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Monograph
Pheidole Westwood, 1839 (Hymenoptera, Formicidae) of Madagascar – an introduction and a taxonomic revision of eleven species groups
expand article infoSebastian Salata, Brian L. Fisher
‡ Department of Entomology, California Academy of Sciences, San Francisco, United States of America
Open Access

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 aelloea sp. nov., P. ala sp. nov., P. andapa sp. nov., P. ankerana sp. nov., P. avaratra sp. nov., P. bemarahaensis sp. nov., P. bemarivoensis sp. nov., P. binara sp. nov., P. boribora sp. nov., P. brevipilosa sp. nov., P. curvistriata sp. nov., P. diakritos sp. nov., P. ehazoara sp. nov., P. ferruginea sp. nov., P. fisaka sp. nov., P. fitarata sp. nov., P. glabra sp. nov., P. goavana sp. nov., P. lamperos sp. nov., P. longipilosa sp. nov., P. lutea sp. nov., P. madinika sp. nov., P. mahaboensis sp. nov., P. maizina sp. nov., P. makaensis sp. nov., P. makirovana sp. nov., P. manantenensis sp. nov., P. mantadia sp. nov., P. marieannae sp. nov., P. masoala sp. nov., P. mavesatra sp. nov., P. miramila sp. nov., P. moramanaensis sp. nov., P. navoatrensis sp. nov., P. ocypodea sp. nov., P. parviocula sp. nov., P. podargea sp. nov., P. praegrandis sp. nov., P. ranohirensis sp. nov., P. rugocephala sp. nov., P. rugofitarata sp. nov., P. typhlos sp. nov., P. vatovavensis sp. nov., P. voasara sp. nov., P. vohemarensis sp. nov., and P. zavamanira sp. 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.

Keywords

endemic species, Malagasy region, Myrmicinae, taxonomy

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 (2009, 2019). The taxonomy of the genus has been revised for the Nearctic (Wilson 2003), Afrotropic (Fischer et al. 2012), Indomalayan (Eguchi 2001, 2006, 2008; Eguchi and Bui 2005; Eguchi et al. 2016), Australasian (Sarnat et al. 2016), 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, 1892, 1895a, b, 1897; Emery 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 Fisher (1997) 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:

CASC California Academy of Sciences, San Francisco, California, USA;

MHNG Museum d’Historie Naturelle, Geneva, Switzerland.

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 (2009, 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 (2009, 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.

Abbreviations:

m. male;

q. gyne;

s major worker;

w. minor worker.

Synopsis of species of Madagascar examined in this study

Pheidole annemariae group

Pheidole annemariae Forel, 1918

Pheidole marieannae sp. nov.

Pheidole curvistriata group

Pheidole curvistriata sp. nov.

Pheidole makirovana sp. nov.

Pheidole mantadia sp. nov.

Pheidole moramanaensis sp. nov.

Pheidole diakritos group

Pheidole diakritos sp. nov.

Pheidole ensifera group

Pheidole ensifera Forel, 1897

Pheidole ocypodea sp. nov.

Pheidole aelloea sp. nov.

Pheidole podargea sp. nov.

Pheidole ferruginea group

Pheidole longipilosa complex

Pheidole longipilosa sp. nov.

Pheidole ferruginea complex

Pheidole ferruginea sp. nov.

Pheidole rugocephala sp. nov.

Pheidole vohemarensis sp. nov.

Pheidole manantenensis sp. nov.

Pheidole longispinosa group

Pheidole scabrata complex

Pheidole scabrata Forel, 1895 stat. nov.

Pheidole maizina sp. nov.

Pheidole longispinosa complex

Pheidole longispinosa Forel, 1891

Pheidole praegrandis sp. nov.

Pheidole mahaboensis sp. nov.

Pheidole lutea group

Pheidole lutea complex

Pheidole lutea sp. nov.

Pheidole ranohirensis sp. nov.

Pheidole voasara sp. nov.

Pheidole navoatrensis complex

Pheidole navoatrensis sp. nov.

Pheidole parviocula sp. nov.

Pheidole typhlos sp. nov.

Pheidole makaensis group

Pheidole makaensis sp. nov.

Pheidole fitarata sp. nov.

Pheidole rugofitarata sp. nov.

Pheidole ehazoara sp. nov.

Pheidole avaratra sp. nov.

Pheidole masoala group

Pheidole masoala complex

Pheidole masoala sp. nov.

Pheidole madinika sp. nov.

Pheidole fisaka sp. nov.

Pheidole binara sp. nov.

Pheidole andapa sp. nov.

Pheidole lamperos complex

Pheidole lamperos sp. nov.

Pheidole zavamanira complex

Pheidole zavamanira sp. nov.

Pheidole nemoralis group

Pheidole bemarivoensis complex

Pheidole bemarahaensis sp. nov.

Pheidole bemarivoensis sp. nov.

Pheidole nemoralis complex

Pheidole nemoralis Forel, 1892

Pheidole ala sp. nov.

Pheidole petax group

Pheidole petax complex

Pheidole petax Forel, 1895 stat. nov.

Pheidole mavesatra complex

Pheidole mavesatra sp. nov.

Pheidole goavana sp. nov.

Pheidole ankerana complex

Pheidole ankerana sp. nov.

Pheidole vatovavensis sp. nov.

Pheidole boribora complex

Pheidole boribora sp. nov.

Pheidole miramila sp. nov.

Pheidole brevipilosa complex

Pheidole brevipilosa sp. nov.

Pheidole glabra sp. nov.

Figure 1. 

Pheidole moramanaensis sp. nov., illustrations of measurements (A–C) A dorsal view B profile C full-face view D inner hypostomal teeth (IHT) and outer hypostomal teeth (OHT).

Taxonomy

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 (Fig. 2B, C) 2
2 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 (Fig. 2A–C). 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 (Fig. 2A–C). 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) 3
3 Minor & major workers. Antennal sockets deep; frontal lobes distinct and lobe-like (Fig. 4A, B). Major workers. Outer hypostomal teeth absent (Fig. 4C) Pheidole diakritos group*
Minor & major workers. Antennal sockets shallow; frontal lobes absent or indistinct (Fig. 3D–F). Major workers. Outer hypostomal teeth present (Fig. 4D–F) 4
4 Major & minor workers. Relatively large species, propodeal spines moderately to very long, thin and acute. Major workers. Inner and outer hypostomal teeth closely spaced and often connected by concavity (Fig. 4E). Minor workers. At least genae and frons with smooth notches (except P. maizina) (Fig. 5C, D, G, H) 5
Major & minor workers. Smaller species, propodeal spines short to moderately long, with wide or narrow base (Figs 7H, J, 9A–F, 10A–F). Major workers. Inner and outer teeth not closely spaced and if closely spaced, then propodeal spines shorter. Minor workers. Head foveolate, if genae and frons with smooth notches then propodeal spines short (Fig. 9A–F) 6
5 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 two-fifths of its length; promesonotum high, short and convex; petiole with shorter and thicker peduncle (Fig. 5G, H) Pheidole ensifera group*
6 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, 7A–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 the posterior margin of head by at least one-third of its length, promesonotum flat and long, posterior mesonotum steep, propodeal spines minute or relatively long, mesosoma predominately smooth (Fig. 8G–I) 7
7 Major workers. Head, in full-face view, elongate without arcuate or transverse rugae on occipital lobes or head oval; frons and lateral sides of head with thick longitudinal rugae; occipital lobes with irregular rugulae and sculpture weakening posteriorly; promesonotum high and arched; propodeal spines short with wide base (Fig. 6C, D, G). Minor workers. Head never foveolate, at least frons, genae, and malar area smooth and shiny smooth, 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; mesosoma almost entirely smooth; body yellow (Fig. 6F) Pheidole lutea group*
Major workers. Head not oval or elongate, if oval at least one of the following characters present: frons or lateral sides of head with thick irregular rugae, occipital lobes smooth, promesonotum low, propodeal spines relatively long (Fig. 7A–D). Minor workers. Head at least partly foveolate, if foveolae absent then at least one of the following characters present: scape longer, when laid back, surpassing the posterior head margin by at least two-fifths of its length, mesosoma at least partially foveolate, promesonotal groove present and deep, body not yellow (Fig. 8A, B, D–F) 8
8 Major workers. Head, in full-face view, sub-oval to cordate; frons and lateral sides of head predominately with irregular, thick rugae, if frons with longitudinal rugae then antennal scrobes present and well delimited or head in full face view square and in lateral view oval to sub-oval, with thick and sparse longitudinal rugae present on frons and lateral sides of head, promesonotum short, steep and high without promesonotal groove (Fig. 7A–F). Minor workers. Promesonotal groove absent or indistinct. Head and mesosoma predominately foveolate; promesonotum, in lateral view, low, long and convex with indistinct to moderately long propodeal spines (Fig. 8C), if posterior mesonotum steep then propodeal spines long (Fig. 8G); if head and mesosoma predominately smooth, promesonotum short and arched and propodeal spines minute to small (Fig. 8D) 9
Major workers. Head, in full-face view square to sub-rectangular, sometimes with lateral sides slightly convex; frons and lateral sides of had predominately with longitudinal rugae, promesonotum never steep or high, predominately low and arched (Fig. 7G–L). Minor workers. Promesonotal groove present and distinct (Fig. 8A, B, E, F), if absent or indistinct then head and mesosoma predominately foveolate; promesonotum, in lateral view, low, short to long with posterior mesonotum steep and propodeal spines minute to small (Fig. 8H, I) 10
9 Major workers. Head, in full-face view, sub-oval to cordate; frons with longitudinal rugae and antennal scrobes present and well delimited (Fig. 7A, B). Minor workers. Promesonotal groove absent; head and mesosoma predominately foveolate; promesonotum, in lateral view, box-like or convex with posterior mesonotum steep; propodeal spines short to long (Fig. 8G, J) Pheidole ferruginea group*
Major workers. Head, in full-face view, sub-oval; frons and lateral sides of had predominately with irregular, thick rugae; antennal scrobes absent or poorly delimited or head in full face view square and in lateral view oval to sub-oval, with thick and sparse longitudinal rugae present on frons and lateral sides of head (Fig. 7C–F). Minor workers. Promesonotal groove absent or indistinct. Head and mesosoma predominately foveolate; promesonotum, in lateral view, low, long and convex with indistinct to short propodeal spines or head and mesosoma predominately smooth, promesonotum short and arched and propodeal spines minute to small (Fig. 8C, D) Pheidole sikorae group
10 Major workers. Head predominately with fine, sparse longitudinal rugae, sometimes fading posteriorly or head in lateral view oval to sub-oval with thick rugae, distinctly irregular on lateral sides of head with smooth to indistinctly foveolate interspaces; antennal scrobes absent or indistinct; promesonotum low; promesonotal groove present deep or indistinct (Fig. 7G–J). Minor workers. Head elongate, the posterior region of the head elongated into short to long neck, if neck absent then promesonotum, in lateral view, relatively long to long, low and convex and promesonotal groove present (Fig. 8A, E, F, K, L) Pheidole bessonii group
Major workers. Head predominately with fine, sparse longitudinal rugae limited to frons, occipital lobes with irregular rugulae and promesonotal groove absent; if occipital lobes with fine longitudinal rugae then antennal scrobes present and distinct, promesonotum low and short with deep promesonotal groove (Fig. 7K, L). Minor workers. Head not elongated; promesonotal groove absent, if present then promesonotum, in lateral view, short, relatively high and with steep posterior mesonotum (Fig. 8B, H, I) 11
11 Major workers. Promesonotal process well developed; promesonotum low and short; antennal scrobes distinct (Fig. 7K, L). Minor workers. Head and mesosoma predominately smooth and shiny; promesonotal groove present (Fig. 8B) Pheidole fervens group
Major workers. Promesonotal process absent or weakly developed; promesonotum moderately high to high; antennal scrobes absent or present but not delimited by carinulae (Figs 10A–F, 11A–D). Minor workers. Head and mesosoma predominately foveolae, sometimes with smooth notches; promesonotal groove absent (Fig. 8H, I) 12
12 Major & minor workers. Mesonotal spines present (sometimes major workers with teeth-like spines), propodeal spines long (Figs 9A, 10A) Pheidole annemariae group*
Major & minor workers. Mesonotal spines absent (sometimes major workers with bulge-like process), propodeal spines short to moderately long (Figs 9B–F, 10B–F) 13
13 Major workers. Small body size (WL < 0.9 mm); head, in lateral view, sub-rectangular; ventral and dorsal faces relatively flat; antennal scrobes, when present, indistinctly to distinctly delimited (Figs 10B,C, 11A, B). Minor workers. Small body size (WL < 0.6mm); body yellow and foveolate with no additional sculpture (except P. lamperos and P. zavamanira); genae always smooth; propodeal spines minute to small. Minor workers are indistinguishable from those of the nemoralis group (Fig. 9B, C) Pheidole masoala group*
Major workers. Body bigger (WL > 1.0 mm), if WL < 0.9 then head in lateral view, sub-rectangular to sub-oval, with ventral and dorsal faces convex and antennal scrobes not delimited (Figs 10D–F, 11C–E). Minor workers. Body bigger (WL > 0.65mm), if WL < 0.6 then body yellow or dark brown; head and mesosoma entirely foveolate or with distinct smooth notches; sometimes head and/or mesosoma with additional rugae. Note: Minor workers of the masoala group are indistinguishable from those of the nemoralis group (Fig. 9D–F) 14
14 Major workers. Head in full face view rectangular, slightly widening posteriorly, in lateral view sub-rectangular with ventral and dorsal faces finely convex (Figs 10F, 11E). Minor workers. Head foveolate, with thick, sparse, and longitudinal rugae on frons and sparse, irregular to arcuate, thick rugae on vertex; mesosoma foveolate or foveolate with additional thick and irregular rugae; body yellow to brown (Fig. 9F) Pheidole curvistriata group*
Major workers. Head in full face view square to sub-rectangular, never widening posteriorly, in lateral view sub-oval or sob-rectangular with ventral and dorsal faces finely convex (Figs 10D, E, 11C, D). Minor workers. Head and mesosoma foveolate with no additional sculpture, if additional rugae present then body never yellow (except Pheidole ankerana) (Fig. 9D, E) 15
15 Major workers. Body size small (WL < 0.9 mm); head in full-face view square or subrectangular with anterior and posterior sides of eyes slightly convex, in lateral view sub-oval or sub-rectangular; body bright brown to dark brown (Figs 10E, 11D). Minor workers. Small body size (WL < 0.6mm); head foveolate with no additional rugae; mesosoma entirely foveolate or foveolate with smooth notches on its lateral sides; body yellow (Fig. 9E) Pheidole nemoralis group*
Major workers. Body size bigger (WL > 1.1 mm); if WL < 0.9 mm then head, in lateral view not sub-oval, if sub-rectangular then in full-face view, square with anterior and posterior sides of eyes relatively straight and body dark brown to black (Figs 10D, 11C). Minor workers. Body larger (WL > 0.6 mm); if WL < 0.6 mm then head foveolate with additional rugae; mesosoma foveolate with smooth notches and additional indistinct rugae; body dark brown (Fig. 9D) Pheidole petax group*
Figure 2. 

