Pheidole Westwood, 1839 (Hymenoptera, Formicidae) of Madagascar – an introduction and a taxonomic revision of eleven species groups

Abstract The present study represents an introduction to the revision of Pheidole Westwood, 1839 from Madagascar. Sixteen species groups are established, of which eleven are revised below, and illustrated identification keys to species groups and species of groups revised in this monograph are presented. Two species are raised to species level: Pheidole petax Forel, 1895 stat. nov., and P. scabrata Forel, 1895 stat. nov. We also redescribe worker castes and designate lectotypes for P. annemariae Forel, 1918, P. nemoralis Forel, 1892, P. petax Forel, 1895, P. ensifera Forel, 1897, P. longispinosa Forel, 1891, and P. scabrata Forel, 1895. The following 46 new species are described: Pheidole aelloeasp. nov., P. alasp. nov., P. andapasp. nov., P. ankeranasp. nov., P. avaratrasp. nov., P. bemarahaensissp. nov., P. bemarivoensissp. nov., P. binarasp. nov., P. boriborasp. nov., P. brevipilosasp. nov., P. curvistriatasp. nov., P. diakritossp. nov., P. ehazoarasp. nov., P. ferrugineasp. nov., P. fisakasp. nov., P. fitaratasp. nov., P. glabrasp. nov., P. goavanasp. nov., P. lamperossp. nov., P. longipilosasp. nov., P. luteasp. nov., P. madinikasp. nov., P. mahaboensissp. nov., P. maizinasp. nov., P. makaensissp. nov., P. makirovanasp. nov., P. manantenensissp. nov., P. mantadiasp. nov., P. marieannaesp. nov., P. masoalasp. nov., P. mavesatrasp. nov., P. miramilasp. nov., P. moramanaensissp. nov., P. navoatrensissp. nov., P. ocypodeasp. nov., P. parvioculasp. nov., P. podargeasp. nov., P. praegrandissp. nov., P. ranohirensissp. nov., P. rugocephalasp. nov., P. rugofitaratasp. nov., P. typhlossp. nov., P. vatovavensissp. nov., P. voasarasp. nov., P. vohemarensissp. nov., and P. zavamanirasp. nov. At present, there are 69 valid species and subspecies of Pheidole known from Madagascar, but this number is expected to increase significantly with upcoming taxonomic revisions of the species groups not revised in this study.


Introduction
The genus Pheidole Westwood, 1839 is one of the most diverse ant genera and contains 1047 valid species and 134 valid subspecies (Bolton 2019). While distribution of the genus is worldwide, its species richness varies with zoogeographic region. The Neotropical region is the most species-rich; the current number of described taxa constitutes nearly 50 % of all known Pheidole species, with far fewer species numbers in other regions. The fauna of this genus was exhaustively studied by Wilson (2003), and later amended and updated by Longino (2009Longino ( , 2019. The taxonomy of the genus has been revised for the Nearctic (Wilson 2003), Afrotropic (Fischer et al. 2012), Indomalayan (Eguchi 2001(Eguchi , 2006(Eguchi , 2008Eguchi and Bui 2005;Eguchi et al. 2016), Australasian , and Oceanian (Sarnat 2008;Fischer et al. 2016) regions. The taxonomy of the Malagasy region has been greatly improved in recent years by Fischer and Fisher (2013). Covering Comoros, Juan de Nova Island, Mauritius, Mayotte, Réunion, and Seychelles, their survey confirmed the presence of thirteen species on these islands, of which seven were newly described.
Very little is known about the Pheidole fauna of Madagascar. Most of the data describing the diversity of this genus on the island can be found in a number of short faunistic notes, with the majority of species descriptions originating in the late 1800s. There are only 18 valid Pheidole taxa described from Madagascar, of which 17 were described before the twentieth century (Forel 1891(Forel , 1892(Forel , 1895a(Forel , b, 1897Emery 1899). The last Madagascar Pheidole species was described a century ago (Forel 1918). Since the last publication of Forel (1918), almost 80 years passed without additional work on the Pheidole of Madagascar. In late 1990s  provided a complete overview of the species described from the Malagasy region and discussed their known distributions. He also listed Pheidole within the four most species-rich and abundant ant genera of Madagascar. Later, Fisher and Peeters (2019) estimated that the number of undescribed endemic species exceeded 100.
Thanks to comprehensive inventories conducted across the island by Fisher and members of the Malagasy Arthropod team at the Madagascar Biodiversity Centre in Madagascar, the Pheidole collection deposited at the California Academy of Sciences (CAS) now contains more than 50,000 specimens from almost 4,500 localities. Recent inventories have yielded a large number of undescribed Pheidole species, confirming assumptions presented by Fisher and Peeters (2019).
Here we present an introduction to Pheidole in Madagascar, define species groups, and provide an illustrated identification key to species groups. This monograph is a first attempt to define species groups for the Malagasy region. Our divisions are based mostly on morphological similarities within taxa. We recognize 16 species groups containing approximately 140 species. In this work we define and revise eleven groups containing 52 species, 46 of which are described as new. This publication is meant to be the first in a series aiming to revise the taxonomy of all Pheidole in Madagascar.
Based on data gathered thus far, Pheidole of Madagascar are distinct from species known from other islands in the Malagasy region. Only three species are confirmed from Madagascar as well as surrounding islands. Two are invasive worldwide: P. megacephala (Fabricius) and P. indica Mayr. The third, P. megatron Fischer & Fisher, is a member of the megacephala group described from Comoros. Here we confirm its presence in urban areas of Antsiranana prefecture. Three other species absent from Madagascar but found on nearby islands can be assigned to Malagasy species groups. Pheidole ragnax Fischer & Fisher is a member of the P. bessonii group and should be assigned to the P. grallatrix complex. Pheidole jonas Forel is similar to members of the P. bemarivoensis complex of the P. nemoralis group. Finally, Pheidole vulcan Fischer & Fisher represents a distinct member of the P. petax group and most likely is closely related to species grouped within P. ankerana complex. Pheidole braueri Forel, P. dodo Fischer & Fisher, and P. komori Fischer & Fisher bear a distinct set of characters unknown from Madagascar. Pheidole loki Fischer & Fisher cannot be assigned to any known species group until its major workers desctiption. However, based on the morphology of minor workers, it appears most similar to members of the P. makaensis group.
Among the 16 species groups known from Madagascar defined here, members of only three are reported from outside the Malagasy region. The P. fervens group is native to Indoaustralia (Sarnat et al 2015), while the P. megacephala group is considered native to the Afrotropics and Malagasy region (Sarnat et al 2015). Members of the P. bessonii group are morphologically reminiscent of the P. longipes group from Indo-Australia. In conclusion, the vast majority of groups and species are endemic to Madagascar, one group is also native to the Afrotropical Region, and two groups are of Indo-Australian origin.

Materials and methods
The majority of the material was collected by Brian L. Fisher and members of the Madagascar Biodiversity Centre from across Madagascar between 1991 and 2018. The study was supported with material deposited in the Museum d'Historie Naturelle, Geneva, Switzerland.
Repositories. Collections are referred to by the following acronyms:
All observations and measurements were taken using a pin-holding stage, permitting rotations around the X, Y, and Z axes at magnifications from 32× to 100× with a Leica MZ12.5 microscope and an orthogonal crosshair micrometre, at an accuracy of 0.01 mm to approximately 0.005 mm. All measurements are presented in mm units as minimum and maximum values, with the arithmetic mean in parentheses. We attach the measurement data in the Supplementary file. Photographs were taken using a JVC KY-75 or Leica DFC450 digital camera with a Leica Z16 APO microscope and Leica Application Suite software (v3.8). Unless stated otherwise, photographs were taken by Michele Esposito. Images of specimens and data of all pinned specimens examined in the present contribution are available online on AntWeb (https://www.AntWeb.org) and accessible using the unique CASENT identifying specimen code. Most measurements and indices are the same as in Longino (2009Longino ( , 2019 and based on several other revisions (Eguchi 2008;Fischer et al. 2012;Fischer and Fisher 2013;Wang et al. 2018). The general morphological terminology follows Wilson (2003) and Longino (2009Longino ( , 2019. As older taxa are often insufficiently characterized by their original describers, diagnoses are provided in the redescriptions for P. annemariae Forel, 1918, P. nemoralis Forel, 1892, P. petax Forel, 1895, P. ensifera Forel, 1897, P. longispinosa Forel, 1891, and P. scabrata Forel, 1895 to make identifications easier. Pilosity inclination degree follows that used in Wilson (1955). Appressed (0-5°) hairs run parallel or nearly parallel to the body surface. Decumbent hairs stand 10-40°, subdecumbent hair stand ~45° from the surface°, suberect hairs bend about 10°-20° from vertical, and erect hairs stand vertical or nearly vertical.
Maps were generated using tmap v2.2 package on R v3.5. R Core Team (2018). The concepts of ecotones follow those used by Yoder and Nowak (2006).

Measurements and indices
Measurements:

EL
eye length; measured along the maximum vertical diameter of eye; HL maximum distance from the midpoint of the anterior clypeal margin to the midpoint of the posterior margin of the head, measured in full-face view; in majors from midpoint of tangent between anteriormost position of clypeus to midpoint of tangent between posteriormost projection of the vertex; HW head width; measured in full-face view, at widest point of the head, directly above the eyes; MTL metatibia length; measured from the junction with femur to the junction with first tarsal segment; PNW pronotum width; maximum width of promesonotum measured in dorsal view; PPW postpetiole width; maximum width of postpetiole in dorsal view; PSL propodeal spine length; measured from the centre of the propodeal spiracle to the tip of the propodeal spine in lateral view; PTW petiole width; maximum width of petiole in dorsal view; SL scape length; maximum straight-line length of scape excluding the basal condylar bulb; WL mesosoma length (Weber's length); diagonal length of mesosoma in lateral view from the anterior point of the pronotal slope and excluding the neck, to the posteroventral margin of the propodeum. Indices: CI cephalic index: HW / HL * 100; MTI tibia index: MTL / HW * 100; SI scape index: SL / HW * 100; PNI pronotum index: PNW / HW * 100; PPI postpetiole width index: PPW / PTW * 100; PSLI propodeal spine index: PSL / HW * 100.

Synopsis of species of Madagascar examined in this study
Pheidole annemariae group Pheidole annemariae Forel, 1918 Pheidole marieannae sp. nov.

Preliminary key to Pheidole species groups of Madagascar (minor and major workers)
Note: This key is preliminary and ongoing revisions of all species groups may lead to further changes. Additionally, because of high infraspecific variability of minor workers we strongly encourage readers to use nest samples consisting of both major and minor workers while using the key. An asterisk (*) denotes groups revised in this monograph.

1
Social parasite of Pheidole sp. Major workers. Absent. Minor workers. Yellow, with smooth body sculpture, promesonotal and metanotal grooves absent, promesonotum evenly arched and big eyes ( Fig. 2A) ....... Pheidole lucida group -Nonparasitic species. Both major and minor workers present. If body colouration of minor workers is yellow and body sculpture smooth them eyes never big ( Minor & major workers. Postpetiole in profile with conspicuous ventral convexity (sometimes subtriangular in major workers) (Fig. 3A, B). Minor workers. Head smooth, scape relatively short, surpassing the posterior head margin by two-fifths of its length, promesonotum in lateral view convex (Fig. 3A). Major workers. Head oval to cordate; occipital lobes shiny and smooth or with indistinct rugulae; frons with sparse and thick longitudinal rugulae and smooth to finely rugulose interspaces; inner hypostomal teeth indistinct or absent (Figs 3B,4D) ... Pheidole megacephala group - Minor & major workers. Postpetiole in profile without conspicuous convexity, ventral process indistinct or, if present, acute and present only in major workers . Minor workers. Head at least partially foveolate or rugulose. If head entirely smooth then scape short and not surpassing the posterior head margin and promesonotum not evenly convex or scape long, surpassing the posterior head margin by at least one-third of its length and promesonotum of different shape . Major workers. Head shape different; if head oval or cordate then at least one of the following characters present: occipital lobes never smooth, inner hypostomal teeth well developed, antennal scrobes present, frons with sculpture other than sparse thick longitudinal rugae (Figs 3E,F,4C,E,F Minor & major workers. Antennal sockets deep; frontal lobes distinct and lobelike (Fig. 4A, B). Major workers. Outer hypostomal teeth absent (Fig. 4C)  Major workers. Head, in full-face view, trapezoid, widening posteriorly; head sculpture fine, occipital lobes smooth or with indistinct microsculpture; promesonotum short, low, and evenly convex (Fig. 5A,B). Minor workers. Scape, when arranged along the anteroposterior axis of the head, surpassing posterior head margin by one third or more than half of its length; promesonotum low, long, and slightly convex; petiole with long and thin peduncle (Fig. 5C,D) ....Pheidole longispinosa group* -Major workers. Head, in full-face view, rectangular; head sculpture strong rugoreticulate, sometimes weakens posteriorly; promesonotum short, angular, and relatively low to high (Fig. 5E,F). Minor workers. Scape, when arranged along the anteroposterior axis of the head, surpassing posterior head margin by one-to twofifths of its length; promesonotum high, short and convex; petiole with shorter and thicker peduncle (Fig. 5G, H)  Major workers. Head in full-face view rectangular and longer than wide; in lateral view sub-oval to sub-rectangular; occipital lobes always with arcuate and/or transverse rugae; genae at least with a smooth notch (Fig. 6A, B). Minor workers. Head foveolate with at least smooth notches on genae; scape short, reaching the posterior margin of head or surpassing it by one-to two-fifths of its length; promesonotum low, convex, short; propodeal spines small and triangular; mesosoma predominately foveolate but always with smooth notches (Fig. 6E) ....Pheidole makaensis group* -Major workers. Head in full-face view not elongated; if elongated then occipital lobes lacking arcuate or transverse rugae (Figs 6C,D,. Minor workers. Head smooth or entirely foveolate; if head foveolate with smooth notches on genae then at least one of the following characters present: scape longer, surpassing

Species accounts
Repetitive characters occurring in the majority of species have been omitted. Unless stated otherwise, the following descriptions apply to all species treated here: Major workers. Antennal sockets shallow; frontal lobes absent; head in full-face view with distinct median concavity; antenna 12-segmented, with 3-segmented club; masticatory margin of mandible with large, stout apical and preapical teeth, followed by a long diastema and then a short and crenulate tooth just before the rounded basal angle; outer surface of mandible mostly smooth and shining, sometimes with weak and sparse foveolae; antennal scrobes present; promesonotum strongly convex, well above the level of propodeum; postpetiole short with slightly convex dorsum; ventral process absent.
Minor workers. Antennal sockets shallow; frontal lobes absent; head in full-face view oval, posterior and anterior of eyes convex; antenna 12-segmented, with 3-segmented club; humeral area not developed; clypeus smooth and shiny; its anterior margin regularly convex; promesonotum well above the level of propodeum; petiole smooth; petiole with node moderately low, triangular and small postpetiole smooth; gaster smooth and shiny.

