ground, and dead twigs above the ground. Workers were collected from sifted litter.	en	Salata, Sebastian, Fisher, Brian L. (2020): The ant genus Pheidole Westwood, 1839 (Hymenoptera: Formicidae) in Madagascar-taxonomic revision of the bessonii species-group. Zootaxa 4843 (1): 1-64, DOI: 10.11646/zootaxa.4843.1.1
03938025FF98A90DA7D8B11EFC50FC34.taxon	discussion	Comments. Pheidole decollata is most similar to sympatric P. oswaldi. Its major workers can be separated based on the short, angular, and moderately low promesonotum and moderately large inner hypostomal tooth. In contrast, majors of P. oswaldi have a moderately long, angular, and very low promesonotum and small inner hypostomal tooth. Minors of P. decollata have the posterior region of the head not elongated and the promesonotum is moderately long, moderately low, and angular. Minor workers of P. oswaldi have slightly elongated posterior region of head, and moderately long, angular, and very low promesonotum.	en	Salata, Sebastian, Fisher, Brian L. (2020): The ant genus Pheidole Westwood, 1839 (Hymenoptera: Formicidae) in Madagascar-taxonomic revision of the bessonii species-group. Zootaxa 4843 (1): 1-64, DOI: 10.11646/zootaxa.4843.1.1
03938025FF95A90BA7D8B212FAFBFA60.taxon	description	Figs 14 A – F, 27 F, 28 F	en	Salata, Sebastian, Fisher, Brian L. (2020): The ant genus Pheidole Westwood, 1839 (Hymenoptera: Formicidae) in Madagascar-taxonomic revision of the bessonii species-group. Zootaxa 4843 (1): 1-64, DOI: 10.11646/zootaxa.4843.1.1
03938025FF95A90BA7D8B212FAFBFA60.taxon	materials_examined	HOLOTYPE: 1 s., Madagascar, Antsiranana, Galoko chain, Mont Galoko, - 13.58487 48.71818, 520 m, 17 - Feb- 2013, rainforest, ex rotten log, B. L. Fisher et al. leg. BLF 30615, CASENT 0923251 (CASC). PARATYPES: 1 w., 1 q., the same data as holotype, CASENT 0298332, CASENT 0298333 (CASC, MHNG). Geographic range. Madagascar, Antsiranana, Mont Galoko.	en	Salata, Sebastian, Fisher, Brian L. (2020): The ant genus Pheidole Westwood, 1839 (Hymenoptera: Formicidae) in Madagascar-taxonomic revision of the bessonii species-group. Zootaxa 4843 (1): 1-64, DOI: 10.11646/zootaxa.4843.1.1
03938025FF95A90BA7D8B212FAFBFA60.taxon	diagnosis	Diagnosis. Major workers. Head in full-face view sub-rectangular, not widening posteriorly, with lateral margins slightly convex; side of head with moderately dense, long, suberect to erect pilosity; medial frons with sparse costulae; interspaces between costulae smooth; frons laterally with thick and dense rugae; interspaces between rugae smooth to indistinctly rugulate; occipital lobes with thinner rugae; interspaces between rugae smooth; scape, when laid back, exceeding the midlength of head by two-fifths of its length; inner hypostomal tooth small, bulge-like, and arching posteroventrally; outer hypostomal tooth lobe-like, high, and wide, with top arching posterolaterally; median tooth absent; promesonotum short, angular, and moderately low; posterior mesonotum moderately steep; mesonotal process distinct and tubercle-like; mesosoma microreticulate; promesonotal dorsum with reduced microreticulae and additional transverse rugae; lateral pronotum with smooth notch; katepisternum and lateral sides of propodeum with additional rugae; gaster indistinctly shagreened; body brown. Minor workers. Occiput elongated forming short and moderately narrow neck; nuchal collar distinct; head sculpture smooth; scape, when laid back, exceeding the posterior head margin by three-fifths of its length; promesonotum low, long, and arched; promesonotal groove present; metanotal groove distinct; propodeal spines moderately large and thin; pronotum, mesonotum, and dorsal propodeum smooth; anepisternum, katepisternum, and lateral sides of propodeum with sparse puncta and rugae; body brown.	en	Salata, Sebastian, Fisher, Brian L. (2020): The ant genus Pheidole Westwood, 1839 (Hymenoptera: Formicidae) in Madagascar-taxonomic revision of the bessonii species-group. Zootaxa 4843 (1): 1-64, DOI: 10.11646/zootaxa.4843.1.1
03938025FF95A90BA7D8B212FAFBFA60.taxon	description	Description. Major workers. Measurements (n = 1): HL: 1.65; HW: 1.5; SL: 1.19; EL: 0.18; WL: 1.45; PSL: 0.3; MTL: 1.21; PNW: 0.56; PTW: 0.18; PPW: 0.46; CI: 110.7; SI: 79.7; PSLI: 17.9; PPI: 38.8; PNI: 37.6; MTI: 81.1. Head. In full-face view sub-rectangular, not widening posteriorly with lateral sides slightly convex (Fig. 14 B). 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, decumbent to erect pilosity. Medial frons with sparse costulae; interspaces between costulae smooth. Frons laterally with thick and dense rugae; interspaces between rugae smooth to indistinctly rugulate. Occipital lobes with thinner rugae; interspaces between rugae smooth. Gena with sparse and moderately thick costulae; interspaces between costulae smooth. Sides posterolateral from eyes with dense but very thin network of rugoreticulae; sculpture weakening posteriorly. 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, exceeding the midlength of head by two-fifths of its length; pilosity subdecumbent to erect (Fig. 14 B, D). Inner hypostomal tooth small, bulge-like, and arching posteroventrally; outer hypostomal tooth lobe-like, high, and wide, with top arching posterolaterally; median tooth absent (Fig. 27 F). Mesosoma. In lateral view, promesonotum short, angular, and moderately low; posterior mesonotum moderately steep; mesonotal process distinct and tuberclelike; promesonotal groove absent; metanotal groove absent; propodeal spines long with narrow base and acute top; humeral tubercle weakly produced (Fig. 14 D). Surface shiny and microreticulate; promesonotal dorsum with reduced microreticulae and additional transverse rugae; lateral pronotum with smooth notch; katepisternum and lateral sides of propodeum with additional rugae. Pilosity dense, long, and erect (Fig. 14 D, F). Gaster. Shiny and indistinctly shagreened; pilosity moderately sparse, long, and erect (Fig. D, F). Color. Brown, antenna, mandibles and legs yellowish (Fig. 14 D, F). Description. Minor workers. Measurements (n = 1): HL: 0.79; HW: 0.49; SL: 1.17; EL: 0.13; WL: 1.03; PSL: 0.14; MTL: 1.03; PNW: 0.36; PTW: 0.08; PPW: 0.13; CI: 161.7; SI: 238.3; PSLI: 17.1; PPI: 60.3; PNI: 73.7; MTI: 208.8. Head. In full-face view oval, posterior region elongated forming short and moderately narrow neck; nuchal collar distinct (Fig. 14 A). Pilosity relatively sparse, long, and subdecumbent to erect. Sculpture shiny and smooth; antennal sockets with few indistinct rugulae that are curved posterolaterally; interspaces between rugulae smooth. Clypeus with median longitudinal carina absent; two lateral longitudinal carinae absent. Scape, when laid back, exceeding the posterior head margin by three-fifths of its length; pilosity dense, subdecumbent to erect (Fig. 14 A, C). Mesosoma. In lateral view, promesonotum low, long, and arched; promesonotal groove present; metanotal groove distinct; propodeal spines moderately large, thin, and triangular (Fig. 14 C). Pronotum, mesonotum, and dorsal propodeum smooth; anepisternum, katepisternum, and lateral sides of propodeum with sparse puncta and rugae. Pilosity moderately sparse, long, and erect (Fig. 14 C, E). Gaster. With sparse and erect pilosity (Fig. 14 C, E). Color. Brown, legs, antennae and mandibles yellowish (Fig. 14 C, E).	en	Salata, Sebastian, Fisher, Brian L. (2020): The ant genus Pheidole Westwood, 1839 (Hymenoptera: Formicidae) in Madagascar-taxonomic revision of the bessonii species-group. Zootaxa 4843 (1): 1-64, DOI: 10.11646/zootaxa.4843.1.1
03938025FF95A90BA7D8B212FAFBFA60.taxon	biology_ecology	Biology. The species was collected at 520 m elevation, in rainforest. A nest was located in a rotten log.	en	Salata, Sebastian, Fisher, Brian L. (2020): The ant genus Pheidole Westwood, 1839 (Hymenoptera: Formicidae) in Madagascar-taxonomic revision of the bessonii species-group. Zootaxa 4843 (1): 1-64, DOI: 10.11646/zootaxa.4843.1.1
03938025FF95A90BA7D8B212FAFBFA60.taxon	discussion	Comments. Pheidole flammea is known only from Galoko Mountain. Its majors are most similar to P. ovalinoda and differ in moderately dense, long and suberect to erect pilosity on the side of the head, large outer hypostomal tooth, and mostly smooth interspaces between rugae on head. Majors of P. ovalinoda have dense, short and decumbent to suberect setae on sides of head, strongly reduced outer hypostomal tooth, and more distinctly sculptured interspaces between rugae on head. Minors of P. flammea are most similar to parapatric P. antsahabe and differ in brown body coloration, smooth promesonotum, and larger propodeal spines. In contrast, minors of P. antsahabe have black body coloration, sculptured promesonotum, and shorter propodeal spines.	en	Salata, Sebastian, Fisher, Brian L. (2020): The ant genus Pheidole Westwood, 1839 (Hymenoptera: Formicidae) in Madagascar-taxonomic revision of the bessonii species-group. Zootaxa 4843 (1): 1-64, DOI: 10.11646/zootaxa.4843.1.1
03938025FF95A90BA7D8B212FAFBFA60.taxon	etymology	Etymology. Latin for flames, in reference to the shape and density of setae on head of minor workers.	en	Salata, Sebastian, Fisher, Brian L. (2020): The ant genus Pheidole Westwood, 1839 (Hymenoptera: Formicidae) in Madagascar-taxonomic revision of the bessonii species-group. Zootaxa 4843 (1): 1-64, DOI: 10.11646/zootaxa.4843.1.1
03938025FF93A909A7D8B3E6FE14FE50.taxon	description	Figs 15 A – F, 27 G, 28 G	en	Salata, Sebastian, Fisher, Brian L. (2020): The ant genus Pheidole Westwood, 1839 (Hymenoptera: Formicidae) in Madagascar-taxonomic revision of the bessonii species-group. Zootaxa 4843 (1): 1-64, DOI: 10.11646/zootaxa.4843.1.1
03938025FF93A909A7D8B3E6FE14FE50.taxon	materials_examined	HOLOTYPE: 1 s., Madagascar, Antsiranana, Makirovana forest, - 14.104 50.03574, 225 m, 4 - May- 2011, rainforest, sifted litter (leaf mold, rotten wood), B. L. Fisher et al. leg. BLF 27044, CASENT 0243625 (CASC). PARATYPES: 1 w., the same data as holotype, CASENT 0230800 (CASC). Other material. Madagascar. Antsiranana: 4 w., 2 s., Makirovana forest, - 14.104 50.03574, 225 m, B. L. Fisher et al. leg. (CASC); 1 w., R. S. Manongarivo, 14.5 km 220 ° SW Antanambao, - 13.99833 48.42833, 1175 m, B. L. Fisher et al. leg. (CASC). Geographic range. Madagascar, Antsiranana, Makirovana forest and R. S. Manongarivo.	en	Salata, Sebastian, Fisher, Brian L. (2020): The ant genus Pheidole Westwood, 1839 (Hymenoptera: Formicidae) in Madagascar-taxonomic revision of the bessonii species-group. Zootaxa 4843 (1): 1-64, DOI: 10.11646/zootaxa.4843.1.1
03938025FF93A909A7D8B3E6FE14FE50.taxon	diagnosis	Diagnosis. Major workers. Head in full-face view sub-rectangular, elongated, not widening posteriorly, lateral margins relatively straight with very deep posteromedian concavity; side of head with moderately dense, long, subdecumbent to suberect pilosity; medial frons with dense and thick costulae; interspaces between costulae distinctly rugulate; frons laterally with thick and dense rugae; interspaces between rugae distinctly rugulate; occipital lobes with thick and dense rugae; interspaces between rugae distinctly rugulate; sculpture not weakening posteriorly; scape, when laid back, exceeding the midlength of head by two-fifths of its length; inner hypostomal tooth indistinct and bulge-like; outer hypostomal tooth lobe-like, high, and wide, with top arching posterolaterally; median tooth absent; promesonotum short, angular, and moderately low; posterior mesonotum smoothly declining towards propodeum; mesonotal process indistinct and tubercle-like; mesosoma punctate; promesonotum with additional thin, dense, and transverse rugae; katepisternum and lateral sides of propodeum with additional thick and dense rugulae; gaster shagreened; body orange. Minor workers. Occiput elongated forming moderately long and narrow neck; nuchal collar distinct; head sculpture smooth; scape, when laid back, exceeding the posterior head margin by three-fifths of its length; promesonotum low, very long, and arched; promesonotal groove present; metanotal groove distinct; propodeal spines long and thin; mesosoma smooth; body yellow.	en	Salata, Sebastian, Fisher, Brian L. (2020): The ant genus Pheidole Westwood, 1839 (Hymenoptera: Formicidae) in Madagascar-taxonomic revision of the bessonii species-group. Zootaxa 4843 (1): 1-64, DOI: 10.11646/zootaxa.4843.1.1
03938025FF93A909A7D8B3E6FE14FE50.taxon	description	Description. Major workers. Measurements (n = 3): HL: 1.86 – 1.89 (1.88); HW: 1.56 – 1.66 (1.6); SL: 1.33 – 1.41 (1.37); EL: 0.2 – 0.23 (0.22); WL: 1.68 – 1.76 (1.71); PSL: 0.4 – 0.42 (0.41); MTL: 1.53 – 1.54 (1.54); PNW: 0.64 – 0.66 (0.65); PTW: 0.18 – 0.2 (0.19); PPW: 0.53 – 0.58 (0.56); CI: 113.7 – 119.3 (117.3); SI: 83.6 – 90.2 (85.8); PSLI: 21.0 – 22.3 (21.9); PPI: 31.2 – 35.8 (33.8); PNI: 39.6 – 42.3 (40.7); MTI: 92.8 – 98.9 (95.9). Head. In full-face view sub-rectangular, elongated, not widening posteriorly, lateral sides relatively straight with very deep posteromedian concavity (Fig. 15 B). In lateral view sub-oval, not depressed posteriorly; ventral and dorsal margins convex; inner hypostomal tooth not visible. Side of head with moderately dense, long, subdecumbent to suberect pilosity; head dorsum with relatively dense, long, suberect to erect pilosity. Medial frons with dense and thick costulae; interspaces between costulae distinctly rugulate. Frons laterally with thick and dense rugae; interspaces between rugae distinctly rugulate. Occipital lobes with thick and dense rugae; interspaces between rugae distinctly rugulate; sculpture not weakening posteriorly. Gena with dense and moderately thick costulae; interspaces between costulae mostly smooth. Sides posterolateral from eyes with dense but very thin network of rugoreticulae; sculpture not weakening posteriorly. Center of clypeus shiny with few rugae, lateral sides with distinct rugulae; median notch present, moderately wide and shallow; median longitudinal carina present; lateral longitudinal carinae present. Scape, when laid back, exceeding the midlength of head by two-fifths of its length; pilosity subdecumbent to erect (Fig. 15 B, D). Inner hypostomal tooth indistinct, bulge-like; outer hypostomal tooth lobe-like, high, and wide, with top arching posterolaterally; median tooth absent (Fig. 27 G). Mesosoma. In lateral view, promesonotum short, angular, and moderately low; posterior mesonotum smoothly declining towards propodeum; mesonotal process indistinct and tubercle-like; promesonotal groove absent; metanotal groove indistinct; propodeal spines long, with narrow base and acute top; humeral tubercle weakly produced (Fig. 15 D). Surface punctate; promesonotum with additional thin, dense, and transverse rugae; katepisternum and lateral sides of propodeum with additional thick and dense rugulae. Pilosity dense, long, and erect (Fig. 15 D, F). Petiole. In rear view node dorsoventrally slightly convex (Fig. 15 D, F). Gaster. Dull and shagreened; pilosity moderately sparse, long, and erect (Fig. 15 D, F). Color. Orange, antennae and legs yellow, gaster and mandibles brownish-orange (Fig. 15 D, F). Description. Minor workers. Measurements (n = 5): HL: 0.84 – 0.91 (0.87); HW: 0.48 – 0.51 (0.49); SL: 1.26 – 1.36 (1.31); EL: 0.14 – 0.16 (0.15); WL: 1.12 – 1.18 (1.14); PSL: 0.2 – 0.24 (0.22); MTL: 1.14 – 1.25 (1.19); PNW: 0.37 – 0.38 (0.38); PTW: 0.07 – 0.09 (0.08); PPW: 0.13 – 0.16 (0.15); CI: 173.5 – 185.3 (177.1); SI: 257.9 – 279.6 (266.9); PSLI: 22.8 – 27.6 (25.7); PPI: 49.0 – 62.0 (56.0); PNI: 74.4 – 79.6 (76.9); MTI: 236.1 – 256.3 (242.2). Head. In full-face view oval, posterior region elongated forming moderately long and narrow neck; nuchal collar distinct (Fig. 15 A). Pilosity relatively sparse, moderately long, subdecumbent to erect. Sculpture shiny and smooth. Clypeus with median longitudinal carina absent; two lateral longitudinal carinae absent. Scape, when laid back, exceeding the posterior head margin by three-fifths of its length; pilosity dense, subdecumbent to erect (Fig. 15 A, C). Mesosoma. In lateral view, promesonotum low, very long, and arched; promesonotal groove present; metanotal groove distinct; propodeal spines long, thin, and triangular (Fig. 15 C). Sculpture smooth. Pilosity very sparse, long, and erect (Fig. 15 C, E). Gaster. With sparse and erect pilosity (Fig. 15 C, E). Color. Yellow (Fig. 15 C, E).	en	Salata, Sebastian, Fisher, Brian L. (2020): The ant genus Pheidole Westwood, 1839 (Hymenoptera: Formicidae) in Madagascar-taxonomic revision of the bessonii species-group. Zootaxa 4843 (1): 1-64, DOI: 10.11646/zootaxa.4843.1.1