Minor worker, profile. Pheidole oculata Forel (A), Pheidole lutea sp. nov. (B), Pheidole madecassa (C).

Figure 3. 

Pheidole megacephala (Fabricius), minor worker, profile (A) (Shannon Hartman). Major worker, profile (B), head (D). Pheidole longispinosa Forel minor worker, profile (C). Pheidole ferruginea sp. nov. major worker, head (E). Pheidole cf. sikorae major worker, head (F).

Figure 4. 

Pheidole diakritos sp. nov. minor worker, head (B). Major worker, head (A), hypostomal teeth (C). Pheidole megacephala (Fabricius) major worker, hypostomal teeth (D). Pheidole ensifera Forel major worker, hypostomal teeth (E). Pheidole nemoralis Forel major worker, hypostomal teeth (F).

Figure 5. 

Pheidole praegrandis sp. nov. major worker, head (B), profile (A). Minor worker, head (C), profile (D). Pheidole ocypodea sp. nov. major worker, head (F), profile (E). Minor worker, head (G), profile (H).

Figure 6. 

Pheidole fitarata sp. nov. major worker, head (A). Minor worker, profile (E). Pheidole makaensis sp. nov. major worker, head (B). Pheidole navoatrensis sp. nov. major worker, head (C). Pheidole typhlos sp. nov. major worker, head (D), profile (G). Minor worker, profile (F).

Figure 7. 

Pheidole ferruginea sp. nov. major worker, head (A). Pheidole manantenensis sp. nov. major worker, head (B). Pheidole cf. sikorae major worker, head (C). Pheidole veteratrix Forel major worker, head (D). Pheidole cf. veteratrix major worker, head (E), profile (F). Pheidole bessonii Forel major worker, head (G), profile (H). Pheidole cf. bessonii major worker, head (I), profile (J). Pheidole indica Mayr major worker, head (K), profile (L).

Figure 8. 

Pheidole cf. madecassa minor worker, profile (A). Pheidole indica Mayr minor worker, profile (B). Pheidole veteratrix Forel minor worker, profile (C). Pheidole cf. veteratrix minor worker, profile (D). Pheidole cf. bessonii minor worker, profile (E). Pheidole bessonii Forel minor worker, profile (F). Pheidole vahamarensis sp. nov. minor worker, profile (G). Pheidole curvistriata sp. nov. minor worker, profile (H). Pheidole mavesatra sp. nov. minor worker, profile (I). Pheidole rugocephala sp. nov. minor worker, profile (J). Pheidole grallatrix Forel minor worker, profile (K). Pheidole cf. grallatrix minor worker, profile (L).

Figure 9. 

Minor worker, profile. Pheidole annemariae Forel (A). Pheidole masoala sp. nov. (B). Pheidole zavamanira sp. nov. (C). Pheidole petax Forel (D). Pheidole bemarahaensis sp. nov. (E). Pheidole curvistriata sp. nov. (F).

Figure 10. 

Major worker, profile. Pheidole annemariae Forel (A). Pheidole masoala sp. nov. (B). Pheidole zavamanira sp. nov. (C). Pheidole petax Forel (D). Pheidole bemarahaensis sp. nov. (E). Pheidole curvistriata sp. nov. (F).

Figure 11. 

Major worker, head. Pheidole masoala sp. nov. (A). Pheidole zavamanira sp. nov. (B). Pheidole petax Forel (C). Pheidole bemarahaensis sp. nov. (D). Pheidole curvistriata sp. nov. (E).

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 sub-rectangular; 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.

Pheidole diakritos sp. nov.

Figs 12A–F, 84M, 86M

Type material

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

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

Diagnosis

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

Description

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

Figure 12. 

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

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

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

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

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.

Key to the Pheidole lutea group

1 Major workers. Head, in full-face view, elongate; occipital lobes smooth, sometimes with indistinct, very sparse, longitudinal to irregular rugoreticulate; genae smooth (Fig. 13). Minor workers. Head vertex smooth and never with few arcuate, interrupted rugae; scape, when laid back, reaching posterior head margin (Fig. 13) 2
Major workers. Head, in full-face view, oval; occipital lobes and genae with thick, sparse, irregular rugae, interspaces with indistinct foveolae (Fig. 14). Minor workers. Vertex smooth and with few arcuate, interrupted rugae; scape, when laid back, surpassing the posterior head margin by one-fifth of its length (Fig. 14) 3
2 Major workers. Genae and antennal scrobes never foveolate, outer hypostomal teeth approximately as high as inner hypostomal teeth, with very wide base and upper half thin, rectangular, and pointed outward, anepisternum, katepisternum, and mesosoma smooth (Fig. 13A, F, I). Minor workers. Promesonotum in later view evenly arched, high, and short (Fig. 13D) P. lutea sp. nov.
Major workers. Genae and antennal scrobes foveolate, outer hypostomal teeth dentate, smaller and thinner than inner hypostomal teeth and never pointed outward; anepisternum, katepisternum, and mesosoma never entirely smooth (Fig. 13B, G, J). Minor workers. Promesonotum in later view low, short, slightly convex, and with relatively steep posterior declivity (Fig. 13E) P. ranohirensis sp. nov.1
3 Major workers. Eyes well-developed (EL > 0.1), posterior mesonotum never concave, outer hypostomal teeth weakly developed (Fig. 14A, D). Minor workers. Eyes well developed (EL > 0.07), promesonotum convex, and propodeal spines small, triangular (Fig. 14G) P. navoatrensis sp. nov.
Major workers. Eyes small and reduced (EL < 0.1), posterior mesonotum concave, outer hypostomal teeth well developed (Fig. 14B, C, E, F). Minor workers. Eyes small and reduced (EL < 0.06), promesonotum box-like, if convex then propodeal spines indistinct (Fig. 14H, I) 4
4 Major workers. Sides of head with sparse and short pilosity, inner hypostomal teeth distinct, triangular, propodeal spines short and triangular (Fig. 14C, F, K). Minor workers. Promesonotum box-like, propodeal spines distinct, short and triangular (Fig. 14I) P. typhlos sp. nov.
Major workers. Sides of head with dense and relatively long pilosity, inner hypostomal teeth indistinct, lobe-like, propodeal spines absent or weakly developed (Fig. 14B, E, J). Minor workers. Promesonotum convex, propodeal spines weakly developed, indistinct (Fig. 14H) P. parviocula sp. nov.
Figure 13. 

Pheidole lutea sp. nov., head of major worker (A), profile of major worker (F) and minor worker (D), and hypostomal teeth (I). Pheidole ranohirensis sp. nov., head of major worker (B), profile of major worker (G) and minor worker (E), and hypostomal teeth (J). Pheidole voasara sp. nov., head of major worker (C), profile of major worker (H), and hypostomal teeth (K).

Figure 14. 

Pheidole navoatrensis sp. nov., profile of major worker (A), minor worker (G) and hypostomal teeth (D). Pheidole typchlos, head of major worker (K), profile of major worker (C), minor worker (I) and hypostomal teeth (F). Pheidole parviocula sp. nov., head of major worker (J), profile of major worker (B), minor worker (H) and hypostomal teeth (E).

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.

Pheidole lutea sp. nov.

Figs 15A–F, 84Z, 87D

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; B.L. Fisher et al. leg.; BLF07348, CASENT0485691 (CASC). Paratypes. Madagascar. •5w.; same data as for holotype; CASENT0485689, CASENT0485690, CASENT0872151CASENT0872153 (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.

Description

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

Figure 15. 

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

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

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.

Comments

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

Pheidole ranohirensis sp. nov.

Figs 16A–F, 85T, 87D

Type material

Holotype. Madagascar. •1 major worker; Fianarantsoa; Parc National d’Isalo, Sahanafa River, 29.2 km 351°N Ranohira; -22.31333, 45.29167; alt. 500 m; 10 Feb 2003; B.L. Fisher et al. leg.; BLF07681, CASENT0490851, top specimen (CASC). Paratypes. Madagascar. •6w., 5s.; same data as for holotype; CASENT0490850, CASENT0490852CASENT0490854, CASENT0490856, CASENT0872083, CASENT0872206CASENT0872214 (CASC).

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

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.

Description

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

Figure 16. 

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

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

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.

Pheidole voasara sp. nov.

Figs 17A–C, 85Z, 88J

Type material

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

Other material

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

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

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

Figure 17. 

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

Minor workers. Unknown.

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.

Pheidole navoatrensis sp. nov.

Figs 18A–F, 85M, 87R

Type material

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

Other material

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

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.

Description

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

Figure 18. 

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

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

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.

Comment

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

Pheidole parviocula sp. nov.

Figs 19A–F, 85P, 87U

Type material

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

Other material

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

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.

Description

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

Figure 19. 

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

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

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 parviocula sp. nov. can be distinguished from P. typhlos sp. nov. by convex shape of promesonotum convex, presence of promesonotal groove and weakly developed, small, triangular, apex acute.

Pheidole typhlos sp. nov.

Figs 20A–F, 85X, 88H

Type material

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

Other material

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

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

Description

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

Figure 20. 

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

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

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.

Key to the Pheidole longispinosa group

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) 2
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, B, G). Minor workers. Head sculpture smooth or indistinctly rugoreticulate on its central part, genae always smooth (Fig. 21D, E) P. scabrata Forel
Major workers. Genae with rugulae, sometimes posterior part with reduced sculpture, tips of outer hypostomal teeth directed outward; inner hypostomal teeth distinctly bigger than outer hypostomal teeth; katepisternum never with smooth area; body dark brown to black (Fig. 21C, F). Minor workers. Head sculpture rugoreticulate, genae never smooth (Fig. 21H, I) P. maizina sp. nov.
3 Major workers. Head and first gastral tergite at least partially shagreened, body dark brown to black (Fig. 22A, H). Minor workers. Mesosoma rugoreticulate, body dark brown to black (Fig. 15E) P. praegrandis sp. nov.
Major workers. Head and first gastral tergite never shagreened, body brown to dark brown (Fig. 22B–D). Minor workers. Mesosoma smooth or with reduced and indistinct rugulae, body brown to dark brown (Fig. 22F, G) 4
4 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) P. longispinosa Forel
Major workers. Metanotal groove present; frons with longitudinal rugae reaching further than midlength of head (Fig. 22B). Minor workers. Promesonotal groove present, metanotal groove narrow and relatively deep, mesosoma with fine superficial rugulae (Fig. 22F) P. mahaboensis sp. nov.
Figure 21. 

Pheidole scabrata, head and profile of major worker (A, B), head of minor worker (D, E), and hypostomal teeth (G). Pheidole maizina sp. nov., head and profile of major worker (C), head of minor worker (H, I), and hypostomal teeth (F).

Figure 22. 

Pheidole praegrandis sp. nov., head of major worker (A), profile of minor worker (E), and gaster (H). Pheidole mahaboensis sp. nov., head of major worker (B), profile of minor worker (F), and gaster (D). Pheidole longispinosa Forel, head of major worker (C), profile of minor worker (G).

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.