Revision of the Pheidole diakritos group
Diagnosis. Major worker. Head, in full-face view rectangular; in lateral view subrectangular; ventral and dorsal faces relatively flat; sides of the head with dense, relatively long, erect pilosity; antennal sockets deep, smooth or with a few rugae; frontal lobes distinct and lobe-like; head shiny, with sparse, thick, irregular rugae, interspaces smooth; inner hypostomal teeth distinct, closely spaced, triangular, with rounded apex and base slightly wider than apex; outer hypostomal teeth absent. Promesonotum short, angular and low; promesonotal groove absent; metanotal groove indistinct; propodeal spines long; mesosoma with thick, dense rugoreticulation; gaster shagreened on the whole surface. Minor worker. Head shiny, with sparse, thick, irregular rugae, interspaces smooth or with rugulae; scape, when laid back, surpassing posterior head margin by two-fifths of its length; promesonotum, in lateral view, convex; promesonotal groove present; metanotal groove present; humeral area with short triangular tubercles; pronotum and mesonotum with thick, dense rugoreticulation, sculpture slightly weakening on dorsum; katepisternum, anepisternum, and propodeum foveolate.
Comments. This species-group can be easily distinguished from others by presence of deep antennal sockets and distinct lobe-like frontal lobes in both, major and minor workers. Major workers also can be distinguished by absence of outer hypostomal teeth, and minor workers by presence of short triangular tubercles on the humeral area.
This group contains only one species: Pheidole diakritos sp. nov. distributed across the evergreen rainforest biome. ; head in full-face view rectangular; in lateral view subrectangular with visible inner hypostomal teeth; antennal scrobes absent; sides of the head with dense, relatively long, erect pilosity; antennal sockets deep; frontal lobes distinct and lobe-like; inner hypostomal teeth distinct, closely spaced, triangular, with rounded apex and base slightly wider than apex; outer hypostomal teeth absent. Minor workers. Head shiny, with sparse, thick, irregular rugae, interspaces smooth or rugulose; frontal lobes present; antennal sockets deep; propodeal spines long and thin (PSL: 0.17-0.21 (0.19)).
Etymology. Greek for distinct [διακριτός], in reference to its being the only known species with distinct frontal lobes and deep antennal sockets.
Biology. The species was collected at elevation between 360-865 m, in rainforest, and in montane rainforest. Nesting preferences are unknown.

Revision of the Pheidole lutea group
Diagnosis. Major workers. Head, in full-face view, oval or elongate, in lateral view sub-oval, ventral and dorsal faces convex or relatively flat; dorsal face not depressed posteriorly; antennal scrobes absent or very indistinct, occipital lobes smooth or with thick, sparse, irregular rugae, sculpture weakening posteriorly; frons with thick rugae, interspaces smooth to rugo-foveolate; genae smooth or with thick, sparse, irregular rugae; promesonotum relatively low to high, and arched; propodeal spines absent, weakly developed, and lobe-like or small and short, triangular; mesosoma surface with fine, sparse to moderately sparse rugoreticulation (dorsal surface with weaker sculpture) or smooth; body yellow to occasionally bright brown. Minor workers. At least frons, genae, and malar area smooth and shiny, sometimes lateral sides of frons with short, indistinct, longitudinal rugulae; scape short, when laid back, reaching posterior head margin or surpassing the posterior head margin by one-fifth of its length; promesonotum, in lateral view, convex or box-like; mesosoma almost entirely smooth; body yellow.
Comments. Major workers of this group can be easily distinguished from others based on elongate to oval head capsule in full-face view and sub-oval in lateral view, head sculpture weakening posteriorly and reduced sculpture of mesosoma, strongly reduced and sometimes absent to small propodeal spines, and bright body colouration. Minor workers can be separated from other species based on smooth and shiny head and mesosoma sculpture, short scape, and yellow body colouration.
The group is divided into two complexes. The P. lutea complex contains three species: P. lutea sp. nov., P. ranohirensis sp. nov., and P. voasara sp. nov., all distributed across dry deciduous forest biome and Sambirano rainforest biome. The P. navoatrensis complex also contains three species: P. navoatrensis sp. nov., P. parviocula sp. nov., and P. typhlos sp. nov. Pheidole navoatrensis sp. nov. and P. parviocula are known from central highlands, and P. navoatrensis additionally expands its range to the dry deciduous biome. Pheidole typhlos is known only from its type locality, the Galoko massif.

Revision of the Pheidole lutea complex
Diagnosis. Major workers. Head, in full-face view, oval; sides of the head with moderately dense to dense, long, erect pilosity; antennal scrobes indistinct and not delimited by carinulae; scrobe surface shiny, with sparse, thick, longitudinal to irregular rugae, interspaces smooth, indistinctly rugulose to foveolate; occipital lobes and genae with thick, sparse, irregular rugae, interspaces with indistinct foveolae, sculpture weakening posteriorly; frons with dense, thick, and longitudinal rugae, interspaces smooth to foveolate; promesonotum, in lateral view, short, angular, and relatively high; promesonotal and metanotal grooves absent; propodeal spines minute and indistinct to small, triangular; mesosoma with fine, sparse rugoreticulation and indistinct, sparse foveolae or foveolate with additional sparse and thick rugae on promesonotal dorsum and propodeum, sculpture slightly weakening on dorsum; anepisternum and mesosoma with indistinct sculpture or smooth; gaster smooth to finely shagreened; body orange to yellowish brown. Minor workers. Frons, genae, and malar area smooth and shiny, vertex smooth and with few arcuate, interrupted rugae; scape, when laid back, surpassing the posterior head margin by one-fifth of its length; promesonotum moderately low to high, short, evenly arched; promesonotal and metanotal grooves absent; propodeal spines very small, indistinct, triangular; mesosoma smooth and shiny, only dorsum with few transverse, thick rugulae; body yellow. Comments. Major workers of this complex can be easily distinguished based on a combination of the following characters: head, in full-face view, oval (but not elongate) and in lateral view sub-oval, lacking smooth patches and predominately covered with longitudinal rugae; minute to small propodeal spines; gaster never distinctly shagreened, and bright body colouration. Minor workers can be distinguished based on smooth sculpture of head and mesosoma, with few additional arcuate rugae on vertex and transverse rugulae on mesosoma, moderately short scape, and yellow body. Type material. Holotype. Madagascar. •1 major worker; Fianarantsoa; Parc National d' Isalo, 9.1 km 354°N Ranohira;-22.48167, 45.46167;alt. 725 m;31 Jan 2003;.; BLF07348, CASENT0485691 (CASC). Paratypes. Madagascar. •5w.; same data as for holotype; CASENT0485689, CASENT0485690, CASENT0872151-CASENT0872153 (CASC).
Diagnosis. Major workers. Head in full-face view oval, with anterior and posterior sides slightly convex; sides of the head with moderately dense, long, erect pilosity; occipital lobes and genae shiny, with thick, sparse, irregular rugae, interspaces with indistinct foveolae, sculpture weakening posteriorly; inner hypostomal teeth distinct, moderately high, closely spaced, triangular, with rounded apex and wide base; outer hypostomal approximately as high as inner hypostomal teeth, dentate, with very wide base, and upper half thin, rectangular with top pointed outward; mesosoma shiny, with fine, sparse rugoreticulation and indistinct, sparse foveolae, sculpture weakening on dorsum, anepisternum, katepisternum, and mesosoma smooth; gaster smooth; body orange to bright brown. Minor workers. Frons, genae, and malar area smooth and shiny; vertex smooth and shiny with few arcuate, interrupted rugae; promesonotum moderately high, short, evenly arched; propodeal spines very small, indistinct, triangular; mesosoma smooth and shiny, only dorsum with few transverse, thick rugulae.
Etymology. Latin for yellow, in reference to body colouration of major and minor workers.
Biology. The species was collected at elevation 725 m, in gallery forest. Nest was located in rotten log.
Diagnosis. Major workers. Head in full-face view oval, with anterior and posterior sides slightly convex; sides of the head with dense, long, erect pilosity; occipital lobes and genae shiny, with thick, sparse, irregular rugae, interspaces with fine but distinct foveolae, sculpture weakening posteriorly; inner hypostomal teeth distinct, moderately high, closely spaced, triangular, with rounded apex and narrow base; outer hypostomal approximately smaller and thinner than inner hypostomal teeth, with moderately wide base, dentate; mesosoma shiny, foveolate with additional sparse and thick rugae on promesonotal dorsum and propodeum, sculpture slightly weakening on dorsum; anepisternum and mesosoma with indistinct sculpture or smooth; body reddish brown. Minor workers. Frons, genae, and malar area smooth and shiny; vertex smooth and shiny with few arcuate, interrupted rugae; promesonotum low, short, slightly convex, with relatively steep posterior declivity; propodeal spines very small, indistinct, triangular; mesosoma smooth and shiny, only dorsum with few transverse, thick rugulae.
Etymology. From the type locality.
Biology. The species was collected at elevation between 30-600 m, in gallery forest, in tropical dry forest, in rainforest. Nests were located in rotten logs and in dead twigs above ground.
Comments. Pheidole ranohirensis sp. nov. is most similar to P. lutea sp. nov. and P. voasara sp. nov. Major workers. Pheidole ranohirensis sp. nov. differs from P. lutea sp. nov. by surface of genae and antennal scrobes foveolate, dentate outer hypostomal teeth, which are smaller and thinner than inner hypostomal teeth and is never pointed outward, and by anepisternum, katepisternum, and mesosoma never entirely smooth; from P. voasara sp. nov. in indistinct or partially smooth sculpture on anepisternum and mesosoma, foveolate frons, and dentate outer hypostomal teeth, which are smaller and thinner than inner hypostomal teeth. Minor workers. Pheidole ranohirensis sp. nov. differs from P. lutea sp. nov. in promesonotum low, short, slightly convex, and with relatively steep posterior declivity. Minor workers of P. voasara sp. nov. are unknown.  -18.26667, 45.40667;alt. 1050 m;13 Jan 2003;CASENT0028086 (CASC).

Pheidole voasara
Diagnosis. Major workers. Head, in full-face view, oval, with anterior and posterior sides slightly convex; sides of the head with sparse, long, erect pilosity; occipital lobes and genae shiny, with sparse, indistinct and irregular rugae, interspaces with fine but distinct foveolae, sculpture weakening posteriorly; inner hypostomal teeth distinct, moderately high, closely spaced, triangular, with rounded apex and wide base, tops directed inward; outer hypostomal bigger and wider than inner hypostomal teeth, with wide base, lobe-like; mesosoma shiny, shiny, foveolate with additional sparse to moderately dense, and moderately thick, rugae, sculpture slightly weakening on dorsum; upper part of mesosoma with indistinct sculpture; body yellowish brown.
Description  3). Head. In full-face view oval, with anterior and posterior sides slightly convex (Fig. 17C). In lateral view sub-oval; ventral and dorsal faces convex; dorsal face not depressed posteriorly; inner hypostomal teeth visible. Sides of the head with moderately dense, long, erect pilosity; whole head with moderately dense, short, suberect to erect pilosity. Antennal scrobes indistinct and not delimited by carinulae; scrobe surface shiny, with sparse, thick, longitudinal to irregular rugae, interspaces with fine but distinct foveolae. Occipital lobes and genae shiny, with sparse, indistinct and irregular rugae, interspaces with fine but distinct foveolae, sculpture weakening posteriorly; frons with dense, thick, and longitudinal rugae, interspaces smooth; malar area with dense, thick, longitudinal rugulae, interspaces smooth. Centre of clypeus smooth and shiny, lateral sides with longitudinal rugae; median notch present, wide and indistinct; median longitudinal carina absent; lateral longitudinal carinae absent. Scape, when laid back, reaching the midlength of head; pilosity suberect to erect . Inner hypostomal teeth distinct, moderately high, closely spaced, triangular, with rounded apex and wide base, tops directed inward; outer hypostomal teeth larger and wider than inner, with wide base, lobe-like (Fig. 85Z). Mesosoma. In lateral view, promesonotum short, angular, and high, posterior mesonotum relatively steep, with small tubercle-like projections; promesonotal groove absent; metanotal groove absent; propodeal spines small, triangular, with acute apex; humeral area laterally weakly produced (Fig. 17B). Surface shiny, foveolate with additional sparse to moderately dense, and moderately thick rugae, sculpture slightly weakening on dorsum; upper part of mesosoma with indistinct sculpture. Pilosity moderately sparse, long, and erect (Fig. 17A, B). Petiole. Shiny and with foveolae; peduncle moderately long, with distinct horizontal lobes on its basal part; node relatively low, triangular, with rounded apex, in rear view node dorsoventrally concave; pilosity moderately dense and erect (Fig. 17A,B). Postpetiole. Shiny and with foveolae, dorsum partially smooth; short and flat; in dorsal view very wide, almost semi-oval; pilosity long, moderately dense, and erect (Fig. 17A, B). Gaster. Shiny and finely shagreened on the basal part of first tergite; pilosity dense, moderately long and erect (Fig. 17A, B). Colour. Unicolourous, orange to yellowish brown; gaster brown; legs dark yellow (Fig. 17A, B).
Minor workers. Unknown. Etymology. Malagasy for orange, in reference to bright body colouration. Biology. The species was collected at elevation between 725-1100 m, in gallery forest, and in montane rainforest. Nesting preferences unknown.
Comments. Pheidole voasara sp. nov. is most similar to P. lutea sp. nov. and P. ranohirensis sp. nov. Major workers. Pheidole voasara sp. nov. differs from P. lutea sp. nov. by surface of genae and antennal scrobes foveolate, lobe-like outer hypostomal teeth, inner hypostomal teeth pointed inward, and by anepisternum, katepisternum, and mesosoma never smooth; from P. ranohirensis sp. nov. in distinct and never smooth sculpture on anepisternum and mesosoma, frons never foveolate, and lobe-like outer hypostomal teeth, which are bigger and wider than inner hypostomal teeth. Minor workers. Unknown.

Revision of the Pheidole navoatrensis complex
Diagnosis. Major workers. Head, in full-face view, elongate; antennal scrobes absent or very indistinct, occipital lobes smooth, sometimes with indistinct, very sparse, longitudinal to irregular rugoreticulations, sculpture, when present, fading anteriorly; frons with thick, sparse, short and interrupted rugae and smooth interspaces or smooth with few short, thick, longitudinal rugae on the anterior part; genae smooth; promesonotum relatively low to high, and arched; propodeal spines absent, weakly developed, and lobe-like or small and short, triangular; mesosoma surface with fine, sparse to moderately sparse rugoreticulation (dorsal surface with weaker sculpture) or smooth with indistinct and sparse sculpture on lateral sides; gaster smooth; body yellow, orange and occasionally bright brown. Minor workers. Head smooth, only lateral sides of frons with short, indistinct, longitudinal rugulae; scape, when laid back, reaching posterior head margin; promesonotum, in lateral view, convex or box-like; mesosoma sculpture smooth and shiny, and sometimes lateral sides with indistinct, irregular, and sparse rugae; body yellow.
Comments. Major workers of this complex can be easily distinguished from others based on elongate head capsule, strongly reduced sculpture of head and mesosoma, strongly reduced or small propodeal spines, and bright body colouration. Minor workers can be separated from other species based on smooth and shiny head and mesosoma sculpture, short scape, and yellow body colouration.  -23.4185, 46.4583;alt. 1365 m;8 Feb 2009;CASENT0149710 (CASC).

Pheidole navoatrensis
Diagnosis. Major workers. Head elongate; sides of the head with very sparse, moderately short, erect pilosity; frons smooth, with few short, thick, longitudinal rugae on the anterior part; genae shiny and smooth; inner hypostomal teeth distinct, small, closely spaced, triangular, with rounded apex; outer hypostomal teeth weakly developed, dentate; body yellow to orange; mesosoma smooth, sometimes with indistinct and sparse foveolae or rugulae on propodeum and promesonotum. Minor workers. Body yellow; head shiny and smooth, only lateral sides of frons with short, indistinct, longitudinal rugulae; propodeal spines short and triangular; mesosoma smooth and shiny.
Etymology. From the type locality. Biology. The species was collected at elevation between 70-1640 m, in dry forest on sandy soil, in Uapaca woodland, in savannah woodland, in tropical dry forest, in gallery forest, in shrubland. Nests were located in soil and under stones.
Diagnosis. Major workers. Eyes small and reduced; body yellow; head rectangular, anterior of eyes relatively straight, posterior of eyes slightly convex; sides of the head with dense, relatively long, erect pilosity; occipital lobes and genae shiny and smooth, sometimes with indistinct, very sparse, longitudinal rugulae; frons and malar area shiny, with thick, sparse, short, and interrupted rugae, interspaces smooth; propodeal spines absent or weakly developed, lobe-like; mesosoma shiny, with fine, sparse to moderately sparse rugoreticulation, sculpture on dorsum weaker or with smooth patches; inner hypostomal teeth indistinct, very low, lobe-like, with rounded apex; outer hypostomal teeth distinct, dentate, and with rounded tops directed outward. Minor workers. Eyes small and reduced; body yellow; head and mesosoma shiny and smooth; propodeal spines indistinct, triangular.
Etymology. Latin for small eyes. Biology. The species was collected at elevation between 1002-1987 m, in savannah grassland, in Uapaca woodland, in savannah woodland, and in grassland. Nests were located in soil, under stones, and in dead tree stumps.
Comments. This species is most similar to P. typhlos sp. nov. Major workers. Pheidole parviocula sp. nov. can be distinguished from P. typhlos sp. nov. by dense, relatively long pilosity on sides of the head, small, lobe-like inner hypostomal teeth and weakly developed or absent propodeal spines. Minor workers. Pheidole (CASC).
Etymology. Greek for blind [τυφλός], in reference to the reduced eyes. Biology. The species was collected at elevation between 520-980 m, in rainforest, in montane forest. Nesting preferences unknown.
Comments. This species is most similar to P. parviocula sp. nov. Major workers. Pheidole typhlos sp. nov. can be distinguished from P. parviocula sp. nov. by sparse and short pilosity on sides of the head, presence of distinct, triangular inner hypostomal teeth and presence of short, triangular propodeal spines. Minor workers. Pheidole typhlos sp. nov. can be distinguished from P. parviocula sp. nov. by box-like promesonotum, absence of promesonotal groove and presence of short and triangular propodeal spines.