03938025FF93A909A7D8B3E6FE14FE50.taxon	biology_ecology	Biology. The species was collected between 225 – 1175 m elevation, in rainforest and montane rainforest. Nesting preferences are unknown. Workers were collected from sifted litter.	en	Salata, Sebastian, Fisher, Brian L. (2020): The ant genus Pheidole Westwood, 1839 (Hymenoptera: Formicidae) in Madagascar-taxonomic revision of the bessonii species-group. Zootaxa 4843 (1): 1-64, DOI: 10.11646/zootaxa.4843.1.1
03938025FF93A909A7D8B3E6FE14FE50.taxon	discussion	Comments. Pheidole flavodepressa is known from two localities in Antsiranana and is most similar to parapatric P. zirafy. Majors of P. flavodepressa can be separated based on orange body coloration, side of head with moderately dense, long and subdecumbent to suberect pilosity, and occipital lobes with thick and dense rugae with distinctly rugulate interspaces between rugae. Majors of P. zirafy have blackish brown body, moderately sparse, short and appressed to decumbent pilosity on side of head, and occipital lobes with rugae that are thin, weakening posteriorly, and archposterolaterally, with mostly smooth interspaces between rugae. Minors of P. flavodepressa have yellow body coloration, shorter and thicker neck, and smooth mesosoma. In contrast, P. zirafy has dark brown minors with longer and thinner neck and mostly sculptured mesosoma. Minor workers of P. flavodepressa may also be confused with those of P. antsahabe, P. similis, P. mantadioflava, and P. clara, but differ from them in distinctly longer and thinner neck.	en	Salata, Sebastian, Fisher, Brian L. (2020): The ant genus Pheidole Westwood, 1839 (Hymenoptera: Formicidae) in Madagascar-taxonomic revision of the bessonii species-group. Zootaxa 4843 (1): 1-64, DOI: 10.11646/zootaxa.4843.1.1
03938025FF93A909A7D8B3E6FE14FE50.taxon	etymology	Etymology. Latin for yellow and deep, in reference to the yellow body coloration and deep posteromedian concavity in major workers.	en	Salata, Sebastian, Fisher, Brian L. (2020): The ant genus Pheidole Westwood, 1839 (Hymenoptera: Formicidae) in Madagascar-taxonomic revision of the bessonii species-group. Zootaxa 4843 (1): 1-64, DOI: 10.11646/zootaxa.4843.1.1
03938025FFAEA935A7D8B69FFC69FE99.taxon	description	Figs 16 A – F, 27 H, 28 H	en	Salata, Sebastian, Fisher, Brian L. (2020): The ant genus Pheidole Westwood, 1839 (Hymenoptera: Formicidae) in Madagascar-taxonomic revision of the bessonii species-group. Zootaxa 4843 (1): 1-64, DOI: 10.11646/zootaxa.4843.1.1
03938025FFAEA935A7D8B69FFC69FE99.taxon	materials_examined	Other material. Madagascar. Antsiranana: 3 w., 6.5 km SSW Befingotra, Rés. Anjanaharibe-Sud, - 14.75 49.5, 875 m, B. L. Fisher et al. leg. (CASC); 1 w., Forêt de Binara, 7.5 km 230 ° SW Daraina, - 13.255 49.61667, 375 m, B. L. Fisher et al. leg. (CASC); 1 w., Makirovana forest, - 14.17066 49.95409, 415 m, B. L. Fisher et al. leg. (CASC); 4 w., 1 s., Masoala National Park, - 15.32331 50.30751, 60 m, B. L. Fisher et al. leg. (CASC); 1 w., 1 s., 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); 1 w., 3 s., Parc National de Marojejy, Manantenina River, 28.0 km 38 ° NE Andapa, 8.2 km 333 ° NNW Manantenina, - 14.43667 49.775, 450 m, B. L. Fisher et al. leg. (CASC); 1 w., RNI Marojejy, 8 km NW Manantenina, - 14.43333 49.78333, 450 m, E. L. Quinter leg. (CASC); 4 w., SAVA Region, District of Sambava, Marojejy National Park, 5 km W of Manantenina village, 1 st Camp site (Mantella), - 14.43817 49.774, 487 m, M. Rin’Ha leg. (CASC); 4 w., 1 s., Sava Region: Parc National de Marojejy, near Manantenina River, - 14.43677 49.77541, 475 m, B. L. Fisher et al. leg. (CASC). Fianarantsoa: 5 w., 1 s., 7.6 km 122 º Kianjavato, Forêt Classée Vatovavy, - 21.4 47.94, 175 m, B. L. Fisher et al. leg. (CASC); 2 s., Forêt d’Ambalagoavy Nord, Ikongo, Ambatombe, - 21.857068 47.37849, 625 m, Harin’Hala & Irwin leg. (CASC); 2 w., JIRAMA water works near river, Ranomafana National Park, Fianarantsoa Prov., - 21.2485 47.45217, 690 m, Irwin et al. leg. (CASC); 1 w., radio tower, Ranomafana National Park, Fianarantsoa Prov., - 21.25833 47.40717, 1130 m, Irwin et al. leg. (CASC); 1 s., Ranomafana, - 21.25 47.36667, A. Pauly leg. (CASC); 1 w., Réserve Speciale Manombo 24.5 km 228 ° Farafangana, - 23.01583 47.719, 30 m, B. L. Fisher et al. leg. (CASC). Toamasina: 1 w., 5.3 km SSE Ambanizana, Andranobe, - 15.67133 49.97395, 425 m, B. L. Fisher et al. leg. (CASC); 2 w., 6.3 km S Ambanizana, Andranobe, - 15.6813 49.958, 25 m, B. L. Fisher et al. leg. (CASC); 1 w., Bevolota 17.1 km N Andasibe, - 18.77071 48.43164, 995 m, B. L. Fisher et al. leg. (CASC); 2 w., F. C. Sandranantitra, - 18.04833 49.09167, 450 m, H. J. Ratsirarson leg. (CASC); 5 w., 4 s., Forêt d’Analava Mandrisy, 5.9 km 195 º Antanambe, - 16.48567 49.847, 10 m, B. L. Fisher et al. leg. (CASC); 7 w., 5 s., Ile Sainte Marie, Forêt Kalalao, 9.9 km 34 ° Ambodifotatra, - 16.9225 49.88733, 100 m, B. L. Fisher et al. leg. (CASC); 2 s., Manakambahiny, near Vavatenina Forest, - 17.46667 49.35, A. Pauly leg. (CASC); 3 w., Montagne d’Akirindro 7.6 km 341 ° NNW Ambinanitelo, - 15.28833 49.54833, 600 m, B. L. Fisher et al. leg. (CASC); 7 w., 4 s., 1 q., Montagne d’Anjanaharibe, 18.0 km 21 ° NNE Ambinanitelo, - 15.18833 49.615, 470 m, B. L. Fisher et al. leg. (CASC); 1 w., Montagne d’Anjanaharibe, 19.5 km 27 ° NNE Ambinanitelo, - 15.17833 49.635, 1100 m, B. L. Fisher et al. leg. (CASC); 1 s., Nosy Mangabe, 7.43 km S Maroantsetra, - 15.4973 49.76223, 3 m, B. L. Fisher et al. leg. (CASC); 1 w., 1 s., Parc National de Zahamena, Besaky River, - 17.75244 48.85321, 760 m, B. L. Fisher et al. leg. (CASC); 6 w., 3 s., Parc National Mananara-Nord, 7.1 km 261 ° Antanambe, - 16.455 49.7875, 225 m, B. L. Fisher et al. leg. (CASC); 5 w., Parcelle E 3 Tampolo, - 17.28104 49.43012, 10 m, B. L. Fisher et al. leg. (CASC); 2 w., Parcelle K 7 Tampolo, - 17.28333 49.41667, 10 m, B. L. Fisher et al. leg. (CASC); 6 w., 3 s., Parcelle K 9 Tampolo, - 17.175 49.268, 10 m, B. L. Fisher et al. leg. (CASC); 4 w., 1 s., Res. Ambodiriana, 4.8 km 306 ° Manompana, along Manompana river, - 16.67233 49.70117, 125 m, B. L. Fisher et al. leg. (CASC); 1 w., Reserve Betampona, Camp Rendrirendry 34.1 km 332 ° Toamasina, - 17.924 49.19967, 390 m, B. L. Fisher et al. leg. (CASC); 1 w., Réserve Spéciale Ambatovaky, Sandrangato river, - 16.7633 49.26692, 520 m, B. L. Fisher et al. leg. (CASC); 4 w., Réserve Spéciale Ambatovaky, Sandrangato river, - 16.81739 49.29402, 360 m, B. L. Fisher et al. leg. (CASC); 3 w., 6 s., 2 m., S. F. Tampolo, 10 km NNE Fenoarivo Atn., - 17.2825 49.43, 10 m, B. L. Fisher et al. leg. (CASC); 5 w., 5 s., Sahafina forest 11.4 km W Brickaville, - 18.81445 48.96205, 140 m, B. L. Fisher et al. leg. (CASC); 1 w., 1 s., Tampolo, - 17.28333 49.41667, 10 m, B. L. Fisher et al. leg. (CASC); 1 w., Torotorofotsy, - 18.77048 48.43043, 1005 m, B. L. Fisher et al. leg. (CASC). Toliara: 6 w., Anosy Region, Anosyenne Mts, 29.33 km NW Manantenina, - 24.13993 47.07418, 540 m, B. L. Fisher et al. leg. (CASC); 1 w., 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 rainforest on the eastern part of the island.	en	Salata, Sebastian, Fisher, Brian L. (2020): The ant genus Pheidole Westwood, 1839 (Hymenoptera: Formicidae) in Madagascar-taxonomic revision of the bessonii species-group. Zootaxa 4843 (1): 1-64, DOI: 10.11646/zootaxa.4843.1.1
03938025FFAEA935A7D8B69FFC69FE99.taxon	diagnosis	Diagnosis. Major workers. Head in full-face view sub-rectangular, not widening posteriorly, with lateral margins slightly convex; side of head with moderately sparse, short, appressed to subdecumbent pilosity; frons with thin and sparse costulae that sometimes arch posterolaterally on the posterior part; interspaces between costulae smooth to indistinctly punctate; occipital lobes sometimes partially smooth, with thin rugopuncta that weaken posteriorly and arch posterolaterally; scape, when laid back, exceeding the midlength of head by two-fifths of its length; inner hypostomal tooth indistinct, low and bulge-like; outer hypostomal tooth lobe-like, higher than inner tooth, top directed posteriorly; inner and outer hypostomal tooth closely spaced and not connected by concavity; promesonotum short, angular and low; posterior mesonotum smoothly declining towards propodeum; mesonotal process distinct and tubercle-like; mesosoma smooth, only posterolateral sides of pronotum, katepisternum, and propodeum sometimes with sparse and thick rugulae; gaster smooth; body reddish brown to brown. Minor workers. Occiput elongated into a long, narrow neck terminating in a distinct nuchal collar; head sculpture smooth; scape, when laid back, exceeding the posterior head margin by half of its length; promesonotum very low, long, and slightly arched; promesonotal groove present; metanotal groove distinct; propodeal spines long and thin; mesosoma smooth; body brown to dark brown.	en	Salata, Sebastian, Fisher, Brian L. (2020): The ant genus Pheidole Westwood, 1839 (Hymenoptera: Formicidae) in Madagascar-taxonomic revision of the bessonii species-group. Zootaxa 4843 (1): 1-64, DOI: 10.11646/zootaxa.4843.1.1
03938025FFAEA935A7D8B69FFC69FE99.taxon	description	Description. Major workers. Measurements (n = 10): HL: 2.04 – 2.36 (2.26); HW: 1.87 – 2.21 (2.09); SL: 1.55 – 1.68 (1.63); EL: 0.25 – 0.28 (0.26); WL: 1.84 – 2.19 (2.08); PSL: 0.39 – 0.48 (0.44); MTL: 1.8 – 2.03 (1.92); PNW: 0.69 – 0.83 (0.76); PTW: 0.21 – 0.24 (0.23); PPW: 0.49 – 0.58 (0.53); CI: 104.1 – 110.5 (107.8); SI: 73.8 – 83.6 (77.8); PSLI: 17.8 – 20.9 (19.6); PPI: 39.4 – 46.2 (42.8); PNI: 34.9 – 38.4 (36.1); MTI: 87.9 – 96.3 (91.5). Head. In full-face view sub-rectangular, not widening posteriorly with lateral sides slightly convex (Fig. 16 B). In lateral view sub-oval, not depressed posteriorly; ventral and dorsal margins convex; inner hypostomal tooth visible. Side of head with moderately sparse, short, appressed to subdecumbent pilosity; head dorsum with relatively dense, long, decumbent to erect pilosity. Frons with thin and sparse costulae that sometimes arch posterolaterally on the posterior part; interspaces between costulae smooth to indistinctly punctate. Occipital lobes sometimes partially smooth; with thin rugopuncta that weaken posteriorly arch posterolaterally. Gena with dense and thick costulae; interspaces between costulae distinctly punctate. Sides posterolateral from eyes with dense and thin microreticulae that weaken posteriorly. Center of clypeus shiny and indistinctly punctate, lateral sides with weakly developed rugulae; median notch present, moderately wide and shallow; median longitudinal carina present; lateral longitudinal carinae present. Scape, when laid back, exceeding the midlength of head by two-fifths of its length; pilosity subdecumbent to erect (Fig. 16 B, D). Inner hypostomal tooth indistinct, low and bulge-like; outer hypostomal tooth lobe-like, higher than inner tooth, top directed posteriorly; inner and outer hypostomal tooth closely spaced and not connected by concavity (Fig. 27 H). Mesosoma. In lateral view, promesonotum short, angular and low; posterior mesonotum smoothly declining towards propodeum; mesonotal process distinct and tubercle-like; promesonotal groove absent; metanotal groove absent; propodeal spines long, narrow and with acute top; humeral tubercle weakly produced (Fig. 16 D). Surface shiny and smooth; posterolateral sides of pronotum, katepisternum, and propodeum sometimes with sparse and thick rugulae. Pilosity moderately dense, short, and erect (Fig. 16 D, F). Petiole. Smooth; in rear view node dorsoventrally convex medially (Fig. 16 D, F). Postpetiole. Shagreened; dorsum with slightly sparser sculpture (Fig. 16 D, F). Gaster. Shiny and smooth; pilosity moderately sparse, moderately long, and erect (Fig. 16 D, F). Color. Reddish brown to brown, antennae and legs yellowish brown (Fig. 16 D, F). Description. Minor workers. Measurements (n = 10): HL: 0.99 – 1.22 (1.04); HW: 0.58 – 0.77 (0.62); SL: 1.51 – 1.9 (1.6); EL: 0.16 – 0.19 (0.17); WL: 1.27 – 1.58 (1.38); PSL: 0.22 – 0.29 (0.25); MTL: 1.4 – 1.77 (1.51); PNW: 0.42 – 0.55 (0.46); PTW: 0.08 – 0.11 (0.1); PPW: 0.14 – 0.2 (0.16); CI: 159.4 – 175.0 (167.8); SI: 246.5 – 269.5 (258.3); PSLI: 21.8 – 25.0 (23.8); PPI: 51.6 – 79.4 (63.5); PNI: 71.0 – 77.1 (74.4); MTI: 230.6 – 258.9 (243.3). Head. In full-face view oval, posterior region elongated into long, narrow neck terminated by distinct nuchal collar (Fig. 16 A). Pilosity relatively sparse, moderately long, subdecumbent to erect. Sculpture shiny and smooth; antennal sockets with few indistinct and curved posterolaterally rugulate; interspaces between rugulae smooth. Clypeus with median longitudinal carina absent; two lateral longitudinal carinae absent. Scape, when laid back, exceeding the posterior head margin, neck not included, by half of its length; pilosity dense, subdecumbent to erect (Fig. 16 A, C). Mesosoma. In lateral view, promesonotum very low, long, and slightly arched; promesonotal groove present; metanotal groove distinct; propodeal spines long, thin, and triangular (Fig. 16 C). Sculpture smooth. Pilosity very sparse, short, and erect (Fig. 16 C, E). Petiole. Peduncle with ventral face straight (Fig. 16 C, E). Gaster. With sparse and erect pilosity (Fig. 16 C, E). Color. Brown to dark brown, legs and antennae brighter (Fig. 16 C, E).	en	Salata, Sebastian, Fisher, Brian L. (2020): The ant genus Pheidole Westwood, 1839 (Hymenoptera: Formicidae) in Madagascar-taxonomic revision of the bessonii species-group. Zootaxa 4843 (1): 1-64, DOI: 10.11646/zootaxa.4843.1.1
03938025FFAEA935A7D8B69FFC69FE99.taxon	biology_ecology	Biology. The species was collected between 3 – 1130 m elevation, in rainforest, littoral forest, montane rainforest, low elevation rainforest, an open area near a stream, tropical dry forest, mixed tropical forest, and on a sandy beach. Nests were located in rotten logs, soil, root mats, and under stones. Workers were collected from sifted litter and tree trunks.	en	Salata, Sebastian, Fisher, Brian L. (2020): The ant genus Pheidole Westwood, 1839 (Hymenoptera: Formicidae) in Madagascar-taxonomic revision of the bessonii species-group. Zootaxa 4843 (1): 1-64, DOI: 10.11646/zootaxa.4843.1.1
03938025FFAEA935A7D8B69FFC69FE99.taxon	discussion	Comments. Majors of P. grallatrix are most similar to P. bessonii, P. atsirakambiaty, and P. voreios and can be easily separated based on smooth mesosoma in which the posterolateral sides of pronotum, katepisternum, and propodeum may be sparsely rugulate. In contrast, P. bessonii, P. atsirakambiaty, and P. voreios have entirely sculptured mesosoma. Minors of P. grallatrix are most similar to P. zirafy and can be separated based on smooth mesosoma. In contrast, P. zirafy has dense and thin rugoreticulae on lateral sides of propodeum, anepisternum, and katepisternum. Distribution of P. grallatrix overlaps only with P. zirafy and P. voreios.	en	Salata, Sebastian, Fisher, Brian L. (2020): The ant genus Pheidole Westwood, 1839 (Hymenoptera: Formicidae) in Madagascar-taxonomic revision of the bessonii species-group. Zootaxa 4843 (1): 1-64, DOI: 10.11646/zootaxa.4843.1.1
03938025FFADA933A7D8B7BEFDC7FC58.taxon	description	Figs 17 A – F, 27 I, 28 I	en	Salata, Sebastian, Fisher, Brian L. (2020): The ant genus Pheidole Westwood, 1839 (Hymenoptera: Formicidae) in Madagascar-taxonomic revision of the bessonii species-group. Zootaxa 4843 (1): 1-64, DOI: 10.11646/zootaxa.4843.1.1
03938025FFADA933A7D8B7BEFDC7FC58.taxon	materials_examined	Other material. Madagascar. Antsiranana: 2 w., 1 s., 9.2 km WSW Befingotra, Rés. Anjanaharibe-Sud, - 14.75 49.46667, 1180 m, B. L. Fisher et al. leg. (CASC); 1 w., 9.2 km WSW Befingotra, Rés. Anjanaharibe-Sud, - 14.75, 49.46667 1200 m, B. L. Fisher et al. leg. (CASC). Toamasina: 3 w., 1 s., 6.9 km NE Ambanizana, Ambohitsitondroina, - 15.58506 50.00952, 825 m, B. L. Fisher et al. leg. (CASC); 6 w., F. C. Didy, - 18.19833 48.57833, 960 m, H. J. Ratsirarson leg. (CASC); 1 s., Forêt Ambatovy, 14.3 km 57 ° Moramanga, - 18.85083 48.32, 1075 m, B. L. Fisher et al. leg. (CASC); 2 w., 1 s., Reserve Betampona, Camp Vohitsivalana, 37.1 km 338 ° Toamasina, - 17.88667 49.2025, 520 m, B. L. Fisher et al. leg. (CASC); 1 w., 2 s., Réserve Spéciale Ambatovaky, Sandrangato river, - 16.76912 49.26704, 475 m, B. L. Fisher et al. leg. (CASC); 3 w., Réserve Spéciale Ambatovaky, Sandrangato river, - 16.80561 49.29507, 480 m, B. L. Fisher et al. leg. (CASC). Geographic range. Madagascar, from Andapa in Antsiranana south to Moramanga in Toamasina.	en	Salata, Sebastian, Fisher, Brian L. (2020): The ant genus Pheidole Westwood, 1839 (Hymenoptera: Formicidae) in Madagascar-taxonomic revision of the bessonii species-group. Zootaxa 4843 (1): 1-64, DOI: 10.11646/zootaxa.4843.1.1