Pheidole longispinosa Forel, 1891

Figs 23A–F, 84Y, 87C

Type material

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

Other material

Madagascar. – Antananarivo: •12w., 2s.; 3 km 41°NE Andranomay, 11.5 km 147°SSE Anjozorobe; -18.47333, 47.96; alt. 1300 m; 5 Dec 2000; B.L. Fisher et al. leg.; CASENT0406929, CASENT0406930, CASENT0413533, CASENT0413537, CASENT0427722, CASENT0427724, CASENT0427725, CASENT0427733. •1 w.; Ambatolaona; -18.928, 47.88283; alt. 1382 m; 19 Feb 2007; B.L. Fisher et al. leg.; CASENT0120674 (CASC). •4w., 1s, 1q.; Mandraka; -18.91813, 47.91717; alt. 1312 m; 20 Mar 2014; B.L. Fisher et al. leg.; CASENT0377078, CASENT0378205, CASENT0378163, CASENT0378568 (CASC). •4w, 1m.; Mandraka Park; -18.9019, 47.90786; 1360 m; 11 Mar 2012; B.L. Fisher et al. leg.; CASENT0275803, CASENT0275804, CASENT0275863 (CASC). •11w., 5s., 1m., 1q; Reg. Analamanga, St. Forestière Mandraka; -18.9183, 47.91687; alt. 1285 m; 25 Mar 2015; B.L. Fisher et al. leg.; CASENT0390404-CASENT0390406, CASENT0390415, CASENT0390429, CASENT0390496, CASENT0390497, CASENT0390506, CASENT0390533, CASENT0720697, CASENT0720833 (CASC); •3w., 1s.; Tsinjoarivo forest, Ankadivory; -19.71572, 47.82084; alt. 1385 m; 23 Aug 2014; B.L. Fisher et al. leg.; CASENT0389650, CASENT0389655, CASENT0389682 (CASC). – Antsiranana: •1 w.; 9.2 km WSW Befingotra, Rés. Anjanaharibe-Sud; -14.75, 49.46667; alt. 1280 m; 5 Nov 1994; B.L. Fisher leg.; CASENT0198233 (CASC). •3w., 1s., 2q; Betaolana Forest, along Bekona River; -14.52996, 49.44039; alt. 880 m; 4 Mar 2009; B.L. Fisher et al. leg.; CASENT0152375, CASENT0152403, CASENT0152410 (CASC).•3w.; Parc National de Marojejy, Manantenina River, 27.6 km 35°NE Andapa, 9.6 km 327°NNW Manantenina; -14.435, 49.76; alt. 775 m; 17 Nov 2003; B.L. Fisher leg.; CASENT0487914 (CASC). – Fianarantsoa: •5w., 2s.; 2 km W Andrambovato, along river Tatamaly; -21.51167, 47.41; alt. 1075 m; 3 Jun 2005; B.L. Fisher et al. leg.; CASENT0060852, CASENT0060853, CASENT0060932, CASENT0060933, CASENT0061669 (CASC). •1w.; 38 km S Ambalavao, Rés. Andringitra; -22.2, 46.96667; alt. 1680 m; 23 Oct 1993; B.L. Fisher leg.; CASENT0198397 (CASC). •3w., 1q.; 40 km S Ambalavao, Rés. Andringitra; -22.21667, 46.96667; alt. 1275 m; 19 Oct 1993; B.L. Fisher leg.; CASENT0198376, CASENT0198377 (CASC). •7w, 1s., 1q.; 8.0 km NE Ivohibe; -22.42167, 46.89833; alt. 1200 m; 3 Nov 1997; B.L. Fisher leg.; CASENT0198369, CASENT0198371, CASENT0198803, CASENT0198868 (CASC). •4w., 3s., 1q.; Parc National Befotaka-Midongy, Papango 28.5 km S Midongy-Sud, Mount Papango; -23.84083, 46.9575; alt. 1250 m; 17 Nov 2006; B.L. Fisher et al. leg.; CASENT0118416, CASENT0119618, CASENT0119592, CASENT0118442 (CASC). •7w., 1s., 1q.; R.S. Ivohibe 8.0 km E Ivohibe; -22.48333, 46.96833; alt. 1200 m; 15 Oct 1997; B.L. Fisher leg.; CASENT0196901, CASENT0198003, CASENT0198004, CASENT0198372, CASENT0198805, CASENT0198807, CASENT0198867 (CASC). – Toamasina: •1w.; [Morano-Chrome forêt, 25 km W]; Morarano Chrome; Amparafaravola; -17.75, 47.98333; alt. 1276 m; 15 Jun 1991; A. Pauly leg.; CASENT0198378 (CASC). •1w., 1s., 1q.; 6.9 km NE Ambanizana, Ambohitsitondroina; -15.56667, 50; alt. 1000 m; 9 Dec 1993; B.L. Fisher leg.; CASENT0198379 (CASC). •1w.; Ambanizana, Parc National Masoala; -15.57167, 50.00611; alt. 800–897 m; 26 Feb 2003; Andriamalala D. et al. leg.; CASENT0047728 (CASC). •8w.; Ankerana; -18.40636, 48.80254; alt. 1108 m; 19 Jan 2012; B.L. Fisher et al. leg.; CASENT0273728, CASENT0274171, CASENT0274172, CASENT0274173, CASENT0274993, CASENT0274994, CASENT0274994 (CASC). •1w., 1s.; Corridor Forestier Analamay-Mantadia, Tsaravoniana; -18.76124, 48.42134; alt. 939 m; 2 Dec 2012; B.L. Fisher et al. leg.; CASENT0300341 (CASC). •3w., 2s.; F.C. Didy; -18.19833, 48.57833; alt. 960 m; 16 Dec 1998; Ratsirarson H.J. leg.; CASENT0198005, CASENT0198006, CASENT0198373 (CASC). •2w., 1s.; Montagne d’Anjanaharibe, 19.5 km 27°NNE Ambinanitelo; -15.17833, 49.635; alt. 1100 m; 12 Mar 2003; B.L. Fisher et al. leg.; CASENT0037455, CASENT0037462, CASENT0048698, CASENT0048727 (CASC). •1w.; P.N. Mantadia; -18.79167, 48.42667; alt. 895 m; 25 Nov 1998; Ratsirarson H.J. leg.; CASENT0198384 (CASC). •1w., 1s.; Parc National d´ Andasibe-Mantadia, Forêt de Mantadia, 25.7 km 248° Moramanga; -18.81402, 48.43028; alt. 1040 m; 13 Jul 2006; Raharimalala & Blaimer leg.; CASENT0117442 (CASC). •1w., 1s.; Parc National de Zahamena, Onibe River; -17.75908, 48.85468; alt. 780 m; 21 Feb 2009; B.L. Fisher et al. leg.; CASENT0152053 (CASC). – Toliara: •7w., 1s., 1q.; 13 km NW Enkara, Rés Andohahela; -24.55, 46.8; alt. 1300 m; 2 Dec 1992; B.L. Fisher leg.; CASENT0198374, CASENT0198375, CASENT0198381, CASENT0198382 (CASC). •31w., 14s., 9q.; Anosy Region, Anosyenne Mts, 29.33 km NW Manantenina; -24.13993, 47.07418; alt. 540 m; 21 Feb 2015; B.L. Fisher et al. leg.; CASENT0704727, CASENT0704727, CASENT0704749, CASENT0704780, CASENT0723465, CASENT0704473, CASENT0704200, CASENT0704201, CASENT0704205, CASENT0704206, CASENT0704217, CASENT0704218, CASENT0704226, CASENT0704227, CASENT0704228, CASENT0704229, CASENT0704242, CASENT0704245, CASENT0704273, CASENT0704353, CASENT0704354, CASENT0704355, CASENT0704393, CASENT0704394, CASENT0704867, CASENT0704874, CASENT0704886, CASENT0704887, CASENT0705875, CASENT0705877, CASENT0721001, CASENT0721005, CASENT0721007, CASENT0721009, CASENT0721013 (CASC). •20w., 2s.; Parc National d’Andohahela, Col du Sedro, 3.8 km 113°ESE Mahamavo, 37.6 km 341°NNW Tolagnaro; -24.76389, 46.75167; alt. 900 m; 21 Jan 2002; B.L. Fisher et al. leg.; CASENT0078380, CASENT0430695, CASENT0430759, CASENT0451281, CASENT0451303, CASENT0451309, CASENT0456249, CASENT0460156, CASENT0460158, CASENT0483964, CASENT0484129, CASENT0484130 (CASC). •3w.; Parc National de Kirindy Mite, 16.3 km 127°SE Belo sur Mer; -20.79528, 44.147; alt. 80 m; 6 Dec 2001; B.L. Fisher et al. leg.; CASENT0477205 (CASC). •5w. 2s.,1q.; Réserve Spéciale Kalambatritra, Ambinanitelo; -23.4502, 46.45658; alt. 1325 m; 11 Feb 2009; B.L. Fisher et al. leg.; CASENT0148863, CASENT0148866, CASENT0149937CASENT0149939 (CASC). •18w., 7s., 3m., 11q.; Réserve Spéciale Kalambatritra, Ampanihy; -23.463, 46.47057; alt. 1269 m; 10 Feb 2009; B.L. Fisher et al. leg.; CASENT0148871, CASENT0148872, CASENT0148875, CASENT0148945, CASENT0148953, CASENT0148964, CASENT0148967, CASENT0148972, CASENT0148983, CASENT0148994, CASENT0148995, CASENT0148996, CASENT0149001, CASENT0149008, CASENT0149801, CASENT0149992, CASENT0150511, CASENT0150535, CASENT0151012, CASENT0151013, CASENT0151019, CASENT0151020, CASENT0152223, CASENT0153227, CASENT0235023 (CASC). •6w., 3s., 2q.; Réserve Spéciale Kalambatritra, Befarara; -23.4178, 46.4478; alt. 1390 m; 7 Feb 2009; B.L. Fisher et al. leg.; CASENT0149598, CASENT0149599, CASENT0149869, CASENT0149871, CASENT0150723 (CASC). •1w., 1s.; Réserve Spéciale Kalambatritra, Betanana; -23.4144, 46.459; alt. 1360 m; 8 Feb 2009; B.L. Fisher et al. leg.; CASENT0148654 (CASC).

Diagnosis

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

Redescription

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

Figure 23. 

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

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

Biology

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

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.

Pheidole mahaboensis sp. nov.