Revision of the Pheidole longispinosa group
Diagnosis. Major workers. Large species; head, in full-face view, trapezoid, widened posteriorly, in lateral view sub-oval, ventral and dorsal faces convex, dorsal face not depressed posteriorly; antennal scrobes absent or indistinct; occipital lobes smooth, at least on the posterior part; genae smooth to finely rugulose; head sculpture weakens posteriorly; promesonotum short, low, and evenly convex; propodeal spines very long; first gastral tergite smooth to shagreened; body brown to black. Minor workers. Head smooth to rugoreticulate, central part of frons usually with smooth notch; scape, when laid back, surpassing posterior head margin by one third or more than half of its length; promesonotum low, long, and slightly convex; promesonotal groove absent or very indistinct; metanotal groove very indistinct; propodeal spines very long; petiole with long and thin peduncle; body brown to black.
Comments. Members of this group are divided into two complexes. The P. longispinosa complex contains three species: P. longispinosa Forel, P. praegrandis sp. nov., and P. mahaboensis sp. nov. Majority of records of Pheidole longispinosa come from central highlands, but the species is known also from dry deciduous biome and evergreen rainforest. Pheidole mahaboensis is distributed cross the evergreen rainforest biome and central highlands, in southern part of its distribution range P. mahaboensis occurs sympatric with P. longispinosa. Pheidole praegrandis is known exclusively from the evergreen rainforest biome and a centre of its distribution is located in its northernmost part. However, the species is sympatric with P. mahaboensis in remaining parts of this biome. The P. scabrata complex contains two species: P. scabrata and P. maizina sp. nov. of sympatric distribution limited to the northern parts of evergreen forest and central highlands biomes.

1
Major workers. Sides of the head with relatively long, erect pilosity; antennal lobes never predominately smooth; promesonotum with strong sculpture (Fig. 21). Minor workers. Scape, when laid back, surpassing posterior head margin by one third of its length; mesosoma with dense rugoreticulation, sometimes dorsum with weaker sculpture, but never smooth (Fig. 21 Major workers. Sides of the head without or with short and decumbent pilosity; promesonotum predominately smooth (Fig. 22). Minor workers. Scape, when laid back, surpassing posterior head margin by more than half of its length; mesosoma smooth, indistinctly rugulose or the whole surface finely rugoreticulate (Fig. 22) .... 3 2 Major workers. Genae smooth and shiny on the entire surface or indistinctly rugulose, tips of outer hypostomal teeth never directed outward, inner hypostomal teeth slightly bigger than outer hypostomal teeth; katepisternum with smooth area; body brown to dark brown (Fig. 21A Major workers. Metanotal groove absent or shallow, frons with longitudinal rugae never reaching further than midlength of head (Fig. 22C). Minor workers. Promesonotal groove absent, metanotal groove shallow and wide, mesosoma with reduced sculpture (Fig. 22G)

Revision of the Pheidole longispinosa complex
Diagnosis. Major workers. Sides of the head with pilosity absent or very sparse, short, and decumbent; antennal scrobes absent or very indistinct; occipital lobes smooth and shiny or partially shagreened; genae smooth or partially superficially rugulose or shagreened; promesonotum low, short, and relatively flat; promesonotal groove absent; metanotal groove absent or distinct; propodeal spines very long; mesosoma shiny and smooth with weak, sparse, and irregular rugae on pronotum and lateral sides of propodeum or rugoreticulate, with weaker sculpture on propodeal dorsum; gaster smooth or first gastral tergite shagreened. Minor workers. Scape, when laid back, surpassing posterior head margin by more than half of its length; promesonotal groove absent; metanotal groove indistinct to distinct; mesosoma smooth, indistinctly rugulose or the whole surface finely rugoreticulate.
Comments. Species of this complex are characterized by a large body size, both in minor and major workers. Major workers can be distinguished by a combination of the following characters: head, in full-face view, trapezoid and widened posteriorly, in lateral view sub-oval; sides of the head with pilosity absent or very sparse and short; predominantly smooth occipital lobes; very long and massive propodeal spines and brown to black body colouration. Minor workers can be distinguished by smooth or finely rugulose head with smooth patches; long scape (surpassing posterior head margin by more than half of its length); very long and massive propodeal spines; smooth or indistinctly rugulose mesosoma, and long peduncle of petiole.
Comments. This species is most similar to P. mahaboensis sp. nov. and P. praegrandis sp. nov. Major workers. Pheidole longispinosa can be distinguished from P. mahaboensis sp. nov. by absence of metanotal groove, lack of pilosity at the sides of head, and longitudinal rugae never reaching further than midlength of head; from P. praegrandis sp. nov. by brighter body colouration, reduced head sculpture, and not shagreened first gastral tergite and head. Minor workers. Pheidole longispinosa can be distinguished from P. mahaboensis sp. nov. by absence of promesonotal groove, shallow and wide metanotal groove and reduced sculpture on mesosoma; from P. praegrandis sp. nov. by mostly smooth and shiny sculpture of head and mesosoma, which is never rugoreticulate.  .7)); head in full-face view trapezoid, widened posteriorly; sides of the head with very sparse, short, decumbent pilosity; frons with longitudinal rugae reaching further than midlength of head; gaster smooth and shiny; body reddish brown to dark brown. Minor workers. Large species: 0.93-1.08 (1.0), HW: 0.79-0.93 (0.88), WL: 1.2-1. 5 (1.4); propodeal spines very long (PSL: 0.53-0.66 (0.61)); scape, when laid back, surpassing posterior head margin by more than half its length; pronotum and mesonotal dorsum shiny, smooth or with indistinct, sparse, superficial rugulae; anepisternum, katepisternum and propodeum shiny, with slightly denser superficial rugulae; promesonotal groove present; metanotal groove narrow and relatively deep. In lateral view sub-oval; ventral and dorsal faces convex; inner hypostomal teeth visible. Sides of the head with very sparse, short, decumbent pilosity; frons and vertex with few additional, long, erect setae. Antennal scrobes absent or very indistinct. Occipital lobes smooth and shiny; genae smooth and shiny or dull, superficially rugulose with median shiny patch widening posteriorly; centre of frons shiny with longitudinal rugae reaching at least midlength of head, sometimes surface between rugae with superficial rugulae; malar area and lateral sides of head superficially rugulose, sometimes with several short, longitudinal rugae; posterior part of frons smooth and shiny or superficially rugulose; head sculpture weakens posteriorly. Centre of clypeus smooth and shiny, lateral sides superficially rugulose; median notch present, shallow, and wide; median longitudinal carina absent; lateral longitudinal carinae absent. Scape, when laid back, reaching midlength of head; pilosity decumbent to subdecumbent (Fig. 24B,D). Inner hypostomal teeth distinct, low, triangular, and thick, with rounded apex, closely spaced; outer hypostomal teeth distinct, slightly smaller and thinner than inner hypostomal teeth, lobe-like (Fig. 85B). Mesosoma. In lateral view, promesonotum low, short, and relatively flat; pronotum slightly convex; mesonotum slightly concave; posterior mesonotum relatively steep; promesonotal groove absent; metanotal groove narrow and relatively deep; propodeal spines very long, massive basally, with acute apex; humeral area laterally weakly or not produced (Fig. 24D). Surface shiny and smooth with weak, sparse, and irregular rugae on pronotum, lateral sides of propodeum, katepisternum, and anepisternum. Pilosity very sparse, short, and decumbent; dorsum with few additional, long erect setae (Fig. 24D, F). Petiole.
Etymology. From the type locality. Biology. The species was collected at between 20-1275 m in elevation, in rainforest and once in open area near stream. Nests were located in rotten logs and tree stumps.
Comments. This species is most similar to P. longispinosa and P. praegrandis sp. nov. Major workers. Pheidole mahaboensis sp. nov. can be distinguished from P. longispinosa by presence of metanotal groove, presence of very sparse, short, decumbent pilosity at the sides of head, and longitudinal rugae reaching at least midlength of head; from P. praegrandis sp. nov. by presence of metanotal groove, presence of very sparse, short, decumbent pilosity at the sides of head, and not shagreened first gastral tergite and head. Minor workers. Pheidole mahaboensis sp. nov. can be distinguished from P. longispinosa by presence of promesonotal groove, narrow and relatively deep metanotal groove, and presence of fine superficial rugulae on most of mesosoma; from P. praegrandis sp. nov. by mostly smooth and shiny sculpture of head and mesosoma, which is never rugoreticulate.  24 (2.11); propodeal spines very long (PSL: 0.81-0.96 (0.89)); head in full-face view trapezoid, widened posteriorly; sides of the head without pilosity; frons with fine, dense, longitudinal rugae reaching at most midlength of head; first gastral tergite shagreened, at least on its basal half; body dark brown to black. Minor workers. Large species: HL: 1.15-1.29 (1.2), HW: 1.04-1.14 (1.1), WL: 1.86-2.01 (1.92); propodeal spines very long (PSL: 0.91-1.0 (0.97)); scape, when laid back, surpassing posterior head margin by more than half its length; mesosoma finely rugoreticulate; promesonotal groove absent or very indistinct; metanotal groove shallow and wide.
Etymology. Latin for huge, in reference to large body size and very long propodeal spines.
Biology. The species was collected between 250-900 m in elevation, in rainforest and montane rainforest. Nests were located in rotten logs and tree stumps.
Comments. This species is most similar to P. longispinosa and P. mahaboensis sp. nov. Major workers. Pheidole praegrandis sp. nov. can be distinguished from P. longispinosa by shagreened surface of head and first gastral tergite, and darker body colouration; from P. mahaboensis sp. nov. by absence of metanotal groove, absence of very sparse, short, decumbent pilosity at the sides of head, and shagreened first gastral tergite and head. Minor workers. Pheidole praegrandis sp. nov. can be distinguished from P. longispinosa by rugoreticulate mesosoma sculpture; from P. mahaboensis sp. nov. by rugoreticulate mesosoma sculpture, and shallow and wide metanotal groove.

Revision of the Pheidole scabrata complex
Diagnosis. Sides of the head with sparse, relatively long, erect pilosity; antennal scrobes absent; occipital lobes smooth, at least on the posterior part; genae smooth to finely rugulose; centre of frons with longitudinal rugae directed outward posteriorly; inner and outer teeth closely spaced and connected by indistinct concavity; promesonotal groove absent; mesosoma with sparse to relatively dense rugoreticulation, mesosomal dorsum and propodeum with weaker sculpture or smooth patches; first gastral tergite smooth to shagreened. Minor workers. Scape, when laid back, surpassing posterior head margin by one third of its length; promesonotal groove absent or very indistinct; metanotal groove very indistinct; mesosoma with dense rugoreticulation, sometimes dorsum with weaker sculpture, but never smooth.
Comments. Species of this complex are characterized by a large body, both in minor and major workers. Major workers can be distinguished by a combination of the following characters: head, in full-face view, trapezoid and widened posteriorly, in lateral view sub-oval; sides of the head with sparse, relatively long, erect pilosity, occipital lobes predominantly smooth; frons with rugae directed outward, and very long, massive propodeal spines. Minor workers can be distinguished based on long scape (surpassing posterior head margin by one-third of its length); very long propodeal spines; mesosoma with dense rugoreticulation, and long and thin peduncle of petiole.  .77); propodeal spines very long (PSL: 0.5-0.63 (0.55)); head in full-face view trapezoid, widened posteriorly; sides of the head with sparse, relatively long, erect pilosity; frons with rugoreticulation and additional longitudi-nal rugae on the whole surface, sculpture weakening posteriorly; first gastral tergite smooth to slightly shagreened; body brown to dark brown; inner hypostomal teeth distinct, triangular and thick, with rounded apex, closely spaced; outer hypostomal teeth distinct, low, lobe-like, with wide base, smaller and thinner than inner hypostomal teeth. Minor workers. Large species: HL: 0.93-1.06 (0.99); HW: 0.9-1.06 (0.96), 1.41-1.65 (1.48); propodeal spines very long (PSL: 0.41-0.48 (0.43)); scape, when laid back, surpassing posterior head margin by one third of its length; head sculpture never rugoreticulate; mesosoma with dense rugoreticulation, sometimes dorsum with weaker sculpture, but never smooth.
Biology. The species was collected at between 825-1220 m in elevation, in montane rainforest, rainforest, transitional humid forest, and tropical dry forest. Nests were located in rotten logs, and in rotten sticks on the ground.
Etymology. Malagasy for dark, in reference to dark body colouration. Biology. The species was collected at between 400-1200 m in elevation, in rainforest, montane rainforest, and transitional humid forest. Nests were located in in rotten logs and stumps, and in rotten sticks on the ground.
Comments. This species is most similar to P. scabrata. Major workers. Pheidole maizina sp. nov. can be distinguished from P. scabrata by having distinct rugulae on at least half of the genae, tips of outer hypostomal teeth directed outward, never smooth katepisternum, densely shagreened surface of first gastral tergite, and darker body colouration. Minor workers. Pheidole maizina sp. nov. can be distinguished from P. scabrata by rugoreticulate head sculpture with never smooth genae.