03938025FFADA933A7D8B7BEFDC7FC58.taxon	diagnosis	Diagnosis. Major workers. Head in full-face view sub-rectangular, slightly widening posteriorly, with lateral margins slightly convex; side of head with dense, long, subdecumbent to suberect pilosity; antennal scrobes very indistinct with sparse and very thin rugocostulae; interspaces between rugocostulae smooth; frons with sparse and thin costulae; interspaces between costulae smooth; occipital lobes smooth; scape, when laid back, not reaching the midlength of head; inner hypostomal tooth indistinct, tooth-like, closely spaced, and directed posteriorly; 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 smooth with very indistinct, and thin rugulae; gaster indistinctly shagreened; body yellow to orange. Minor workers. Occiput not elongated; nuchal collar absent; head sculpture smooth; scape, when laid back, exceeding the posterior head margin by one-fifth of its length; promesonotum moderately low, moderately short, and arched; promesonotal groove present; metanotal groove distinct; propodeal spines indistinct; mesosoma smooth; body yellow.	en	Salata, Sebastian, Fisher, Brian L. (2020): The ant genus Pheidole Westwood, 1839 (Hymenoptera: Formicidae) in Madagascar-taxonomic revision of the bessonii species-group. Zootaxa 4843 (1): 1-64, DOI: 10.11646/zootaxa.4843.1.1
03938025FFADA933A7D8B7BEFDC7FC58.taxon	description	Description. Major workers. Measurements (n = 6): HL: 1.87 – 2.38 (2.15); HW: 1.88 – 2.35 (2.07); SL: 0.94 – 0.99 (0.96); EL: 0.22 – 0.26 (0.23); WL: 1.45 – 1.67 (1.55); PSL: 0.2 – 0.3 (0.24); MTL: 1.07 – 1.21 (1.14); PNW: 0.69 – 0.8 (0.74); PTW: 0.18 – 0.28 (0.23); PPW: 0.55 – 0.73 (0.63); CI: 99.7 – 107.9 (104.0); SI: 39.9 – 52.4 (46.8); PSLI: 9.7 – 13.0 (11.2); PPI: 32.9 – 41.1 (37.4); PNI: 34.2 – 36.6 (35.7); MTI: 51.4 – 61.6 (55.3). Head. In full-face view sub-rectangular, slightly widening posteriorly, with lateral sides slightly convex (Fig. 17 B). In lateral view sub-oval, not depressed posteriorly; ventral and dorsal margins convex; inner hypostomal tooth visible. Side of head with dense, long, subdecumbent to suberect pilosity; head dorsum with relatively dense, long, suberect to erect pilosity. Antennal scrobes very indistinct; with sparse and very thin rugocostulae; interspaces between rugocostulae smooth. Frons with sparse and thin costulae; interspaces between costulae smooth. Occipital lobes smooth. Gena with moderately dense and thin costulae; interspaces between costulae smooth. Sides posterolateral from eyes smooth; anterior parts indistinctly shagreened. 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. 17 B, D). Inner hypostomal tooth indistinct, tooth-like, closely spaced, and directed posteriorly; outer hypostomal tooth lobe-like, high, and moderately wide; median tooth absent (Fig. 27 I). 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. 17 D). Surface mostly smooth, with very indistinct, and thin rugulae. Pilosity moderately dense, long, and erect (Fig. 17 D, F). Gaster. Shiny and indistinctly shagreened; pilosity moderately dense, long, and erect (Fig. 17 D, F). Color. Yellow to orange, gaster and mandibles darker (Fig. 17 D, F). Description. Minor workers. Measurements (n = 7): HL: 0.68 – 0.77 (0.73); HW: 0.6 – 0.66 (0.63); SL: 0.85 – 0.96 (0.9); EL: 0.12 – 0.14 (0.13); WL: 0.86 – 1.04 (0.96); PSL: 0.09 – 0.11 (0.1); MTL: 0.68 – 0.89 (0.78); PNW: 0.37 – 0.42 (0.4); PTW: 0.1 – 0.11 (0.1); PPW: 0.17 – 0.2 (0.18); CI: 113.3 – 126.3 (117.0); SI: 132.1 – 153.6 (143.4); PSLI: 12.9 – 15.3 (13.9); PPI: 53.3 – 63.3 (58.1); PNI: 61.3 – 65.1 (63.3); MTI: 110.5 – 144.6 (124.6). Head. In full-face view oval, posterior region not elongated; nuchal collar absent (Fig. 17 A). Pilosity relatively sparse, moderately long, subdecumbent to erect. Sculpture smooth; anterolateral frons with weakly developed puncta. 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. 17 A, C). Mesosoma. In lateral view, promesonotum moderately low, moderately short, and arched; promesonotal groove present; metanotal groove distinct; propodeal spines weakly developed and triangular (Fig. 17 C). Sculpture smooth. Pilosity very sparse, long, and erect (Fig. 17 C, E). Gaster. With sparse and erect pilosity (Fig. 17 C, E). Color. Yellow (Fig. 17 C, E).	en	Salata, Sebastian, Fisher, Brian L. (2020): The ant genus Pheidole Westwood, 1839 (Hymenoptera: Formicidae) in Madagascar-taxonomic revision of the bessonii species-group. Zootaxa 4843 (1): 1-64, DOI: 10.11646/zootaxa.4843.1.1
03938025FFADA933A7D8B7BEFDC7FC58.taxon	biology_ecology	Biology. The species was collected between 475 – 1200 m elevation, in rainforest and montane rainforest. Nests were located in rotten logs, dead branches above the ground, and root mats. Workers were collected from sifted litter and on low vegetation.	en	Salata, Sebastian, Fisher, Brian L. (2020): The ant genus Pheidole Westwood, 1839 (Hymenoptera: Formicidae) in Madagascar-taxonomic revision of the bessonii species-group. Zootaxa 4843 (1): 1-64, DOI: 10.11646/zootaxa.4843.1.1
03938025FFADA933A7D8B7BEFDC7FC58.taxon	discussion	Comments. Pheidole madecassa is most similar to sympatric P. maro. Major workers of P. madecassa can be separated by 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. Majors of P. maro have indistinctly punctate interspaces between rugocostulae on antennal scrobes, sculptured mesosoma, large inner hypostomal tooth, and reddish-brown body. Minor workers of P. madecassa can be distinguished by smooth mesosoma and mostly smooth frons with weakly punctuated anterolateral areas. In contrast, minors of P. maro have indistinctly sculptured frons and mesosoma.	en	Salata, Sebastian, Fisher, Brian L. (2020): The ant genus Pheidole Westwood, 1839 (Hymenoptera: Formicidae) in Madagascar-taxonomic revision of the bessonii species-group. Zootaxa 4843 (1): 1-64, DOI: 10.11646/zootaxa.4843.1.1
03938025FFABA931A7D8B5FEFED8FEE0.taxon	description	Figs 18 A – F, 27 J, 28 J	en	Salata, Sebastian, Fisher, Brian L. (2020): The ant genus Pheidole Westwood, 1839 (Hymenoptera: Formicidae) in Madagascar-taxonomic revision of the bessonii species-group. Zootaxa 4843 (1): 1-64, DOI: 10.11646/zootaxa.4843.1.1
03938025FFABA931A7D8B5FEFED8FEE0.taxon	materials_examined	HOLOTYPE: 1 s., Madagascar, Toamasina, Corridor Forestier Analamay-Mantadia, Ambatoharanana, - 18.80424 48.40081, 968 m, 12 - Dec- 2012, rainforest, ex dead bamboo above ground, B. L. Fisher et al. leg. BLF 30377, CASENT 0923250 (CASC). PARATYPES: 1 w., the same data as holotype, CASENT 0301826 (CASC). Other material. Madagascar. Toamasina: 1 w., Ambatovy, 12.4 km NE Moramanga, - 18.84963 48.2947, 1010 m, B. L. Fisher et al. leg. (CASC); 2 w., Ankerana, - 18.4017 48.80605, 1035 m, B. L. Fisher et al. leg. (CASC); 1 w., 1 s., Bevolota 17.1 km N Andasibe, - 18.77071 48.43164, 995 m, B. L. Fisher et al. leg. (CASC); 1 w., 1 s., Corridor Forestier Analamay-Mantadia, Ambatoharanana, - 18.80388 48.40506, 1013 m, B. L. Fisher et al. leg. (CASC); 1 w., 1 s., Corridor Forestier Analamay-Mantadia, Ambatoharanana, - 18.80424 48.40081, 968 m, B. L. Fisher et al. leg. (CASC); 1 w., 1 s., Corridor Forestier Analamay-Mantadia, Ambatoharanana, - 18.79956 48.4028, 1058 m, B. L. Fisher et al. leg. (CASC); 1 w., 1 s., Corridor Forestier Analamay-Mantadia, Ambatoharanana, - 18.80438 48.40735, 960 m, B. L. Fisher et al. leg. (CASC); 1 w., 1 s., Corridor Forestier Analamay-Mantadia, Ambohibolakely, - 18.77898 48.36375, 918 m, B. L. Fisher et al. leg. (CASC); 1 w., 1 s., Corridor Forestier Analamay-Mantadia, Tsaravoniana, - 18.76124 48.42134, 939 m, B. L. Fisher et al. leg. (CASC); 1 w., 1 s., Corridor Forestier Analamay-Mantadia, Tsaravoniana, - 18.75737 48.42302, 1018 m, B. L. Fisher et al. leg. (CASC); 1 w., 2 s., F. C. Didy, - 18.19833 48.57833, 960 m, H. J. Ratsirarson leg. (CASC); 1 w., 1 q., Réserve Spéciale Ambatovaky, Sandrangato river, - 16.76912 49.26704, 475 m, B. L. Fisher et al. leg. (CASC); 2 w., Réserve Spéciale Ambatovaky, Sandrangato river, - 16.7633 49.26692, 520 m, B. L. Fisher et al. leg. (CASC). Geographic range. Madagascar, distributed in mountains and highlands between Moramanga and Antenina in Toamasina.	en	Salata, Sebastian, Fisher, Brian L. (2020): The ant genus Pheidole Westwood, 1839 (Hymenoptera: Formicidae) in Madagascar-taxonomic revision of the bessonii species-group. Zootaxa 4843 (1): 1-64, DOI: 10.11646/zootaxa.4843.1.1
03938025FFABA931A7D8B5FEFED8FEE0.taxon	diagnosis	Diagnosis. Major workers. Head in full-face view sub-rectangular, not widening posteriorly, with lateral margins slightly convex; side of head with moderately dense, long, suberect to erect pilosity; antennal scrobes present, very indistinct, and mostly punctate, with additional network of thin and sparse rugae; frons with moderately dense and thick costulae; interspaces between costulae smooth to indistinctly punctate; occipital lobes with very indistinct and short rugae; sculpture weakening posteriorly and posterior part mostly smooth; scape, when laid back, exceeding the midlength of head by one-fifth of its length; inner hypostomal tooth distinct and bulge-like; outer hypostomal tooth lobe-like, high, and wide, with top directed posteriorly; median tooth absent; promesonotum short, angular, and moderately low, anterior mesonotum placed slightly higher than pronotum; posterior mesonotum moderately steep; mesonotal process distinct and tubercle-like; mesosoma microreticulate; promesonotal dorsum with sculpture reduced and additional transverse rugae; gaster indistinctly shagreened; body orange. Minor workers. Occiput elongated forming short and moderately narrow neck; nuchal collar distinct; head sculpture mostly smooth; frons with sparse puncta; scape, when laid back, exceeding the posterior head margin by three-fifths of its length; promesonotum low, long, and arched; promesonotal groove present; metanotal groove distinct; propodeal spines small with wide base; mesosoma punctate; body yellow.	en	Salata, Sebastian, Fisher, Brian L. (2020): The ant genus Pheidole Westwood, 1839 (Hymenoptera: Formicidae) in Madagascar-taxonomic revision of the bessonii species-group. Zootaxa 4843 (1): 1-64, DOI: 10.11646/zootaxa.4843.1.1
03938025FFABA931A7D8B5FEFED8FEE0.taxon	description	Description. Major workers. Measurements (n = 10): HL: 1.57 – 1.82 (1.69); HW: 1.47 – 1.8 (1.65); SL: 0.95 – 1.06 (1.0); EL: 0.21 – 0.25 (0.23); WL: 1.35 – 1.53 (1.47); PSL: 0.23 – 0.28 (0.25); MTL: 0.78 – 1.17 (1.06); PNW: 0.54 – 0.64 (0.6); PTW: 0.18 – 0.21 (0.2); PPW: 0.39 – 0.51 (0.44); CI: 98.5 – 107.0 (102.6); SI: 57.5 – 64.8 (60.7); PSLI: 13.5 – 16.9 (14.8); PPI: 40.1 – 51.3 (45.5); PNI: 32.5 – 37.7 (36.0); MTI: 52.9 – 71.3 (64.2). Head. In full-face view sub-rectangular, not widening posteriorly, with lateral sides slightly convex (Fig. 18 B). 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, decumbent to erect pilosity. Antennal scrobes present and very indistinct; mostly punctate with additional network of thin and sparse rugae. Frons with moderately dense and thick costulae; interspaces between costulae smooth to indistinctly punctate. Occipital lobes with very indistinct and short rugae; sculpture weakening posteriorly and posterior part mostly smooth. Gena with sparse and moderately thick costulae; interspaces between costulae smooth. Sides posterolateral from eyes with dense but very thin network of rugoreticulae; sometimes with smooth notches; sculpture weakening posteriorly. 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, exceeding the midlength of head by one-fifth of its length; pilosity subdecumbent to erect (Fig. 18 B, D). Inner hypostomal tooth distinct and bulge-like; outer hypostomal tooth lobe-like, high, and wide, with top directed posteriorly; median tooth absent (Fig. 27 J). Mesosoma. In lateral view, promesonotum short, angular, and moderately low, anterior mesonotum placed slightly higher than pronotum; posterior mesonotum moderately steep; mesonotal process distinct and tubercle-like; promesonotal groove absent; metanotal groove absent; propodeal spines moderately long, with narrow base and acute top; humeral tubercle weakly produced (Fig. 18 D). Surface shiny and microreticulate; promesonotal dorsum with microreticulae reduced and additional transverse rugae. Pilosity moderately dense, long, and erect (Fig. 18 D, F). Gaster. Shiny and indistinctly shagreened; pilosity moderately sparse, long, and erect (Fig. 18 D, F). Color. Orange, legs and occipital lobes brighter, yellow (Fig. 18 D, F). Description. Minor workers. Measurements (n = 10): HL: 0.78 – 0.84 (0.81); HW: 0.53 – 0.58 (0.55); SL: 0.97 – 1.1 (1.04); EL: 0.14 – 0.17 (0.15); WL: 0.99 – 1.11 (1.06); PSL: 0.07 – 0.11 (0.1); MTL: 0.83 – 0.96 (0.9); PNW: 0.36 – 0.41 (0.38); PTW: 0.08 – 0.11 (0.1); PPW: 0.14 – 0.19 (0.17); CI: 145.6 – 154.1 (147.6); SI: 180.7 – 194.6 (188.9); PSLI: 9.5 – 14.1 (12.2); PPI: 51.3 – 70.1 (62.3); PNI: 66.9 – 71.3 (69.4); MTI: 154.7 – 169.7 (164.1). Head. In full-face view oval, posterior region elongated forming short and moderately narrow neck; nuchal collar distinct (Fig. 18 A). Pilosity relatively sparse, long, subdecumbent to erect. Sculpture shiny and mostly smooth; frons with sparse puncta. Clypeus with median longitudinal carina absent; two lateral longitudinal carinae absent. Scape, when laid back, exceeding the posterior head margin by three-fifths of its length; pilosity dense, subdecumbent to erect (Fig. 18 A, C). Mesosoma. In lateral view, promesonotum low, long, and arched; promesonotal groove present; metanotal groove distinct; propodeal spines small and triangular, with wide base (Fig. 18 C). Sculpture punctate. Pilosity very sparse, moderately long, and erect (Fig. 18 C, E). Gaster. With sparse and erect pilosity (Fig. 18 C, E). Color. Yellow, mandibles and gaster usually darker (Fig. 18 C, E).	en	Salata, Sebastian, Fisher, Brian L. (2020): The ant genus Pheidole Westwood, 1839 (Hymenoptera: Formicidae) in Madagascar-taxonomic revision of the bessonii species-group. Zootaxa 4843 (1): 1-64, DOI: 10.11646/zootaxa.4843.1.1
03938025FFABA931A7D8B5FEFED8FEE0.taxon	biology_ecology	Biology. The species was collected between 475 – 1035 m elevation, in rainforest, montane rainforest, and montane forest. Nests were located in rotten logs and sticks on the ground and dead bamboo above ground. Workers were collected from sifted litter.	en	Salata, Sebastian, Fisher, Brian L. (2020): The ant genus Pheidole Westwood, 1839 (Hymenoptera: Formicidae) in Madagascar-taxonomic revision of the bessonii species-group. Zootaxa 4843 (1): 1-64, DOI: 10.11646/zootaxa.4843.1.1
03938025FFABA931A7D8B5FEFED8FEE0.taxon	discussion	Comments. Pheidole mantadioflava, known from the northeastern part of Toamasina, is most similar to P. clara, recorded from southeastern Antsiranana and the northernmost part of Toamasina. Majors of P. mantadioflava differ in the presence of antennal scrobes and anterior mesonotum placed slightly higher than pronotum. In contrast, majors of P. clara lack antennal scrobes and their anterior mesonotum is not placed higher than the pronotum. Minors of P. mantadioflava can be separated based on presence of sparse puncta on the entire surface of frons and small propodeal spines with a wide base. Minor workers of P. clara have sparse puncta restricted only to anterolateral frons, and their propodeal spines are small and thin.	en	Salata, Sebastian, Fisher, Brian L. (2020): The ant genus Pheidole Westwood, 1839 (Hymenoptera: Formicidae) in Madagascar-taxonomic revision of the bessonii species-group. Zootaxa 4843 (1): 1-64, DOI: 10.11646/zootaxa.4843.1.1