Figs 24A–F, 85B, 87F

Type material

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

Other material

Madagascar. – Antsiranana: •7w., 4s, 2q., 1m; Masoala National Park; -15.3014, 50.22776; alt. 280 m; 7 Mar 2014; B.L. Fisher et al. leg.; CASENT0376080, CASENT0376081, CASENT0377018, CASENT0377035, CASENT0377036, CASENT0377038, CASENT0377568 (CASC). – Fianarantsoa: •1w., 1s.; 2 km W Andrambovato, along river Tatamaly; -21.51167, 47.41; alt. 1075 m; 3 Jun 2005; B.L. Fisher et al. leg.; CASENT0060947 (CASC). •4w., 2q; 40 km S Ambalavao, Rés. Andringitra; -22.21667, 46.96667; alt. 1275 m; 19 Oct 1993; B.L. Fisher leg.; CASENT0198822, CASENT0198823, CASENT0198815, CASENT0198831 (CASC). •5w., 3s., 2q.; 45 km S Ambalavao; -22.21667, 47.01667; alt. 720 m; 26 Sep 1993; B.L. Fisher leg.; CASENT0198830, CASENT0198832, CASENT0198824, CASENT0198829 (CASC). •14w., 7s., 1q.; 7.6 km 122° Kianjavato, Forêt Classée Vatovavy; -21.4, 47.94; alt. 175 m; 6 Jun 2005; B.L. Fisher et al. leg.; CASENT0059894, CASENT0059895, CASENT0059940, CASENT0059941, CASENT0059944, CASENT0060019, CASENT0060020, CASENT0060166, CASENT0060167, CASENT0060298, CASENT0060495, CASENT0061087, CASENT0061148, CASENT0061233 (CASC). •10w., 1s.; 8.0 km NE Ivohibe; -22.42167, 46.89833; alt. 1200 m; 3 Nov 1997; B.L. Fisher et al. leg.; CASENT0198821, CASENT0009788, CASENT0198231, CASENT0198806, CASENT0198825, CASENT0198871, CASENT0198874 (CASC). •2w., 1s.; Belle Vue trail, Ranomafana National Park, Fianarantsoa Prov.; -21.2665, 47.42017; alt. 1020 m; 23 May 2002; Harin’Hala leg.; CASENT0079551, CASENT0080544, CASENT0080545, CASENT0080546, CASENT0112227, CASENT0112304, CASENT0112699, CASENT0112768, CASENT0112778, CASENT0113028, CASENT0113154, CASENT0113191, CASENT0113192, CASENT0113227, CASENT0113325, CASENT0113341, CASENT0113369, CASENT0113399, CASENT0113408, CASENT0113411, CASENT0113816, CASENT0113817, CASENT0114099, CASENT0114211, CASENT0114327, CASENT0114751, CASENT0114767 (CASC). •4w.; Fitovavy Fitovinany Region, District of Ifanadiana Belle vue area1200 m S of Ranomafana National Park entrance; -21.2665, 47.42017; alt. 1018 m; 28 May 2003; Rin’ha leg.; CASENT0114066, CASENT0114710, CASENT0112925, CASENT0113635 (CASC). •8w., 6s.; Forêt de Vevembe, 66.6 km 293° Farafangana; -22.791, 47.18183; alt. 600 m; 24 Apr 2006; B.L. Fisher et al. leg.; CASENT0070669, CASENT0070706, CASENT0070718, CASENT0070740, CASENT0070753, CASENT0071145, CASENT0108002, CASENT0108034 (CASC). •1w.; Ifanadiana Pref: Ranomafana S.-Pref: Ranomafana, Talatakely site near R. Namorona; -21.26806, 47.4247; alt. 967 m; 5 Dec 2004; Lees et al. leg.; CASENT0056933 (CASC). •3w.; JIRAMA water works near river, Ranomafana National Park; -21.2485, 47.45217; alt. 690 m, 28 Jan 2002; Harin’Hala leg.; CASENT0114140, CASENT0114293, CASENT0114295 (CASC). •12w., 6s., 1q.; Parc National Befotaka-Midongy, Papango 27.7 km S Midongy-Sud, Mount Papango; -23.83517, 46.96367, alt. 940 m, 14 Nov 2006; B.L. Fisher et al. leg.; CASENT0119097, CASENT0119108, CASENT0119110, CASENT0119111, CASENT0119113, CASENT0119121, CASENT0119424, CASENT0119441, CASENT0125642, CASENT0128545, CASENT0128694, CASENT0128699 (CASC). •1w.; Parc National Befotaka-Midongy, Papango 28.5 km S Midongy-Sud, Mount Papango; -23.84083, 46.9575; alt. 1250 m; 17 Nov 2006; B.L. Fisher et al. leg.; CASENT0128569 (CASC). •129w., 19s., 1q.; Parc National de Ranomafana, Vatoharanana River, 4.1 km 231°SW Ranomafana; -21.29, 47.43333; alt. 1100 m; 27 Mar 2003; Fisher et al. leg.; CASENT0039679, CASENT0039915, CASENT0039916, CASENT0039920, CASENT0039923, CASENT0039930, CASENT0039941, CASENT0039951, CASENT0039954, CASENT0039959, CASENT0039963, CASENT0039967, CASENT0039967, CASENT0039970, CASENT0039978, CASENT0040051, CASENT0040056, CASENT0040058, CASENT0040075, CASENT0040079, CASENT0040084, CASENT0040181, CASENT0040200, CASENT0040215, CASENT0040232, CASENT0040235, CASENT0040238, CASENT0040240, CASENT0040244, CASENT0040251, CASENT0040253, CASENT0040264, CASENT0040270, CASENT0040326, CASENT0040331, CASENT0040336, CASENT0040337, CASENT0040344, CASENT0040350, CASENT0040360, CASENT0040415, CASENT0040421, CASENT0049613, CASENT0050290, CASENT0073552, CASENT0488608, CASENT0488609, CASENT0488627, CASENT0488640, CASENT0488641, CASENT0488642, CASENT0488643, CASENT0488659, CASENT0488662, CASENT0488677, CASENT0488678, CASENT0488679, CASENT0488680, CASENT0497159, CASENT0497161, CASENT0497211CASENT0497216, CASENT0497327CASENT0497333, CASENT0497372CASENT0497377, CASENT0497412CASENT0497416, CASENT0497428, CASENT0497429, CASENT0497595CASENT0497598, CASENT0497643CASENT0497646, CASENT0497685 (CASC). •11w., 1s.; Parc Nationale Ranomafana: Talatakely; -21.24833, 47.42667; 9 Apr 1998; Griswold et al. leg.; CASENT0096608, CASENT0096609, CASENT0096712, CASENT0096713, CASENT0096714, CASENT0096765, CASENT0096770, CASENT0096771, CASENT0096772, CASENT0096774, CASENT0096775, CASENT0097967 (CASC). •3w., 4s.; R.S. Ivohibe 8.0 km E Ivohibe; -22.48333, 46.96833; alt. 1200 m; 15 Oct 1997; B.L. Fisher et al. leg.; CASENT0198804, CASENT0198872 (CASC). •2w., 1s.; R.S. Ivohibe, 7.5 km ENE Ivohibe; -22.47,46.96; alt. 900 m; 7 Oct 1997; B.L. Fisher et al. leg.; CASENT0198232, CASENT0198802, CASENT0198869 (CASC). •21w., 1s.; radio tower, Ranomafana National Park; -21.25833, 47.40717; alt. 1130 m; 14 Jun 2002; Irwin & Harin’Hala leg.; CASENT0052853, CASENT0053113, CASENT0053232, CASENT0053238, CASENT0078772, CASENT0079894, CASENT0080504, CASENT0111901, CASENT0112824, CASENT0112862, CASENT0113134, CASENT0113293, CASENT0113643, CASENT0113751, CASENT0114008, CASENT0114062, CASENT0114517, CASENT0114536, CASENT0114537, CASENT0114642, CASENT0114661 (CASC). •3w.; Ranomafana; -21.25, 47.36667; 1 Mar 1994; A. Pauly leg.; CASENT0096206-CASENT0096208 (CASC). •2w.; Ranomafana National Park, Talatakely area, 0.4 km WSW of Park Entrance; -21.41667, 47.68333; alt. 900 m; 2 Jan 2001; Kavanaugh leg.; CASENT0007643, CASENT0007644 (CASC). •3w.; Ranomafana National Park, Talatakely; Sahambavy; Fianarantsoa Rural; -21.451179, 47.3023894; alt. 1139 m; 30 Oct 1998; Lee & Ribardo leg.; CASENT0198826 (CASC). •1w.; research cabin at Talatakely, Ranomafana National Park; -21.25041, 47.41945; alt. 900 m; 11 Apr 1998; Irwin & Schlinger leg.; CASENT0198817 (CASC). •9w., 4s., 2q.; Réserve Forestière d’Agnalazaha, Mahabo, 42.9 km 215° Farafangana; -23.19383, 47.723; alt. 20 m; 19 Apr 2006; B.L. Fisher et al. leg.; CASENT0070855CASENT0070858, CASENT0070976, CASENT0070977, CASENT0071584, CASENT0072846, CASENT0072853 (CASC). •17w., 8s., 1q.; Réserve Spéciale Manombo 24.5 km 228° Farafangana; -23.01583, 47.719; alt. 30 m; 21 Apr 2006; B.L. Fisher et al. leg.; CASENT0071522CASENT0071525, CASENT0071528, CASENT0071547, CASENT0071605, CASENT0071606, CASENT0072089, CASENT0072093, CASENT0072098, CASENT0073134, CASENT0073136, CASENT0073142, CASENT0073153, CASENT0073157, CASENT0108512, CASENT0108514, CASENT0108519, CASENT0108527 (CASC). •13w., 1s.; Vohiparara broken bridge; -21.22617, 47.36983; alt. 1110 m; 25 Jul 2002; Harin’Hala leg.; CASENT0052215, CASENT0078742, CASENT0079658, CASENT0079845, CASENT0111141, CASENT0111855, CASENT0111955, CASENT0111993, CASENT0112046, CASENT0112070, CASENT0112157, CASENT0112176, CASENT0112606, CASENT0113065 (CASC). – Toamasina: •2w., 1s.; 6.2 km SSE Ambanizana, Be Dinta; -15.66667, 49.99806; alt. 600 m; 20 Nov 1993; B.L. Fisher leg.; CASENT0198816 (CASC). •2w., 1s., 1q.; Analalava, 7.0 km 255° Mahavelona; -17.7095, 49.454; alt. 50 m; 27 Nov 2005; B.L. Fisher et al. leg.; CASENT0067391, CASENT0067392 (CASC). •1w.; F.C. Andriantantely; -18.695, 48.81333; alt. 530 m; 4 Dec 1998; Ratsirarson leg.; CASENT0198230 (CASC). •2w.; F.C. Sandranantitra; -18.04833, 49.09167; alt. 450 m; 21 Jan 1999; Ratsirarson leg.; CASENT0198808 (CASC). •2w., 2s.; Mahavelona (Foulpointe); -17.66667, 49.5; 25 Dec 1993; A. Pauly leg.; CASENT0096124, CASENT0096181, CASENT0096365, CASENT0096366 (CASC). •1w., 1s., 1 q.; Manakambahiny, near Vavatenina Forest; -17.46667, 49.35; 9 Feb 1995; A. Pauly leg.; CASENT0094829, CASENT0095095, CASENT0095102 (CASC). •3q.; Montagne d’Anjanaharibe, 18.0 km 21°NNE Ambinanitelo; -15.18833, 49.615; alt. 470 m; 8 Mar 2003; Fisher et al. leg.; CASENT0495177 (CASC). •1w.; Parc National de Zahamena, Onibe River; -17.75908, 48.85468; alt. 780 m; 21 Feb 2009; B.L. Fisher et al. leg.; CASENT0152016 (CASC). •6w., 4s.; Parc National Mananara-Nord, 7.1 km 261° Antanambe; -16.455, 49.7875; alt. 225 m; 15 Nov 2005; B.L. Fisher et al. leg.; CASENT0067356, CASENT0067462, CASENT0067464, CASENT0067473, CASENT0067601, CASENT0071327 (CASC). •4w., 2s.; Reserve Betampona, Camp Rendrirendry 34.1 km 332° Toamasina; -17.924, 49.19967; alt. 390 m; 30 Nov 2005; B.L. Fisher et al. leg.; CASENT0067806, CASENT0068019, CASENT0071898, CASENT0072529 (CASC). •40w., 7s., 5q.; Réserve Spéciale Ambatovaky, Sandrangato River; -16.81745, 49.2925; alt. 400 m; 26 Feb 2010 B.L. Fisher et al. leg.; CASENT0160439, CASENT0160440, CASENT0161071, CASENT0161072, CASENT0161084, CASENT0161085, CASENT0161474, CASENT0161475, CASENT0161806-CASENT0161808; CASENT0161881, CASENT0161953, CASENT0161954, CASENT0162120, CASENT0162121, CASENT0162670, CASENT0163018, CASENT0163071, CASENT0163072, CASENT0163114, CASENT0163138, CASENT0163160, CASENT0163552, CASENT0163614, CASENT0163667, CASENT0163871, CASENT0163941, CASENT0163985, CASENT0164288, CASENT0164294, CASENT0164299, CASENT0164304, CASENT0164319, CASENT0164433, CASENT0164441, CASENT0164456, CASENT0164469, CASENT0164487, CASENT0164496 (CASC). – Toliara: •1w., 1s.; 10 km NW Enakara, Rés. Andohahela; -24.56667, 46.81667; alt. 430 m; 22 Nov 1992; B.L. Fisher leg.; CASENT0198809 (CASC). •4w., 1s.; 10 km NW Enakara, Rés. Andohahela; -24.56667, 46.81667; alt. 425 m; 24 Nov 1992; B.L. Fisher leg.; CASENT0198811, CASENT0709102 (CASC). •4w., 1s.; 11 km NW Enakara, Rés. Andohahela; -24.56667, 46.83333; alt. 800 m; 20 Nov 1992; B.L. Fisher leg.; CASENT0198812, CASENT0198813 (CASC). •2w.; 13 km NW Enakara, Rés. Andohahela; -24.55, 46.8; alt. 1250 m; 30 Nov 1992; B.L. Fisher leg.; CASENT0198380, CASENT0198814 (CASC). •6w., 2s., 1q.; 2.7 km WNW 302° Ste. Luce; -24.77167, 47.17167; alt. 20 m; 9 Dec 1998; B.L. Fisher et al. leg.; CASENT0198818CASENT0198820, CASENT0198870 (CASC). •12w., 4s., 2m.; Anosy Region, Anosyenne Mts, 29.33 km NW Manantenina; -24.13993, 47.07418; alt. 540 m; 21 Feb 2015; B.L. Fisher et al. leg.; CASENT0704710, CASENT0705814, CASENT0723419, CASENT0723782, CASENT0723788, CASENT0723798, CASENT0723801, CASENT0723811, CASENT0723833, CASENT0704455, CASENT0704456, CASENT0704499, CASENT0704520 (CASC). •2w., 1s., 1m.; Anosy Region, Parc National Andohahela, Col de Tanatana; -24.7585, 46.85367; alt. 275 m; 7 Mar 2015; B.L. Fisher et al. leg.; CASENT0724259, CASENT0724260 (CASC). •5w., 1s.; Cul du Marosohy; -24.55, 46.83333; alt. 600 m; 14 Nov 1992; B.L. Fisher leg.; CASENT0198810, CASENT0198827 (CASC). •8w., 6s., 1q.; Forêt Ivohibe 55.0 km N Tolagnaro; -24.569, 47.204; alt. 200 m; 3 Dec 2006; B.L. Fisher et al. leg.; CASENT0122618, CASENT0121993, CASENT0122577, CASENT0122635, CASENT0122790, CASENT0122792, CASENT0122793, CASENT0122800 (CASC). •4w., 4s.; Grand Lavasoa, 25.9 km W Tolagnaro; -25.08767, 46.749; alt. 450 m; 30 Nov 2006; B.L. Fisher et al. leg.; CASENT0122875, CASENT0122896, CASENT0122907, CASENT0122908 (CASC). •1w., 1q.; Manatantely, 8.9 km NW Tolagnaro; -24.9815, 46.92567; alt. 100 m; 27 Nov 2006; B.L. Fisher et al. leg.; CASENT0125791, CASENT0125793 (CASC). •3w., 2s.; Parc National Andohahela, Col de Tanatana, 33.3 km NW Tolagnaro; -24.7585, 46.85367; alt. 275 m; 22 Nov 2006; B.L. Fisher et al. leg. CASENT0121898, CASENT0121900, CASENT0121917 (CASC).

Diagnosis

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

Description

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

Figure 24. 

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

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

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.

Pheidole praegrandis sp. nov.

Figs 25A–F, 85S, 88C

Type material

Holotype. Madagascar. •1 major worker; Antsiranana; Parc National de Marojejy, Manantenina River, 27.6 km 35°NE Andapa, 9.6 km 327°NNW Manantenina; -14.435, 49.76; alt. 775 m; 15 Nov 2003; B.L. Fisher et al. leg.; BLF08889, CASENT0494942, top specimen on the pin (CASC). Paratypes. Madagascar. •9w., 2s.; same data as for holotype; CASENT0494952, CASENT0872086, CASENT0494943-CASENT0494945, CASENT0872223CASENT0872229 (CASC).

Other material

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

Diagnosis

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

Description

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

Figure 25. 

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

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

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.

Pheidole scabrata Forel, 1895, stat. nov.

Figs 26A–F, 85W, 88G

Type material

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

Other material

Madagascar. –Antsiranana: •4w., 4q.; 9.2 km WSW Befingotra, Rés. Anjanaharibe-Sud; -14.75, 49.46667; alt. 1200 m; 9 Nov 1994; B.L. Fisher leg.; CASENT0198388, CASENT0198392, CASENT0198395, CASENT0198396 (CASC). •32w., 7s.; Forêt de Binara, 9.4 km 235°SW Daraina; -13.26333, 49.6; alt. 1100 m; 5 Dec 2003; B.L. Fisher leg.; CASENT0043382, CASENT0043383, CASENT0494064CASENT0494066, CASENT0494103CASENT0494105, CASENT0494126CASENT0494128, CASENT0494134CASENT0494136, CASENT0494152 (CASC). •4w.; R.S. Manongarivo, 14.5 km 220°SW Antanambao; -13.99833, 48.42833; alt. 1175 m; 20 Oct 1998; B.L. Fisher leg.; CASENT0198007, CASENT0198386 (CASC). •3w.; R.S. Manongarivo, 14.5 km 220°SW Antanambao; -14, 48.43167; alt. 1220 m; 18 Oct 1998; B.L. Fisher leg.; CASENT0198387 (CASC). •1w.; R.S. Manongarivo, 14.5 km 220°SW Antanambao; -13.99833, 48.42833; alt. 1175 m; 20 Oct 1998; B.L. Fisher leg.; CASENT0198767 (CASC). –Mahajanga: •2w.; Réserve Spéciale Marotandrano, Marotandrano 48.3km S Mandritsara; -16.28322, 48.81443; alt. 865 m; 6 Dec 2007; B.L. Fisher et al. leg.; CASENT0140632, CASENT0140653 (CASC). –Toamasina: •1w., 1s.; 23 km E Moramanga; -18.98028, 48.45306; alt. 900 m; 10 Nov 1996; B.L. Fisher leg.; CASENT0198385 (CASC). •1w.; Ambatovy, 12.4 km NE Moramanga; -18.85813, 48.28488; alt. 1040 m; 5 Mar 2007; B.L. Fisher et al. leg.; CASENT0124242 (CASC). •2w., 2s.; Bevolota 17.1 km N Andasibe; -18.77071, 48.43164; alt. 995 m; 12 Dec 2007; B.L. Fisher et al. leg.; CASENT0135100, CASENT0235126 (CASC). •2w., 1s.; Forêt Ambatovy, 14.3 km 57° Moramanga; -18.85083, 48.32; alt. 1075 m; 21 Mar 2004; B.L. Fisher et al. leg.; CASENT0047076, CASENT0052866 (CASC). –Toliara: •4w., 3s.; Réserve Spéciale d’Ambohijanahary, Forêt d’Ankazotsihitafototra, 34.6 km 314°NW Ambaravaranala; -18.26, 45.41833; alt. 1100 m; 16 Jan 2003; B.L. Fisher et al. leg.; CASENT0029499, CASENT0029619, CASENT0029620, CASENT0029719CASENT0029722 (CASC). •22w., 5s.; Réserve Spéciale d’Ambohijanahary, Forêt d’Ankazotsihitafototra, 35.2 km 312°NW Ambaravaranala; -18.26667, 45.40667; alt. 1050 m; 13 Jan 2003; B.L. Fisher et al. leg.; CASENT0028299, CASENT0028313, CASENT0028333, CASENT0485910, CASENT0485911, CASENT0485912, CASENT0485913, CASENT0496690, CASENT0496691, CASENT0496743, CASENT0496744, CASENT0496745, CASENT0496747 (CASC).

Diagnosis

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

Redescription

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

Figure 26. 

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

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

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.

Comments

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

Pheidole maizina sp. nov.

Figs 27A–F, 85C, 87G

Type material

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

Other material

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

Diagnosis

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

Description

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

Figure 27. 

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

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

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.

Key to the Pheidole ensifera group

1 Major workers. Occipital lobes and genae shiny, smooth or with very fine and sparse rugoreticulation; promesonotum, in lateral view, short, angular, and relatively low (Fig. 28A, E). Minor workers. Pronotum and mesonotum with fine to moderately dense foveolae, sometimes foveola weakening on the dorsal surface; head predominately smooth (Fig. 28I) P. ensifera Forel
Major workers. Occipital lobes shiny, with sparse and thick rugoreticulation; genae shiny, with moderately dense and fine rugoreticulation or smooth with anterior part with sparse and fine rugulae (Fig. 28B–D, F–H). Minor workers. Pronotum and at least dorsal surface of mesonotum smooth, with indistinct, sparse rugulae or with fine, dense rugoreticulation. If mesosoma entirely sculptured on the head finely rugo-punctate and smooth area, if present, limited to basal area of frons and genae (Fig. 28J–L) 2
2 Major workers. Posterior declivity of promesonotum steep to relatively steep, inner hypostomal teeth wide and pointed outward (Figs 28G, 29B). Minor workers. Petiolar peduncle long, postpetiole approximately 1.5 times longer that wide, head almost always smooth (Fig. 29E, I–J) P. ocypodea sp. nov.
Major workers. Posterior declivity of promesonotum convex, inner hypostomal teeth narrower and never pointed outward (Figs 28F, H, 29A, C). Minor workers. Petiolar peduncle short, postpetiole approximately as long as wide, at least lateral sides of head and malar area with fine rugae (Fig. 29D, F–H, K, L) 3
3 Major workers. Sides of head with sparse and long pilosity (distance between setae approximately as long as their length), genae and propodeum with reduced or partially absent sculpture (Fig. 28B, F). Minor workers. Vertex, genae, frons and mesosomal dorsum at least partially smooth, promesonotal and metanotal grooves absent (Figs 28J, 29D) P. aelloea sp. nov.
Major workers. Sides of head with denser and shorter pilosity (distance between shorter than their length), genae and propodeum never smooth (Fig. 28D, H). Minor workers. Head finely rugo-punctate and smooth area, if present, limited to basal area of frons and genae, promesonotal and metanotal grooves indistinct but always present (Figs 28L, 29F) P. podargea sp. nov.
Figure 28. 