Revision of the Pheidole ensifera group
Diagnosis. Major workers. Head, in full face view, rectangular, in lateral view elongate and oval, ventral and dorsal faces slightly convex, dorsal face not depressed posteriorly; sides of the head with sparse, relatively long to long, suberect pilosity; antennal scrobes indistinct and not delimited by carinulae; scrobe surface shiny, with dense to sparse, fine, longitudinal to irregular rugulae; occipital lobes and genae smooth or with fine to thick, sparse rugoreticulation; frons with sparse, thick, and longitudinal rugae, sometimes rugae on posterior part of frons more irregular and directed outward; sculpture weakening posteriorly; inner and outer teeth closely spaced and connected by concavity; promesonotum short, angular, and relatively low to high; promesonotal and metanotal grooves absent; propodeal spines long to very long, thin; mesosoma with fine to thin, sparse to dense rugoreticulation, sculpture weakening on dorsum, sometimes propodeum with a smooth patch on its dorsal surface; gaster shagreened, at least at the basal part of first tergite; body reddish brown to dark brown. Minor workers. Head smooth or finely rugo-punctuate, lateral sides of head sometimes finely rugulose to rugo-punctate; scape, when laid back, surpassing posterior head margin by one-to two-fifths of its length; promesonotum high, short and convex; propodeal spines very to moderately long; whole mesosoma rugo-punctate, or smooth to moderately foveolate with sometimes thicker and denser sculpture on lateral sides; body bright brown to dark brown.
Comments. Species of this group are characterized by a large body size, both in minor and major workers. Major workers can be distinguished from others based on a combination of the following characters: head in full-face view rectangular, in lateral view elongate and oval; antennal scrobes with fine, longitudinal to irregular rugulae; occipital lobes and genae smooth or with sparse rugoreticulation; inner and outer teeth closely spaced and connected by concavity, and long to very long propodeal spines. Minor workers can be distinguished based on smooth to rugo-punctuate surface of frons; relatively short scape (surpassing posterior head margin by one-to two-fifths of its length); high, short, and convex promesonotum; very to moderately long propodeal spines, and bright brown to dark brown body colouration.
This group includes four species: P. ensifera Forel, P. ocypodea sp. nov., P. aelloea sp. nov., and P. podargea sp. nov. The distribution of the whole group is limited to the Northernmost part of the island (Antsiranana district) and all species are sympatric. Minor workers of members of this group manifest high infraspecific variability, therefore we encourage to use major workers in species determination.
Biology. The species was collected between 30-1343 m in elevation, in littoral and tropical dry rainforest and in-transition humid forest. Nests were located in litter (leaf mould, rotten wood), rotten logs and branches on the ground, and rotting tree stumps.
Comments. Major workers. Pheidole ensifera differs from other members of the group in shiny and smooth to finely rugoreticulate occipital lobes and genae and relatively low and short promesonotum. Minor workers. Pheidole ensifera differs from other members of the group in surface of pronotum and mesonotum never smooth and with fine to moderately dense foveolae. ; head in full-face view rectangular, with lateral sides relatively straight, only their posteriormost part slightly convex; sides of the head with sparse, relatively long, erect pilosity; occipital lobes shiny, with sparse and thick rugoreticulation; inner hypostomal teeth distinct, triangular, and moderately thin, with rounded apex, closely spaced; outer hypostomal teeth distinct, approximately as high as inner hypostomal teeth, lobe-like, with base wider than inner hypostomal teeth, inner and outer teeth closely spaced and connected by indistinct concavity. Minor workers. Body size moderate: HL: 0.63-0.78 (0.68), HW: 0.61-0.79 (0.67), WL: 0.83-0.98 (0.9); propodeal spines very long (PSL: 0.3-0.37 (0.32)); scape, when laid back, surpassing posterior head margin by one-fifth of its length; lateral sides of head and malar area finely rugulose, sculpture weakening posteriorly; vertex, genae, and frons smooth; pronotum, dorsal surface of mesonotum, and dorsal and posterior surface of propodeum smooth, sometimes propodeum with indistinct, sparse rugulae; katepisternum, anepisternum, and lateral sides of propodeum with thick and dense rugo-punctae. Head. In full-face view rectangular, lateral sides relatively straight, only their posteriormost part slightly convex (Fig. 31B). In lateral view oval; ventral and dorsal faces finely convex; inner hypostomal teeth visible. Sides of the head with sparse, relatively long, erect pilosity; whole head with sparse, relatively long, erect pilosity. Antennal scrobes indistinct and not delimited by carinulae. Occipital lobes shiny, with sparse and thick rugoreticulation; genae shiny, with fine and sparse rugoreticulation, sometimes posterior part with reduced sculpture; malar area and frons with longitudinal and sparse rugae, on posterior part of frons rugae directed outward and more irregular, interspaces smooth; lateral sides of head shiny, with irregular, moderately dense and thick rugoreticulation. Centre of clypeus smooth and shiny, lateral sides with longitudinal rugae; median notch present, moderately wide, and narrow; median longitudinal carina present; lateral longitudinal carinae absent. Scape, when laid back, reaching mid-length of head; pilosity suberect to erect (Fig. 31B, D). Inner hypostomal teeth distinct, triangular, and moderately thin, with rounded apex, closely spaced; outer hypostomal teeth distinct, approximately as high as inner hypostomal teeth, lobe-like, with base wider than inner hypostomal teeth, inner and outer teeth closely spaced and connected by indistinct concavity (Fig. 84A). Mesosoma. In lateral view, promesonotum short, angular, and high, with slightly convex posterior declivity; promesonotal groove absent; metanotal groove absent; propodeal spines very long, narrow, massive basally, with acute apex; humeral area laterally absent to weakly produced (Fig. 31D). Surface shiny, pronotum, dorsal surface of mesonotum, and dorsal and posterior surface of propodeum with fine, sparse to moderately sparse rugoreticulation, sometimes sculpture on propodeum strongly reduced and absent on its dorsal surface; katepisternum, anepisternum, and lateral sides of propodeum with thick and sparse rugoreticulation. Pilosity moderately sparse, long, and erect (Fig. 31D, F). Petiole. Shagreened; peduncle relatively long, with triangular apex, and thick horizontal lobes on its basal part; node triangular with rounded apex, in rear view node dorsoventrally slightly depressed; pilosity moderately sparse and erect (Fig. 31D,F). Postpetiole. Shagreened; in dorsal view sides with acute, moderately long, relatively narrow, and angular projections; pilosity long, moderately sparse, and erect (Fig. 31D,F). Petiole. First gastral tergite shagreened, at least on its basal part; pilosity moderately sparse, long, and erect (Fig. 31D, F). Colour. Unicolourous, ocherous to reddish-brown, sometimes head brighter than the rest of the body (Fig. 31D, F).  suberect to erect (Fig. 31A, C). Mesosoma. In lateral view, promesonotum high, short, and convex; promesonotal groove absent; metanotal groove absent; propodeal spines very long, massive basally, with acute apex (Fig. 31C). Surface shiny, pronotum, dorsal surface of mesonotum, and dorsal and posterior surface of propodeum smooth, sometimes propodeum with indistinct, sparse rugulae; katepisternum, anepisternum, and lateral sides of propodeum with thick and dense rugose punctae. Pilosity sparse, long, and erect (Fig. 31C, E). Petiole. Vertex of peduncle finely shagreened; peduncle long and moderately thin; node low, bulge-like; with few long, erect setae (Fig. 31C, E). Postpetiole. Moderately long, low, and slightly convex; with few long, erect setae at the anterior edge (Fig. 31C, E). Petiole. With few long, erect setae (Fig. 31C, E). Colour. Unicolourous, brown to dark brown (Fig. 31C, E).
Etymology. Named after Aello, a harpy from Greek mythology, in reference to the long, sharp propodeal spines of minor workers reminiscent of claws.
Biology. The species was collected between 10-2100 m in elevation, in rainforest, tropical dry forest, littoral rainforest, montane rainforest, and montane shrubland. Nests were located in rotten logs, rotten sticks on ground, dead twigs above ground, and soil.
Comments. This species is most similar to P. ocypodea sp. nov. and P. podargea sp. nov. Major workers. Pheidole aelloea sp. nov. can be distinguished from P. ocypodea sp. nov. by slightly convex posterior declivity of promesonotum and inner and outer hypostomal teeth not pointed inward or outward; it differs from P. podargea sp. nov. by longer and sparser pilosity on sides of head and reduced to absent sculpture on genae and propodeum. Minor workers. Pheidole aelloea sp. nov. can be distinguished from P. ocypodea sp. nov. by finely rugulose and never smooth lateral sides of head and malar area, absence of metanotal groove, long petiolar peduncle, and short postpetiole which is approximately as long as high; it differs from P. podargea sp. nov. in smooth sculpture of vertex, genae, and area between frontal carinae, absence of promesonotal and metanotal groove, and at least partially smooth surface of pronotum, dorsal surface of mesonotum, and dorsal and posterior surface of propodeum.  .41)); head in full-face view rectangular, with lateral sides relatively straight, only their posteriormost part slightly convex; sides of the head with sparse, relatively long, erect pilosity; occipital lobes shiny, with sparse and thick rugoreticulation; inner hypostomal teeth distinct, low, closely spaced, triangular, with rounded apex directed outward; outer hypostomal teeth distinct, low, lobe-like, with base wide and tops directed inward; inner and outer teeth closely spaced and connected by concavity. Minor workers. Body size moderate: HL: 0.63-0.72 (0.66); HW: 0.6-0.7 (0.64), WL: 0.8-0.93 (0.85); propodeal spines very long (PSL: 0.26-0.32 (0.28)); scape, when laid back, surpassing posterior head margin by two-fifths of its length; lateral sides of head and malar area smooth and shiny or with indistinct, sparse rugulae, sculpture weakening posteriorly; vertex, genae, and frons smooth; pronotum, dorsal surface of mesonotum, and dorsal and posterior surface of propodeum smooth; katepisternum, anepisternum, and lateral sides of propodeum with thick and sparse rugae.
Etymology. Named after Ocypode, a harpy from Greek mythology, in reference to the long and sharp propodeal spines of minor workers reminiscent of claws.
Biology. The species was collected between 400-980 m in elevation, in rainforest and montane rainforest. Nests were located in in rotten logs and rotten sticks on ground.
Comments. This species is most similar to P. aelloea sp. nov. and P. podargea sp. nov. Major workers. Pheidole ocypodea sp. nov. can be distinguished from P. aelloea sp. nov. by steep to relatively steep posterior declivity of promesonotum, inner hypostomal teeth pointed outward and outer hypostomal teeth pointed inward; from P. podargea sp. nov. it differs in longer and sparser pilosity on sides of head, reduced to absent sculpture on genae and propodeum, and inner hypostomal teeth pointed outward. Minor workers. Pheidole ocypodea sp. nov. can be distinguished from P. aelloea sp. nov. by smooth lateral sides of head and malar area, presence of metanotal groove, short petiolar peduncle, and long postpetiole which is approximately 1.5 times longer than high; from P. podargea sp. nov. it differs in smooth sculpture of vertex, genae, and area between frontal carinae, absence of promesonotal groove and at least partially smooth surface of pronotum, dorsal surface of mesonotum, and dorsal and posterior surface of propodeum.  .42)); head in full-face view rectangular, with lateral sides relatively straight, only their posteriormost part slightly convex; sides of the head with moderately dense, moderately long, erect pilosity; occipital lobes shiny, with sparse and thick rugoreticulation; inner hypostomal teeth distinct, closely spaced, triangular, with rounded apex; outer hypostomal teeth distinct, as high as inner hypostomal teeth, lobe-like, with base wide and tops directed slightly inward; inner and outer teeth closely spaced and connected by concavity. Minor workers. Body size moderate: HL: 0.61-0.7 (0.66), HW: 0.62-0.69 (0.65), WL: 0.81-0.9 (0.86); propodeal spines very long (PSL: 0.25-0.31 (0.28)); scape, when laid back, surpassing posterior head margin by two-fifths of its length; whole head with fine and dense rugo-punctuation, only basal area of frons and genae with sculpture reduced or sometimes absent; mesosoma rugo-punctate, sometimes sculpture weakening on dorsal surface and pronotum.
Etymology. Named after Podarge, a harpy from Greek mythology, in reference to long and sharp propodeal spines of minor workers reminiscent of claws.
Biology. The species was collected between 210-800 m in elevation, in rainforest and tropical dry forest. Nests were located in rotten logs.
Comments. This species is most similar to P. aelloea sp. nov. and P. ocypodea sp. nov. Major workers. Pheidole podargea sp. nov. can be distinguished from P. aelloea sp. nov. and P. ocypodea sp. nov. by outer hypostomal teeth approximately the same size as the inner ones and lack of smooth sculpture on genae and propodeum. Minor workers. Pheidole podargea sp. nov. can be distinguished from P. aelloea sp. nov. and P. ocypodea sp. nov. by smooth area limited to basal area of frons and genae or the whole head finely rugo-punctate, presence of promesonotal and metanotal groove, and absence of smooth surfaces on mesosoma.

Revision of the Pheidole ferruginea group
Diagnosis. Major workers. Head in full-face view cordate or sub-oval, widened posteriorly, in lateral view oval to sub-oval, ventral and dorsal faces strongly convex, dorsal face not depressed posteriorly; antennal scrobes developed, well delimited by carinulae (except P. longipilosa); occipital lobes with sparse and irregular rugoreticulation; promesonotum short, angular, and low to high; promesonotal and metanotal grooves absent; propodeal spines moderately long to long, with base wide or narrow; mesosoma predominately sculptured sometimes with smooth notches; first gastral tergite smooth or shagreened, at least on its basal part; body bright brown to brown. Minor workers. Whole head foveolate, sometimes with additional longitudinal rugae on frons; scape, when laid back, surpassing posterior head margin by one-to two-fifths of its length; promesonotum box-like or convex; propodeal spines short, moderately long or very long; promesonotal and metanotal grooves absent or present; mesosoma predominately foveolate, sometimes with additional rugae on mesosomal dorsum; body yellow to brown.
Comments. Major workers of this group can be distinguished based on the combination of following characters: cordate to sub-oval head in lateral view, oval to suboval in dorsal view; well-developed antennal scrobes delimited by carinulae (except P. longipilosa); never smooth occipital lobes; lack of promesonotal and metanotal grooves; moderately long to long propodeal spines; strong sculpture of mesosoma, occasionally with smooth notches and bright brown to brown body. Minor workers can be separated entirely foveolate head, sometimes with additional longitudinal rugae on frons; short scape surpassing posterior head margin by one-to two-fifths of its length; and moderately long or very long propodeal spines (except P. longipilosa).
The group is divided into two complexes. The P. ferruginea complex includes four species: P. ferruginea sp. nov., P. rugocephala sp. nov., P. vohemarensis sp. nov., and P. manantenensis sp. nov. Pheidole ferruginea is relatively common across the evergreen forest biome and is sympatric with the remaining three members of the complex. The distribution range of P. rugocephala sp. nov., P. vohemarensis sp. nov., and P. manantenensis sp. nov. is predominately limited to the Antsiranana prefecture. Pheidole rugocephala is known from the evergreen forest biome located between Toamasina and Andapa, while P. vohemarensis, known from the same biome, inhabits area between Ambinaelo and Antsirabe. Pheidole vohemarensis sp. nov. is known from area spread between Andapa and Antisianana and its distribution predominately covers the dry deciduous forest biome. Pheidole longipilosa creates a single-species complex and is known from Forêt Classée d'Analavelona and Parc National d'Isalo in the Toliara prefecture.
Key to the Pheidole ferruginea group.

1
Major workers. Head in full-face view sub-oval; sides of head with dense, very long, erect pilosity; antennal scrobes indistinct and not delimited by carinulae; propodeal spines triangular, with wide base (Fig. 34O). Minor workers. Propodeal spines short (

Revision of the Pheidole ferruginea complex
Diagnosis. Major workers. Head in full-face view cordate, widened posteriorly; antennal scrobes developed, well delimited by carinulae; occipital lobes with sparse and thick rugoreticulation or foveolae, with additional thick, sparse, irregular rugae; frons with sparse, thick, and longitudinal to irregular rugae, interspaces smooth, or with fine rugulae or superficially foveolate; promesonotum short, angular, and low to moderately high; propodeal spines long, with base wide or narrow; mesosoma with thick and sparse to dense rugoreticulation, sometimes sculpture weakening on dorsum, or foveolate, with additional sparse to dense rugoreticulation; first gastral tergite shagreened, at least on its basal part; body reddish brown to brown. Minor workers. Scape, when laid back, surpassing posterior head margin by one-to two-fifths of its length; promesonotum box-like or convex; propodeal spines moderately long to very long; promesonotal groove absent or present; metanotal groove absent or present; mesosoma foveolate, sometimes with additional rugae on mesosomal dorsum; sometimes katepisternum smooth; body yellow to brown. Comments. Major workers of this complex can be distinguished based on a combination of the following characters: head in full-face view cordate, in lateral view oval to sub-oval; antennal scrobes developed and well delimited; never smooth occipital lobes and frons; long propodeal spines; and first gastral tergite shagreened, at least on its basal part. Minor workers can be distinguished based on foveolate head and mesosoma, sometimes with additional longitudinal rugae on frons or promesonotum; moderately long to long propodeal spines; and body yellow to brown. Diagnosis. Head in full-face view cordate, widened posteriorly; sides of the head with sparse, long, suberect pilosity; antennal scrobes strongly developed, well delimited and forming distinct dorsal concavity beneath frontal carina; scrobe surface shiny, with sparse, thick, longitudinal and sometimes additional irregular rugae; delimited ventrally and posteriorly by carinulae; propodeal spines long (PSL: 0.23-0.28 (0.26)); first gastral tergite shagreened, at least on its basal part. Minor workers. Head foveolate, sometimes with additional longitudinal rugae on frons; propodeal spines very long (PSL: 0.15-0.18 (0.16)); promesonotum box-like; mesosoma foveolate, sometimes with additional rugae on mesosomal dorsum, and katepisternum with smooth notch.
Etymology. Latin for rusty, in reference to body colouration of major workers. Biology. The species was collected between 20-1312 m in elevation, in rainforest, montane rainforest, and littoral rainforest. Nests were located in rotten logs and tree stumps, rotten sticks on ground, and the petioles of Melastomataceae.
Comments. This species is most similar to P. rugocephala sp. nov. Major workers. Pheidole ferruginea sp. nov. can be distinguished from P. rugocephala sp. nov. by the surface of antennal scrobes never foveolate; presence of longitudinal rugae on frons; inner hypostomal teeth newer pointed inward; never smooth surface of promesonotum; and lack of horizontal lobes on the basal part of petiolar peduncle. Minor workers. Pheidole ferruginea sp. nov. can be distinguished from P. rugocephala sp. nov. by long propodeal spines and foveolate clypeus, and lack of promesonotal and metanotal grooves.  (CASC).
Etymology. Latin for strong and distinct head sculpture on major workers. Biology. The species was collected between 15-1200 m in elevation, in rainforest, montane rainforest, and littoral rainforest. Nesting preferences unknown.
Etymology. From the type locality. Biology. The species was collected between 225-775 m in elevation, in rainforest. Nest was located in the petiole of Melastomataceae.
Comments. Pheidole manantenensis sp. nov. is most similar to P. vohemarensis sp. nov. Majors workers. It differs from P. vohemarensis sp. nov. in presence of irregular rugae on antennal scrobes, frons with foveolae, lower promesonotum, and petiolar peduncle with short horizontal lobes on its basal part. Minor workers. It differs from P. vohemarensis sp. nov. in promesonotum with posterior declivity smoothly declining towards propodeum, and foveolate katepisternum and mesonotum.  (CASC).
Etymology. From type locality. Biology. The species was collected between 10-775 m in elevation, in tropical dry forest, littoral rainforest, and rainforest. Nests were located in rotten logs.
Comments. Pheidole vohemarensis sp. nov. is most similar to P. manantenensis sp. nov. Major workers. It differs from P. manantenensis sp. nov. in presence of longitudinal rugae on antennal scrobes, frons never with foveolae, higher promesonotum and petiolar peduncle with wide and distinct horizontal lobes on its basal part. Minor workers. It differs from P. vohemarensis sp. nov. in promesonotum with posterior declivity relatively steep and katepisternum and mesonotum with smooth notches.