03938025FFABA931A7D8B5FEFED8FEE0.taxon	etymology	Etymology. Combination of the name of the type locality and a Latin world for yellow, in reference to the body colouration.	en	Salata, Sebastian, Fisher, Brian L. (2020): The ant genus Pheidole Westwood, 1839 (Hymenoptera: Formicidae) in Madagascar-taxonomic revision of the bessonii species-group. Zootaxa 4843 (1): 1-64, DOI: 10.11646/zootaxa.4843.1.1
03938025FFA6A93DA7D8B69FFB3EFC11.taxon	description	Figs 19 A – F, 27 K, 28 K	en	Salata, Sebastian, Fisher, Brian L. (2020): The ant genus Pheidole Westwood, 1839 (Hymenoptera: Formicidae) in Madagascar-taxonomic revision of the bessonii species-group. Zootaxa 4843 (1): 1-64, DOI: 10.11646/zootaxa.4843.1.1
03938025FFA6A93DA7D8B69FFB3EFC11.taxon	materials_examined	HOLOTYPE: 1 s., 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. BLF 30097, CASENT 0923256 (CASC). PARATYPES: 1 w., the same data as holotype, CASENT 0299307 (CASC). Other material. Antananarivo: 1 w., Forêt de galerie, Telomirahavavy, 23.4 km NNE Ankazobe, - 18.12167 47.20627, 1520 m, B. L. Fisher et al. leg. (CASC). Antsiranana: 1 w., Ampasindava, Forêt d’Ambilanivy, 3.9 km 181 ° S Ambaliha, - 13.79861 48.16167, 600 m, B. L. Fisher et al. leg. (CASC); 1 w., Forêt Ambanitaza, 26.1 km 347 ° Antalaha, - 14.67933 50.18367, 240 m, B. L. Fisher et al. leg. (CASC); 1 w., 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); 1 w., Forêt de Binara, 9.1 km 233 ° SW Daraina, - 13.26333 49.60333, 800 m, B. L. Fisher et al. leg. (CASC); 1 w., Forêt de Binara, 9.1 km 233 ° SW Daraina, - 13.26333 49.60333, 650 - 800 m, B. L. Fisher et al. leg. (CASC); 1 w., Makirovana forest, - 14.16506 49.9477, 900 m, B. L. Fisher et al. leg. (CASC); 1 w., Montagne des Français, 7.2 km 142 ° SE Antsiranana (= Diego Suarez), - 12.32278 49.33817, 180 m, B. L. Fisher et al. leg. (CASC); 1 w., 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); 1 w., 1 s., R. S. Manongarivo, 14.5 km 220 ° SW Antanambao, - 13.99833 48.42833, 1175 m, B. L. Fisher et al. leg. (CASC); 2 w., Réserve Spéciale d’Ambre, 3.5 km 235 ° SW Sakaramy, - 12.46889 49.24217, 325 m, B. L. Fisher et al. leg. (CASC); 1 w., 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); 2 w., 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); 3 w., Sakalava Beach [vegetated beach dunes], - 12.26972 49.39167, 10 m, R. Harin’Hala leg. (CASC); 2 w., SAVA Region, District of Sambava, Marojejy National Park, 5 km W of Manantenina village, 1 st Camp site (Mantella), - 14.43817 49.774, 487 m, M. Rin’Ha leg. (CASC). Fianarantsoa: 2 w., 43 km S Ambalavao, Rés. Andringitra, - 22.23333 47, 825 m, B. L. Fisher et al. leg. (CASC); 4 w., 45 km S. Ambalavao, - 22.21667 47.01667, 785 m, B. L. Fisher et al. leg. (CASC); 2 w., 9.0 km NE Ivohibe, - 22.42667 46.93833, 900 m, B. L. Fisher et al. leg. (CASC); 21 w., Belle Vue trail, Ranomafana National Park, Fianarantsoa Prov., - 21.2665 47.42017, 1020 m, M. E. Irwin et. al. leg. (CASC); 2 w., Fitovavy Fitovinany Region, District of Ifanadiana, 12 km W of Ranomafana, - 21.25083 47.40717, 1127 m, M. Rin’Ha leg. (CASC); 2 w., 2 s., 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); 1 w., R. S. Ivohibe 8.0 km E Ivohibe, - 22.48333 46.96833, 1200 m, B. L. Fisher et al. leg. (CASC); 1 w., 1 s., R. S. Ivohibe, 7.5 km ENE Ivohibe, - 22.47 46.96, 900 m, B. L. Fisher et al. leg. (CASC); 18 w., radio tower, Ranomafana National Park, Fianarantsoa Prov., - 21.25833 47.40717, 1130 m, M. E. Irwin et. al. leg. (CASC); 10 w., Vohiparara broken bridge, Fianarantsoa Prov., - 21.22617 47.36983, 1110 m, R. Harin’Hala leg. MA- 02 - 09 A- 06 (CASC). Mahajanga: 1 w., Réserve Spéciale Marotandrano, Marotandrano 48.3 km S Mandritsara, - 16.28322 48.81443, 865 m, B. L. Fisher et al. leg. (CASC). Toamasina: 2 w., 7 km SE Andasibe National Park Headquarters, - 18.969856 48.465894, 1050 m, M. E. Irwin et. al. leg. (CASC); 1 s., Ambanizana, Parc National Masoala, - 15.57167 50.00611, 800 - 897 m, D. Andriamalala et. al. leg. (CASC); 1 w., 1 s., Ambanizana, Parc National Masoala, - 15.57167 50.00611, 900 - 950 m, D. Andriamalala et. al. leg. (CASC); 2 w., 2 s., Ambatovy, 12.4 km NE Moramanga, - 18.84963 48.2947, 1010 m, B. L. Fisher et al. leg. (CASC); 2 w., 2 s., Ambatovy, 12.4 km NE Moramanga, - 18.83937 48.30842, 1080 m, B. L. Fisher et al. leg. (CASC); 3 w., Ambatovy, 12.4 km NE Moramanga, - 18.84773 48.29568, 1000 m, B. L. Fisher et al. leg. (CASC); 1 w., Ambatovy, 12.4 km NE Moramanga, - 18.85813 48.28488, 1040 m, B. L. Fisher et al. leg. (CASC); 1 w., 1 s., Analamay, - 18.80623 48.33707, 1068 m, B. L. Fisher et al. leg. (CASC); 2 w., Andasibe National Park, botanic garden near entrance, West of ANGAP office, - 18.925172 48.418651, 1025 m, M. E. Irwin et. al. leg. (CASC); 2 w., Ankerana, - 18.40636 48.80254, 1108 m, B. L. Fisher et al. leg. (CASC); 1 w., Ankerana, - 18.4017 48.80605, 1035 m, B. L. Fisher et al. leg. (CASC); 1 s., Bevolota 17.1 km N Andasibe, - 18.77071 48.43164, 995 m, B. L. Fisher et al. leg. (CASC); 1 s., Corridor Forestier Analamay-Mantadia, Ambatoharanana, - 18.80398 48.40358, 1064 m, B. L. Fisher et al. leg. (CASC); 2 w., 1 s., Corridor Forestier Analamay-Mantadia, Ambatoharanana, - 18.79956 48.4028, 1058 m, B. L. Fisher et al. leg. (CASC); 2 w., 2 s., Corridor Forestier Analamay-Mantadia, Ambatoharanana, - 18.80438 48.40735, 960 m, B. L. Fisher et al. leg. (CASC); 2 w., Corridor Forestier Analamay-Mantadia, Ambohibolakely, - 18.77898 48.36375, 918 m, B. L. Fisher et al. leg. (CASC); 2 w., Corridor Forestier Analamay-Mantadia, Ambohibolakely, - 18.76087 48.37128, 1044 m, B. L. Fisher et al. leg. (CASC); 1 w., 1 s., Corridor Forestier Analamay-Mantadia, Tsaravoniana, - 18.76124 48.42134, 939 m, B. L. Fisher et al. leg. (CASC); 4 w., 4 s., Corridor Forestier Analamay-Mantadia, Tsaravoniana, - 18.76465 48.41938, 1039 m, B. L. Fisher et al. leg. (CASC); 1 w., Corridor Forestier Analamay-Mantadia, Tsaravoniana, - 18.75641 48.42195, 1036 m, B. L. Fisher et al. leg. (CASC); 1 w., Corridor Forestier Analamay-Mantadia, Tsaravoniana, - 18.75737 48.42302, 1018 m, B. L. Fisher et al. leg. (CASC); 3 w., F. C. Andriantantely, - 18.695 48.81333, 530 m, H. J. Ratsirarson leg. (CASC); 2 w., 1 s., F. C. Didy, - 18.19833 48.57833, 960 m, H. J. Ratsirarson leg. (CASC); 1 w., F. C. Sandranantitra, - 18.04833 49.09167, 450 m, H. J. Ratsirarson leg. (CASC); 2 w., Forêt Ambatovy, 14.3 km 57 ° Moramanga, - 18.85083 48.32, 1075 m, B. L. Fisher et al. leg. (CASC); 2 w., Montagne d’Akirindro 7.6 km 341 ° NNW Ambinanitelo, - 15.28833 49.54833, 600 m, B. L. Fisher et al. leg. (CASC); 2 w., 2 s., Montagne d’Anjanaharibe, 18.0 km 21 ° NNE Ambinanitelo, - 15.18833 49.615, 470 m, B. L. Fisher et al. leg. (CASC); 5 w., 2 s., Montagne d’Anjanaharibe, 19.5 km 27 ° NNE Ambinanitelo, - 15.17833 49.635, 1100 m, B. L. Fisher et al. leg. (CASC); 3 w., P. N. Mantadia, - 18.79167 48.42667, 895 m, H. J. Ratsirarson leg. (CASC); 1 w., Parc National de Zahamena, - 17.73359 48.72625, 950 m, B. L. Fisher et al. leg. (CASC); 1 w., 1 s., Parc National de Zahamena, Onibe River, - 17.75908 48.85468, 780 m, B. L. Fisher et al. leg. (CASC); 1 s., Parc National Mananara-Nord, 7.1 km 261 ° Antanambe, - 16.455 49.7875, 225 m, B. L. Fisher et al. leg. (CASC); 1 w., Parcelle K 9 Tampolo, - 17.175 49.268, 10 m, B. L. Fisher et al. leg. (CASC); 4 w., Réserve Spéciale Ambatovaky, Sandrangato river, - 16.77274 49.26551, 450 m, B. L. Fisher et al. leg. (CASC); 3 w., Réserve Spéciale Ambatovaky, Sandrangato river, - 16.81739 49.29402, 360 m, B. L. Fisher et al. leg. (CASC); 2 w., 1 s., Sahafina forest 11.4 km W Brickaville, - 18.81445 48.96205, 140 m, B. L. Fisher et al. leg. (CASC); 1 w., 1 s., Station forestière Analamazaotra, Analamazaotra 1.3 km S Andasibe, - 18.38466 48.41271, 980 m, B. L. Fisher et al. leg. (CASC); 4 w., 1 s., Torotorofotsy, - 18.87082 48.34737, 1070 m, B. L. Fisher et al. leg. (CASC); 1 w., Torotorofotsy, - 18.77048 48.43043, 1005 m, B. L. Fisher et al. leg. (CASC); 1 w., Toliara, 13 km NW Enakara, Rés. Andohahela, - 24.55 46.8, 1250 m, B. L. Fisher et al. leg. (CASC); 1 w., 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.	en	Salata, Sebastian, Fisher, Brian L. (2020): The ant genus Pheidole Westwood, 1839 (Hymenoptera: Formicidae) in Madagascar-taxonomic revision of the bessonii species-group. Zootaxa 4843 (1): 1-64, DOI: 10.11646/zootaxa.4843.1.1
03938025FFA6A93DA7D8B69FFB3EFC11.taxon	diagnosis	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.	en	Salata, Sebastian, Fisher, Brian L. (2020): The ant genus Pheidole Westwood, 1839 (Hymenoptera: Formicidae) in Madagascar-taxonomic revision of the bessonii species-group. Zootaxa 4843 (1): 1-64, DOI: 10.11646/zootaxa.4843.1.1
03938025FFA6A93DA7D8B69FFB3EFC11.taxon	description	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. 19 B). 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. 19 B, 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. 27 K). 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. 19 D). Surface indistinctly microreticulate. Pilosity moderately dense, long, and erect (Fig. 19 D, F). Postpetiole. In dorsal view trapezoidal and distinctly wider than long (Fig. 19 D, F). Gaster. Shiny and shagreened; pilosity moderately sparse, long, and erect (Fig. 19 D, F). Color. Reddish brown, antenna, legs and propodeum yellowish, gaster usually darker than mesosoma (Fig. 19 D, F). 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. 19 A). 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. 19 A, 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. 19 C). Sculpture with very sparse and weakly developed puncta. Pilosity sparse, moderately long, and erect (Fig. 19 C, E). Gaster. With sparse and erect pilosity (Fig. 19 C, E). Color. Yellowish brown, mandibles, antenna, and gaster usually brighter than the rest of the body (Fig. 19 C, E).	en	Salata, Sebastian, Fisher, Brian L. (2020): The ant genus Pheidole Westwood, 1839 (Hymenoptera: Formicidae) in Madagascar-taxonomic revision of the bessonii species-group. Zootaxa 4843 (1): 1-64, DOI: 10.11646/zootaxa.4843.1.1
03938025FFA6A93DA7D8B69FFB3EFC11.taxon	biology_ecology	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.	en	Salata, Sebastian, Fisher, Brian L. (2020): The ant genus Pheidole Westwood, 1839 (Hymenoptera: Formicidae) in Madagascar-taxonomic revision of the bessonii species-group. Zootaxa 4843 (1): 1-64, DOI: 10.11646/zootaxa.4843.1.1
03938025FFA6A93DA7D8B69FFB3EFC11.taxon	discussion	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.	en	Salata, Sebastian, Fisher, Brian L. (2020): The ant genus Pheidole Westwood, 1839 (Hymenoptera: Formicidae) in Madagascar-taxonomic revision of the bessonii species-group. Zootaxa 4843 (1): 1-64, DOI: 10.11646/zootaxa.4843.1.1
03938025FFA6A93DA7D8B69FFB3EFC11.taxon	etymology	Etymology. Malagasy for many or plentiful, in reference to the wide distribution of the species.	en	Salata, Sebastian, Fisher, Brian L. (2020): The ant genus Pheidole Westwood, 1839 (Hymenoptera: Formicidae) in Madagascar-taxonomic revision of the bessonii species-group. Zootaxa 4843 (1): 1-64, DOI: 10.11646/zootaxa.4843.1.1
03938025FFA5A938A7D8B236FCE4FDE4.taxon	description	Figs 20 A – F, 27 L, 28 L	en	Salata, Sebastian, Fisher, Brian L. (2020): The ant genus Pheidole Westwood, 1839 (Hymenoptera: Formicidae) in Madagascar-taxonomic revision of the bessonii species-group. Zootaxa 4843 (1): 1-64, DOI: 10.11646/zootaxa.4843.1.1
03938025FFA5A938A7D8B236FCE4FDE4.taxon	materials_examined	Other material. Madagascar. Fianarantsoa: 3 w., 1 s., Parc National de Ranomafana, Vatoharanana River, 4.1 km 231 ° SW Ranomafana, - 21.29 47.43333, 1100 m, Fisher et al. leg. (CASC). Toamasina: 2 w., 1 s., 5.3 km SSE Ambanizana, Andranobe, - 15.67133 49.97395, 425 m, Fisher et al. leg. (CASC); 8 w., 3 q., Ankerana, - 18.4061 48.82029, 725 m, Fisher et al. leg. (CASC); 1 w., 1 s., Ankerana, - 18.40062 48.81311, 865 m, Fisher et al. leg. (CASC); 8 w., Ankerana, - 18.40636 48.80254, 1108 m, Fisher et al. leg. (CASC); 6 w., 2 s., 1 m., Ankerana, - 18.4061 48.82029, 725 m, Fisher et al. leg. (CASC); 1 w., 1 s., Ankerana, - 18.40672 48.82281, 681 m, Fisher et al. leg. (CASC); 2 w., 1 s., 1 m., Ankerana, - 18.40636 48.80254, 1108 m, Fisher et al. leg. (CASC); 1 w., F. C. Andriantantely, - 18.695 48.81333, 530 m, H. J. Ratsirarson leg. (CASC); 6 w., F. C. Sandranantitra, - 18.04833 49.09167, 450 m, H. J. Ratsirarson leg. (CASC); 2 w., Montagne d’Akirindro 7.6 km 341 ° NNW Ambinanitelo, - 15.28833 49.54833, 600 m, Fisher et al. leg. (CASC); 26 w., 12 s., Montagne d’Anjanaharibe, 18.0 km 21 ° NNE Ambinanitelo, - 15.18833 49.615, 470 m, Fisher et al. leg. (CASC); 5 s., Montagne d’Anjanaharibe, 19.5 km 27 ° NNE Ambinanitelo, - 15.17833 49.635, 1100 m, Fisher et al. leg. (CASC); 1 s., Nosy Mangabe, 7.43 km S Maroantsetra, - 15.4973 49.76223, 3 m, Fisher et al. leg. (CASC); 1 w., 1 s., Parc National de Zahamena, - 17.73359 48.72625, 950 m, Fisher et al. leg. (CASC); 2 w., 1 s., Parc National de Zahamena, Tetezambatana forest, near junction of Nosivola and Manakambahiny Rivers, - 17.74298 48.72936, 860 m, Fisher et al. leg. (CASC); 1 w., 1 s., Parc National Mananara-Nord, 7.1 km 261 ° Antanambe, - 16.455 49.7875, 225 m, Fisher et al. leg. (CASC); 2 w., 2 s., Res. Ambodiriana, 4.8 km 306 ° Manompana, along Manompana river, - 16.67233 49.70117, 125 m, Fisher et al. leg. (CASC); 2 w., 2 s., Reserve Betampona, Camp Rendrirendry 34.1 km 332 ° Toamasina, - 17.924 49.19967, 390 m, Fisher et al. leg. (CASC); 2 w., 2 s., Reserve Betampona, Camp Vohitsivalana, 37.1 km 338 ° Toamasina, - 17.88667 49.2025, 520 m, Fisher et al. leg. (CASC); 2 w., 1 s., 1 m., Réserve Spéciale Ambatovaky, Sandrangato river, - 16.81753 49.29498, 360 m, Fisher et al. leg. (CASC); 2 w., 1 m., Réserve Spéciale Ambatovaky, Sandrangato river, - 16.76912 49.26704, 475 m, Fisher et al. leg. (CASC); 1 w., 1 s., Réserve Spéciale Ambatovaky, Sandrangato river, - 16.77468 49.26551, 355 m, Fisher et al. leg. (CASC); 1 w., 1 s., 1 q., Réserve Spéciale Ambatovaky, Sandrangato river, - 16.7633 49.26692, 520 m, Fisher et al. leg. (CASC); 1 w., 1 s., Réserve Spéciale Ambatovaky, Sandrangato river, - 16.7702 49.26638, 470 m, Fisher et al. leg. (CASC); 1 w., 1 s., Réserve Spéciale Ambatovaky, Sandrangato river, - 16.7755 49.26427, 430 m, Fisher et al. leg. (CASC); 2 w., 2 s., Réserve Spéciale Ambatovaky, Sandrangato river, - 16.81739 49.29402, 360 m, Fisher et al. leg. (CASC); 4 w., 3 s., 1 q., Réserve Spéciale Ambatovaky, Sandrangato river, - 16.81745 49.2925, 400 m, Fisher et al. leg. (CASC); 3 w., 1 m., Réserve Spéciale Ambatovaky, Sandrangato river, - 16.80561 49.29507, 480 m, Fisher et al. leg. (CASC). Geographic range. Madagascar, widely distributed in the central and eastern parts of the island.	en	Salata, Sebastian, Fisher, Brian L. (2020): The ant genus Pheidole Westwood, 1839 (Hymenoptera: Formicidae) in Madagascar-taxonomic revision of the bessonii species-group. Zootaxa 4843 (1): 1-64, DOI: 10.11646/zootaxa.4843.1.1
03938025FFA5A938A7D8B236FCE4FDE4.taxon	diagnosis	Diagnosis. Major workers. Head in full-face view sub-rectangular, not widening posteriorly, with lateral margins relatively straight; side of head with dense, long, suberect to erect pilosity; medial frons with moderately dense and thick costulae; interspaces between costulae distinctly punctate; frons laterally with thick and sparse rugae; interspaces between rugae distinctly punctate; occipital lobes punctate, sometimes with additional sparse and weakly developed rugae; scape, when laid back, exceeding the midlength of head by one-fifth of its length; inner hypostomal tooth distinct, small, and bulge-like, with top arching posteroventrally; outer hypostomal tooth lobe-like, moderately high, and wide, with top directed posteriorly; median tooth absent; promesonotum moderately long, angular, and very low; posterior mesonotum smoothly declining towards propodeum; mesonotal process weakly developed and tubercle-like; mesosoma punctate; lateral sides of pronotum, propodeum, and katepisternum with additional thin rugae; gaster indistinctly shagreened; body yellowish brown to brown. Minor workers. Occiput slightly elongated; nuchal collar indistinct; head sculpture punctate, gena mostly or entirely smooth; scape, when laid back, exceeding the posterior head margin by two-fifths of its length; promesonotum low, long, and arched; promesonotal groove present; metanotal groove distinct; propodeal spines small and thin; mesosoma punctate; body yellowish brown.	en	Salata, Sebastian, Fisher, Brian L. (2020): The ant genus Pheidole Westwood, 1839 (Hymenoptera: Formicidae) in Madagascar-taxonomic revision of the bessonii species-group. Zootaxa 4843 (1): 1-64, DOI: 10.11646/zootaxa.4843.1.1