Pheidole ensifera Forel, head and profile of major worker (A, E), profile of minor worker (I). Pheidole aelloea sp. nov., head and profile of major worker (B, F), profile of minor worker (J). Pheidole ocypodea sp. nov., head and profile of major worker (C, G), profile of minor worker (K). Pheidole podargea sp. nov., head and profile of major worker (D, H), profile of minor worker (L).

Figure 29. 

Pheidole aelloea sp. nov., hypostomal teeth (A), head of minor worker (D), petiole and postpetiole (G, H). Pheidole ocypodea sp. nov., hypostomal teeth (B), head of minor worker (E), petiole and postpetiole (I, J). Pheidole podargea sp. nov., hypostomal teeth (C), head of minor worker (F), petiole and postpetiole (K, L).

Pheidole ensifera Forel, 1897

Figs 30A–F, 84O, 86O

Type material

Pheidole ensifera Forel, 1897: 197 (s.w.). Lectotype [designated here]: major worker (top specimen, CASENT0101780): Madagascar, Antsiranana, Nosy Be, coll. Voeltzkow (MHNG) [examined]. Paralectotypes: 1 major worker (CASENT0810540, bottom specimen, the same pin as lectotype) (MHNG) [examined], 2 minor workers (1 pin, CASENT0101650) (MHNG) [examined], 2 minor workers (1 pin, CASENT0923207) (MHNG) [examined], 3 major workers (1 pin, CASENT0923208) (MHNG) [examined], 3 major workers (1 pin, CASENT0923209) (MHNG) [examined]: the same data as lectotype.

Other material

Madagascar. –Antsiranana: •130w., 28s.; Ampasindava, Forêt d’Ambilanivy, 3.9 km 181°S Ambaliha; -13.79861, 48.16167; alt. 600 m; 4 Mar 2001; B.L. Fisher et al. leg.; CASENT0044202, CASENT0044202, CASENT0406703, CASENT0420010, CASENT0420016, CASENT0420048, CASENT0420052, CASENT0420054, CASENT0420057, CASENT0420059, CASENT0427703, CASENT0427705, CASENT046136, CASENT0464031, CASENT0464033, CASENT0464037, CASENT0464039, CASENT0464041, CASENT0464045, CASENT0464049, CASENT0464050, CASENT0464052, CASENT0464055, CASENT0464056, CASENT0464057, CASENT0464061, CASENT0464063, CASENT0464064, CASENT0464066, CASENT0464071, CASENT0464075, CASENT0464080, CASENT0464081, CASENT0464083, CASENT0464088, CASENT0464091, CASENT0464095, CASENT0464099, CASENT0464102, CASENT0464104CASENT0464107, CASENT0464113, CASENT0464115, CASENT0464119, CASENT0464121, CASENT0464124, CASENT0464127, CASENT0464129, CASENT0464132, CASENT0464141, CASENT0464142, CASENT0464145, CASENT0464146, CASENT0464149, CASENT0464150, CASENT0464151, CASENT0464153, CASENT0464154, CASENT0464155, CASENT0464158, CASENT0464170, CASENT0464176, CASENT0464178, CASENT0464180, CASENT0464182, CASENT0464183, CASENT0464185, CASENT0464186, CASENT0464189, CASENT0464190, CASENT0464192, CASENT0464194, CASENT0464196, CASENT0464200CASENT0464203, CASENT0464207CASENT0464212, CASENT0464214, CASENT0464215, CASENT0464219, CASENT0464222, CASENT0464223, CASENT0464224, CASENT0464231, CASENT0464234,, CASENT0464237, CASENT0464240, CASENT0464240, CASENT0464243, CASENT0464244, CASENT0464250, CASENT0464253CASENT0464256, CASENT0464258, CASENT0464259, CASENT0464261, CASENT0464262, CASENT0464264, CASENT0464266, CASENT0464273, CASENT0464279, CASENT0464289, CASENT0464292, CASENT0464296, CASENT0464298, CASENT0464307, CASENT0464315, CASENT0464317, CASENT0464326, CASENT0464332, CASENT0464342, CASENT0464357, CASENT0464364, CASENT0464367, CASENT0464368, CASENT0464371, CASENT0464381, CASENT0464382, CASENT0464383, CASENT0464391, CASENT0464393, CASENT0464396, CASENT046439, CASENT0464400, CASENT0464402, CASENT0464403, CASENT0464405, CASENT0464411, CASENT0464412, CASENT0464413, CASENT0464413, CASENT0464414, CASENT0464415 (CASC). •3w., 2s.; Forêt d’ Andavakoera, 21.4 km 75°ENE Ambilobe; 4.6 km 356°N Betsiaka; -13.11833, 49.23; alt. 425 m; 15 Dec 2003; B.L. Fisher leg.; CASENT0044099, CASENT0044127, CASENT0044204, CASENT0044205 (CASC). •1w., 1s.; Galoko chain, Mont Galoko; -13.58745, 48.71419; alt. 380 m; 23 Feb 2013; B.L. Fisher et al. leg.; CASENT0303020 (CASC). •1w., 3s.; Galoko chain, Mont Galoko; -13.58487, 48.71818; alt. 520 m; 17 Feb 2013; B.L. Fisher et al. leg.; CASENT0298364, CASENT0305044, CASENT0305049 (CASC). •1w., 1s.; Galoko chain, Mont Kalabenono; -13.63999, 48.67374; alt. 498 m; 15 Oct 2013; B.L. Fisher et al. leg.; CASENT0370653 (CASC). •41w., 20s., 3q.; Nosy Be, Réserve Naturelle Intégrale de Lokobe, 6.3 km 112°ESE Hellville; -13.41933, 48.33117; alt. 30 m; 19 Mar 2001; B.L. Fisher et al. leg.; CASENT0403272, CASENT0421458, CASENT0427835, CASENT0427836, CASENT0427876, CASENT0427879, CASENT0427887, CASENT0462766, CASENT0462792, CASENT0462797, CASENT0462798, CASENT0462803, CASENT0462805, CASENT0462806, CASENT0462807, CASENT0462810, CASENT0462880, CASENT0462881, CASENT0462884, CASENT0462886CASENT0462891, CASENT0462896, CASENT0462913, CASENT0462914, CASENT0462916, CASENT0462924, CASENT0462930, CASENT0462933CASENT0462935, CASENT0462938CASENT0462942, CASENT0462963, CASENT0462988CASENT0462991, CASENT0462993, CASENT0463018CASENT0463021, CASENT0463044, CASENT0463060, CASENT0463063, CASENT0463067, CASENT0463075, CASENT0463101, CASENT0463118, CASENT0466246, CASENT0466297, CASENT0466299 (CASC). •5w., 2s.; R.S. Manongarivo, 10.8 km 229°SW Antanambao; -13.96167, 48.43333; alt. 400 m; 8 Nov 1998; B.L. Fisher leg.; CASENT0198013, CASENT0198014, CASENT0198522, CASENT0198524 (CASC). •4w., 1s; R.S. Manongarivo, 12.8 km 228°SW Antanambao; -13.97667, 48.42333; alt. 780 m; 11 Oct 1998; B.L. Fisher leg.; CASENT0198015, CASENT0198521, CASENT0198523 (CASC). •1w., 1s.; Sakaramy; -12.44114, 49.23197; alt. 260 m; 12 May 2011; B.L. Fisher et al. leg.; CASENT0261332 (CASC). –Mahajanga: •6w., 2s.; Réserve Spéciale Marotandrano, Marotandrano 48.3 km S Mandritsara; -16.28322, 48.81443; alt. 865 m; 6 Dec 2007; B.L. Fisher et al. leg.; CASENT0140640, CASENT0140643, CASENT0140667, CASENT0140672, CASENT0140682, CASENT0140685, CASENT0140687, CASENT0140692 (CASC).

Diagnosis

Major workers. Body size moderate: HL: 1.63–1.74 (1.69), HW: 1.43–1.51 (1.46), WL: 1.05–1.2 (1.11); propodeal spines very long (PSL: 0.31–0.37 (0.34)); head in full-face view rectangular, with lateral sides relatively straight, only their posteriormost part slightly convex; sides of the head with sparse, long, suberect pilosity; occipital lobes shiny, smooth or with very fine and sparse rugoreticulation; inner hypostomal teeth distinct, closely spaced, lobe-like, with rounded apex and wide base; outer hypostomal teeth bigger and wider than inner hypostomal teeth, lobe-like, with tops directed outward; inner and outer teeth closely spaced and connected by concavity. Minor workers. Body size moderate: HL: 0.58–0.69 (0.63), HW: 0.56–0.67 (0.6), WL: 0.76–0.87 (0.8); propodeal spines long (PSL: 0.18–0.22 (0.2)); scape, when laid back, surpassing posterior head margin by one-fifth of its length; lateral sides of head and malar area shiny, smooth or with indistinct, sparse rugulae, sculpture weakening posteriorly; vertex, genae and frons smooth; mesosoma foveolate.

Redescription

Major workers. Measurements (N = 10): HL: 1.63–1.74 (1.69); HW: 1.43–1.51 (1.46); SL: 0.67–0.73 (0.7); EL: 0.17–0.19 (0.18); WL: 1.05–1.2 (1.11); PSL: 0.31–0.37 (0.34); MTL: 0.66–0.71 (0.69); PNW: 0.58–0.67 (0.61); PTW: 0.15–0.19 (0.17); PPW: 0.46–0.54 (0.5); CI: 85.6–88.8 (87.0); SI: 46.5–50.1 (47.9); PSLI: 18.8–22.5 (20.4); PPI: 30.0–36.9 (34.1); PNI: 40.2–44.6 (41.7); MTI: 44.3–48.7 (47.2). Head. In full-face view rectangular, with lateral sides relatively straight, only their posteriormost part slightly convex (Fig. 30B). In lateral view elongate and oval; ventral and dorsal faces slightly convex; inner hypostomal teeth visible. Sides of the head with sparse, long, suberect pilosity; whole head with moderately sparse, long, suberect to erect pilosity. Antennal scrobes indistinct and not delimited; scrobe surface shiny, with dense to sparse, fine, longitudinal to irregular rugulae. Occipital lobes and genae shiny, smooth or with very fine and sparse rugoreticulation; frons with sparse, thick, and longitudinal rugae, interspaces with very fine and sparse rugulae, sculpture weakening posteriorly; malar area with sparse to moderately sparse, thick, and longitudinal rugae, interspaces with fine and dense rugulae. Centre of clypeus smooth and shiny, lateral sides with longitudinal rugae; median notch present, narrow and shallow to moderate; median longitudinal carina absent; lateral longitudinal carinae absent. Scape, when laid back, slightly surpass the midlength of head; pilosity decumbent to erect (Fig. 30B, D). Inner hypostomal teeth distinct, closely spaced, lobe-like, with rounded apex and wide base; outer hypostomal teeth bigger and wider than inner hypostomal teeth, lobe-like, with tops directed outward; inner and outer teeth closely spaced and connected by concavity (Fig. 84O). Mesosoma. In lateral view, promesonotum short, angular and relatively low, posterior mesonotum convex, with low tubercle-like projection; promesonotal groove absent; metanotal groove absent; propodeal spines long, thin, massive basally, with acute apex; humeral area laterally weakly produced (Fig. 30D). Surface shiny, with fine to thin, dense rugoreticulation, sculpture weakening on dorsum, sometimes propodeum with smooth patch on its dorsal surface. Pilosity sparse, long, and erect (Fig. 30D, F). Petiole. Shiny; peduncle shagreened, long, without horizontal lobes on its basal part; node smooth, low, and thick, triangular, with rounded apex, in rear view node slightly convex; pilosity sparse and erect (Fig. 30D, F). Postpetiole. Shagreened; in dorsal view sides with short, acute, and triangular projections; pilosity long, sparse and erect (Fig. 30D, F). Petiole. Shagreened, most often on the whole surface; pilosity moderately sparse, long, and erect (Fig. 30D, F). Colour. Unicolourous, reddish brown to dark brown, malar area and lower parts of frons with colouration brighter that the rest of the body (Fig. 30D, F).

Figure 30. 

Pheidole ensifera Forel, full-face view (A), profile (C), and dorsal view (E) of minor worker (CASENT0298364) and full-face view (B), profile (D), and dorsal view (F) of major worker (CASENT0923226).

Minor workers. Measurements (N = 10): HL: 0.58–0.69 (0.63); HW: 0.56–0.67 (0.6); SL: 0.6–0.67 (0.63); EL: 0.1–0.12 (0.11); WL: 0.76–0.87 (0.8); PSL: 0.18–0.22 (0.2); MTL: 0.46–0.52 (0.49); PNW: 0.37–0.44 (0.39); PTW: 0.08–0.12 (0.09); PPW: 0.13–0.18 (0.14); CI: 92.5–100.9 (95.4); SI: 101.4–107.1 (104.5); PSLI: 29.0–34.1 (31.4); PPI: 56.3–67.0 (61.2); PNI: 63.0–67.6 (65.2); MTI: 79.2–83.2 (81.7). Head. Occipital margin straight or indistinctly concave; occipital carina indistinct, weakly developed (Fig. 30A). Pilosity moderately dense, long, suberect to erect. Lateral sides of head and malar area shiny, smooth or with indistinct, sparse rugulae, sculpture weakening posteriorly; vertex, genae, and frons smooth; antennal sockets with sparse, sometimes interrupted carinae curved outward. Clypeus with median longitudinal carina absent; two lateral longitudinal carinae present. Scape, when laid back, surpassing posterior head margin by one-fifth of its length; pilosity suberect to erect (Fig. 30A, C). Mesosoma. In lateral view, promesonotum convex; promesonotal groove absent; metanotal groove indistinct; propodeal spines moderately long, massive basally, with acute apex (Fig. 30C). Pronotum and mesonotum with fine to moderately dense foveolate, sometimes foveolae weakening on the dorsal surface; katepisternum, anepisternum, and propodeum with thicker and denser foveolae. Pilosity moderately sparse, long and erect (Fig. 30C, E). Petiole. Shiny; peduncle rugulae relatively long and thin; node smooth, relatively high, triangular; with few long, erect setae (Fig. 30C, E). Postpetiole. With indistinct rugulae; short and convex; with few long, erect setae at the anterior edge (Fig. 30C, E). Petiole. With sparse and erect setae (Fig. 30C, E). Colour. Unicolourous, brown to dark brown (Fig. 30C, E).