Revision of the Pheidole longipilosa complex
Diagnosis. Major workers. Head in full-face view sub-oval; sides of head with dense, very long, erect pilosity; antennal scrobes indistinct and not delimited by carinulae; occipital lobes with thick, irregular rugae, interspaces with distinct, irregular rugulae; genae shiny, with dense and thin, irregular rugulae, central part with smooth notch; promesonotum relatively high and arched; propodeal spines small to moderately long, triangular; promesonotum foveolate with additional indistinct, sparsely rugoreticulate, sometimes sculpture fading on dorsal surface and lower parts of lateral sides; gaster smooth; body reddish brown to bright brown. Minor workers. Scape, when laid back, surpassing the posterior head margin by one-fifth of its length; promesonotum low, convex, short; promesonotal and metanotal grooves absent; propodeal spines short, tri-angular; mesosoma foveolate; katepisternum smooth; anepisternum and mesonotum with fading sculpture and sometimes smooth; body yellow.
Comments. Major workers of this complex can be distinguished based on a combination of the following characters: head in full-face view elongated, in lateral view sub-oval; sides of the head with dense, long, erect pilosity; head sculptured but never with arcuate or transverse rugulae, central part of frons smooth; propodeal spines small to moderately long, triangular, and smooth gaster. Minor workers can be separated based on foveolate head and mesosoma with additional rugae on frons and smooth katepisternum, short and triangular propodeal spines, and yellow body colouration.  (CASC).

Pheidole longipilosa
Diagnosis. Major workers. Head in full-face view sub-oval; sides of the head with dense, long, erect pilosity; occipital lobes shiny, with thick, irregular rugae, interspaces with distinct, irregular rugulae not fading posteriorly; inner hypostomal teeth distinct, high, closely spaced, triangular, with rounded apex directed inward, and wide base; outer hypostomal teeth taller and wider than inner hypostomal teeth, lobe-like, directed outward; gaster smooth. Minor workers. Head shiny, foveolate, with additional short, longitudinal, thick rugae on frons, genae with fading sculpture and smooth notch; scape, when laid back, surpassing the posterior head margin by one-fifth of its length; promesonotum low, convex, short, with posterior declivity smoothly declining towards propodeum; mesosoma foveolate, katepisternum smooth; propodeal spines short, triangular.
Etymology. Latin for long pilosity, in reference to the very long pilosity on sides of head in major workers.
Biology. The species was collected between 500-1100 m in elevation, in montane and gallery forests. Nests were located in rotten logs.

Revision of the Pheidole annemariae group
Diagnosis. Major workers. Head in full-face view rectangular, slightly longer than wide, in lateral view sub-rectangular; ventral and dorsal faces finely convex; dorsal face finely depressed posteriorly, forming shallow transverse depression; sides of the head with moderately dense, long, erect pilosity; antennal scrobes indistinct and not delimited by carinulae; scrobe surface foveolate to rugo-foveolate, with distinct, thin, moderately sparse to dense, longitudinal rugae; occipital lobes with thick, sparse, irregular rugae, interspaces with fine rugulae fading posteriorly; frons with moderately sparse to dense, thick, longitudinal, and sometimes interrupted rugae, interspaces smooth to rugulose; promesonotum low and arched; posterior mesonotum with distinct teeth-like projections; promesonotal and metanotal grooves absent; propodeal spines long, triangular; mesosoma sculpture well developed, foveolate to rugo-foveolate; gaster finely shagreened; body brown to brownish black. Minor workers. Head foveolate, genae smooth; scape, when laid back, reaching the posterior head margin or surpassing it by two-fifths of its length; promesonotum low, slightly convex, short; mesonotal spines present, small, and triangular; promesonotal groove absent; metanotal groove absent or indistinct; propodeal spines long, triangular; mesosoma foveolate; katepisternum and mesonotum with smooth notches or smooth; body yellow.
Comments. Major workers of this group can be distinguished based on a combination of the following characters: head in full-face view rectangular, and in lateral view sub-rectangular; scrobe surface foveolate to rugo-foveolate, with distinct and thin longitudinal rugae; posterior mesonotum with distinct teeth-like projections, and long, triangular propodeal spines. Minor workers can be separated based on foveolate head and mesosoma, with at least partly smooth genae, katepisternum, and mesonotum; presence of mesonotal spines and long propodeal spines.
Etymology. Conversion of the name of the most similar species, P. annemariae Forel. Biology. The species was collected between 10-550 m in elevation, in rainforest and littoral rainforest. Nests were located in rotten logs, rotten sticks on ground, and soil.
Comments. Pheidole marieannae sp. nov. is most similar to P. annemariae. Major workers. It differs from P. annemariae in absence of smooth notch on genae, inner and outer teeth never connected by indistinct concavity, posterior mesonotum with tubercle-like projections and shorter propodeal spines. Minor workers. It differs from P. annemariae in presence of indistinct mesonotal spines and katepisternum, anepisternum, and mesonotum entirely smooth.

Revision of the Pheidole makaensis group
Diagnosis. Major workers. Head in full-face view elongated, in lateral view suboval to sub-rectangular; ventral and dorsal faces convex; dorsal face not or finely depressed posteriorly; sides of the head with moderately dense, long to very long, erect pilosity; antennal scrobes indistinct and not delimited by carinulae; scrobe surface with thick, sparse, longitudinal rugae, interspaces smooth, rugo-foveolate, or foveolate, with thick, longitudinal, short, and interrupted rugae; occipital lobes always with arcuate and/or transverse rugae; genae smooth or with dense, thin, longitudinal rugulae and smooth notch in the centre; inner and outer teeth closely spaced and connected by concavity; promesonotum relatively low and arched; promesonotal groove absent; metanotal groove absent or indistinct; propodeal spines short to moderately long, triangular; mesosoma foveolate with additional sculpture; promesonotum with sculpture reduced; gaster finely shagreened, at least on the basal part of the first tergite; body dark yellow to brown. Minor workers. Head foveolate; genae smooth or with smooth notch; sometimes frons with additional longitudinal rugae; scape, when laid back, reaching the posterior margin of head or surpassing it by oneto two-fifths of its length; promesonotum low, convex, short; promesonotal groove absent; metanotal groove indistinct; propodeal spines small, triangular; mesosoma foveolate; anepisternum, katepisternum, and mesonotum sometimes with fading sculpture; body yellow to dark yellow.
Comments. Major workers can be distinguished based on a combination of the following characters: head in full-face view elongated; sides of the head with moderately dense, long to very long, erect pilosity; occipital lobes always with arcuate and/ or transverse rugae; genae at least with a smooth notch; inner and outer teeth closely spaced and connected by concavity. Minor workers can be distinguished based on foveolate head and mesosoma, with genae at least with a smooth notch; small, triangular propodeal spines; yellow to dark yellow body colouration.
This species group contains five species: P. makaensis sp. nov., P. fitarata sp. nov., P. rugofitarata sp. nov., P. ehazoara sp. nov., and P. avaratra sp. nov. There are two species of this group known only from their type localities: Pheidole ehazoara sp. nov. collected in Ehazoara Canyon in Toliara and P. makaensis sp. nov. described from Makay Mts. in Toliara. Pheidole fitarata sp. nov. is sympatric with P. avaratra sp. nov. and distribution of both taxa is limited to northernmost parts of the Sambirano rainforest and dry deciduous forest biomes in the Antsiranana prefecture. Pheidole rugofitarata sp. nov. is known from area spread between Belo and Antonibe.

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Major workers. Body yellowish brown to brown, promesonotum high and arched, dorsal mesonotum slightly concave, posterior mesonotum steep, frons with interspaces rugo-foveolae, outer hypostomal teeth dentate with relatively wide base, never directed outward ( (CASC). Diagnosis. Major workers. Head in full-face view elongated; sides of the head with moderately dense, very long, erect pilosity; occipital lobes with thick, sparse, short, irregular, and slightly arcuate rugae, interspaces smooth; inner hypostomal teeth distinct, moderately high, closely spaced, triangular, with rounded apex and wide base; outer hypostomal teeth bigger and wider than inner hypostomal teeth, lobe-like; inner and outer teeth closely spaced and connected by moderately deep concavity. Minor workers. Head foveolate, genae smooth; scape, when laid back, surpassing the posterior head margin by two-fifths of its length; promesonotum low, convex, short, with posterior declivity smoothly declining towards propodeum; mesosoma foveolate, anepisternum, katepisternum and mesonotum with fading sculpture; propodeal spines small, triangular.
Etymology. From the type locality. Biology. The species was collected at an elevation of 500 m, in gallery forest with bamboo. Nest was located in dead twig above ground.
Comments. Pheidole makaensis sp. nov. is most similar to P. avaratra sp. nov. Major workers. It differs from P. avaratra sp. nov. in very sparse and interrupted rugae on frons and frons with mostly smooth interspaces; smooth genae, lobe-like outer hypostomal teeth which are distinctly bigger than inner hypostomal teeth, and katepisternum with smooth notch. Minor workers. It differs from P. avaratra sp. nov. in denser and longer pilosity on head, mesosoma, and gaster. ing indistinct transverse depression between frons and occipital lobes; inner hypostomal teeth visible. Sides of the head with moderately dense, long, erect pilosity; whole head with dense, moderately long, suberect to erect pilosity. Antennal scrobes indistinct and not delimited by carinulae; scrobe surface shiny, foveolate, with thick, longitudinal, short, and interrupted rugae. Occipital lobes shiny, smooth to indistinctly foveolate, with thick, sparse, short, transverse, and sometimes arcuate rugae; frons with moderately dense, thick, longitudinal and interrupted rugae, interspaces smooth and indistinctly foveolate on the posterior part, rugae weakening posteriorly and never connected with rugae on the occipital lobes; genae shiny and smooth; malar area with thick, longitudinal, moderately sparse rugae, interspaces smooth. Centre of clypeus shiny and smooth, lateral sides with longitudinal rugae; median notch present, wide, and shallow; median longitudinal carina absent; lateral longitudinal carinae absent. Scape, when laid back, not reaching the mid-length of head; pilosity suberect to erect (Fig. 46B, D). Inner hypostomal teeth distinct, high, closely spaced, triangular, with rounded apex directed inward; outer hypostomal teeth thinner than inner hypostomal teeth and approximately the same height, triangular (Fig. 84S). Mesosoma. In lateral view, promesonotum relatively low and arched, posterior mesonotum relatively steep, with tubercle-like projections; promesonotal groove absent; metanotal groove indistinct; propodeal spines moderately long, triangular, with sharp apex and wide base; humeral area laterally weakly produced (Fig. 46D). Surface finely foveolate, dorsal surface of promesonotum with fading sculpture and sometimes with smooth notch; katepisternum with smooth notch; lower half of lateral sides of propodeum with additional longitudinal rugae. Pilosity sparse, very long, and erect (Fig. 46D, F). Petiole. Shiny; peduncle moderately long, finely foveolate, with indistinct horizontal lobes on its basal part; node smooth, relatively high, triangular, with rounded apex, in rear view node slightly convex; pilosity moderately sparse and erect (Fig. 46D, F). Postpetiole. Shiny and finely shagreened, dorsum with fading sculpture; in dorsal view sides with moderately short, acute, and triangular projections; pilosity long to moderately long, and erect (Fig. 46D, F). Petiole. Shiny and finely shagreened; pilosity moderately dense, long, and erect (Fig. 46D, F). Colour. Bright brown to brown; legs, lower frons, malar area and lower parts of lateral sides of mesosoma yellowish brown (Fig. 46D, F).  (Fig. 46A). Pilosity moderately dense, long, suberect. Head shiny, foveolate; genae smooth. Clypeus smooth or finely foveolate; median longitudinal carina absent; two lateral longitudinal carinae absent. Scape, when laid back, surpassing the posterior head margin by two-fifths of its length; pilosity suberect to erect (Fig. 46A, C).
Etymology. Malagasy for mirror, in reference to smooth and shiny genae of major workers.
Biology. The species was collected between 10-780 m in elevation, in rainforest, in tropical dry forest, and in littoral rainforest. Nests were located rotten logs.
Comments. Pheidole fitarata sp. nov. is most similar to P. rugofitarata sp. nov. and P. avaratra sp. nov. Major workers. Pheidole fitarata sp. nov. differs from P. rugofitarata sp. nov. in less distinct and interrupted longitudinal rugae on frons and antennal scrobes, lack of connection between rugae on frons and occipital lobes, smooth genae, and never lobe-like outer hypostomal teeth; from P. avaratra sp. nov. in presence of longitudinal, thick, and interrupted rugae on frons and antennal scrobes, frons with rugae weakening posteriorly and never connected with rugae on the occipital lobes, genae smooth, and smaller and directed inward inner hypostomal teeth. Minor workers. Pheidole fitarata sp. nov. differs from P. avaratra sp. nov. in longer and lower promesonotum and katepisternum with smooth notch.  (CASC).
Diagnosis. Major workers. Head in full-face view elongated; sides of the head with moderately dense, long, erect pilosity; occipital lobes indistinctly foveolate, with thick, sparse, short, transverse, and sometimes arcuate rugae; inner hypostomal teeth distinct, high, closely spaced, triangular, with rounded apex; outer hypostomal teeth wider and higher than inner hypostomal teeth, lobe-like.
Minor workers. Unknown. Etymology. Similar to P. fitarata, but with more distinct head sculpture in major workers.
Biology. The species was collected between 10-250 m in elevation, in rainforest and tropical dry forest. Nests were located in rotten logs.
Diagnosis. Major workers. Head in full-face view elongated; sides of the head with moderately dense, long, erect pilosity; occipital lobes with thick, sparse, short, irregular, and arcuate rugae, interspaces superficially rugulae; inner hypostomal teeth distinct, high, closely spaced, triangular, with rounded apex; outer hypostomal teeth thinner than inner hypostomal teeth and approximately the same height, triangular, but thin. Minor workers. Head foveolate, genae with smooth notch; scape, when laid back, surpassing the posterior head margin by one-fifth of its length; promesonotum low, convex, short, with posterior declivity smoothly declining towards propodeum; mesosoma foveolate; propodeal spines short, triangular.
Etymology. Malagasy for north, in reference to the distribution of the species. Biology. The species was collected between 15-210 m in elevation, in rainforest, tropical dry forest, and open secondary vegetation. Nests were located in rotten sticks on ground, under moss, and in soil.
Comments. Pheidole avaratra sp. nov. is most similar to P. fitarata sp. nov. Major workers. It differs from P. fitarata sp. nov. in presence of longitudinal, very thick, and never interrupted rugae on frons and antennal scrobes, presence of rugae connecting frons and occipital lobes, never entirely smooth genae, and bigger and never directed inward inner hypostomal teeth. Minor workers. Pheidole (CASC).
Etymology. After the locus typicus. Biology. The species was collected at 175 m in elevation, in tropical dry rainforest. Nest was located under a stone.
Comments. Pheidole ehazoara sp. nov. is most similar to P. fitarata sp. nov. Major workers. It differs from P. fitarata sp. nov. in dark yellow body colouration, frons with rugae connected with rugae on the occipital lobes, absence of foveolae on head, shorter propodeal spines with wider base, and dentate outer hypostomal teeth directed outward. Minor workers. It differs from P. fitarata sp. nov. in presence of the additional longitudinal rugae on frons and malar area, longer postpetiole, and never smooth genae.