03938025FFA5A938A7D8B236FCE4FDE4.taxon	description	Description. Major workers. Measurements (n = 10): HL: 1.56 – 1.73 (1.62); HW: 1.49 – 1.66 (1.55); SL: 1.04 – 1.18 (1.11); EL: 0.16 – 0.21 (0.19); WL: 1.54 – 1.68 (1.6); PSL: 0.19 – 0.27 (0.24); MTL: 1.14 – 1.38 (1.24); PNW: 0.52 – 0.65 (0.59); PTW: 0.15 – 0.18 (0.16); PPW: 0.36 – 0.45 (0.4); CI: 101.9 – 106.7 (104.4); SI: 69.3 – 75.6 (71.4); PSLI: 11.9 – 15.9 (15.1); PPI: 36.6 – 45.6 (40.9); PNI: 34.8 – 41.1 (38.0); MTI: 74.5 – 87.9 (79.8). Head. In full-face view sub-rectangular, not widening posteriorly, with lateral sides relatively straight (Fig. 20 B). In lateral view sub-oval, not depressed posteriorly; ventral and dorsal margins convex; inner hypostomal tooth visible. Side of head with dense, long, suberect to erect pilosity; head dorsum with relatively dense, long, suberect to erect pilosity. Medial frons with moderately dense and thick costulae; interspaces between costulae distinctly punctate. Frons laterally with thick and sparse rugae; interspaces between rugae distinctly punctate. Occipital lobes punctate, with sparse and sometimes weakly developed rugae. Gena with dense and moderately thick costulae; interspaces between costulae punctate. Sides posterolateral from eyes punctate and sometimes with additional network of thin and sparse rugulae. Center of clypeus shiny with weakly developed puncta, lateral sides with distinct rugulae; median notch present, moderately wide, and shallow; median longitudinal carina present; lateral longitudinal carinae present. Scape, when laid back, exceeding the midlength of head by one-fifth of its length; pilosity subdecumbent to erect (Fig. 20 B, D). Inner hypostomal tooth distinct, small, and bulge-like, with top arching posteroventrally; outer hypostomal tooth lobe-like, moderately high, and wide, with top directed posteriorly; median tooth absent (Fig. 27 L). Mesosoma. In lateral view, promesonotum moderately long, angular, and very low; posterior mesonotum smoothly declining towards propodeum; mesonotal process weakly developed and tuberclelike; promesonotal groove absent; metanotal groove indistinct; propodeal spines moderately long with narrow base, and acute top; humeral tubercle weakly produced (Fig. 20 D). Surface punctate; lateral sides of pronotum, propodeum, and katepisternum with additional thin rugae. Pilosity dense, long, and erect (Fig. 20 D, F). Gaster. Shiny and indistinctly shagreened; pilosity moderately sparse, long, and erect (Fig. 20 D, F). Color. Yellowish brown to brown, antennae and legs usually brighter (Fig. 20 D, F). Description. Minor workers. Measurements (n = 10): HL: 0.73 – 0.82 (0.78); HW: 0.49 – 0.57 (0.53); SL: 1.03 – 1.17 (1.12); EL: 0.13 – 0.15 (0.14); WL: 1.02 – 1.14 (1.07); PSL: 0.11 – 0.14 (0.13); MTL: 0.9 – 1.07 (0.99); PNW: 0.37 – 0.42 (0.39); PTW: 0.07 – 0.11 (0.08); PPW: 0.14 – 0.2 (0.17); CI: 141.8 – 149.8 (147.0); SI: 200.6 – 216.2 (209.8); PSLI: 14.9 – 17.9 (16.2); PPI: 44.7 – 54.0 (50.4); PNI: 68.8 – 75.7 (72.6); MTI: 164.2 – 198.3 (185.6). Head. In full-face view oval, posterior region slightly elongated; nuchal collar weakly developed (Fig. 20 A). Pilosity relatively sparse, moderately long, subdecumbent to erect. Sculpture punctate, gena mostly or entirely smooth. 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. 20 A, C). Mesosoma. In lateral view, promesonotum low, long, and arched; promesonotal groove present; metanotal groove distinct; propodeal spines small, thin, and triangular (Fig. 20 C). Sculpture punctate. Pilosity sparse, long, and erect (Fig. 20 C, E). Gaster. With sparse and erect pilosity (Fig. 20 C, E). Color. Yellowish brown (Fig. 20 C, E).	en	Salata, Sebastian, Fisher, Brian L. (2020): The ant genus Pheidole Westwood, 1839 (Hymenoptera: Formicidae) in Madagascar-taxonomic revision of the bessonii species-group. Zootaxa 4843 (1): 1-64, DOI: 10.11646/zootaxa.4843.1.1
03938025FFA5A938A7D8B236FCE4FDE4.taxon	biology_ecology	Biology. The species was collected between 3 – 1490 m elevation, in rainforest, montane forest, littoral rainforest, montane rainforest, and montane shrubland. Nests were located in rotten logs, rotting tree stumps, and rotten sticks on the ground. Workers were collected from sifted litter, on low vegetation, and on tree trunks.	en	Salata, Sebastian, Fisher, Brian L. (2020): The ant genus Pheidole Westwood, 1839 (Hymenoptera: Formicidae) in Madagascar-taxonomic revision of the bessonii species-group. Zootaxa 4843 (1): 1-64, DOI: 10.11646/zootaxa.4843.1.1
03938025FFA5A938A7D8B236FCE4FDE4.taxon	discussion	Comments. Pheidole oswaldi is most similar to sympatric P. decollata. Its major workers can be separated by moderately long, angular, and very low promesonotum and small inner hypostomal tooth. In contrast, majors of P. decollata have short, angular, and moderately low promesonotum and moderately large inner hypostomal tooth. Minors of P. oswaldi differ in slightly elongated posterior region of head and moderately long, angular, and very low promesonotum. Minor workers of P. decollata have posterior region of the head not elongated and the promesonotum is moderately long, moderately low, and angular.	en	Salata, Sebastian, Fisher, Brian L. (2020): The ant genus Pheidole Westwood, 1839 (Hymenoptera: Formicidae) in Madagascar-taxonomic revision of the bessonii species-group. Zootaxa 4843 (1): 1-64, DOI: 10.11646/zootaxa.4843.1.1
03938025FFA0A926A7D8B46DFA9AFBED.taxon	description	Figs 21 A – F, 27 M, 28 M	en	Salata, Sebastian, Fisher, Brian L. (2020): The ant genus Pheidole Westwood, 1839 (Hymenoptera: Formicidae) in Madagascar-taxonomic revision of the bessonii species-group. Zootaxa 4843 (1): 1-64, DOI: 10.11646/zootaxa.4843.1.1
03938025FFA0A926A7D8B46DFA9AFBED.taxon	materials_examined	HOLOTYPE: 1 s., Madagascar, Antsiranana, Sava Region: Parc National de Marojejy, Manantenina River, 27.9 km 24.3 ° NE Andapa, - 14.43462 49.75853, 850 m, 9 - Feb- 2018, rainforest, ex tunelle on clay bark, B. L. Fisher et al. leg. BLF 40902, CASENT 0808080 (CASC). PARATYPES: 1 w., the same data as holotype, CASENT 0923277 (CASC). Other material. Madagascar. Antsiranana: 3 w., 2 s., Sava Region: Parc National de Marojejy, Manantenina River, 27.9 km 24.3 ° NE Andapa, - 14.43462 49.75853, 850 m, B. L. Fisher et al., (CASC). Geographic range. Madagascar, Antsiranana, Parc National de Marojejy.	en	Salata, Sebastian, Fisher, Brian L. (2020): The ant genus Pheidole Westwood, 1839 (Hymenoptera: Formicidae) in Madagascar-taxonomic revision of the bessonii species-group. Zootaxa 4843 (1): 1-64, DOI: 10.11646/zootaxa.4843.1.1
03938025FFA0A926A7D8B46DFA9AFBED.taxon	diagnosis	Diagnosis. Major workers. Head in full-face view sub-rectangular, not widening posteriorly, with lateral margins relatively straight; side of head with dense, short, decumbent to suberect pilosity; anteromedial frons with moderately dense and thick costulae; interspaces between costulae indistinctly rugulate; posteromedial frons rugulate; interspaces between rugulae more distinctly rugulate; frons laterally with thick and moderately dense rugae; interspaces between rugae indistinctly rugulate; occipital lobes with thick and moderately dense rugae; interspaces between rugae mostly smooth; scape, when laid back, exceeding the midlength of head by one-fifth of its length; inner hypostomal tooth distinct and bulge-like; outer hypostomal tooth absent to very indistinct; median tooth absent; promesonotum short, angular, and moderately low; posterior mesonotum moderately steep; mesonotal process distinct; mesosoma microreticulate, lateral sides of pronotum, propodeum, and katepisternum with additional thin rugae; pronotal dorsum with additional rugae; gaster shagreened; body brown. Minor workers. Occiput not elongated; nuchal collar indistinct; head sculpture smooth; scape, when laid back, exceeding the posterior head margin by two-fifths of its length; promesonotum moderately low, moderately long, and arched; promesonotal groove present; metanotal groove distinct; propodeal spines very small and thin; promesonotum smooth; anepisternum, katepisternum, and propodeum punctate; body yellowish brown; petiolar node high, triangular, and thick.	en	Salata, Sebastian, Fisher, Brian L. (2020): The ant genus Pheidole Westwood, 1839 (Hymenoptera: Formicidae) in Madagascar-taxonomic revision of the bessonii species-group. Zootaxa 4843 (1): 1-64, DOI: 10.11646/zootaxa.4843.1.1
03938025FFA0A926A7D8B46DFA9AFBED.taxon	description	Description. Major workers. Measurements (n = 4): HL: 1.35 – 1.5 (1.41); HW: 1.3 – 1.43 (1.36); SL: 0.92 – 0.98 (0.95); EL: 0.18 – 0.19 (0.18); WL: 1.25 – 1.35 (1.29); PSL: 0.2 – 0.23 (0.22); MTL: 0.95 – 1.02 (0.99); PNW: 0.53 – 0.57 (0.56); PTW: 0.17 – 0.19 (0.18); PPW: 0.33 – 0.42 (0.38); CI: 101.9 – 105.3 (103.9); SI: 68.5 – 71.1 (70.2); PSLI: 14.8 – 16.6 (15.7); PPI: 43.3 – 50.5 (46.3); PNI: 39.9 – 41.7 (41.0); MTI: 71.4 – 74.3 (73.1). Head. In full-face view sub-rectangular, not widening posteriorly, with lateral sides relatively straight (Fig. 21 B). In lateral view sub-oval, not depressed posteriorly; ventral and dorsal margins convex; inner hypostomal tooth visible. Side of head with dense, short, decumbent to suberect pilosity; head dorsum with relatively dense, long, suberect to erect pilosity. Anteromedial frons with moderately dense and thick costulae; interspaces between costulae indistinctly rugulate; posteromedial frons rugulate; interspaces between rugulae more distinctly rugulate. Frons laterally with thick and moderately dense rugae; interspaces between rugae indistinctly rugulate. Occipital lobes with thick and moderately dense rugae; interspaces between rugae mostly smooth. Gena with dense and moderately thick costulae; interspaces between costulae mostly punctate. Sides posterolateral from eyes with thick rugulae; interspaces between rugulae distinctly rugulate. Center of clypeus shiny with weakly developed puncta, lateral sides with distinct rugulae; median notch present, moderately wide and shallow; median longitudinal carina present; lateral longitudinal carinae present. Scape, when laid back, exceeding the midlength of head by one-fifth of its length; pilosity subdecumbent to erect (Fig. 21 B, D). Inner hypostomal tooth distinct, bulge-like; outer hypostomal tooth absent to very indistinct; median tooth absent (Fig. 21 M). 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 indistinct; propodeal spines moderately long with moderately wide base and acute top; humeral tubercle weakly produced (Fig. 21 D). Surface microreticulate; lateral sides of pronotum, propodeum, and katepisternum with additional thin rugae; pronotal dorsum with additional rugae. Pilosity moderately dense, long, and erect (Fig. 21 D, F). Gaster. Shiny and shagreened; pilosity moderately sparse, long, and erect (Fig. 21 D, F). Color. Brown, legs yellowish (Fig. 21 D, F). Description. Minor workers. Measurements (n = 4): HL: 0.6 – 0.64 (0.62); HW: 0.46 – 0.49 (0.47); SL: 0.77 – 0.8 (0.79); EL: 0.11 – 0.13 (0.13); WL: 0.81 – 0.87 (0.84); PSL: 0.09 – 0.1 (0.09); MTL: 0.64 – 0.67 (0.65); PNW: 0.3 – 0.35 (0.33); PTW: 0.08 – 0.11 (0.09); PPW: 0.12 – 0.14 (0.13); CI: 127.6 – 134.7 (130.5); SI: 162.9 – 170.7 (166.1); PSLI: 14.1 – 15.7 (14.8); PPI: 59.4 – 79.9 (69.6); PNI: 64.4 – 73.8 (70.0); MTI: 134.5 – 138.6 (137.3). Head. In full-face view oval, posterior region not elongated; nuchal collar weakly developed (Fig. 21 A). Pilosity relatively sparse, moderately short, subdecumbent to erect. Sculpture mostly smooth; anterolateral frons punctate. 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. 21 A, C). Mesosoma. In lateral view, promesonotum moderately low, moderately long, and arched; promesonotal groove present; metanotal groove distinct; propodeal spines very small, thin, and triangular (Fig. 21 C). Promesonotum smooth; anepisternum, katepisternum, and propodeum punctate. Pilosity sparse, moderately long, and erect (Fig. 21 C, E). Gaster. With sparse and erect pilosity (Fig. 21 C, E). Color. Yellowish brown, gaster and head darker than mesosoma (Fig. 21 C, E).	en	Salata, Sebastian, Fisher, Brian L. (2020): The ant genus Pheidole Westwood, 1839 (Hymenoptera: Formicidae) in Madagascar-taxonomic revision of the bessonii species-group. Zootaxa 4843 (1): 1-64, DOI: 10.11646/zootaxa.4843.1.1
03938025FFA0A926A7D8B46DFA9AFBED.taxon	biology_ecology	Biology. The species was collected at 850 m elevation, in rainforest. Nests were located in the soil and on clay banks.	en	Salata, Sebastian, Fisher, Brian L. (2020): The ant genus Pheidole Westwood, 1839 (Hymenoptera: Formicidae) in Madagascar-taxonomic revision of the bessonii species-group. Zootaxa 4843 (1): 1-64, DOI: 10.11646/zootaxa.4843.1.1
03938025FFA0A926A7D8B46DFA9AFBED.taxon	discussion	Comments. Pheidole ovalinoda is known only from Parc National de Marojejy in Antsiranana. Its major workers can be confused with parapatric P. flammea, recorded only from Galoko Mountain. Majors of P. ovalinoda differ from P. flammea in the presence of dense, short and decumbent to suberect setae on the sides of the head, strongly reduced outer hypostomal tooth, and more distinctly sculptured interspaces between rugae on head. Majors of P. flammea have moderately dense, long and suberect to erect pilosity on the side of the head, large outer hypostomal tooth, and mostly smooth interspaces between rugae on head. Minors of P. ovalinoda are easy to separate from other Pheidole known from the island based on the combination of the following characters: petiolar node high, triangular, and thick; postpetiole short and moderately high; yellowish brown body coloration; neck absent, and nuchal collar indistinct; head and promesonotum mostly smooth; promesonotal and metanotal grooves distinct.	en	Salata, Sebastian, Fisher, Brian L. (2020): The ant genus Pheidole Westwood, 1839 (Hymenoptera: Formicidae) in Madagascar-taxonomic revision of the bessonii species-group. Zootaxa 4843 (1): 1-64, DOI: 10.11646/zootaxa.4843.1.1
03938025FFA0A926A7D8B46DFA9AFBED.taxon	etymology	Etymology. Latin for oval and node, in reference to the high and spherical postpetiole of minor workers.	en	Salata, Sebastian, Fisher, Brian L. (2020): The ant genus Pheidole Westwood, 1839 (Hymenoptera: Formicidae) in Madagascar-taxonomic revision of the bessonii species-group. Zootaxa 4843 (1): 1-64, DOI: 10.11646/zootaxa.4843.1.1
03938025FFBEA924A7D8B26AFAE7F9F5.taxon	description	Figs 22 A – F, 27 N, 28 N	en	Salata, Sebastian, Fisher, Brian L. (2020): The ant genus Pheidole Westwood, 1839 (Hymenoptera: Formicidae) in Madagascar-taxonomic revision of the bessonii species-group. Zootaxa 4843 (1): 1-64, DOI: 10.11646/zootaxa.4843.1.1
03938025FFBEA924A7D8B26AFAE7F9F5.taxon	materials_examined	HOLOTYPE: 1 s., Madagascar, Antsiranana, Nosy Be, Réserve Naturelle Intégrale de Lokobe, 6.3 km 112 ° ESE Hellville, - 13.41933 48.33117, 30 m, 19 - Mar- 2001, rainforest, ex rotten log, Fisher et al. leg. BLF 03447, CASENT 0427852 (CASC). PARATYPES: 2 w., the same data as holotype, CASENT 0427853, CASENT 0427854 (CASC, MHNG). Other material. Madagascar. Antsiranana: 3 w., Nosy Be, Réserve Naturelle Intégrale de Lokobe, 6.3 km 112 ° ESE Hellville, - 13.41933 48.33117, 30 m, Fisher et al. leg. (CASC); 7 w., 1 s., Nosy Be, Réserve Naturelle Intégrale de Lokobe, 6.3 km 112 ° ESE Hellville, - 13.41933 48.33117, 30 m, Fisher et al. leg. (CASC). Geographic range. Madagascar, Antsiranana, Nosy Be.	en	Salata, Sebastian, Fisher, Brian L. (2020): The ant genus Pheidole Westwood, 1839 (Hymenoptera: Formicidae) in Madagascar-taxonomic revision of the bessonii species-group. Zootaxa 4843 (1): 1-64, DOI: 10.11646/zootaxa.4843.1.1