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.

Pheidole aelloea sp. nov.

Figs 31A–F, 84A, 86A

Type material

Holotype. Madagascar. •1 major worker; Antsiranana; Makirovana Forest; -14.17066, 49.95409; alt. 415 m; 29 Apr 2011; B.L. Fisher et al. leg.; BLF26671, CASENT0236213 (CASC). Paratypes. Madagascar. •2w., 1q.; same data as for holotype; CASENT0923221, CASENT0236212, CASENT0872221 (CASC).

Other material

Madagascar. –Antsiranana: •10w., 11s., 1m., 1q.; Ambondrobe, 41.1 km 175° Vohemar; -13.71533, 50.10167; alt. 10 m; 29 Nov 2004; B.L. Fisher leg.; CASENT0056070, CASENT0056092, CASENT0056307, CASENT0056312, CASENT0056322, CASENT0056517, CASENT0056522, CASENT0056529, CASENT0056534, CASENT0056536, CASENT0056662, CASENT0056663, CASENT0056666, CASENT0056666, CASENT0056679, CASENT0107922, CASENT0110497, CASENT0110498, CASENT0110545, CASENT0235068 (CASC). •1w., 1s.; Binara Forest; -13.26207, 49.60505; alt. 692 m; 18 Oct 2013; B.L. Fisher et al. leg.; CASENT0369418, CASENT0369423 (CASC). •16w., 5s., 1q.; Forêt Ambanitaza, 26.1 km 347° Antalaha; -14.67933, 50.18367; alt. 240 m; 26 Nov 2004; B.L. Fisher leg.; CASENT0054846, CASENT0054850, CASENT0054851, CASENT0054853, CASENT0054893, CASENT0054894, CASENT0054951, CASENT0054955, CASENT0054968, CASENT0054972, CASENT0055042, CASENT0055061, CASENT0055550, CASENT0055558, CASENT0055562, CASENT0055562, CASENT0055592, CASENT0058967, CASENT0058977, CASENT0109535 (CASC). •10w., 9s., 3q.; Forêt d’ Antsahabe, 11.4 km 275°W Daraina; -13.21167, 49.55667; alt. 550 m; 12 Dec 2003; B.L. Fisher leg.; CASENT0042268, CASENT0042329, CASENT0042334, CASENT0042335, CASENT0053689, CASENT0053690, CASENT0053936, CASENT0053937, CASENT0053941, CASENT0053942, CASENT0076806, CASENT0076807 (CASC). •21w., 1s., 2q.; Forêt de Binara, 7.5 km 230°SW Daraina; -13.255, 49.61667; alt. 375 m; 1 Dec 2003; B.L. Fisher leg.; CASENT0041716, CASENT0041724, CASENT0041734, CASENT0041742, CASENT0041795, CASENT0041797, CASENT0041800, CASENT0041805, CASENT0041808, CASENT0041811, CASENT0041815, CASENT0041823, CASENT0041828, CASENT0041885, CASENT0041887, CASENT0041889, CASENT0041897, CASENT0041902, CASENT0041906, CASENT0041907, CASENT0041909, CASENT0041909, CASENT0073337, CASENT0076448 (CASC). •1s.; Forêt de Binara, 9.1 km 233°SW Daraina; -13.26333, 49.60333; alt. 650–800 m; 19 Nov 2004; B.L. Fisher leg.; CASENT0053842 (CASC). •6w., 5s., 2q.; Galoko chain, Mont Galoko; -13.59358, 48.73157; alt. 1100 m; 22 Feb 2013; B.L. Fisher et al. leg.; CASENT0300979, CASENT0300983, CASENT0301038, CASENT0301040, CASENT0301059, CASENT0301069, CASENT0301073 (CASC). •1w., 1s.; Galoko chain, Mont Galoko; -13.58745, 48.71419; alt. 380 m; 23 Feb 2013; B.L. Fisher et al. leg.; CASENT0303019 (CASC). •5w., 3s.; Galoko chain, Mont Galoko; -13.58487, 48.71818; alt. 520 m; 19 Feb 2013; B.L. Fisher et al. leg.; CASENT0302940, CASENT0304461, CASENT0304986, CASENT0304992, CASENT0305000, CASENT0305011 (CASC). •1w., 1s.; Galoko chain, Mont Galoko; -13.5888, 48.72864; alt. 980 m; 20 Feb 2013; B.L. Fisher et al. leg.; CASENT0300190 (CASC). •2w., 1s.; Galoko chain, Mont Galoko; -13.5888, 48.72864; alt. 980 m; 20 Feb 2013; B.L. Fisher et al. leg.; CASENT0304408, CASENT0304755 (CASC). •4w., 3s.; Galoko chain, Mont Kalabenono; -13.64179, 48.67282; alt. 643 m; 10 Oct 2013; B.L. Fisher et al. leg.; CASENT0366980, CASENT0369681, CASENT0369743, CASENT0369780 (CASC). •2w.; Makirovana Forest; -14.104, 50.03574; alt. 225 m; 4 May 2011; B.L. Fisher et al. leg.; CASENT0231485, CASENT0243618 (CASC). •14w., 1s., 2q.; Makirovana Forest; -14.17066, 49.95409; alt. 415 m; 29 Apr 2011; B.L. Fisher et al. leg.; CASENT0231284, CASENT0231285, CASENT0231304, CASENT0231305, CASENT0236148, CASENT0236154, CASENT0243267, CASENT0243271, CASENT0243278, CASENT0243280, CASENT0243284, CASENT0243301, CASENT0243305, CASENT0243325 (CASC). •1w.; Makirovana Forest; -14.16044, 49.95216; alt. 550 m; 1 May 2011; B.L. Fisher et al. leg.; CASENT0230611 (CASC). •2w.; Masoala National Park; -15.3014, 50.22776; alt. 280 m; 7 Mar 2014; B.L. Fisher et al. leg.; CASENT0377714, CASENT0377725 (CASC). •2w.; Masoala National Park; -15.32331, 50.30751; alt. 60 m; 10 Mar 2014; B.L. Fisher et al. leg.; CASENT0376961, CASENT0378012 (CASC). •11w., 18s.; Parc National de Marojejy, Manantenina River, 28.0 km 38°NE Andapa, 8.2 km 333°NNW Manantenina; -14.43667, 49.775; alt. 450 m; 12 Nov 2003; B.L. Fisher et al. leg.; CASENT0045743, CASENT0045810, CASENT0045818, CASENT0045826, CASENT0045846, CASENT0045866, CASENT0045869, CASENT0045895, CASENT0045901, CASENT0045909, CASENT0045979, CASENT0046010, CASENT0046075, CASENT0077158, CASENT0077159, CASENT0077160, CASENT0077161, CASENT0077162, CASENT0077163 (CASC). •1s.; R.S. Manongarivo, 14.5 km 220°SW Antanambao; -13.99833, 48.42833; alt. 1175 m; 20 Oct 1998; B.L. Fisher leg.; CASENT0198835 (CASC). •2w., 1s.; Réserve Spéciale d’Ambre, 3.5 km 235°SW Sakaramy; -12.46889, 49.24217; alt. 325 m; 26 Jan 2001; B.L. Fisher et al. leg.; CASENT0423920, CASENT0423924, CASENT0484626 (CASC). •1w., 1s.; Sava Region: Parc National de Marojejy, Manantenina River, 28.1 km 25.7NE Andapa; -14.43553, 49.76463; alt. 680 m; 11 Feb 2018; B.L. Fisher et al. leg.; CASENT0809603 (CASC). •1w.; Sava Region: Parc National de Marojejy, near Manantenina River; -14.43677, 49.77541; alt. 475 m; 7 Feb 2018; B.L. Fisher et al. leg.; CASENT0825163 (CASC). •3w., 1s., 1m.; Sava Region: Parc National de Marojejy, near Manantenina tributary, 28.3 km 28.5NE Andapa, forest along trail below Camp 1; -14.43934, 49.77689; alt. 450 m; 8 Feb 2018; B.L. Fisher et al. leg.; CASENT0825240, CASENT0825241, CASENT0825254 (CASC). •1w., 1q.; Sava Region: Parc National de Marojejy, near summit, 25.4 km 20.1NE Andapa; -14.44918, 49.73243; alt. 2100 m; 10 Feb 2018; B.L. Fisher et al. leg.; CASENT0809554 (CASC). –Toamasina: •1q.; 6.3 km S Ambanizana, Andranobe; -15.6813, 49.958; alt. 25 m; 14 Nov 1993; B.L. Fisher leg.; CASENT0198118 (CASC).

Diagnosis

Major workers. Body size moderate: HL: 1.85–2.45 (2.0), HW: 1.56–2.02 (1.64), WL: 1.17–1.35 (1.25); propodeal spines very long (PSL: 0.38–0.48 (0.42)); 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.

Description

Major workers. Measurements (N = 10): HL: 1.85–2.45 (2.0); HW: 1.56–2.02 (1.64); SL: 0.73–0.89 (0.77); EL: 0.14–0.18 (0.16); WL: 1.17–1.35 (1.25); PSL: 0.38–0.48 (0.42); MTL: 0.67–0.89 (0.75); PNW: 0.7–0.8 (0.73); PTW: 0.2–0.22 (0.2); PPW: 0.59–0.7 (0.66); CI: 79.6–85.9 (83.2); SI: 44.1–49.3 (46.9); PSLI: 19.8–25.0 (21.5); PPI: 28.3–34.5 (31.0); PNI: 39.5–46.8 (44.8); MTI: 41.8–50.6 (45.8). 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).

Figure 31. 

Pheidole aelloea sp. nov., full-face view (A), profile (C), and dorsal view (E) of paratype minor worker (CASENT0923221) and full-face view (B), profile (D), and dorsal view (F) of holotype major worker (CASENT0236213).

Minor workers. Measurements (N = 10): HL: 0.63–0.78 (0.68); HW: 0.61–0.79 (0.67); SL: 0.66–0.8 (0.7); EL: 0.09–0.13 (0.11); WL: 0.83–0.98 (0.9); PSL: 0.3–0.37 (0.32); MTL: 0.52–0.66 (0.57); PNW: 0.39–0.49 (0.42); PTW: 0.09–0.14 (0.1); PPW: 0.15–0.25 (0.19); CI: 95.9–102.7 (98.3); SI: 100.5–119.0 (105.9); PSLI: 43.0–49.5 (46.7); PPI: 48.5–64.9 (54.3); PNI: 60.3–65.6 (63.0); MTI: 83.3–88.1 (85.5). Head. In full-face view square, posterior of eyes slightly convex, anterior of eyes relatively straight, occipital margin straight or indistinctly convex; occipital carina indistinct, weakly developed (Fig. 31A). Pilosity sparse, long, and erect. Lateral sides of head and malar area finely rugulose, sculpture weakening posteriorly; vertex, genae, and frons smooth; antennal sockets with sparse carinae curved outward. Clypeus with median longitudinal carina present; two lateral longitudinal carinae present. Scape, when laid back, surpassing posterior head margin by one-fifth of its length; pilosity 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.

Pheidole ocypodea sp. nov.

Figs 32A–F, 85O, 87T

Type material

Holotype. Madagascar. •1 major worker; Antsiranana; Galoko chain, Mont Galoko; -13.5888, 48.72864; alt. 980 m; 20 Feb 2013; B.L. Fisher et al. leg.; BLF30960, CASENT0304390 (CASC). Paratype. Madagascar. •1 minor worker; same data as for holotype; CASENT0923220 (CASC).

Other material

Madagascar. –Antsiranana: •8w., 11s.; R.S. Manongarivo, 10.8 km 229°SW Antanambao; -13.96167, 48.43333; alt. 400 m; 8 Nov 1998; B.L. Fisher leg.; CASENT0198625, CASENT0198626, CASENT0198834, CASENT0198836, CASENT0846551CASENT0846562 (CASC). •32w., 24s.; R.S. Manongarivo, 12.8 km 228°SW Antanambao; -13.97667, 48.42333; alt. 780 m; 11 Sep 1998; B.L. Fisher leg.; CASENT0196883, CASENT0198627, CASENT0198837, CASENT0846501CASENT0846550 (CASC).

Diagnosis

Major workers. Body size moderate: HL: 1.82–2.01 (1.95); HW: 1.52–1.66 (1.6), WL: 1.16–1.31 (1.26); propodeal spines very long (PSL: 0.38–0.45 (0.41)); 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.

Description

Major workers. Measurements (N = 10): HL: 1.82–2.01 (1.95); HW: 1.52–1.66 (1.6); SL: 0.71–0.8 (0.75); EL: 0.15–0.19 (0.17); WL: 1.16–1.31 (1.26); PSL: 0.38–0.45 (0.41); MTL: 0.69–0.79 (0.75); PNW: 0.68–0.78 (0.75); PTW: 0.18–0.25 (0.21); PPW: 0.62–0.73 (0.65); CI: 79.8–84.5 (82.3); SI: 44.5–50.1 (46.8); PSLI: 20.3–22.7 (21.3); PPI: 28.0–37.7 (32.4); PNI: 44.7–48.9 (46.6); MTI: 44.5–48.3 (46.5). Head. In full-face view rectangular, with lateral sides relatively straight, only their posteriormost part slightly convex (Fig. 32B). In lateral view oval; ventral and dorsal faces convex; inner hypostomal teeth visible. Sides of the head with sparse, relatively long, erect pilosity; whole head with moderately dense, relatively long, erect pilosity. Antennal scrobes indistinct and not delimited by carinulae. Occipital lobes shiny, with sparse and thick rugoreticulation; genae smooth and shiny, only anterior part sometimes with sparse and fine rugulae; frons with longitudinal and sparse rugae, on posterior part of frons rugae directed outward and more irregular, interspaces with sparse to moderately sparse rugose foveolae; malar area and lateral sides of head shiny, with longitudinal rugae, surface between rugae with dense rugose foveolae. 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 midlength of head; pilosity suberect to erect (Fig. 32B, D). 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 (Fig. 85O). Mesosoma. In lateral view, promesonotum short, angular, and high, with relatively steep posterior declivity; promesonotal groove absent; metanotal groove absent; propodeal spines very long, narrow, with acute apex; humeral area laterally absent to weakly produced (Fig. 32D). Surface shiny, pronotum and dorsal surface of mesonotum smooth or sometimes with indistinct, sparse rugulae; propodeum, katepisternum, and anepisternum with thick and dense rugoreticulation. Pilosity sparse, long, and erect (Fig. 32D, F). Petiole. Shagreened; peduncle relatively long, without horizontal lobes on its basal part; node triangular and thin, with rounded apex, in rear view node dorsoventrally slightly depressed; pilosity moderately sparse and erect (Fig. 32D, F). Postpetiole. Shagreened; in dorsal view sides with acute, horn-like, moderately long projections; pilosity long, moderately sparse, and erect (Fig. 32D, F). Petiole. First gastral tergite shagreened, at least on its basal part; pilosity moderately dense, long, and erect (Fig. 32D, F). Colour. Unicolourous, ocherous to reddish-brown (Fig. 32D, F).