Revision of the Pheidole curvistriata group
Diagnosis. Major workers. Head in full face view rectangular, slightly widening posteriorly, in lateral view sub-rectangular with ventral and dorsal faces finely convex, dorsal face finely depressed posteriorly; antennal scrobes shallowly impressed; occipital lobes with thick, sparse, irregular rugae, interspaces smooth to finely rugulose; frons with thin to thick, sparse to dense, longitudinal rugae, interspaces smooth to indistinctly rugulose; lateral sides of head with fine, irregular rugoreticulation or dense and thin longitudinal rugulae, interspaces finely foveolate; promesonotum short, angular, and relatively low; promesonotal groove absent; metanotal groove absent or indistinct; propodeal spines moderately long, narrow, with base slightly wider than top; mesosoma finely rugoreticulate or foveolate, katepisternum and at least lower anepisternum smooth, lateral sides of propodeum with few thick, longitudinal rugae; first gastral tergite finely shagreened; body brown to dark brown. Minor workers. Head foveolate, with thick, sparse, and longitudinal rugae on frons and sparse, irregular to arcuate, thick rugae on vertex; scape, when laid back, surpassing the posterior head margin by two-fifths of its length; promesonotum low, short, flat, or slightly convex, with steep posterior declivity; promesonotal groove absent; metanotal groove absent or indistinct; propodeal spines short or moderately long, triangular; mesosoma foveolate or foveolate with additional thick and irregular rugae, sometimes anepisternum and katepisternum smooth; body yellow to brown.

Comments.
Major workers of this group can be distinguished based on a combination of the following characters: head in full face view rectangular, slightly widening posteriorly, in lateral view sub-rectangular; occipital lobes with thick, sparse, irregular rugae, interspaces smooth to finely rugulose; moderately long and narrow propodeal spines; finely rugoreticulate or foveolate mesosoma with smooth katepisternum and at least lower part of anepisternum; finely shagreened first gastral tergite, and brown to dark brown body. Minor workers can be easily distinguished based on foveolate head with additional rugae on frons and irregular to arcuate rugae on vertex; foveolate mesosoma usually with additional irregular rugae and yellow to brown body.
Major workers of this group are extremely similar and sometimes the key features overlap. Therefore, we recommend using minor workers, which possess more distinct and stable morphological characters, for species determination within this group.
This group contains four species: P. curvistriata sp. nov., P. makirovana sp. nov., P. mantadia sp. nov., and P. moramanaensis sp. nov. All members of this group are sympatric and, except P. mantadia sp. nov. known only from its type locality (Corridor Forestier Analamay-Mantadia, Toamasina), are distributed across central highlands and evergreen forest.
Diagnosis. Major workers. Head in full face view rectangular, slightly widening posteriorly; sides of the head with dense, relatively long, erect pilosity; frons with fine, irregularly rugoreticulate, interspaces finely foveolate; inner hypostomal teeth distinct, moderately high, thick and triangular, with rounded apex; outer hypostomal teeth thinner but higher, with rounded tips, and wide base; inner and outer teeth closely spaced and connected by indistinct concavity; propodeal spines moderately long, narrow, with base slightly wider than top; first gastral tergite finely shagreened. Minor workers. Whole head foveolate, thick, sparse and longitudinal rugae overlie foveolate sculpture on frons and malar area; sparse, irregular to arcuate, thick rugae overlie foveolate sculpture on vertex; promesonotum low, short, flat or slightly convex, with steep posterior declivity; mesosoma foveolate, with sparse, thick, and irregular rugae overlying foveolae, katepisternum smooth.
Etymology. Latin for arcuate rugae, in reference to head sculpture in minor workers. Biology. The species was collected between 530-1150 m in elevation, in rainforest and montane rainforest. Nests were located in rotten logs, twigs, and stumps.
Etymology. From the type locality. Biology. The species was collected between 865-1075 m in elevation, in transition humid forest, rainforest, and montane rainforest. Nests were located in rotten logs, rotted twigs on the ground, and dead bamboo above the ground.
Comments. Pheidole moramanaensis sp. nov. is most similar to P. curvistriata sp. nov. and P. mantadia sp. nov. Major workers. Pheidole moramanaensis sp. nov. can be distinguished from P. curvistriata sp. nov. by presence of fine foveolate sculpture on frons and mesosoma, low and thick inner hypostomal teeth which are wider than high, and thick outer hypostomal teeth; form P. mantadia sp. nov. by denser and longer pilosity on head sides, and higher and narrower outer hypostomal teeth. Minor workers. Pheidole moramanaensis sp. nov. can be distinguished from P. curvistriata sp. nov. by frons and malar area with thin, longitudinal to irregular rugae, and vertex with thin irregular and never arcuate rugae, absence of irregular rugae on mesosoma, and smooth mesonotum; from P. mantadia sp. nov. by frons and malar area with thin but distinct, irregular rugae and smooth clypeus.  (CASC).
Etymology. From the type locality. Biology. The species was collected between 450-1200 m in elevation, in transitional humid forest, rainforest, and montane rainforest. Nests were located in rotten logs and rotten twigs on the ground.
Etymology. From the type locality. Biology. The species was collected at 939 m in elevation, in rainforest. Nest was located in rotten log.
Comments. Pheidole mantadia sp. nov. is most similar to P. curvistriata sp. nov. and P. moramanaensis sp. nov. Major workers. Pheidole mantadia sp. nov. can be distinguished from P. curvistriata sp. nov. by presence of fine foveolate sculpture on frons and mesosoma, low and thick inner hypostomal teeth, which are wider than high and thick, high outer hypostomal teeth; from P. moramanaensis sp. nov. by sparser and shorter pilosity on sides of head. Minor workers. Pheidole mantadia sp. nov. can be distinguished from P. curvistriata sp. nov. by frons and malar with indistinct, longitudinal rugae, and vertex with indistinct irregular and never arcuate rugae, absence of irregular rugae on mesosoma, and smooth mesonotum; from P. moramanaensis sp. nov. by frons and malar area with indistinct, longitudinal rugae, and never smooth clypeus.

Revision of the Pheidole nemoralis group
Diagnosis. Major workers. Body size small; head in full-face view square or subrectangular, anterior and posterior sides of eyes slightly convex, in lateral view sub-oval or sub-rectangular, dorsal face not depressed or finely depressed posteriorly; antennal scrobes indistinct and not delimited by carinulae; scrobe surface foveolate with sparse, thick, longitudinal to irregular rugae; occipital lobes with indistinct to distinct, sparse, irregular rugae, interspaces smooth to foveolate; frons with dense, thick, and longitudinal rugae, interspaces smooth to rugo-foveolate; promesonotum moderately short, relatively low and convex; promesonotal and metanotal groove absent; propodeal spines small to moderately long, triangular; mesosoma with fine foveolae or rugo-foveolae; gaster smooth or indistinctly shagreened; body bright brown to dark brown. Minor workers. Head foveolate, sometimes with reduced or absent sculpture on genae; scape short, when laid back surpassing the posterior head margin by one to two-fifths of its length; promesonotum low, short or long, slightly convex, with relatively steep posterior declivity or declivity smoothly declining towards propodeum; promesonotal groove absent; mesosoma entirely foveolate or foveolate with smooth notches on its lateral sides; propodeal spines minute to moderately long, triangular; body yellow.
Comments. Major workers of this group can be distinguished based on the combination of the following characters: relatively small body size; square or subrectangular head in full-face view, indistinct and not delimited by carinulae antennal scrobes with scrobe surface foveolate with sparse, thick, longitudinal to irregular rugae; occipital lobes with indistinct to distinct, sparse, irregular rugae with smooth to foveolate interspaces; frons with dense, thick, and longitudinal rugae, moderately short and relatively low promesonotum, lack of promesonotal and metanotal grooves, entirely sculptured mesosoma and bright brown to dark brown body.
The group is divided into two complexes. The P. nemoralis complex contains two sympatric species distributed across evergreen forest and central highlands biomes: P. nemoralis and P. ala sp. nov. The P. bemarivoensis complex also contains two species: P. bemarahaensis sp. nov., and P. bemarivoensis sp. nov. Pheidole bemarahaensis is known only from its type locality (Parc National Tsingy de Bemaraha, Mahajanga) and is sympatric with P. bemarivoensis distributed in lowlands spread between Belo and Ambilobe.

Revision of the Pheidole nemoralis complex
Diagnosis. Major workers. Head in full-face view sub-rectangular, in lateral view subrectangular, ventral and dorsal faces relatively convex and dorsal face finely depressed posteriorly; scrobe surface foveolate with sparse, thick, irregular rugae; occipital lobes with indistinct to distinct, sparse, irregular rugae, interspaces sometimes foveolate, and sculpture sometimes weakening posteriorly; frons with dense, thick, and longitudinal rugae, interspaces distinctly to finely foveolate or rugo-foveolate; propodeal spines small; mesosoma with fine but distinct foveolae or rugo-foveolae, and with additional sparse, fine, and irregular rugae; gaster smooth or finely shagreened; body bright brown to dark brown. Minor workers. Head foveolate; genae with reduced sculpture or smooth; scape, when laid back, surpassing the posterior head margin by one-fifth of its length; promesonotum low, short, slightly convex, with relatively steep posterior declivity or declivity smoothly declining towards propodeum; metanotal groove absent or indistinct; propodeal spines minute to small, triangular; mesosoma foveolate, katepisternum, anepisternum, and mesonotum smooth.
Comments. Major workers of this complex can be distinguished based on a combination of the following characters: head in full-face and lateral views sub-rectangular; foveolate scrobe surface with sparse, thick, irregular rugae; small and triangular propodeal spines; mesosoma with fine but distinct foveolae or rugo-foveolae, and additional sparse, irregular rugae; bright brown to dark brown body. Minor workers can be separated based on foveolate head with genae smooth or with reduced sculpture and smooth katepisternum, anepisternum, and mesonotum; minute to small propodeal spines, and yellow body.
Biology. The species was collected between 10-1620 m in elevation, in montane rainforest, rainforest, montane shrubland, open secondary vegetation, gallery forest, and littoral rainforest. Nests were located in rotten logs and tree stumps, and in dead twigs above ground.
Etymology. Malagasy for forest, in reference to habitat of the species. Biology. The species was collected between 520-1410 m in elevation, in rainforest and montane rainforest. Nests were located in rotten logs, and in rotten twigs on the ground. Diagnosis. Pheidole ala sp. nov. is most similar to P. nemoralis. Major workers. Pheidole ala sp. nov. differs from P. nemoralis in frons with rugo-foveolate sculpture between rugae and never shagreened gaster. Minor workers. Pheidole ala sp. nov. differs from P. nemoralis in mesosoma with posterior declivity smoothly declining towards propodeum, smooth genae, and minute propodeal spines.

Revision of the Pheidole bemarivoensis complex
Diagnosis. Major workers. Body size small; head in full-face view square, in lateral view sub-oval, ventral and dorsal faces convex, dorsal face not depressed posteriorly; scrobe foveolate with sparse, thick, longitudinal to irregular rugulae; occipital lobes foveolate, with indistinct, sparse, irregular rugae; sculpture weakening posteriorly; frons with dense, thick, and longitudinal rugae, interspaces smooth to foveolate; propodeal spines small to moderately long; mesosoma with fine foveolae and with additional rugae; gaster indistinctly shagreened; body brown to dark brown. Minor workers. Head foveolate; scape, when laid back, surpassing the posterior head margin by one to two-fifths of its length; promesonotum low, long, slightly convex, with relatively steep posterior declivity; metanotal groove present; propodeal spines very small or moderately long, triangular; mesosoma foveolate.
Comments. Major workers of this group can be distinguished based on a combination of the following characters: small body size; head in full-face view square and in lateral view sub-oval; foveolate scrobes with additional sparse rugulae; foveolate occipital lobes, with additional sparse, irregular rugae; sculpture weakening posteriorly; indistinctly shagreened gaster and brown to dark brown body. Minor workers can be separated based on small body size, entirely foveolate head and mesosoma; scape, when laid back, surpassing the posterior head margin by one-to two-fifths of its length; long and low promesonotum with relatively steep posterior declivity, and yellow body. 44.9-49.3 (47.1). Head. In full-face view square, anterior and posterior sides of eyes slightly convex (Fig. 60B). In lateral view sub-oval; ventral and dorsal faces convex; dorsal face not depressed posteriorly; inner hypostomal teeth visible. Sides of the head with sparse, long, erect pilosity; whole head with dense, short, suberect to erect pilosity. Antennal scrobes very indistinct and not delimited by carinulae; scrobe surface shiny, finely foveolate with sparse, thick, longitudinal to irregular rugulae in posterior part. Occipital lobes shiny, with fine foveolae and indistinct, sparse, irregular rugae, sculpture weakening posteriorly; frons and malar area with dense, thick, and longitudinal rugae, interspaces superficially foveolate; genae shiny, with fine to indistinct foveolae. Centre of clypeus smooth and shiny, lateral sides with longitudinal rugae; median notch present, narrow, and shallow; median longitudinal carina present; lateral longitudinal carinae present. Scape, when laid back, slightly exceeding the midlength of head; pilosity suberect to erect (Fig. 60B, D). Inner and outer hypostomal teeth distinct, approximately the same size, closely spaced, dentate, with rounded apex and wide base; outer hypostomal teeth with base slightly wider than inner hypostomal teeth (Fig. 84H). Mesosoma. In lateral view, promesonotum moderately short, relatively low, and convex, posterior mesonotum steep, with small tubercle-like projections; promesonotal groove absent; metanotal groove absent; propodeal spines moderately long, triangular, with rounded apex; humeral area laterally well produced (Fig. 60D). Surface shiny, with fine but distinct foveolae, additional sparse, fine, transverse to irregular rugae on promesonotal dorsum and sometimes propodeum, sculpture slightly weakening on dorsal surface. Pilosity moderately sparse, very long, and erect (Fig. 60D, F). Petiole. Shiny and with fine foveolae; peduncle short, with short horizontal lobes on its basal part; node relatively high, triangular, with rounded apex, in rear view node straight or dorsoventrally slightly concave; pilosity moderately sparse and erect (Fig. 60D, F). Postpetiole. Shiny, with fine and sparse foveolae, smooth on dorsum; in dorsal view sides with moderately short, acute, and triangular projections; pilosity moderately long, dense, and erect (Fig. 60D, F). Petiole. Shiny and indistinctly shagreened; pilosity moderately dense, short, and erect (Fig. 60D, F). Colour. Unicolourous, brown to dark brown; legs yellow ( Fig. 60D, F).  (Fig. 60A). Pilosity moderately sparse, long, suberect to erect. Head foveolate. Clypeus with fine and sometimes reduced foveolae; median longitudinal carina absent; two lateral longitudinal carinae absent. Scape, when laid back, surpassing the posterior head margin by one-fifth of its length; pilosity suberect to erect (Fig. 60A, C). Mesosoma. In lateral view, promesonotum low, long, slightly convex, with relatively steep posterior declivity; promesonotal groove absent; metanotal groove indistinct; propodeal spines very small, triangular, apex acute (Fig. 60C). Sculpture foveolate. Pilosity sparse, mod-erately long, and erect (Fig. 60C, E). Petiole. Shiny and with fine foveolae; peduncle short and thin with ventral face slightly convex; node moderately high, triangular, and small; with few short, erect setae (Fig. 60C, E). Postpetiole. Moderately long, low, and slightly convex; with few short, erect setae (Fig. 60C, E). Petiole. With sparse, erect pilosity (Fig. 60C, E). Colour. Unicolourous, yellow (Fig. 60C, E).
Etymology. From the type locality. Biology. The species was collected between 30-600 m in elevation, in rainforest, tropical dry forest, and disturbed dry forest. Nests were located in rotten logs, branches on ground, and soil.
Etymology. From the type locality. Biology. The species was collected at 100 m in elevation, in tropical dry forest. Nest was located in a rotten log.
Comments. Pheidole bemarahaensis sp. nov. is most similar to P. bemarivoensis sp. nov. Major workers. Pheidole bemarahaensis sp. nov. differs from P. bemarivoensis sp. nov. in at least lower part of frons smooth between rugae, more distinct rugo-foveolate sculpture on genae, and thicker dorsal surface of promesonotum. Minor workers. P. bemarahaensis sp. nov. differs from P. bemarivoensis sp. nov. in moderately long propodeal spines.