03938025FFBEA924A7D8B26AFAE7F9F5.taxon	diagnosis	Diagnosis. Major workers. Head in full-face view sub-rectangular, not widening posteriorly, with lateral margins relatively straight; side of head with moderately sparse, short, subdecumbent to suberect pilosity; medial frons with moderately sparse and thick costulae; interspaces between costulae smooth; frons laterally with thick rugocostulae; interspaces between rugocostulae rugulate; occipital lobes with thick rugae; interspaces between rugae smooth; scape, when laid back, exceeding the midlength of head by two-fifths of its length; inner hypostomal tooth distinct, small, bulge-like, with top arching posteroventrally; outer hypostomal tooth lobe-like, high, and wide, with top directed posteriorly; median tooth absent; promesonotum short, angular, and moderately low; posterior mesonotum moderately steep; mesonotal process distinct and tubercle-like; mesosoma microreticulate; promesonotal dorsum with sculpture reduced; gaster shagreened; body orange. Minor workers. Occiput elongated forming short and moderately narrow neck; nuchal collar distinct; head sculpture smooth; scape, when laid back, exceeding the posterior head margin by two-fifths of its length; promesonotum low, long, and arched; promesonotal groove present; metanotal groove distinct; propodeal spines moderate with narrow base; promesonotum smooth; katepisternum and propodeum with sparse puncta; body yellow.	en	Salata, Sebastian, Fisher, Brian L. (2020): The ant genus Pheidole Westwood, 1839 (Hymenoptera: Formicidae) in Madagascar-taxonomic revision of the bessonii species-group. Zootaxa 4843 (1): 1-64, DOI: 10.11646/zootaxa.4843.1.1
03938025FFBEA924A7D8B26AFAE7F9F5.taxon	description	Description. Major workers. Measurements (n = 2): HL: 1.55, 1.58; HW: 1.38, 1.47; SL: 1.18, 1.14; EL: 0.21, 0.21; WL: 1.46, 1.44; PSL: 0.3, 0.31; MTL: 1.17, 1.17; PNW: 0.56, 0.58; PTW: 0.21, 0.21; PPW: 0.43, 0.5; CI: 112.2, 107.7; SI: 85.5, 77.7; PSLI: 19.2, 19.3; PPI: 48.1, 41.8; PNI: 40.3, 39.7; MTI: 84.8, 79.9. Head. In full-face view sub-rectangular, not widening posteriorly, with lateral sides relatively straight (Fig. 22 B). In lateral view sub-oval, not depressed posteriorly; ventral and dorsal margins convex; inner hypostomal tooth not visible. Side of head with moderately sparse, short, subdecumbent to suberect pilosity; head dorsum with relatively dense, long, decumbent to erect pilosity. Medial frons with moderately sparse and thick costulae; interspaces between costulae smooth. Frons laterally with thick rugocostulae; interspaces between rugocostulae rugulate. Occipital lobes with thick rugae; interspaces between rugae smooth. Gena with dense and thick costulae; interspaces between costulae smooth. Sides posterolateral from eyes with dense but very thin network of rugoreticulae; sculpture slightly weakening posteriorly. Center of clypeus shiny and smooth, lateral sides with distinct 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. 22 B, D). Inner hypostomal tooth distinct, small, bulge-like, with top arching posteroventrally; outer hypostomal tooth lobe-like, high, and wide, with top directed posteriorly; median tooth absent (Fig. 27 N). 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 present; propodeal spines moderately long, with narrow base and acute top; humeral tubercle weakly produced (Fig. 22 D). Surface shiny and microreticulate; promesonotal dorsum with sculpture reduced. Pilosity moderately dense, moderately long, and erect (Fig. 22 D, F). Petiole. In rear view node dorsoventrally slightly convex (Fig. 22 D, F). Gaster. Shiny and shagreened; pilosity moderately sparse, long, and erect (Fig. 22 D, F). Color. Orange, legs yellow, gaster darker than mesosoma (Fig. 22 D, F). Description. Minor workers. Measurements (n = 10): HL: 0.7 – 0.89 (0.81); HW: 0.46 – 0.57 (0.52); SL: 1.01 – 1.28 (1.18); EL: 0.13 – 0.17 (0.14); WL: 0.89 – 1.24 (1.06); PSL: 0.12 – 0.17 (0.15); MTL: 0.79 – 1.2 (0.99); PNW: 0.32 – 0.44 (0.37); PTW: 0.07 – 0.12 (0.09); PPW: 0.12 – 0.21 (0.15); CI: 152.4 – 161.2 (156.0); SI: 220.8 – 237.2 (228.0); PSLI: 15.1 – 20.7 (18.2); PPI: 52.4 – 70.6 (61.6); PNI: 68.2 – 77.4 (71.5); MTI: 161.5 – 211.0 (190.5). Head. In full-face view oval, posterior region elongated forming short and moderately narrow neck; nuchal collar distinct (Fig. 22 A). Pilosity relatively sparse, long, and subdecumbent to erect. Sculpture shiny and mostly smooth; antennal sockets with few indistinct rugulae that are curved posterolaterally; interspaces between rugulae smooth. 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. 22 A, C). Mesosoma. In lateral view, promesonotum low, long, and arched; promesonotal groove present; metanotal groove distinct; propodeal spines moderate and triangular, with narrow base (Fig. 22 C). Promesonotum smooth; katepisternum and propodeum sparsely punctate. Pilosity very sparse, long, and erect (Fig. 22 C, E). Gaster. With sparse and erect pilosity (Fig. 22 C, E). Color. Yellow, gaster and head darker than mesosoma (Fig. 22 C, E).	en	Salata, Sebastian, Fisher, Brian L. (2020): The ant genus Pheidole Westwood, 1839 (Hymenoptera: Formicidae) in Madagascar-taxonomic revision of the bessonii species-group. Zootaxa 4843 (1): 1-64, DOI: 10.11646/zootaxa.4843.1.1
03938025FFBEA924A7D8B26AFAE7F9F5.taxon	biology_ecology	Biology. The species was collected at 30 m elevation, in rainforest. Nests were located in rotten logs. Workers were collected from sifted litter.	en	Salata, Sebastian, Fisher, Brian L. (2020): The ant genus Pheidole Westwood, 1839 (Hymenoptera: Formicidae) in Madagascar-taxonomic revision of the bessonii species-group. Zootaxa 4843 (1): 1-64, DOI: 10.11646/zootaxa.4843.1.1
03938025FFBEA924A7D8B26AFAE7F9F5.taxon	discussion	Comments. Among species known from Nosy Be islet, Pheidole similis is the only member of the bessonii group with both minor and major workers yellow to orange. It is most similar to two species known from the eastern parts of Antsiranana and the northernmost part of Toamasina: P. clara and P. mantadioflava. Its major workers differ from these two taxa in frons laterally with thick rugocostulae, and with rugulate interspaces between rugulae. Majors of P. clara and P. mantadioflava have frons laterally entirely punctate or rugulate with punctate interspaces between rugulae. Minor workers can be easily separated based on smooth promesonotum. Pheidole similis is morphologically most similar to P. ragnax, described by Fischer & Fisher from Mayotte island. Minors of P. similis can be separated based on longer and more narrow neck, and presence of sparse but distinct puncta on katepisternum and propodeum. Majors differ in more rugulate lateral frons, sparser setae on sides of head, more distinct and dense mesosomal sculpture, and shagreened gaster.	en	Salata, Sebastian, Fisher, Brian L. (2020): The ant genus Pheidole Westwood, 1839 (Hymenoptera: Formicidae) in Madagascar-taxonomic revision of the bessonii species-group. Zootaxa 4843 (1): 1-64, DOI: 10.11646/zootaxa.4843.1.1
03938025FFBEA924A7D8B26AFAE7F9F5.taxon	etymology	Etymology. Latin for similar, in reference to the morphological similarity of P. similis to P. ragnax.	en	Salata, Sebastian, Fisher, Brian L. (2020): The ant genus Pheidole Westwood, 1839 (Hymenoptera: Formicidae) in Madagascar-taxonomic revision of the bessonii species-group. Zootaxa 4843 (1): 1-64, DOI: 10.11646/zootaxa.4843.1.1
03938025FFBCA923A7D8B052FF0CFA60.taxon	description	Figs 23 A – F, 27 O, 28 O	en	Salata, Sebastian, Fisher, Brian L. (2020): The ant genus Pheidole Westwood, 1839 (Hymenoptera: Formicidae) in Madagascar-taxonomic revision of the bessonii species-group. Zootaxa 4843 (1): 1-64, DOI: 10.11646/zootaxa.4843.1.1
03938025FFBCA923A7D8B052FF0CFA60.taxon	materials_examined	HOLOTYPE: 1 s., Madagascar, Antananarivo, Ambohidratrimo, - 18.821 47.44183, 1362 m, 8 - May- 2007, Urban garden, under stone, B. L. Fisher et al. leg. BLF 17538, CASENT 0128508 (CASC). PARATYPES: 1 w., the same data as holotype, CASENT 0235036 (CASC). Other material. Madagascar. Antananarivo: 1 w., Ilafy, - 18.85415 47.56575, 1385 m, B. L. Fisher et al. leg. (CASC); 4 w., Kaloy, - 18.59568 47.65333, 1338 m, B. L. Fisher et al. leg. (CASC); 1 w., Tsimbazaza, - 18.928 47.527, 1300 m, B. L. Fisher et al. leg. (CASC). Antsiranana: 1 w., Ampasindava, Forêt d’Ambilanivy, 3.9 km 181 ° S Ambaliha, - 13.79861 48.16167, 600 m, B. L. Fisher et al. leg. (CASC); 3 w., 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); 1 w., 1 q., Forêt d’Ampondrabe, 26.3 km 10 ° NNE Daraina, - 12.97 49.7, 175 m, B. L. Fisher et al. leg. (CASC); 1 w., Forêt d’Analabe, 30.0 km 72 ° ENE Daraina, - 13.08333 49.90833, 30 m, B. L. Fisher et al. leg. (CASC); 1 w., Forêt de Bekaraoka, 6.8 km 60 ° ENE Daraina, - 13.16667 49.71, 150 m, B. L. Fisher et al. leg. (CASC); 1 w., 1 s., Masoala National Park, - 15.32331 50.30751, 60 m, B. L. Fisher et al. leg. (CASC); 1 w., 1 s., Masoala, Cap Est, Forêt d’Andranoanala, - 15.26158 50.4758, 15 m, B. L. Fisher et al. leg. (CASC); 6 w., Nosy Be, Réserve Naturelle Intégrale de Lokobe, 6.3 km 112 ° ESE Hellville, - 13.41933 48.33117, 30 m, B. L. Fisher et al. leg. (CASC); 2 w., Parc National Montagne d’Ambre, 12.2 km 211 ° SSW Joffreville, - 12.59639 49.1595, 1300 m, Alpert et al. leg. (CASC); 4 w., 1 s., R. S. Manongarivo, 10.8 km 229 ° SW Antanambao, - 13.96167 48.43333, 400 m, B. L. Fisher et al. leg. (CASC); 1 w., 1 s., 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); 2 w., 2 s., Sahamalaza Peninsula, Forêt d’Anabohazo, 21.6 km 247 ° WSW Maromandia, - 14.30889 47.91433, 120 m, B. L. Fisher et al. leg. (CASC). Fianarantsoa: 1 w., Belle Vue trail, Ranomafana National Park, Fianarantsoa Prov., - 21.2665 47.42017, 1020 m, R. Harin’Hala leg. (CASC); 4 w., dry wash, 1 km E of Isalo National Park Interpretive Center, Fianarantsoa Prov., - 22.62667 45.35817, 885 m, R. Harin’Hala leg. (CASC); 7 w., 3 s., Parc National d’Isalo, 9.1 km 354 ° N Ranohira, - 22.48167 45.46167, 725 m, B. L. Fisher et al. leg. (CASC); 3 w., Parc National d’Isalo, Sahanafa River, 29.2 km 351 ° N Ranohira, - 22.31333 45.29167, 500 m, B. L. Fisher et al. leg. (CASC); 1 s., R. S. Ivohibe, 6.5 km ESE Ivohibe, - 22.49667 46.955, 1575 m, B. L. Fisher et al. leg. (CASC); 1 w., Réserve Forestière d’Agnalazaha, Mahabo, 42.9 km 215 ° Farafangana, - 23.19383 47.723, 20 m, B. L. Fisher et al. leg. (CASC); 1 w., stream area, 900 m E of Isalo National Park Interpretive Center, Fianarantsoa Prov., - 22.62667 45.35817, 750 m, R. Harin’Hala leg. (CASC). Mahajanga: 2 w., Parc National de Baie de Baly, 12.4 km 337 ° NNW Soalala, - 16.01 45.265, 10 m, B. L. Fisher et al. leg. (CASC); 8 w., 5 s., Parc National de Namoroka, 16.9 km 317 ° NW Vilanandro, - 16.40667 45.31, 100 m, B. L. Fisher et al. leg. (CASC); 2 w., Parc National Tsingy de Bemaraha, 10.6 km ESE 123 ° Antsalova, - 18.70944 44.71817, 150 m, B. L. Fisher et al. leg. (CASC). Toamasina: 2 w., 1 s., Ambatovy, 12.4 km NE Moramanga, - 18.85813 48.28488, 1040 m, B. L. Fisher et al. leg. (CASC); 2 w., F. C. Didy, - 18.19833 48.57833, 960 m, H. J. Ratsirarson leg. (CASC); 1 w., 1 s., Forêt d’Analava Mandrisy, 5.9 km 195 º Antanambe, - 16.48567 49.847, 10 m, B. L. Fisher et al. leg. (CASC); 2 w., 2 s., 1 m., Menafotaka, village near Onibe, - 17.75872 48.85178, 785 m, B. L. Fisher et al. leg. (CASC); 1 w., 1 s., Moramanga, - 18.94417 48.23067, 922 m, B. L. Fisher et al. leg. (CASC); 1 w., Reserve Betampona, Camp Rendrirendry 34.1 km 332 ° Toamasina, - 17.924 49.19967, 390 m, B. L. Fisher et al. leg. (CASC). Toliara: 4 w., Makay Mts., - 21.13513 45.33098, 650 m, B. L. Fisher et al. leg. (CASC); 1 w., Parc National de Zombitse, 17.7 km 98 ° E Sakaraha, - 22.88833 44.70167, 760 m, B. L. Fisher et al. leg. (CASC); 4 w., Parc National de Zombitse, 19.8 km 84 ° E Sakaraha, - 22.84333 44.71, 770 m, B. L. Fisher et al. leg. (CASC); 1 w., Réserve Spéciale d’Ambohijanahary, Forêt d’Ankazotsihitafototra, 34.6 km 314 ° NW Ambaravaranala, - 18.26 45.41833, 1100 m, B. L. Fisher et al. leg. (CASC); 1 w., southern Isoky-Vohimena Forest, 59 km NE Sakaraha, - 22.46667 44.85, 730 m, B. L. Fisher et al. leg. (CASC). Geographic range. Madagascar, species recorded from most of the island.	en	Salata, Sebastian, Fisher, Brian L. (2020): The ant genus Pheidole Westwood, 1839 (Hymenoptera: Formicidae) in Madagascar-taxonomic revision of the bessonii species-group. Zootaxa 4843 (1): 1-64, DOI: 10.11646/zootaxa.4843.1.1
03938025FFBCA923A7D8B052FF0CFA60.taxon	diagnosis	Diagnosis. Major workers. Head in full-face view sub-rectangular, not widening posteriorly, with lateral margins relatively straight; side of head with moderately dense, short, and appressed to subdecumbent pilosity; medial frons with dense and thick costulae; interspaces between costulae smooth; frons laterally rugulate; interspaces between rugulae punctate; occipital lobes with thin rugae; interspaces between rugae indistinctly punctate to smooth; scape, when laid back, exceeding the midlength of head by one-fifth of its length; inner hypostomal tooth indistinct, small and bulge-like, with top directed posteriorly; outer hypostomal tooth lobe-like, high, and wide, with top directed posteriorly; median tooth absent; promesonotum short, angular, and moderately low; posterior mesonotum moderately steep; mesonotal process distinct; mesosoma punctate; promesonotum with additional transverse and thin rugae; katepisternum and lateral sides of propodeum with additional rugae; gaster shagreened; body dark brown. Minor workers. Occiput not elongated; nuchal collar distinct; head sculpture punctate; sometimes frons and vertex with puncta reduced or absent; scape, when laid back, exceeding the posterior head margin by two-fifths of its length; promesonotum moderately low, moderately long, and arched; promesonotal groove absent to indistinct; metanotal groove distinct; propodeal spines moderate and with narrow base; mesosoma punctate; body dark brown.	en	Salata, Sebastian, Fisher, Brian L. (2020): The ant genus Pheidole Westwood, 1839 (Hymenoptera: Formicidae) in Madagascar-taxonomic revision of the bessonii species-group. Zootaxa 4843 (1): 1-64, DOI: 10.11646/zootaxa.4843.1.1
03938025FFBCA923A7D8B052FF0CFA60.taxon	description	Description. Major workers. Measurements (n = 10): HL: 1.65 – 1.85 (1.74); HW: 1.68 – 1.8 (1.74); SL: 0.92 – 0.99 (0.95); EL: 0.22 – 0.26 (0.23); WL: 1.38 – 1.53 (1.45); PSL: 0.25 – 0.31 (0.28); MTL: 0.94 – 1.1 (1.02); PNW: 0.7 – 0.78 (0.73); PTW: 0.21 – 0.31 (0.24); PPW: 0.58 – 0.71 (0.65); CI: 98.2 – 103.1 (100.6); SI: 52.6 – 57.6 (54.9); PSLI: 14.0 – 17.8 (15.8); PPI: 32.8 – 43.7 (37.2); PNI: 40.7 – 44.4 (42.2); MTI: 54.8 – 65.2 (58.4). Head. In full-face view sub-rectangular, not widening posteriorly, with lateral sides relatively straight (Fig. 23 B). In lateral view sub-oval, not depressed posteriorly; ventral and dorsal margins convex; inner hypostomal tooth not visible. Side of head with moderately dense, short, and appressed to subdecumbent pilosity; head dorsum with relatively dense, long, and decumbent to erect pilosity. Medial frons with dense and thick costulae; interspaces between costulae smooth. Frons laterally rugulate; interspaces between rugulae punctate. Occipital lobes with thin rugae; interspaces between rugae indistinctly punctate to smooth. Gena with dense and thick costulae; interspaces between costulae distinctly punctate. Sides posterolateral from eyes with dense but 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, exceeding the midlength of head by one-fifth of its length; pilosity appressed and subdecumbent (Fig. 23 B, D). Inner hypostomal tooth indistinct, small and bulge-like, with top directed posteriorly; outer hypostomal tooth lobe-like, high, and wide, with top directed posteriorly; median tooth absent (Fig. 27 O). 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 moderately wide base and acute top; humeral tubercle weakly produced (Fig. 23 D). Surface shiny and punctate; promesonotum with additional transverse and thin rugae; katepisternum and lateral sides of propodeum with additional rugae. Pilosity moderately dense, moderately long, and erect (Fig. 23 D, F). Postpetiole. In dorsal view distinctly wider than long (Fig. 23 D, F). Gaster. Dull and shagreened; pilosity moderately sparse, long, and erect (Fig. 23 D, F). Color. Dark brown, legs slightly brighter, antennae yellowish (Fig. 23 D, F). Description. Minor workers. Measurements (n = 10): HL: 0.62 – 0.79 (0.66); HW: 0.55 – 0.68 (0.59); SL: 0.73 – 0.85 (0.79); EL: 0.13 – 0.15 (0.14); WL: 0.78 – 0.93 (0.86); PSL: 0.13 – 0.19 (0.15); MTL: 0.58 – 0.75 (0.64); PNW: 0.36 – 0.48 (0.39); PTW: 0.07 – 0.11 (0.09); PPW: 0.13 – 0.2 (0.14); CI: 100.8 – 118.5 (133.3); SI: 115.6 – 142.8 (133.4); PSLI: 20.6 – 25.4 (22.8); PPI: 49.7 – 74.0 (62.0); PNI: 59.1 – 70.3 (66.7); MTI: 92.8 – 117.9 (109.7). Head. In full-face view oval, posterior region not elongated; nuchal collar distinct (Fig. 23 A). Pilosity relatively sparse, long, subdecumbent to absent. Sculpture shiny and punctate; sometimes frons and vertex with puncta reduced or absent. 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. 23 A, C). Mesosoma. In lateral view, promesonotum moderately low, moderately long, and arched; promesonotal groove absent to indistinct; metanotal groove distinct; propodeal spines moderate and triangular, with narrow base (Fig. 23 C). Sculpture punctate. Pilosity very sparse, short, and erect (Fig. 23 C, E). Gaster. With sparse and erect pilosity (Fig. 23 C, E). Color. Dark brown, legs and antennae brighter (Fig. 23 C, E).	en	Salata, Sebastian, Fisher, Brian L. (2020): The ant genus Pheidole Westwood, 1839 (Hymenoptera: Formicidae) in Madagascar-taxonomic revision of the bessonii species-group. Zootaxa 4843 (1): 1-64, DOI: 10.11646/zootaxa.4843.1.1