Figure 32. 

Pheidole ocypodea sp. nov., full-face view (A), profile (C), and dorsal view (E) of paratype minor worker (CASENT0923220) and full-face view (B), profile (D), and dorsal view (F) of holotype major worker (CASENT0304390).

Minor workers. Measurements (N = 10): HL: 0.63–0.72 (0.66); HW: 0.6–0.7 (0.64); SL: 0.63–0.69 (0.66); EL: 0.09–0.13 (0.11); WL: 0.8–0.93 (0.85); PSL: 0.26–0.32 (0.28); MTL: 0.49–0.57 (0.53); PNW: 0.38–0.45 (0.42); PTW: 0.09–0.12 (0.1); PPW: 0.18–0.24 (0.19); CI: 93.6–98.2 (96.3); SI: 98.3–106.9 (102.9); PSLI: 40.4–45.5 (42.6); PPI: 45.1–57.6 (52.4); PNI: 62.5–68.1 (65.2); MTI: 78.9–86.9 (83.3). Head. In full-face view square, posterior of eyes slightly convex, anterior of eyes relatively straight, occipital margin straight or indistinctly convex; occipital carina indistinct, weakly developed (Fig. 32A). Pilosity sparse, long, and erect. Lateral sides of head and malar area smooth and shiny or with indistinct, sparse rugulae, sculpture weakening posteriorly; vertex, genae, and frons smooth; antennal sockets with sparse, interrupted carinae curved outward. Clypeus with median longitudinal carina present; two lateral longitudinal carinae present. Scape, when laid back, surpassing posterior head margin by two-fifths of its length; pilosity suberect to erect (Fig. 32A, C). Mesosoma. In lateral view, promesonotum high, short, and convex; promesonotal groove absent; metanotal groove shallow and indistinct; propodeal spines very long, massive basally, with acute apex (Fig. 32C). Surface shiny, 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. Pilosity sparse, long, and erect (Fig. 32C, E). Petiole. Peduncle moderately long and thin; node low, bulge-like; with few long, erect setae (Fig. 32C, E). Postpetiole. Long, low, and slightly convex; with few long, erect setae at the anterior edge (Fig. 32C, E). Petiole. Pilosity sparse and erect (Fig. 32C, E). Colour. Unicolourous, bright brown to brown (Fig. 32C, E).

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.

Pheidole podargea sp. nov.

Figs 33A–F, 85R, 88B

Type material

Holotype. Madagascar. •1 major worker; Antsiranana; Forêt d’ Andavakoera, 21.4 km 75°ENE Ambilobe; 4.6 km 356°N Betsiaka; -13.11833, 49.23; alt. 425 m; 16 Dec 2003; B.L. Fisher leg.; BLF10317, CASENT0487570, middle specimen on the pin (CASC). Paratypes. Madagascar. •9w., 8s.; same data as for holotype; CASENT0487566, CASENT0872085, CASENT0487569, CASENT0487568, CASENT0487567 (CASC).

Other material

Madagascar. –Antsiranana: •9w., 4s.; Forêt de Binara, 7.5 km 230°SW Daraina; -13.255, 49.61667; alt. 375 m; 1 Dec 2003; B.L. Fisher leg.; CASENT0041888, CASENT0043218, CASENT0043224, CASENT0043226, CASENT0043235, CASENT0043241, CASENT0043253, CASENT0043256, CASENT0043259, CASENT0043317, CASENT0043319, CASENT0043331, CASENT0043402 (CASC). •5w., 1s., 2q.; Réserve Spéciale d’Ambre, 3.5 km 235°SW Sakaramy; -12.46889, 49.24217; alt. 325 m; 26 Jan 2001; Fisher et al. leg.; CASENT0406663, CASENT0406673, CASENT0423867, CASENT0427613, CASENT0427696, CASENT0427699 (CASC). •32w., 9s., 3q.; Réserve Spéciale de l’Ankarana, 13.6 km 192°SSW Anivorano Nord; -12.86361, 49.22583; alt. 210 m; 16 Feb 2001; Fisher et al. leg.; CASENT0440586, CASENT0440686, CASENT0440729, CASENT0440730, CASENT0440731, CASENT0440735, CASENT0440736, CASENT0440739, CASENT0440740, CASENT0440743, CASENT0440745, CASENT0440746, CASENT0440748, CASENT0440750, CASENT0440752, CASENT0440754, CASENT0440755, CASENT0440756, CASENT0440758, CASENT0440762CASENT0440765, CASENT0440769, CASENT0440771, CASENT0440773, CASENT0440774, CASENT0440776, CASENT0440780, CASENT0440782, CASENT0440786CASENT0440788, CASENT0440790, CASENT0440792, CASENT0440803, CASENT0440808, CASENT0440813, CASENT0441204CASENT0441206, CASENT0441208, CASENT0441211, CASENT0441212, CASENT0441214, CASENT0441216 (CASC).

Diagnosis

Major workers. Body size moderate: HL: 1.85–2.0 (1.9), HW: 1.55–1.65 (1.59), WL: 1.19–1.35 (1.26); propodeal spines very long (PSL: 0.39–0.46 (0.42)); 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.

Description

Major workers. Measurements (N = 10): HL: 1.85–2.0 (1.9); HW: 1.55–1.65 (1.59); SL: 0.67–0.76 (0.72); EL: 0.16–0.2 (0.17); WL: 1.19–1.35 (1.26); PSL: 0.39–0.46 (0.42); MTL: 0.7–0.76 (0.73); PNW: 0.71–0.81 (0.76); PTW: 0.15–0.22 (0.19); PPW: 0.59–0.71 (0.66); CI: 80.1–87.6 (83.4); SI: 42.6–47.7 (45.6); PSLI: 20.9–24.4 (21.8); PPI: 25.4–32.7 (29.1); PNI: 45.5–51.0 (48.2); MTI: 42.6–48.8 (46.4). Head. In full-face view rectangular, with lateral sides relatively straight, only their posteriormost part slightly convex (Fig. 33B). In lateral view oval; ventral and dorsal faces convex; inner hypostomal teeth visible. Sides of the head with moderately dense, moderately long, erect pilosity. Antennal scrobes indistinct and not delimited by carinulae. Occipital lobes shiny, with sparse and thick rugoreticulation; genae shiny, with moderately dense to dense, fine rugoreticulation; frons with longitudinal, sparse to moderately sparse rugae, on posterior part of frons rugae directed outward and more irregular, interspaces with sparse rugulae, sometimes indistinct; malar area and lateral sides of head shiny, with longitudinal rugoreticulation, surface between rugae with sparse and indistinct rugulae. Centre of clypeus smooth and shiny or with fine longitudinal rugulae, lateral sides with longitudinal rugae; median notch present, wide, and shallow; median longitudinal carina present but sometimes indistinct; lateral longitudinal carinae absent. Scape, when laid back, reaching midlength of head; pilosity suberect to erect (Fig. 33B, D). 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 (Fig. 85R). Mesosoma. In lateral view, promesonotum short, angular, and high, with slightly convex posterior declivity; promesonotal groove absent; metanotal groove absent; propodeal spines very long, massive basally, with acute apex; humeral area laterally weakly produced (Fig. 33D). Surface shiny, with fine, dense rugoreticulation, sometimes sculpture weakening on dorsum. Pilosity sparse, long, and erect (Fig. 33D, F). Petiole. Shagreened; peduncle relatively long, with small tooth-like horizontal lobes on its basal part; node triangular, with rounded apex, in rear view node convex; pilosity moderately sparse and erect (Fig. 33D, F). Postpetiole. Shagreened; in dorsal view sides with acute, horn-like, moderately long projections; pilosity long, moderately sparse, and erect (Fig. 33D, F). Petiole. First gastral tergite shagreened, at least on its basal part; pilosity moderately dense, long, and erect (Fig. 33D, F). Colour. Unicolourous, ocherous to reddish-brown (Fig. 33D, F).

Figure 33. 

Pheidole podargea sp. nov., full-face view (A), profile (C), and dorsal view (E) of paratype minor worker (CASENT0487567) and full-face view (B), profile (D), and dorsal view (F) of holotype major worker (CASENT0487570).

Minor workers. Measurements (N = 10): HL: 0.61–0.7 (0.66); HW: 0.62–0.69 (0.65); SL: 0.63–0.7 (0.67); EL: 0.11–0.12 (0.11); WL: 0.81–0.9 (0.86); PSL: 0.25–0.31 (0.28); MTL: 0.49–0.58 (0.53); PNW: 0.39–0.45 (0.43); PTW: 0.09–0.11 (0.1); PPW: 0.17–0.21 (0.19); CI: 94.3–100.7 (97.2); SI: 97.5–107.9 (103.1); PSLI: 38.8–44.3 (41.7); PPI: 46.9–56.9 (51.6); PNI: 63.4–69.7 (66.2); MTI: 78.4–85.0 (82.3). Head. In full-face view square, posterior of eyes slightly convex, anterior of eyes relatively straight, occipital margin straight or indistinctly convex; occipital carina indistinct, weakly developed (Fig. 33A). Pilosity moderately dense, long, and erect. Whole head with fine and dense rugo-punctuation, only basal area of frons and genae with sculpture reduced or sometimes absent; antennal sockets with dense carinae curved outward. Clypeus with median longitudinal carina present; two lateral longitudinal carinae absent. Scape, when laid back, surpassing posterior head margin by two-fifths of its length; pilosity suberect to erect (Fig. 33A, C). Mesosoma. In lateral view, promesonotum high, short, and convex; promesonotal groove present, indistinct; metanotal groove shallow and indistinct; propodeal spines very long, massive basally, with acute apex (Fig. 33C). Whole surface rugo-punctate, sometimes sculpture weakening on dorsal surface and pronotum. Pilosity sparse, long, and erect (Fig. 33C, E). Petiole. Peduncle relatively long and thin; node low, triangular; with few long, erect setae (Fig. 33C, E). Postpetiole. Short, low, and slightly convex; with few long, erect setae at the anterior edge (Fig. 33C, E). Petiole. With few long, erect setae (Fig. 33C, E). Colour. Unicolourous, bright brown to brown (Fig. 33C, E).

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 sub-oval 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 (Fig. 34P) P. longipilosa sp. nov.
Major workers. Head in full-face view cordate, widening posteriorly; sides of head never with dense, very long, erect pilosity; antennal scrobes developed, well delimited by carinulae; propodeal spines thin, with narrow base (Fig. 34A–D). Minor workers. Propodeal spines needle-like, moderately long to very long (Fig. 34I–L) 2
2 Major workers. Antennal scrobes never with foveolate surface; outer hypostomal teeth pointed inward (Fig. 34A, E). Minor workers. Propodeal spines long, promesonotum flat, long, and box-like (Fig. 34I) P. ferruginea sp. nov.
Major workers. Antennal scrobes with foveolate surface; outer hypostomal teeth pointed outward (Fig. 34B–D, F–H). Minor workers. Propodeal spines shorter, if long then promesonotum short, higher, and not box-like (Fig. 34J–L) 3
3 Major workers. Antennal scrobes deep, predominantly foveolate; frons with thick, sparse, longitudinal to irregular rugae, interspaces predominantly smooth (Fig. 34B). Minor workers. Propodeal spines moderately long (Fig. 34J) P. rugocephala sp. nov.
Major workers. Antennal scrobes shallow, foveolate with additional sculpture; frons with longitudinal rugae and never smooth interspaces (Fig. 34C, D). Minor workers. Propodeal spines long (Fig. 34K, L) 4
4 Major workers. Antennal scrobes foveolate with additional longitudinal rugae, frons with area between rugae with fine and dense rugulae, sometimes rugulae fading on the central part of frons, petiolar peduncle with wide and distinct horizontal lobes (Fig. 34C, M). Minor workers. Promesonotum with posterior declivity relatively steep, katepisternum and mesonotum with smooth notches (Fig. 34K) P. vohemarensis sp. nov.
Major workers. Antennal scrobes foveolate with additional irregular and indistinct rugae, frons with interspaces superficially foveolate, petiolar peduncle with shorter and less distinct horizontal lobes (Fig. 34D, N). Minor workers. Promesonotum with posterior declivity smoothly declining towards propodeum, katepisternum and mesonotum foveolate (Fig. 34L) P. manantenensis sp. nov.
Figure 34. 

Pheidole ferruginea sp. nov., head of major worker (A), hypostomal teeth (E), profile of minor worker (I). Pheidole rugocephala sp. nov., head of major worker (B), hypostomal teeth (F), profile of minor worker (J). Pheidole vohemarensis sp. nov., head of major worker (C), hypostomal teeth (G), profile of minor worker (K), dorsal view of petiole of major worker (M). Pheidole manantenensis sp. nov., head of major worker (D), hypostomal teeth (H), profile of minor worker (L), dorsal view of petiole of major worker (N).

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.

Pheidole ferruginea sp. nov.

Figs 35A–F, 84P, 86P

Type material

Holotype. Madagascar. •1 major worker; Antsiranana; Parc National de Marojejy, Manantenina River, 27.6 km 35°NE Andapa, 9.6 km 327°NNW Manantenina; -14.435, 49.76; alt. 775 m; 16 Nov 2003; B.L. Fisher leg.; BLF08998, CASENT0494997, middle specimen on the pin (CASC). Paratypes. Madagascar. •9w., 5s.; same data as for holotype; CASENT0494993CASENT0494996, CASENT0872087, CASENT0872230CASENT0872238 (CASC).