Revision of the Pheidole petax group
Diagnosis. Major workers. Head in full-face view rectangular or square (except P. boribora), anterior and posterior of eyes relatively straight, in lateral view sub-rectangular, ventral and dorsal faces finely convex or flat and dorsal face finely depressed posteriorly; antennal scrobes shallowly impressed and not delimited (except P. ankerana complex); occipital lobes with thick, sparse, irregular rugae; frons with thick, sparse to moderately sparse, longitudinal rugae; antennal scrobes with finely foveolate interspaces (except P. glabra sp. nov. and P. ankerana complex); promesonotum relatively low and angular; propodeal spines small to moderately long, with wide base; body yellowish brown to black. Minor workers. Head foveolate, sometimes with few additional rugae; genae and frons sometimes with smooth notches; scape short, when laid back, surpassing the posterior head margin by one-to two-fifths of its length; promesonotum slightly convex, with relatively steep posterior declivity; mesosoma foveolate, sometimes with smooth notches on the lateral sides; propodeal spines minute to relatively long, triangular or narrow; body yellow to yellowish brown (except P. brevipilosa complex).
Comments. Major workers of this group can be separated by the combination of the following characters: rectangular or square head in full-face view (except P. boribora) in lateral view sub-rectangular, shallowly impressed antennal scrobes usually not delimited by carinulae (except P. ankerana complex), thick, sparse, irregular rugae on occipital lobes and frons always with thick longitudinal rugae; foveolate antennal scrobes (except P. glabra sp. nov. and P. ankerana complex); relatively low and angular promesonotum and yellowish brown to black body. Minor workers can be separated based on foveolate head, sometimes with few additional rugae and smooth notches on genae and frons, short scape which, when laid back, surpass the posterior head margin by one-to two-fifths of its length, foveolate mesosoma, sometimes with smooth notches on the lateral sides and yellow to yellowish brown body (except P. brevipilosa complex).
The group consists of five complexes. Pheidole petax Forel creates a single species complex common across evergreen forests and central highlands, and is sympatric with all members of the group. The P. brevipilosa complex contains two species: P. brevipilosa sp. nov. and P. glabra sp. nov. Pheidole brevipilosa sp. nov. is known from two localities: Réserve Spéciale Marotandrano, Mahajanga and Forêt de Petriky, Toliara. While P. glabra sp. nov. is known from the area surrounding Tolagnaro, Toliara. The P. mavesatra complex contains two species: P. mavesatra sp. nov., and P. goavana sp. nov. distributed in the northern part of the island. Pheidole mavesatra sp. nov. is known only from its type locality (Tampolo, Toamasina), and P. goavana is distributed exclusively in the Antsiranana prefecture, in the area spread between Ambalabe and Antisiranana. The P. ankerana complex contains two species: P. ankerana sp. nov. known from two localities: Ankerana and Parc National de Zahamena, Toamasina, and P. vatovavensis sp. nov. so far recorded only from Forêt Classée Vatovavy, Fianarantsoa. Finally, the P. boribora complex contains two species known only from their type localities: P. boribora sp. nov. and P. miramila sp. nov. Pheidole boribora sp. nov. was described from Parc National d'Andohahela, Toliara, and P. miramila sp. nov. from Ambalavao, Fianarantsoa.

Revision of the Pheidole petax complex
Diagnosis. Major workers. Head in full-face view rectangular, anterior and posterior of eyes relatively straight, in lateral view sub-rectangular, ventral and dorsal faces finely convex; sides of the head with sparse, short, decumbent pilosity; occipital lobes with interspaces smooth; frons with interspaces smooth to finely foveolate; antennal scrobes with dense, fine rugulae, and sometimes with additional sparse and fine, longitudinal rugae; promesonotal and metanotal grooves absent; propodeal spines small, with wide base; pronotum with sparse, irregular, thin rugae, interspaces smooth to finely foveolate; katepisternum smooth; first gastral tergite shagreened; body brown to dark brown. Minor workers. Head foveolate, genae with smooth notches; scape, when laid back, surpassing the posterior head margin by two-fifths of its length; promesonotum slightly convex, with relatively steep posterior declivity; mesosoma foveolate, katepisternum and, sometimes, lateral sides of propodeum smooth; promesonotal groove absent; metanotal groove indistinct; propodeal spines very short and triangular. Comments. Major workers of this complex can be distinguished based on a compilation of the following characters: head in full-face view rectangular and in lateral view sub-rectangular; sides of the head with sparse, short, and decumbent pilosity; occipital lobes with thick and irregular rugae with smooth interspaces; frons with thick and longitudinal rugae and smooth to finely foveolate interspaces; small propodeal spines with wide base; smooth katepisternum; shagreened first gastral tergite, and brown to dark brown body. Minor workers can be separated based on foveolate head and mesosoma, with smooth katepisternum and genae with smooth notches; promesonotum with relatively steep posterior declivity; very short propodeal spines and yellowish-brown body.
This complex complies only one species, P. petax Forel.
Biology. The species was collected between 64-1343 m in elevation, in rainforest, montane rainforest, transitional humid forest, an urban garden, and grassland. Nests were located in rotten logs and tree stumps, and in dead twigs above ground.
Comments. This species in most similar to P. brevipilosa sp. nov. and P. glabra sp. nov. Major workers. Pheidole petax can be distinguished from P. brevipilosa sp. nov. and P. glabra sp. nov. by never erect pilosity on sides of head, and more rectangular head shape in full-face view. Minor workers. Pheidole petax can be distinguished from P. brevipilosa sp. nov. by yellowish brown body colouration, lack of additional indistinct, longitudinal rugae on frons and malar area, lack of additional sparse, irregular rugae on promesonotal dorsum, and foveolate clypeus; from P. glabra sp. nov. by yellowish brown body colouration, sparser pilosity on head, lack of additional indistinct, longitudinal rugae on frons and malar area, never reduced or absent sculpture on the central part of frons, and dorsal surface of pronotum with never reduced sculpture.

Revision of the Pheidole brevipilosa complex
Diagnosis. Major workers. Head in full-face view square, anterior and posterior of eyes relatively straight, in lateral view sub-rectangular; ventral and dorsal faces relatively flat; occipital lobes with interspaces smooth or superficially rugulose; frons with interspaces smooth to finely rugo-foveolate; promesonotal groove absent; metanotal groove absent or indistinct; propodeal spines moderately long, with base slightly wider than top; mesosoma with fine and sparse to dense rugoreticulation, katepisternum smooth, sometimes pronotum and propodeum partially smooth; first gastral tergite finely shagreened; body brownish black to black. Minor workers. Head foveolate, frons and malar area with few additional, indistinct, longitudinal rugae, genae and sometimes frons with weaker sculpture or smooth; scape, when laid back, surpassing the posterior head margin by one-fifth of its length; promesonotum low, slightly convex, with relatively steep posterior declivity; promesonotal groove absent; metanotal groove indistinct; propodeal spines very short, triangular; mesosoma foveolate; katepisternum smooth and promesonotal dorsum with additional sparse, irregular rugae or katepisternum, anepisternum, and parts of propodeum and pronotum smooth.
Comments. Major workers of this complex can be distinguished based on a combination of the following characters: head in full-face view square and in lateral view sub-rectangular; occipital lobes with thick and irregular rugae with smooth or superficially rugulae interspaces; frons with thick and longitudinal rugae with smooth to finely rugo-foveolate interspaces; moderately long propodeal spines, with base slightly wider than top; katepisternum, pronotum, and propodeum smooth or with reduced sculpture; finely shagreened first gastral tergite; and brownish black to black body. Minor workers can be separated based on the following characters: head foveolate with few additional, indistinct, longitudinal rugae on frons and malar area; genae and sometimes frons smooth or with reduced sculpture; very short and triangular propodeal spines; never entirely foveolate mesosoma; and dark brown body colouration.  -16.28322, 48.81443;alt. 865 m;7 Dec 2007;BLF19163, CASENT0235034 (CASC). Paratype. Madagascar. •1 w.; same data as for holotype; CASENT0134271 (CASC).
Diagnosis. Major workers. Head in full-face view square, anterior and posterior of eyes relatively straight; dorsal face indistinctly depressed posteriorly; sides of the head with dense, short, erect pilosity; malar area, lateral sides of head, and genae with dense, fine rugulae and foveolae; inner hypostomal teeth distinct, big, thick, triangular, with rounded apex; outer hypostomal teeth small and thin, with rounded tips directed outward, lobe-like; propodeal spines moderately long, with base slightly wider than top; first gastral tergite shiny and finely shagreened; body brownish black to black. Minor workers. Whole head foveolate, frons and malar area with few additional, indistinct, longitudinal rugae, genae with weaker sculpture; promesonotum low, slightly convex, with relatively steep posterior declivity; mesosoma foveolate, only katepisternum smooth and promesonotal dorsum with additional sparse, irregular rugae; propodeal spines very small, triangular; body dark brown.
Etymology. Latin for short setae, in reference to short setosity on the head sides in major workers.
Biology. The species was collected between 10-467 m in elevation, in transition humid forest, rainforest, and littoral rainforest. Nests were located in rotten logs.
Comments. This species in most similar to P. petax and P. glabra sp. nov. Major workers. Pheidole brevipilosa sp. nov. can be distinguished from P. petax by presence of erect pilosity on sides of head and lateral sides of head, and square head shape in full-face view; from P. glabra sp. nov. by presence of foveolae sculpture on head, weakly developed outer hypostomal teeth, pronotum with sculpture never reduced and lacking smooth notches on its lateral or dorsal surfaces, and lateral sides of propodeum without smooth notches. Minor workers. Pheidole brevipilosa sp. nov. can be distinguished from P. petax by dark brown body colouration, presence of additional indistinct, longitudinal rugae on frons and malar area, presence of additional sparse, irregular rugae on promesonotal dorsum, and clypeus never foveolate; from P. glabra sp. nov. by sparser and shorter pilosity on head, and frons, dorsal surface of pronotum, promesonotal declivity, and lateral surfaces of propodeum with never reduced sculpture.  -25.08767, 46.749;alt. 450 m;30 Nov 2006;BLF15399, CASENT0122911 (CASC). Paratype. Madagascar. •1 w.; same data as for holotype; CASENT0235037 (CASC).
Diagnosis. Major workers. Head in full-face view square, anterior and posterior of eyes relatively straight; sides of the head with dense, short, erect pilosity; malar area with sparse to moderately dense longitudinal rugae, interspaces smooth to finely rugulose; inner hypostomal teeth distinct, big, thick, triangular, with rounded apex; outer hypostomal teeth slightly but distinctly smaller and thin, with rounded tips, and wide base; propodeal spines moderately long, with base slightly wider than top; first gastral tergite shiny and finely shagreened; body brownish black to black. Minor workers. Whole head with sparse foveolae, frons and malar area with few additional, indistinct, longitudinal rugae, genae and centre of frons with sculpture reduced to absent; promesonotum low, slightly convex, with relatively steep posterior declivity; mesosoma foveolate, reduced on dorsal surface of pronotum, with katepisternum, anepisternum, promesonotal declivity, and lateral surfaces of propodeum smooth; propodeal spines very short, triangular; body dark brown.
Description  (Fig. 68B). In lateral view sub-rectangular; ventral and dorsal faces relatively flat; dorsal face indistinctly depressed posteriorly, forming indistinct transverse depression between frons and occipital lobes; inner hypostomal teeth visible. Sides of the head with dense, short, erect pilosity; whole head with dense, short, suberect to erect pilosity. Antennal scrobes indistinct and not delimited by carinulae; scrobe surface shiny, with dense to sparse, fine, longitudinal to irregular rugae, interspaces smooth to rugulae. Occipital lobes with thick, sparse, irregular rugae, interspaces smooth; frons with thick, sparse to moderately sparse, longitudinal rugae, interspaces smooth; malar area with sparse to moderately dense longitudinal rugae, interspaces smooth to finely rugulose; genae with fine, thin to indistinct rugulae. Clypeus shiny and smooth, with thin, longitudinal rugulae on the lateral sides; median notch present, wide and shallow; median longitudinal carina absent; lateral longitudinal carinae absent. Scape, when laid back, slightly surpassing the mid-length of head; pilosity decumbent to erect (Fig. 68B, D). Inner hypostomal teeth distinct, big, thick, triangular, with rounded apex; outer hypostomal teeth slightly but distinctly smaller and thin, with rounded tips, and wide base (Fig. 84T).
Etymology. Latin for smooth, in reference to reduced sculpture of minor workers. Biology. The species was collected at 450 m in elevation, in rainforest. Nest was located in rotten log.
Comments. This species is most similar to P. petax and P. brevipilosa sp. nov. Major workers. Pheidole glabra sp. nov. can be distinguished from P. petax by presence of erect pilosity on sides of head and lateral sides of head, and square head shape in full-face view; from P. brevipilosa sp. nov. by absence of foveolate sculpture on head, well-developed outer hypostomal teeth, pronotum with reduced sculpture and presence of smooth notches on its lateral and dorsal surfaces, and propodeum with pres-ence of smooth notches on lateral sides. Minor workers. Pheidole glabra sp. nov. can be distinguished from P. petax by dark brown body colouration, presence of additional indistinct, longitudinal rugae on frons and malar area, clypeus never foveolate, and reduced sculpture on frons, dorsal surface of pronotum, promesonotal declivity, and lateral surfaces of propodeum; from P. brevipilosa sp. nov. by denser and longer pilosity on head, and frons, dorsal surface of pronotum, promesonotal declivity, and lateral surfaces of propodeum with sculpture reduced or absent.