03938025FFBCA923A7D8B052FF0CFA60.taxon	biology_ecology	Biology. The species was collected between 10 – 1575 m elevation, in tropical dry forest, rainforest, gallery forest, gardens, and littoral rainforest. Nests were located in rotten logs, in soil, under stones, under root mats, and in dead twigs above the ground. Workers were collected from sifted litter and on tree bark.	en	Salata, Sebastian, Fisher, Brian L. (2020): The ant genus Pheidole Westwood, 1839 (Hymenoptera: Formicidae) in Madagascar-taxonomic revision of the bessonii species-group. Zootaxa 4843 (1): 1-64, DOI: 10.11646/zootaxa.4843.1.1
03938025FFBCA923A7D8B052FF0CFA60.taxon	discussion	Comments. Pheidole tenebrovulgaris is a common species widespread across the entire island. Its majors are most similar to majors of sympatric P. bessonii, P. atsirakambiaty, and P. voreios, and can be separated based on presence of dense and thick costulae with smooth interspaces on medial frons, and occipital lobes with rugae that never arch posterolaterally and most often not weaken posteriorly. Minors can be separated based on the combination of the following characters: lack of neck and distinct nuchal collar, entirely or predominantly punctate head, scape exceeding the posterior head margin by two-fifths of its length, moderate propodeal spines with narrow base, entirely punctate mesosoma, and dark brown body.	en	Salata, Sebastian, Fisher, Brian L. (2020): The ant genus Pheidole Westwood, 1839 (Hymenoptera: Formicidae) in Madagascar-taxonomic revision of the bessonii species-group. Zootaxa 4843 (1): 1-64, DOI: 10.11646/zootaxa.4843.1.1
03938025FFBCA923A7D8B052FF0CFA60.taxon	etymology	Etymology. Latin for dark and widespread, in reference to body coloration and distribution range of the species.	en	Salata, Sebastian, Fisher, Brian L. (2020): The ant genus Pheidole Westwood, 1839 (Hymenoptera: Formicidae) in Madagascar-taxonomic revision of the bessonii species-group. Zootaxa 4843 (1): 1-64, DOI: 10.11646/zootaxa.4843.1.1
03938025FFBBA921A7D8B3E6FA9EFEBC.taxon	description	Figs 24 A – F, 27 P, 28 P	en	Salata, Sebastian, Fisher, Brian L. (2020): The ant genus Pheidole Westwood, 1839 (Hymenoptera: Formicidae) in Madagascar-taxonomic revision of the bessonii species-group. Zootaxa 4843 (1): 1-64, DOI: 10.11646/zootaxa.4843.1.1
03938025FFBBA921A7D8B3E6FA9EFEBC.taxon	materials_examined	HOLOTYPE: 1 s., Madagascar, Toliara, Anosy Region, Anosyenne Mts, 31.2 km NW Manantenina, - 24.13894 47.06804, 1125 m, 26 - Feb- 2015, rainforest, ex root mat, B. L. Fisher et al. leg. BLF 36505, CASENT 0704257 (CASC). PARATYPES: 1 w., the same data as holotype, CASENT 0923260 (CASC). Other material. Madagascar. Toliara: 4 w., 2 s., 1 m., 1 q., Anosy Region, Anosyenne Mts, 31.2 km NW Manantenina, - 24.13894 47.06804, 1125 m, B. L. Fisher et al. leg. (CASC); 2 w., 1 s., 1 m., Anosy Region, Anosyenne Mts, 31.2 km NW Manantenina, - 24.13632 47.05485, 1315 m, B. L. Fisher et al. leg. (CASC). Geographic range. Madagascar, Toliara, Anosyenne Mts.	en	Salata, Sebastian, Fisher, Brian L. (2020): The ant genus Pheidole Westwood, 1839 (Hymenoptera: Formicidae) in Madagascar-taxonomic revision of the bessonii species-group. Zootaxa 4843 (1): 1-64, DOI: 10.11646/zootaxa.4843.1.1
03938025FFBBA921A7D8B3E6FA9EFEBC.taxon	diagnosis	Diagnosis. Major workers. Head in full-face view sub-rectangular, not widening posteriorly, with lateral margins slightly convex; side of head with dense, long, erect pilosity; medial frons with moderately sparse and thick rugae; interspaces between rugae smooth; frons laterally with dense rugae; interspaces between rugae reticulate; occipital lobes mostly smooth, posteriorly with sparse and weakly developed rugae; scape, when laid back, slightly exceeding the midlength of head; inner hypostomal tooth distinct, moderate and bulge-like, with top arching posteroventrally; outer hypostomal tooth bulge-like, high, and wide, with top arching posteroventrally; median tooth absent; promesonotum short, angular, and moderately low; posterior mesonotum smoothly descending towards propodeum; mesonotal process very distinct and tubercle-like; mesosoma rugoreticulate; promesonotal dorsum with reduced rugoreticulae and few additional transverse and thick rugae; gaster indistinctly shagreened at base of first tergite; body brown. Minor workers. Occiput slightly elongated; nuchal collar indistinct; head sculpture mostly smooth; frons with weakly developed and sparse puncta; scape, when laid back, exceeding the posterior head margin by three-fifths of its length; promesonotum low, moderately long, and arched; promesonotal groove distinct; metanotal groove distinct; propodeal spines small and with wide base; mesosoma mostly smooth; pronotum, mesonotum, and propodeum with weakly developed and sparse puncta; body brown.	en	Salata, Sebastian, Fisher, Brian L. (2020): The ant genus Pheidole Westwood, 1839 (Hymenoptera: Formicidae) in Madagascar-taxonomic revision of the bessonii species-group. Zootaxa 4843 (1): 1-64, DOI: 10.11646/zootaxa.4843.1.1
03938025FFBBA921A7D8B3E6FA9EFEBC.taxon	description	Description. Major workers. Measurements (n = 4): HL: 2.08 – 2.16 (2.12); HW: 2.09 – 2.14 (2.11); SL: 1.19 – 1.24 (1.21); EL: 0.21 – 0.25 (0.23); WL: 1.82 – 1.91 (1.86); PSL: 0.33 – 0.37 (0.35); MTL: 1.38 – 1.41 (1.4); PNW: 0.81 – 0.97 (0.86); PTW: 0.26 – 0.29 (0.27); PPW: 0.69 – 0.76 (0.72); CI: 99.0 – 103.3 (100.4); SI: 56.5 – 59.0 (57.5); PSLI: 15.6 – 17.5 (16.6); PPI: 35.6 – 38.5 (37.3); PNI: 38.8 – 45.4 (40.8); MTI: 65.8 – 67.3 (66.5). Head. In full-face view sub-rectangular, not widening posteriorly, with lateral sides slightly convex (Fig. 24 B). In lateral view sub-oval, not depressed posteriorly; ventral and dorsal margins convex; inner hypostomal tooth visible. Side of head with dense, long, and erect pilosity; head dorsum with relatively dense, long, decumbent to erect pilosity. Medial frons with moderately sparse and thick rugae; interspaces between rugae smooth. Frons laterally with dense rugae; interspaces between rugae reticulate. Occipital lobes mostly smooth; posteriorly with sparse and weakly developed rugae. Gena with dense and thick costulae; interspaces between costulae distinctly rugulate. Sides posterolateral from eyes with dense and thinner network of rugoreticulae. Center of clypeus shiny and smooth, lateral sides with distinct rugulae; median notch present, moderately wide and shallow; median longitudinal carina absent; lateral longitudinal carinae present. Scape, when laid back, slightly exceeding the midlength of head; pilosity appressed and subdecumbent (Fig. 24 B, D). Inner hypostomal tooth distinct, moderate and bulge-like, with top arching posteroventrally; outer hypostomal tooth bulge-like, high, and wide, with top arching posteroventrally; median tooth absent (Fig. 27 P). Mesosoma. In lateral view, promesonotum short, angular, and moderately low; posterior mesonotum smoothly descending towards propodeum; mesonotal process very distinct and tubercle-like; promesonotal groove absent; metanotal groove indistinct; propodeal spines long with narrow base and acute top; humeral tubercle weakly produced (Fig. 24 D). Surface shiny and rugoreticulate; promesonotal dorsum with reduced rugoreticulae and few additional transverse, thick rugae. Pilosity moderately dense, moderately long, and erect (Fig. 24 D, F). Gaster. Shiny and indistinctly shagreened at base of first tergite; pilosity moderately sparse, long, and erect (Fig. 24 D, F). Color. Brown, legs and antennae slightly brighter (Fig. 24 D, F). Description. Minor workers. Measurements (n = 5): HL: 0.79 – 0.95 (0.87); HW: 0.57 – 0.67 (0.62); SL: 1.11 – 1.19 (1.15); EL: 0.15 – 0.17 (0.16); WL: 1.13 – 1.32 (1.2); PSL: 0.1 – 0.14 (0.12); MTL: 0.95 – 1.09 (1.04); PNW: 0.41 – 0.51 (0.46); PTW: 0.09 – 0.13 (0.11); PPW: 0.17 – 0.22 (0.19); CI: 132.0 – 143.4 (139.3); SI: 178.8 – 193.7 (185.7); PSLI: 12.8 – 15.5 (14.2); PPI: 50.6 – 70.1 (57.4); PNI: 70.7 – 77.0 (73.5); MTI: 164.0 – 177.1 (167.3). Head. In full-face view oval, posterior region slightly elongated; nuchal collar weakly developed (Fig. 24 A). Pilosity sparse, moderately long and subdecumbent. Sculpture shiny and mostly smooth; frons with weakly developed and sparse puncta. Clypeus with median longitudinal carina absent; two lateral longitudinal carinae absent. Scape, when laid back, exceeding the posterior head margin by three-fifths of its length; pilosity dense, subdecumbent to erect (Fig. 24 A, C). Mesosoma. In lateral view, promesonotum low, moderately long, and arched; promesonotal groove distinct; metanotal groove distinct; propodeal spines small and triangular, with wide base (Fig. 24 C). Sculpture mostly smooth; pronotum, mesonotum, and propodeum with weakly developed and sparse puncta. Pilosity very sparse, short, and erect (Fig. 24 C, E). Gaster. With sparse and erect pilosity (Fig. 24 C, E). Color. Bright brown (Fig. 24 C, E).	en	Salata, Sebastian, Fisher, Brian L. (2020): The ant genus Pheidole Westwood, 1839 (Hymenoptera: Formicidae) in Madagascar-taxonomic revision of the bessonii species-group. Zootaxa 4843 (1): 1-64, DOI: 10.11646/zootaxa.4843.1.1
03938025FFBBA921A7D8B3E6FA9EFEBC.taxon	biology_ecology	Biology. The species was collected between 1125 – 1315 m elevation, in rainforest. Nests were located in rotten logs and root mats.	en	Salata, Sebastian, Fisher, Brian L. (2020): The ant genus Pheidole Westwood, 1839 (Hymenoptera: Formicidae) in Madagascar-taxonomic revision of the bessonii species-group. Zootaxa 4843 (1): 1-64, DOI: 10.11646/zootaxa.4843.1.1
03938025FFBBA921A7D8B3E6FA9EFEBC.taxon	discussion	Comments. Pheidole uranus is known only from the Anosyenne Mts. in Toliara. Its major workers are most similar to two species known from northern parts of the island: P. madecassa and sympatric P. maro. Majors of P. uranus can be easily separated based on presence of dense rugae with reticulate interspaces on lateral frons, sparse and weakly developed rugae on posterior occipital lobes, dense pilosity on side of head, and darker body coloration. Majors of P. madecassa and P. maro have lateral frons with weaker sculpture, posterior occipital lobes with reduced or absent sculpture, sparser pilosity on side of head, and orange to brownish orange body. Minors of P. uranus are most similar to P. ovalinoda and can be separated based on more elongated posterior portion of head, and mostly smooth mesosoma. Minors of P. ovalinoda have less elongated posterior portion of head, and partially sculptured mesosoma.	en	Salata, Sebastian, Fisher, Brian L. (2020): The ant genus Pheidole Westwood, 1839 (Hymenoptera: Formicidae) in Madagascar-taxonomic revision of the bessonii species-group. Zootaxa 4843 (1): 1-64, DOI: 10.11646/zootaxa.4843.1.1
03938025FFBBA921A7D8B3E6FA9EFEBC.taxon	etymology	Etymology. Named after the Greek god of the sky, in reference to the high elevation of the type locality.	en	Salata, Sebastian, Fisher, Brian L. (2020): The ant genus Pheidole Westwood, 1839 (Hymenoptera: Formicidae) in Madagascar-taxonomic revision of the bessonii species-group. Zootaxa 4843 (1): 1-64, DOI: 10.11646/zootaxa.4843.1.1
03938025FFB6A92CA7D8B69FFC2AFC34.taxon	description	Figs 25 A – F, 27 Q, 28 Q	en	Salata, Sebastian, Fisher, Brian L. (2020): The ant genus Pheidole Westwood, 1839 (Hymenoptera: Formicidae) in Madagascar-taxonomic revision of the bessonii species-group. Zootaxa 4843 (1): 1-64, DOI: 10.11646/zootaxa.4843.1.1
03938025FFB6A92CA7D8B69FFC2AFC34.taxon	materials_examined	HOLOTYPE: 1 s., Madagascar, Antsiranana, Réserve Spéciale de l’Ankarana, 22.9 km 224 ° SW Anivorano Nord, - 12.90889 49.10983, 80 m, 10 - Feb- 2001, tropical dry forest, ex rotten log, Fisher et al. leg. BLF 02932, CASENT 0406635 (CASC). PARATYPES: 6 w., 2 s., the same data as holotype, CASENT 0406634, CASENT 0406636, CASENT 0872257, CASENT 0406633 (CASC, MHNG, PBZT). Other material. Madagascar, Antsiranana: 1 w., 1 s., Forêt d’ Andavakoera, 21.4 km 75 ° ENE Ambilobe; 4.6 km 356 ° N Betsiaka, - 13.11833 49.23, 425 m, Fisher et al. leg. (CASC); 1 w., Forêt d’Ampombofofo, - 12.09949 49.33874, 25 m, Fisher et al. leg. (CASC); 1 w., Réserve Spéciale de l’Ankarana, 13.6 km 192 ° SSW Anivorano Nord, - 12.86361 49.22583, 210 m, Fisher et al. leg. (CASC); 2 w., Réserve Spéciale de l’Ankarana, 13.6 km 192 ° SSW Anivorano Nord, - 12.86361 49.22583, 210 m, Fisher et al. leg. (CASC); 3 w., Réserve Spéciale de l’Ankarana, 13.6 km 192 ° SSW Anivorano Nord, - 12.86361 49.22583, 210 m, Fisher et al. leg. (CASC); 1 w., Réserve Spéciale de l’Ankarana, 13.6 km 192 ° SSW Anivorano Nord, - 12.86361 49.22583, 210 m, Fisher et al. leg. (CASC); 2 w., Réserve Spéciale de l’Ankarana, 22.9 km 224 ° SW Anivorano Nord, - 12.90889 49.10983, 80 m, Fisher et al. leg. (CASC); 1 w., Réserve Spéciale de l’Ankarana, 22.9 km 224 ° SW Anivorano Nord, - 12.90889 49.10983, 80 m, Fisher et al. leg. (CASC); 6 w., 4 s., Réserve Spéciale de l’Ankarana, 22.9 km 224 ° SW Anivorano Nord, - 12.90889 49.10983, 80 m, Fisher et al. leg. (CASC); 1 w., Réserve Spéciale de l’Ankarana, 22.9 km 224 ° SW Anivorano Nord, - 12.90889 49.10983, 80 m, Fisher et al. leg. (CASC); 1 w., Sahamalaza Peninsula, Forêt d’Anabohazo, 21.6 km 247 ° WSW Maromandia, - 14.30889 47.91433, 120 m, Fisher et al. leg. (CASC); 2 w., Sahamalaza Peninsula, Forêt d’Anabohazo, 21.6 km 247 ° WSW Maromandia, - 14.30889 47.91433, 120 m, Fisher et al. leg. (CASC) Geographic range. Madagascar, Antsiranana, Réserve Spéciale de l’Ankarana, Forêt d’ Andavakoera, Forêt d’Ampombofofo, and Forêt d’Anabohazo.	en	Salata, Sebastian, Fisher, Brian L. (2020): The ant genus Pheidole Westwood, 1839 (Hymenoptera: Formicidae) in Madagascar-taxonomic revision of the bessonii species-group. Zootaxa 4843 (1): 1-64, DOI: 10.11646/zootaxa.4843.1.1
03938025FFB6A92CA7D8B69FFC2AFC34.taxon	diagnosis	Diagnosis. Major workers. Head in full-face view sub-rectangular, not widening posteriorly, with lateral margins relatively straight; side of head with sparse, long, suberect to erect pilosity; medial frons with moderately thick and dense costulae; interspaces between costulae distinctly punctate; frons laterally with sparser costulae; interspaces between costulae distinctly punctate; occipital lobes with thin and sparse rugulae that arch posterolaterally; interspaces between rugulae punctate; sculpture weakening posteriorly; scape, when laid back, exceeding the midlength of head by two-fifths of its length; inner hypostomal tooth very small and bulge-like; outer hypostomal tooth lobe-like, high, and wide, with top arching posteroventrally; median tooth present and indistinct; promesonotum short, angular, and moderately low; posterior mesonotum moderately steep; mesonotal process distinct and tubercle-like; mesosoma punctate; promesonotum with additional thin, sparse, and transverse rugae; katepisternum and propodeum with additional sparse rugae; gaster shagreened; body brown. Minor workers. Occiput elongated; nuchal collar distinct; head sculpture mostly smooth; anterior frons with sparse puncta; scape, when laid back, exceeding the posterior head margin by half of its length; promesonotum low, long, and arched; promesonotal groove present; metanotal groove distinct; propodeal spines small and narrow; mesosoma microreticulate; promesonotum with sparser sculpture; promesonotal dorsum mostly to entirely smooth; body brown.	en	Salata, Sebastian, Fisher, Brian L. (2020): The ant genus Pheidole Westwood, 1839 (Hymenoptera: Formicidae) in Madagascar-taxonomic revision of the bessonii species-group. Zootaxa 4843 (1): 1-64, DOI: 10.11646/zootaxa.4843.1.1
03938025FFB6A92CA7D8B69FFC2AFC34.taxon	description	Description. Major workers. Measurements (n = 9): HL: 1.65 – 1.81 (1.74); HW: 1.5 – 1.68 (1.6); SL: 1.15 – 1.24 (1.19); EL: 0.23 – 0.27 (0.24); WL: 1.62 – 1.74 (1.68); PSL: 0.21 – 0.31 (0.27); MTL: 1.34 – 1.42 (1.37); PNW: 0.61 – 0.66 (0.63); PTW: 0.19 – 0.27 (0.22); PPW: 0.49 – 0.63 (0.57); CI: 106.4 – 112.1 (109.2); SI: 69.7 – 79.0 (74.9); PSLI: 13.0 – 17.4 (15.4); PPI: 34.4 – 42.9 (38.0); PNI: 37.4 – 41.3 (39.3); MTI: 80.9 – 93.7 (86.0). Head. In full-face view sub-rectangular, not widening posteriorly, with lateral sides relatively straight (Fig. 25 B). In lateral view sub-oval, not depressed posteriorly; ventral and dorsal margins convex; inner hypostomal tooth not visible. Side of head with sparse, long, suberect to erect pilosity; head dorsum with relatively dense, long, decumbent to erect pilosity. Medial frons with moderately thick and dense costulae; interspaces between costulae distinctly punctate. Frons laterally with sparser costulae; interspaces between costulae distinctly punctate. Occipital lobes with thin and sparse rugulae that arch posterolaterally; interspaces between rugulae punctate; sculpture weakening posteriorly. Gena with dense and thick costulae; interspaces between costulae distinctly punctate. Sides posterolateral from eyes with dense but thin rugoreticulae. Center of clypeus shiny and mostly 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, exceeding the midlength of head by two-fifths of its length; pilosity subdecumbent to erect (Fig. 25 B, D). Inner hypostomal tooth very small and bulge-like; outer hypostomal tooth lobe-like, high, and wide, with top arching posteroventrally; median tooth present and weakly developed (Fig. 27 Q). 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 present; propodeal spines moderately long with wide base and acute top; humeral tubercle weakly produced (Fig. 25 D). Surface shiny and punctate; promesonotum with additional, thin, sparse, and transverse rugae; katepisternum and propodeum with additional sparse rugae. Pilosity moderately dense, long, and erect (Fig. 25 D, F). Petiole. In rear view node dorsoventrally slightly convex (Fig. 25 D, F). Gaster. Dull and shagreened; pilosity moderately sparse, long, and erect (Fig. 25 D, F). Color. Brown, antennae and legs yellowish brown (Fig. 25 D, F). Description. Minor workers. Measurements (n = 8): HL: 0.77 – 0.85 (0.81); HW: 0.53 – 0.59 (0.56); SL: 1.09 – 1.2 (1.15); EL: 0.16 – 0.18 (0.17); WL: 1.08 – 1.21 (1.15); PSL: 0.11 – 0.13 (0.12); MTL: 1.01 – 1.1 (1.05); PNW: 0.39 – 0.44 (0.42); PTW: 0.09 – 0.11 (0.1); PPW: 0.14 – 0.18 (0.17); CI: 142.6 – 148.3 (145.3); SI: 202.6 – 209.9 (206.3); PSLI: 12.8 – 16.6 (15.4); PPI: 52.9 – 66.4 (59.4); PNI: 71.2 – 77.3 (74.6); MTI: 184.7 – 194.9 (188.8). Head. In full-face view oval, posterior region elongated; nuchal collar distinct (Fig. 25 A). Pilosity relatively sparse, moderately long, subdecumbent to erect. Sculpture shiny and mostly smooth; anterior frons with sparse puncta. Clypeus with median longitudinal carina absent; two lateral longitudinal carinae absent. Scape, when laid back, exceeding the posterior head margin by half of its length; pilosity dense, subdecumbent to erect (Fig. 25 A, C). Mesosoma. In lateral view, promesonotum low, long, and arched; promesonotal groove present; metanotal groove distinct; propodeal spines small and triangular, narrow (Fig. 25 C). Sculpture microreticulate; promesonotum with sparser sculpture and mostly to entirely smooth dorsum. Pilosity very sparse, long, and erect (Fig. 25 C, E). Gaster. With sparse and erect pilosity (Fig. 25 C, E). Color. Brown, legs and antennae yellowish (Fig. 25 C, E).	en	Salata, Sebastian, Fisher, Brian L. (2020): The ant genus Pheidole Westwood, 1839 (Hymenoptera: Formicidae) in Madagascar-taxonomic revision of the bessonii species-group. Zootaxa 4843 (1): 1-64, DOI: 10.11646/zootaxa.4843.1.1
03938025FFB6A92CA7D8B69FFC2AFC34.taxon	biology_ecology	Biology. The species was collected between 25 – 487 m elevation, in tropical dry forest and rainforest. Nests were located in rotten logs and rotten sticks on the ground. Workers were collected from sifted litter and on low vegetation.	en	Salata, Sebastian, Fisher, Brian L. (2020): The ant genus Pheidole Westwood, 1839 (Hymenoptera: Formicidae) in Madagascar-taxonomic revision of the bessonii species-group. Zootaxa 4843 (1): 1-64, DOI: 10.11646/zootaxa.4843.1.1
03938025FFB6A92CA7D8B69FFC2AFC34.taxon	discussion	Comments. Pheidole voreios is known from the northern part of the island and is most similar to the widespread and parapatric P. bessonii. Majors of P. voreios can be distinguished based on rugulate occipital lobes, and frons laterally with sparse costulae and distinctly punctate interspaces. In contrast, P. bessonii has punctate occipital lobes and frons laterally rugocostulate. Minors can be separated based on more elongated posterior region of head, microreticulate mesosoma with sculpture weakening on promesonotum, and small and narrow propodeal spines. In contrast, P. bessonii has less elongated posterior region of head, mostly smooth promesonotum, and very small propodeal spines.	en	Salata, Sebastian, Fisher, Brian L. (2020): The ant genus Pheidole Westwood, 1839 (Hymenoptera: Formicidae) in Madagascar-taxonomic revision of the bessonii species-group. Zootaxa 4843 (1): 1-64, DOI: 10.11646/zootaxa.4843.1.1
03938025FFB6A92CA7D8B69FFC2AFC34.taxon	etymology	Etymology. Greek for northern, in reference to the range of the species.	en	Salata, Sebastian, Fisher, Brian L. (2020): The ant genus Pheidole Westwood, 1839 (Hymenoptera: Formicidae) in Madagascar-taxonomic revision of the bessonii species-group. Zootaxa 4843 (1): 1-64, DOI: 10.11646/zootaxa.4843.1.1
03938025FFB4A928A7D8B21DFB54FD30.taxon	description	Figs 26 A – F, 27 R, 28 R	en	Salata, Sebastian, Fisher, Brian L. (2020): The ant genus Pheidole Westwood, 1839 (Hymenoptera: Formicidae) in Madagascar-taxonomic revision of the bessonii species-group. Zootaxa 4843 (1): 1-64, DOI: 10.11646/zootaxa.4843.1.1
03938025FFB4A928A7D8B21DFB54FD30.taxon	materials_examined	HOLOTYPE: 1 s., Madagascar, Antsiranana, Sava Region: Parc National de Marojejy, Manantenina River, 28.0 km 24.8 ° NE Andapa, - 14.43461 49.76074, 780 m, 13 - Feb- 2018, rainforest, ex rotten log, B. L. Fisher et al. leg. BLF 41182, CASENT 0923249 (CASC). PARATYPES: 3 w., 1 m.: the same data as holotype, CASENT 0807266, CASENT 0807267, CASENT 0807268 (CASC, MHNG, PBZT). Other material. Madagascar. Antsiranana: 4 w., Parc National de Marojejy, Manantenina River, 28.0 km 38 ° NE Andapa, 8.2 km 333 ° NNW Manantenina, - 14.43667 49.775, 450 m, B. L. Fisher et al. leg. (CASC); 6 w., SAVA Region, District of Sambava, Marojejy National Park, 5 km W of Manantenina village, 1 st Camp site (Mantella), - 14.43817 49.774, 487 m, M. Rin’Ha leg. (CASC); 1 w., Sava Region: Parc National de Marojejy, near Manantenina River, - 14.43677 49.77541, 475 m, B. L. Fisher et al. leg. (CASC). Geographic range. Madagascar, Antsiranana, Sava Region.	en	Salata, Sebastian, Fisher, Brian L. (2020): The ant genus Pheidole Westwood, 1839 (Hymenoptera: Formicidae) in Madagascar-taxonomic revision of the bessonii species-group. Zootaxa 4843 (1): 1-64, DOI: 10.11646/zootaxa.4843.1.1
03938025FFB4A928A7D8B21DFB54FD30.taxon	diagnosis	Diagnosis. Major workers. Head in full-face view sub-rectangular, widening posteriorly, with lateral margins relatively straight and very deep posteromedian concavity; side of head with moderately sparse, short, appressed to decumbent pilosity; anteromedial part of frons with moderately thick and dense costulae; interspaces between costulae smooth; posteromedial frons with thinner rugocostulae that arch posterolaterally; interspaces between rugocostulae rugulate; frons laterally with denser and thicker rugae; interspaces between rugae with dense and thin rugulae; occipital lobes with thin rugocostulae that weaken posteriorly and arch posterolaterally, interspaces between rugocostulae mostly smooth; scape, when laid back, exceeding the midlength of head by three-fifths of its length; inner hypostomal tooth distinct, low, wide, closely spaced and bulge-like; outer hypostomal tooth lobe-like and higher than inner tooth, with top directed posteriorly; inner and outer hypostomal tooth closely spaced and not connected by concavity; promesonotum short, angular and low; posterior mesonotum smoothly declining towards propodeum; mesonotal process distinct and tubercle-like; mesosoma with dense and thin rugopuncta; pronotum with additional thin, moderately dense, and transverse rugae; anepisternum, katepisternum, and lateral sides of propodeum with rugae that are more vertical; gaster shagreened; body brownish black. Minor workers. Occiput elongated, terminated by distinct nuchal collar; head sculpture mostly smooth with few rugae on gena and transverse rugae on neck; scape, when laid back, exceeding the posterior head margin by half of its length; promesonotum very low, long, and slightly arched; promesonotal groove present; metanotal groove indistinct; propodeal spines long and thin; promesonotal and propodeal dorsum mostly smooth with sparse and transverse rugae, and lateral sides of propodeum, anepisternum, and katepisternum with dense and thin rugoreticulae; body dark brown.	en	Salata, Sebastian, Fisher, Brian L. (2020): The ant genus Pheidole Westwood, 1839 (Hymenoptera: Formicidae) in Madagascar-taxonomic revision of the bessonii species-group. Zootaxa 4843 (1): 1-64, DOI: 10.11646/zootaxa.4843.1.1
03938025FFB4A928A7D8B21DFB54FD30.taxon	description	Description. Major workers. Measurements (n = 1): HL: 2.44; HW: 2.21; SL: 1.8; EL: 0.246; WL: 2.19; PSL: 0.57; MTL: 2.22; PNW: 0.8; PTW: 0.28; PPW: 0.69; CI: 110.5; SI: 81.4; PSLI: 23.4; PPI: 40.6; PNI: 36.0; MTI: 100.7. Head. In full-face view sub-rectangular, side relatively straight, posteromedian concavity very deep (Fig. 26 B). In lateral view sub-oval, not depressed posteriorly, ventral and dorsal margin convex; inner hypostomal tooth visible. Side of head with moderately sparse, short, appressed to decumbent pilosity; head dorsum with relatively dense, long, decumbent to erect pilosity. Anteromedial part of frons with moderately thick and dense costulae; interspaces between costulae smooth; posteromedial frons with rugocostulae that are thinner and arch posterolaterally; interspaces between rugocostulae rugulate; frons laterally with dense and thick rugae; interspaces between rugae rugulate. Occipital lobes with thin rugocostulae that weaken posteriorly and arch posterolaterally; interspaces between rugocostulae mostly smooth. Gena with dense and thick costulae; interspaces between costulae distinctly rugulate. Sides posterolateral from eyes with dense and thin rugae; interspaces between rugae distinctly rugulate. Center of clypeus shiny and indistinctly rugulate, lateral sides with weakly developed rugulae; median notch present, moderately wide and shallow; median longitudinal carina present; lateral longitudinal carinae present. Scape, when laid back, exceeding midlength of head by three-fifths its length; pilosity subdecumbent to erect (Fig. 26 B, D). Inner hypostomal tooth distinct, low, wide, closely spaced and bulge-like; outer hypostomal tooth lobe-like, higher than inner tooth, top directed posteriorly; inner and outer hypostomal tooth closely spaced and not connected by concavity (Fig. 27 R). Mesosoma. In lateral view, promesonotum short, angular and low; posterior mesonotum smoothly declining towards propodeum; mesonotal process distinct and tubercle-like; promesonotal groove absent; metanotal groove present; propodeal spines long, narrow and with acute top; humeral tubercle weakly produced (Fig. 26 D). Surface shiny and with dense and thin rugopuncta; pronotum with additional thin, moderately dense, and transverse rugae; anepisternum, katepisternum, and lateral sides of propodeum with rugae that are more vertical. Pilosity moderately dense, short, and erect (Fig. 26 D, F). Petiole. Shagreened (Fig. 26 D, F). Postpetiole. Shagreened; dorsum with slightly sparser sculpture (Fig. 26 D, F). Gaster. Dull and shagreened; pilosity moderately sparse, long, and erect (Fig. 26 D, F). Color. Brownish black, antennae and legs dark brown, mandibles reddish brown (Fig. 26 D, F). Description. Minor workers. Measurements (n = 10): HL: 1.17 – 1.44 (1.29); HW: 0.6 – 0.75 (0.66); SL: 1.83 – 2.13 (1.95); EL: 0.14 – 0.22 (0.18); WL: 1.53 – 1.91 (1.69); PSL: 0.34 – 0.45 (0.38); MTL: 1.85 – 2.37 (2.03); PNW: 0.45 – 0.58 (0.52); PTW: 0.11 – 0.16 (0.13); PPW: 0.19 – 0.24 (0.22); CI: 188.4 – 200.0 (193.8); SI: 282.2 – 302.6 (293.2); PSLI: 26.7 – 31.3 (29.2); PPI: 49.6 – 70.6 (58.1); PNI: 74.5 – 79.5 (77.5); MTI: 290.9 – 314.9 (305.2). Head. In full-face view oval, posterior region elongated into long and narrow neck terminated by distinct nuchal collar (Fig. 26 A). Pilosity relatively sparse, moderately long, and subdecumbent to erect. Sculpture shiny and mostly smooth with few rugae on gena and transverse rugae on neck; antennal sockets with few weakly developed rugulae that curve posterolaterally; interspaces between rugulae smooth. Clypeus with median longitudinal carina absent; two lateral longitudinal carinae absent. Scape, when laid back, exceeding the posterior head margin, neck not included, by half of its length; pilosity dense and subdecumbent to erect (Fig. 26 A, C). Mesosoma. In lateral view, promesonotum very low, long, and slightly arched; promesonotal groove present; metanotal groove in distinct; propodeal spines long, thin, and triangular (Fig. 26 C). Promesonotal and propodeal dorsum mostly smooth with sparse and transverse rugae; lateral sides of propodeum, anepisternum, and katepisternum, with dense and thin rugoreticulae. Pilosity very sparse, short, and erect (Fig. 26 C, E). Petiole. Peduncle with ventral face straight (Fig. 26 C, E). Gaster. With sparse and erect pilosity (Fig. 26 C, E). Color. Dark brown, legs and antennae brighter (Fig. 26 C, E).	en	Salata, Sebastian, Fisher, Brian L. (2020): The ant genus Pheidole Westwood, 1839 (Hymenoptera: Formicidae) in Madagascar-taxonomic revision of the bessonii species-group. Zootaxa 4843 (1): 1-64, DOI: 10.11646/zootaxa.4843.1.1
03938025FFB4A928A7D8B21DFB54FD30.taxon	biology_ecology	Biology. The species was collected between 450 – 780 m elevation, in low elevation rainforest and rainforest. Nests were located in rotten logs, and workers were collected from sifted litter and on the ground.	en	Salata, Sebastian, Fisher, Brian L. (2020): The ant genus Pheidole Westwood, 1839 (Hymenoptera: Formicidae) in Madagascar-taxonomic revision of the bessonii species-group. Zootaxa 4843 (1): 1-64, DOI: 10.11646/zootaxa.4843.1.1
03938025FFB4A928A7D8B21DFB54FD30.taxon	discussion	Comments. Majors of P. zirafy are most similar to parapatric P. flavodepressa and differ in blackish brown body, presence of moderately sparse, short, and appressed to decumbent pilosity on side of head, occipital lobes with thin rugae that weaken posteriorly and arch posterolaterally, and mostly smooth interspaces between rugae. In contrast, P. flavodepressa has orange body coloration, side of head with moderately dense, long, subdecumbent to suberect pilosity, and occipital lobes with thick and dense rugae and distinctly rugulate interspaces between rugae. Minors of P. zirafy are most similar to sympatric P. grallatrix and differ in presence of dense and thin rugoreticulae on lateral sides of propodeum, anepisternum, and katepisternum. Minors of P. grallatrix have entirely smooth mesosoma.	en	Salata, Sebastian, Fisher, Brian L. (2020): The ant genus Pheidole Westwood, 1839 (Hymenoptera: Formicidae) in Madagascar-taxonomic revision of the bessonii species-group. Zootaxa 4843 (1): 1-64, DOI: 10.11646/zootaxa.4843.1.1
03938025FFB4A928A7D8B21DFB54FD30.taxon	etymology	Etymology. Malagasy for giraffe, in reference to the very long neck of minor workers.	en	Salata, Sebastian, Fisher, Brian L. (2020): The ant genus Pheidole Westwood, 1839 (Hymenoptera: Formicidae) in Madagascar-taxonomic revision of the bessonii species-group. Zootaxa 4843 (1): 1-64, DOI: 10.11646/zootaxa.4843.1.1