Revision of the Pheidole mavesatra complex
Diagnosis. Major workers. Head in full-face view rectangular, anterior of eyes relatively straight, posterior of eyes slightly convex; in lateral view sub-rectangular; ventral and dorsal faces finely convex or relatively flat; scrobe surface shiny with sparse, thin, longitudinal rugae; occipital lobes with interspaces foveolae; frons with interspaces superficially foveolate; genae shiny, with distinct and thin rugulae; promesonotal groove absent; metanotal groove absent or indistinct; propodeal spines moderately long, triangular, with very wide base; mesosoma foveolate with additional thick, sparse, and irregular rugae on promesonotum and propodeum, and katepisternum smooth or mesosoma with thin, moderately dense to dense rugoreticulation with foveolate interspaces; gaster finely shagreened; body brown to brownish black. Minor workers. Head foveolate and genae with reduced sculpture; scape, when laid back, surpassing the posterior head margin by one-fifth of its length; promesonotum low, long, relatively flat or slightly convex, with steep posterior declivity; promesonotal groove absent; metanotal groove indistinct; propodeal spines moderately long to long, triangular or narrow; mesosoma foveolate; katepisternum at least partly, and sometimes mesosoma, smooth.
Comments. Major workers of this complex can be distinguished based on a combination of the following characters: head in full-face view rectangular and in lateral view sub-rectangular; foveolate antennal scrobes with additional thin, longitudinal rugae; occipital lobes with thick and irregular rugae and foveolate interspaces, foveolae fading posteriorly; moderately long and triangular propodeal spines; finely shagreened gaster and brown to brownish black body. Minor workers can be separated based on: foveolate head and mesosoma with reduced sculpture on genae and katepisternum; low and long promesonotum, with steep posterior declivity, and yellow body.  (CASC).
Etymology. Malagasy for heavy, in reference to massive head of major workers. Biology. The species was collected at 218 m in elevation, in disturbed rainforest. Nests were located under moss.
Comments. Pheidole mavesatra sp. nov. is most similar to P. goavana sp. nov. Major workers. Pheidole mavesatra sp. nov. differs from P. goavana sp. nov. in brown to dark brown body colouration, smooth katepisternum, and promesonotum and lateral surfaces of propodeum with additional thick, sparse, and irregular rugae. Minor workers. Pheidole (CASC).
Diagnosis. Major workers. Head in full-face view rectangular, anterior of eyes relatively straight, posterior of eyes slightly convex; sides of the head with sparse, relatively long, suberect pilosity; scrobe surface shiny, foveolate with moderately dense, thin, longitudinal rugae; inner hypostomal teeth distinct, closely spaced, moderately high, triangular, with rounded apex; outer hypostomal teeth thinner and approximately as high as outer hypostomal teeth, dentate, and with relatively narrow base; gaster finely shagreened; body brownish black. Minor workers. Head foveolate, genae with reduced sculpture; promesonotum low, slightly convex, with steep posterior declivity; mesosoma foveolate; lower half of katepisternum smooth; propodeal spines moderately long, triangular, narrow; body yellow.
Etymology. Malagasy for massive, in reference to massive head of major workers.
Biology. The species was collected between 60-1135 m in elevation, in rainforest, and in montane rainforest. Nests were located in rotten logs.
Comments. Pheidole goavana sp. nov. is most similar to P. mavesatra sp. nov. Major workers. Pheidole mavesatra sp. nov. differs from P. goavana sp. nov. in brownish black body colouration, katepisternum never smooth, and promesonotum and lateral surfaces of propodeum with additional thin, moderately dense to dense rugoreticulation. Minor workers. Pheidole mavesatra sp. nov. differs from P. goavana sp. nov. in promesonotum low, long, slightly convex, and triangular propodeal spines.

Revision of the Pheidole ankerana complex
Diagnosis. Major workers. Head in full-face view rectangular, in lateral view sub-rectangular, ventral and dorsal faces finely convex; sides of the head with moderately dense to dense, long, suberect to erect pilosity; scrobe surface shiny, with dense, thick, and irregular rugoreticulate or foveolate with thick, longitudinal rugae; occipital lobes and frons with interspaces smooth or finely rugulose; inner and outer teeth closely spaced and connected by concavity; promesonotal and metanotal grooves absent; propodeal spines moderately long, triangular; mesosoma with fine to thin, dense rugoreticulation and sometimes additional foveolae, sculpture weakening on dorsum, katepisternum smooth; gaster smooth; body yellowish to reddish brown. Minor worker. Whole head foveolate, sometimes with additional thin, sparse rugae; scape, when laid back, surpassing the posterior head margin by one to two-fifths of its length; promesonotum low, short, slightly convex; propodeal spines minute or relatively long, triangular; mesosoma foveolate, sometimes with additional thin, sparse rugae, katepisternum smooth.
Comments. Major workers of this complex can be distinguished based on a combination of the following characters: head in full-face view rectangular, in lateral view sub-rectangular; occipital lobes with thick, sparse, irregular rugae with at least partially smooth interspaces; inner and outer teeth closely spaced and connected by concavity; moderately long propodeal spines; sculpture of mesosoma weakening on dorsum; smooth gaster and yellowish to reddish brown body colouration. Minor workers have foveolate head and mesosoma, sometimes with additional thin rugae, smooth katepisternum, and yellow body.  (CASC).

Pheidole ankerana
Diagnosis. Major workers. Head in full-face view rectangular, with lateral sides relatively straight, only posteriormost part slightly convex; sides of the head with dense, long, suberect pilosity; occipital lobes and genae shiny, with thick, sparse, irregular rugae; frons with sparse, thick, and longitudinal rugae, interspaces smooth; inner hypostomal teeth distinct, small, and low, closely spaced, lobe-like, with rounded apex and wide base; outer hypostomal teeth bigger and wider than inner hypostomal teeth, triangular; inner and outer teeth closely spaced and connected by concavity; propodeal spines moderately long, triangular, with acute apex; gaster smooth; body reddish brown. Minor workers. Head foveolate with additional thin, sparse rugae on the whole surface; promesonotum low, short, slightly convex, with relatively steep posterior declivity; propodeal spines minute, triangular; mesosoma foveolate with additional thin, sparse rugae, katepisternum smooth. In full-face view rectangular, with lateral sides relatively straight, only posteriormost part slightly convex (Fig. 71B). In lateral view sub-rectangular; ventral and dorsal faces finely convex; dorsal face finely depressed posteriorly, forming shallow transverse depression between frons and occipital lobes; inner hypostomal teeth invisible. Sides of the head with dense, long, suberect pilosity; whole head with dense, moderately long, suberect to erect pilosity. Antennal scrobes indistinct and not delimited by carinulae; scrobe surface shiny, with dense, thick, and irregular rugoreticulation. Occipital lobes and genae shiny, with thick, sparse, irregular rugae; frons with sparse, thick, and longitudinal rugae, interspaces smooth; malar area with dense, thick, and longitudinal rugae, interspaces smooth to finely rugulose. Centre of clypeus smooth and shiny, lateral sides with longitudinal rugae; median notch present, narrow and shallow to moderate; median longitudinal carina present; lateral longitudinal carinae absent. Scape, when laid back, slightly exceeding the midlength of head; pilosity decumbent to erect (Fig. 71B, D). Inner hypostomal teeth distinct, small and low, closely spaced, lobe-like, with rounded apex and wide base; outer hypostomal teeth bigger and wider than inner hypostomal teeth, triangular; inner and outer teeth closely spaced and connected by concavity (Fig. 84D).
Etymology. From the type locality. Biology. The species was collected at 175 m in elevation, in rainforest. Nests were located in rotten logs.
Comments. Pheidole vatovavensis sp. nov. is most similar to P. ankerana sp. nov. Major workers. Pheidole vatovavensis sp. nov. differs from P. ankerana sp. nov. in sparser and shorter pilosity on the sides of head, frons with interspaces never smooth, foveolate sculpture on scrobes, higher promesonotum, and triangular shape of inner hypostomal teeth. Minor workers. Pheidole vatovavensis sp. nov. differs from P. ankerana sp. nov. in lack of additional thin, sparse rugae on the head and mesosoma and presence of relatively long propodeal spines.

Revision of the Pheidole boribora complex
Diagnosis. Major workers. Head in full-face view oval, in lateral view sub-rectangular, ventral and dorsal faces finely convex; sides of the head with sparse to moderately dense, short or long, erect pilosity; occipital lobes with interspaces smooth to finely foveolate; frons with interspaces smooth to finely foveolate; lateral sides of head with additional thin, sparse, longitudinal rugae or with thin, dense irregular rugoreticulation; promesonotal groove absent; metanotal groove absent or indistinct; propodeal spines small or moderately long, triangular; mesosoma with very sparse, transverse to irregular thin rugae, sometimes pronotum and propodeum with additional indistinct and sparse foveolae; gaster finely shagreened; body yellowish to reddish brown. Minor workers. Head foveolate; scape, when laid back, surpassing the posterior head margin by two-fifths of its length; promesonotum low, short, slightly convex; mesosoma foveolate, but mesonotum, anepisternum, katepisternum and lateral surfaces of propodeum smooth.
Comments. Major workers of this complex can be distinguished based on a combination of the following characters: head in full-face view oval, in lateral view subrectangular; occipital lobes with thick, sparse, irregular rugae, interspaces smooth to finely foveolate; mesosoma predominately with very sparse, transverse to irregular, thin rugae; finely shagreened gaster and yellowish to reddish brown body. Minor workers can be separated based on the following characters: foveolate head and mesosoma, but sculpture of mesosoma strongly reduced to absent on its lateral sides and yellow body.
Etymology. Malagasy for oval, in reference to the head shape of major workers. Biology. The species was collected at 900 m in elevation, in montane rainforest. Nest was located in a rotten log. Comments. Major workers. Pheidole boribora sp. nov. is most similar to P. miramila sp. nov. and differs from it by presence of foveolae on head, dense and long pilosity of sides of head, dentate inner hypostomal teeth, and small propodeal spines. Minor workers. Pheidole boribora sp. nov. is most similar to P. petax and differs from it by smooth mesonotum, anepisternum, and lateral surfaces of propodeum.  (CASC). Paratype. Madagascar. •1s.; same data as for holotype; CASENT0872079 (CASC).
Diagnosis. Major workers. Head in full-face view oval, relatively as long as wide, anterior and posterior of eyes moderately convex; sides of the head with sparse, short, erect pilosity; genae smooth and shiny; inner hypostomal teeth distinct, low, thick, and triangular, with rounded apex; outer hypostomal teeth higher, thick, dentate, with rounded tips; propodeal spines moderately long, triangular; first gastral tergite finely shagreened; body yellowish brown to brown.
Minor workers. Unknown. Etymology. Malagasy for a soldier, in reference to the fact that this species is known only from its major workers.
Biology. The species was collected at 785 m in elevation, in rainforest. Nesting preferences unknown.
Comments. Pheidole miramila sp. nov. is most similar to P. boribora sp. nov. Major workers. Pheidole miramila sp. nov. differs from P. boribora sp. nov. by absence of foveolae on head, sparse and short pilosity of sides of head, triangular inner hypostomal teeth, and propodeal spines moderately long. Minor workers. Unknown.

Revision of the Pheidole masoala group
Diagnosis. Major workers. Small species; head in full-face view sub-rectangular or oval, anterior and posterior sides of eyes relatively straight or convex; in lateral view sub-rectangular; ventral and dorsal faces relatively flat; dorsal face finely depressed posteriorly (except Pheidole zavamanira); antennal scrobes absent, indistinct or well developed; scrobe surface and lateral sides of head foveolate with or without additional sparse, thick, and irregular to longitudinal rugae or with thick, longitudinal, and long rugae with smooth to indistinctly rugulose interspaces; frons always with thick to thin, longitudinal rugae; occipital lobes with thick irregular rugae, interspaces smooth to rugulae; promesonotum relatively low and evenly angular; promesonotal groove absent; mesosoma never entirely smooth, most often foveolate with additional indistinct, irregular, short rugulae; gaster smooth to finely shagreened; body yellow to black. Minor workers. Head and mesosoma at least partly foveolate, sometimes with additional rugae; genae always with reduced sculpture or smooth; scape, when laid back, reaching the posterior head margin or surpassing it by one-to two-fifths of its length; promesonotum low, long, flat, or slightly convex, with relatively convex to steep posterior declivity; promesonotal groove absent; metanotal groove present; body yellow to dark brown.
Comments. Species of this group have small body size, in both major and minor workers. Major workers can be distinguished by a combination of the following characters: head in full-face and lateral views sub-rectangular or oval with flat ventral and dorsal faces; in most cases antennal scrobes, when present, are indistinctly to distinctly delimited; foveolate sculpture on the lateral sides of head, sometimes with additional rugae (except P. lamperos); frons always with longitudinal rugae and occipital lobes with irregular rugae; relatively low and evenly angular promesonotum; lack of promesonotal groove, and never entirely smooth mesosoma. Minor workers can be distinguished based on at least partly foveolate head and mesosoma, with smooth genae; short scape reaching the posterior head margin or surpassing it by at most two-fifths of its length; low, long, flat, or slightly convex promesonotum, with relatively convex to steep posterior declivity; and lack of promesonotal groove.
Etymology. From the type locality. Biology. The species was collected between 30-520 m in elevation, in rainforest and disturbed rainforest. Nests were located in rotten logs and sticks on the ground.
Etymology. Malagasy for flat, in reference for shape of the head in major workers. Biology. The species was collected between 725-865 m in elevation, in rainforest. Nests were located in rotten logs.

Pheidole andapa
Diagnosis. Major workers. Head in full-face view sub-rectangular, anterior of eyes straight, posterior of eyes convex; sides of the head with sparse, long, erect pilosity; scrobe surface shiny, foveolate with moderately dense, thick, longitudinal rugae; inner hypostomal teeth distinct, closely spaced, moderately high, triangular, with rounded apex; outer hypostomal teeth thinner and approximately as high as outer hypostomal teeth, triangular, and with relatively wide base; propodeal spines moderately long, with base slightly wider than top; gaster smooth; body black. Minor workers. Head foveolate, genae smooth; promesonotum low, slightly convex, short, with posterior declivity steep; mesosoma foveolate; katepisternum and mesosoma smooth; propodeal spines very short, triangular; body dark yellow.
Etymology. From the type locality. Biology. The species was collected at 1325 m in elevation, in montane rainforest. Nest was located in the petiole of Melastomataceae.

Revision of the Pheidole lamperos complex
Diagnosis. Major worker. Head, in full-face, view sub-rectangular, in lateral view sub-rectangular, ventral and dorsal faces relatively flat, dorsal face finely depressed posteriorly; antennal scrobes indistinct and not delimited by carinulae; occipital lobes with thick, irregular rugae, interspaces distinctly rugulose; antennal scrobes and frons with thick, longitudinal, and long rugae, interspaces smooth to indistinctly rugulose; genae with smooth posterior part; metanotal groove absent; propodeal spines moderately long; promesonotum, katepisternum, anepisternum, and dorsoventral surface of propodeum predominantly smooth, with very indistinct, short irregular rugulae; gaster finely shagreened; body black. Minor worker. Head with sparse but distinct foveolae and smooth interspaces, frons with additional longitudinal to irregular, thick rugae, vertex with thick, sparse, and arcuate rugae, genae smooth; scape, when laid back, surpassing the posterior head margin by one-fifth of its length; promesonotum, in lateral view, low, long, and slightly convex; propodeal spines short, triangular; mesosoma with thick and sparse foveolae with smooth interspaces, promesonotum with additional sparse, thick, transverse rugae; body dark brown.
Etymology. Greek for shiny [λαμπερός], in reference to dark and very shiny sculpture of minor and major workers.
Biology. The species was collected at elevation between 980-1100 meters, in montane forest. Nests were located in rotten logs.

Revision of the Pheidole zavamanira complex
Diagnosis. Major workers. Head in full-face view oval, in lateral view sub-rectangular; ventral and dorsal faces relatively flat and dorsal face not depressed posteriorly; antennal scrobes absent; lateral sides of head foveolate with a few thin, irregular to longitudinal short rugae; occipital lobes shiny, with fine and sparse foveolae and sparse, thick, irregular rugae, foveolae fading posteriorly; genae shiny, with dense and thin rugulae, central part smooth; metanotal groove absent; propodeal spines moderately long; mesosoma with fine and dense foveolae and additional indistinct, irregular, short rugulae on promesonotum, katepisternum smooth; first gastral tergite finely shagreened; body bright brown. Minor workers. Head foveolate, frons with a few indistinct, short, longitudinal rugulae; scape, when laid back, surpassing the posterior head margin by two-fifths of its length; promesonotum low, convex, moderately long, with posterior declivity steep; propodeal spines short, triangular; mesosoma foveolate; body smoky yellow.
Etymology. Malagasy for plants, in reference to nesting habits of this species. Biology. The species was collected between 1050-1100 m in elevation, in montane rainforest. Nests were located in rotten logs.