Other material

Madagascar. –Antsiranana: •15w., 6s., 1m., 1q.; Mandraka; -18.91813, 47.91717; alt. 1312 m; 20 Mar 2014; B.L. Fisher et al. leg.; CASENT0375496, CASENT0377119, CASENT0377122, CASENT0377125, CASENT0377128, CASENT0377134, CASENT0377137, CASENT0377144, CASENT0377148, CASENT0377153, CASENT0377156, CASENT0377159, CASENT0378177, CASENT0378178, CASENT0378179, CASENT0378299, CASENT0378301, CASENT0378305, CASENT0378566, CASENT0378569 (CASC). •16w., 5s.; Reg. Analamanga, St. Forestière Mandraka; -18.9183, 47.91687; alt. 1285 m; 25 Mar 2015; B.L. Fisher et al. leg.; CASENT0390402, CASENT0390414, CASENT0390440, CASENT0390443, CASENT0390444, CASENT0390507, CASENT0390537, CASENT0703931, CASENT0720608, CASENT0720646, CASENT0720649, CASENT0720831, CASENT0720836, CASENT0720837, CASENT0728120, CASENT0728121 (CASC). •6w., 3s.; 6.5 km SSW Befingotra, Rés. Anjanaharibe-Sud; -14.75, 49.5; alt. 875 m; 19 Oct 1994; B.L. Fisher leg.; CASENT0198459CASENT0198461 (CASC). •2s.; 9.2 km WSW Befingotra, Rés. Anjanaharibe-Sud; -14.75, 49.46667; alt. 1200 m; 9 Nov 1994; B.L. Fisher leg.; CASENT0198916 (CASC). •3w., 1s.; Binara Forest; -13.26392, 49.59919; alt. 1065 m; 18 Oct 2013; B.L. Fisher et al. leg.; CASENT0369870, CASENT0369873, CASENT0369876, CASENT0369878 (CASC). •3w., 1s.; Forêt Ambanitaza, 26.1 km 347° Antalaha; -14.67933, 50.18367; alt. 240 m; 26 Nov 2004; B.L. Fisher leg.; CASENT0055590, CASENT0055598 (CASC). •3w., 1s.; Forêt de Binara, 9.1 km 233°SW Daraina; -13.26333, 49.60333; alt. 800 m; 3 Dec 2003; B.L. Fisher leg.; CASENT0043315, CASENT0043364, CASENT0043367, CASENT0043447 (CASC). •12w., 3s., 1m.; Makirovana Forest; -14.17066, 49.95409; alt. 415 m; 28 Apr 2011; B.L. Fisher et al. leg.; CASENT0231230, CASENT0231233, CASENT0231258, CASENT0231270, CASENT0231278, CASENT0236129, CASENT0236143, CASENT0243310, CASENT0243314, CASENT0243315, CASENT0243348 (CASC). •2w.; Masoala National Park; -15.32331, 50.30751; alt. 60 m; 10 Mar 2014; B.L. Fisher et al. leg.; CASENT0376974, CASENT0376995 (CASC). •1s.; Masoala, Cap Est, Forêt d’Andranoanala; -15.26158, 50.4758; alt. 15 m; 15 Mar 2014; B.L. Fisher et al. leg.; CASENT0377900 (CASC). •39w., 7s., 1q.; Parc National de Marojejy, Manantenina River, 27.6 km 35°NE Andapa, 9.6 km 327°NNW Manantenina; -14.435, 49.76; alt. 775 m; 15 Nov 2003; B.L. Fisher et al. leg.; CASENT0045227, CASENT0045231, CASENT0045238, CASENT0045240, CASENT0045243, CASENT0045248, CASENT0045250, CASENT0045252, CASENT0045254, CASENT0045260, CASENT0045264, CASENT0045267, CASENT0045272, CASENT0045334, CASENT0045351, CASENT0045363, CASENT0045366, CASENT0045744, CASENT0045904, CASENT0045987, CASENT0068463, CASENT0068464, CASENT0494859, CASENT0494860, CASENT0494917, CASENT0846566CASENT0846569, CASENT0846571 CASENT0846572, CASENT0846574, CASENT0846577CASENT0846587 (CASC). •20w., 4s.; Parc National de Marojejy, Manantenina River, 28.0 km 38°NE Andapa, 8.2 km 333°NNW Manantenina; -14.43667, 49.775; alt. 450 m; 12 Nov 2003; B.L. Fisher et al. leg.; CASENT0045803, CASENT0045805, CASENT0045811, CASENT0045832, CASENT0045845, CASENT0045867, CASENT0045876, CASENT0045886, CASENT0045889, CASENT0045899, CASENT0045902, CASENT0045908, CASENT0046015, CASENT0046021, CASENT0047669, CASENT0077149, CASENT0077150, CASENT0077152 (CASC). –Fianarantsoa: •6w., 4s., 1q.; 2 km W Andrambovato, along river Tatamaly; -21.51167, 47.41; alt. 1075 m; 3 Jun 2005; B.L. Fisher et al. leg.; CASENT0060943, CASENT0060944, CASENT0061397, CASENT0061581, CASENT0061621, CASENT0061677, CASENT0109000 (CASC). •1w.; 43 km S Ambalavao, Rés. Andringitra; -22.23333, 47; alt. 825 m; 5 Oct 1993; B.L. Fisher leg.; CASENT0198537 (CASC). •1w.; 45 km S. Ambalavao; -22.21667, 47.01667; alt. 785 m; 25 Sep 1993; B.L. Fisher leg.; CASENT0198538 (CASC). •1w.; 7.6 km 122° Kianjavato, Forêt Classée Vatovavy; -21.4, 47.94; alt. 175 m; 6 Jun 2005; B.L. Fisher et al. leg.; CASENT0061235 (CASC). •3w., 1s.; 9.0 km NE Ivohibe; -22.42667, 46.93833; alt. 900 m; 12 Nov 1997; B.L. Fisher et al. leg.; CASENT0198539, CASENT0198541, CASENT0198918 (CASC). •5w., 2s., 1q.; Ambinanindranomena Non-Protected Area, 39.45 km SE Ambalavao; -21.95386, 47.29427; alt. 1069 m; 1 Feb 2012; Andrianjaka & Ravelomanana leg.; CASENT0293842, CASENT0293876, CASENT0293878, CASENT0293916, CASENT0293917, CASENT0293935 (CASC). •1w., 1s.; Forêt d’Ambalagoavy Nord, Ikongo, Ambatombe; -21.857068, 47.37849; alt. 625 m; 1 Dec 2000; Harin’Hala & Irwin leg.; CASENT0009589, CASENT0009636 (CASC). •2w.; P.N. Ranomafana, Sahavondrona-Ampitamarivo; -21.2575, 47.36015; alt. 1100 m; 18 Mar 2003; Clark leg.; CASENT0052899, CASENT0052904 (CASC). •1w.; P.N. Ranomafana, Tolongoina-Ampasimpotsy 3; -21.47412, 47.55742; alt. 520 m; 11 Apr 2003; Clark leg.; CASENT0052880 (CASC). •1w., 1s., 1q.; Parc National Befotaka-Midongy, Papango 27.7 km S Midongy-Sud, Mount Papango; -23.83517, 46.96367; alt. 940 m; 13 Nov 2006; B.L. Fisher et al. leg.; CASENT0128696, CASENT0128700, CASENT0128704 (CASC). •1w., 1s.; Parc National Befotaka-Midongy, Papango 28.5 km S Midongy-Sud, Mount Papango; -23.84083, 46.9575; alt. 1250 m; 17 Nov 2006; B.L. Fisher et al. leg.; CASENT0118391 (CASC). •1w., 1s.; Parc National de Ranomafana, Vatoharanana River, 4.1 km 231°SW Ranomafana; -21.29, 47.43333; alt. 1100 m; 27 Mar 2003; B.L. Fisher et al. leg.; CASENT0039949, CASENT0040207 (CASC). •2w., 1s.; R.S. Ivohibe 8.0 km E Ivohibe; -22.48333, 46.96833; alt. 1200 m; 15 Oct 1997; B.L. Fisher et al. leg.; CASENT0198540, CASENT0198542, CASENT0198919 (CASC). •1w.; Vohiparara Kidonavo 1; -21.22632, 47.37007; alt. 1100 m; 13 Mar 2003; Clark leg.; CASENT0052978 (CASC). –Toamasina: •3w.; Ambanizana, Parc National Masoala; -15.57167, 50.00611; alt. 900–950 m; 26 Feb 2003; Andriamalala et al. leg.; CASENT0008765, CASENT0073478, CASENT0073478 (CASC). •3w., 1s.; Ambatovy, 12.4 km NE Moramanga; -18.83937, 48.30842; alt. 1080 m; 4 Mar 2007; B.L. Fisher et al. leg.; CASENT0123826, CASENT0123867, CASENT0123870, CASENT0123997 (CASC). •1w.; Analamay; -18.80623, 48.33707; alt. 1068 m; 21 Mar 2004; B.L. Fisher et al. leg.; CASENT0046556 (CASC). •2w.; Ankerana; -18.4017, 48.80605; alt. 1035 m; 24 Jan 2012; B.L. Fisher et al. leg.; CASENT0274446, CASENT0274448 (CASC). •6w.; Ankerana; -18.4061, 48.82029; alt. 725 m; 16 Jan 2012; B.L. Fisher et al. leg.; CASENT0274767, CASENT0275345, CASENT0275347, CASENT0275348, CASENT0275462 (CASC). •2w., 2q.; Ankerana; -18.40829, 48.82107; alt. 750 m; 21 Jan 2012; B.L. Fisher et al. leg.; CASENT0274977, CASENT0274978 (CASC). •1w.; Ankerana; -18.4104, 48.8189; alt. 855 m; 22 Jan 2012; B.L. Fisher et al. leg.; CASENT0274417 (CASC). •1w.; Corridor Forestier Analamay-Mantadia, Tsaravoniana; -18.75641, 48.42195; alt. 1036 m; 2 Dec 2012; B.L. Fisher et al. leg.; CASENT0301883 (CASC). •3w., 2s.; F.C. Sandranantitra; -18.04833, 49.09167; alt. 450 m; 18 Jan 1999; Ratsirarson leg.; CASENT0198543, CASENT0198920 (CASC). •2w., 1s.; Ile Sainte Marie, Forêt Kalalao, 9.9 km 34° Ambodifotatra; -16.9225, 49.88733; alt. 100 m; 24 Nov 2005; B.L. Fisher et al. leg.; CASENT0069837, CASENT0069839, CASENT0069844 (CASC). •1w., 1s.; Mahavelona (Foulpointe); -17.66667, 49.5; 1 Nov 1985; A. Pauly leg.; CASENT0095644, CASENT0095650 (CASC). •11w., 11s.; Montagne d’Akirindro 7.6 km 341°NNW Ambinanitelo; -15.28833, 49.54833; alt. 600 m; 17 Mar 2003; B.L. Fisher et al. leg.; CASENT0038892, CASENT0038907, CASENT0039031, CASENT0039045, CASENT0039049, CASENT0039079, CASENT0039080, CASENT0039090, CASENT0039094, CASENT0039109, CASENT0039113, CASENT0039121, CASENT0039194, CASENT0039205, CASENT0039213, CASENT0039217, CASENT0039224, CASENT0039226, CASENT0039233, CASENT0496453 (CASC). •2w.; Montagne d’Anjanaharibe, 18.0 km 21°NNE Ambinanitelo; -15.18833, 49.615; alt. 470 m; 8 Mar 2003; B.L. Fisher et al. leg.; CASENT0037612, CASENT0037639 (CASC). •1w., 3s.; Parc National de Zahamena, Onibe River; -17.75908, 48.85468; alt. 780 m; 21 Feb 2009; B.L. Fisher et al. leg.; CASENT0150453, CASENT0151642, CASENT0153420, CASENT0153422 (CASC). •4w.; Parc National de Zahamena, Tetezambatana forest, near junction of Nosivola and Manakambahiny Rivers; -17.74298, 48.72936; alt. 860 m; 18 Feb 2009; B.L. Fisher et al. leg.; CASENT0151078, CASENT0153357, CASENT0153371, CASENT0153374 (CASC). •4w., 4s.; Parc National Mananara-Nord, 7.1 km 261° Antanambe; -16.455, 49.7875; alt. 225 m; 16 Nov 2005; B.L. Fisher et al. leg.; CASENT0067223, CASENT0068923, CASENT0069441, CASENT0069445 CASENT0069451, CASENT0069605, CASENT0071324 (CASC). •1w., 2s.; Reserve Betampona, Camp Rendrirendry 34.1 km 332° Toamasina; -17.924, 49.19967; alt. 390 m; 28 Nov 2005; B.L. Fisher et al. leg.; CASENT0071885, CASENT0071912, CASENT0069185 (CASC). •1w.; Réserve Spéciale Ambatovaky, Sandrangato River; -16.81739, 49.29402; alt. 360 m; 25 Feb 2010; B.L. Fisher et al. leg.; CASENT0163669 (CASC). •8w., 1q.; Réserve Spéciale Ambatovaky, Sandrangato River; -16.77274, 49.26551; alt. 450 m; 20 Feb 2010; B.L. Fisher et al. leg.; CASENT0164042, CASENT0164417, CASENT0164422, CASENT0164440, CASENT0164468, CASENT0164483, CASENT0164485, CASENT0164492, CASENT0164498 (CASC). •5w., 1q.; Réserve Spéciale Ambatovaky, Sandrangato River; -16.7633, 49.26692; alt. 520 m; 22 Feb 2010; B.L. Fisher et al. leg.; CASENT0163718, CASENT0163722, CASENT0163723, CASENT0163833, CASENT0163873, CASENT0163877 (CASC). –Toliara: •1w., 3s.; 10 km NW Enakara, Rés Andohahela; -24.56667, 46.81667; alt. 420 m; 15 Nov 1992; B.L. Fisher leg.; CASENT0198037, CASENT0196935 (CASC). •6w., 1s., 1q.; 11 km NW Enakara, Rés. Andohahela; -24.56667, 46.83333; alt. 800 m; 17 Nov 1992; B.L. Fisher leg.; CASENT0196936, CASENT0198050, CASENT0217984, CASENT0217992, CASENT0217993 (CASC). •5w.; 2.7 km WNW 302° Ste. Luce; -24.77167, 47.17167; alt. 20 m; 9 Dec 1998; B.L. Fisher et al. leg.; CASENT0196937CASENT0196939, CASENT0198049, CASENT0198051 (CASC). •8w., 2s.; Anosy Region, Anosyenne Mts, 31.2 km NW Manantenina; -24.13894, 47.06804; alt. 1125 m; 26 Feb 2015; B.L. Fisher et al. leg.; CASENT0704213, CASENT0704214, CASENT0704872, CASENT0704873, CASENT0704876, CASENT0704877, CASENT0721011 (CASC). •2w., 1s.; Parc National Andohahela, Col de Tanatana, 33.3 km NW Tolagnaro; -24.7585, 46.85367; alt. 275 m; 24 Nov 2006; B.L. Fisher et al. leg.; CASENT0122564, CASENT0129922, CASENT0129927 (CASC). •1w., 1s.; Parc National Andohahela, Manangotry, 33.8 km NW Tolagnaro; -24.75117, 46.85783; alt. 575 m; 24 Nov 2006; B.L. Fisher et al. leg.; CASENT0121878 (CASC). •21w., 6s.; Parc National d’Andohahela, Col du Sedro, 3.8 km 113°ESE Mahamavo, 37.6 km 341°NNW Tolagnaro; -24.76389, 46.75167; alt. 900 m; 21 Jan 2002; B.L. Fisher et al. leg.; CASENT0078392, CASENT0078409, CASENT0078410, CASENT0078412, CASENT0479172, CASENT0479174, CASENT0479187, CASENT0479188, CASENT0479239, CASENT0479260, CASENT0479261, CASENT0479262, CASENT0479299, CASENT0483039, CASENT0483924, CASENT0483951, CASENT0483967, CASENT0483968, CASENT0483980, CASENT0484029, CASENT0484043, CASENT0484060, CASENT0484062, CASENT0484063, CASENT0484111, CASENT0484183 (CASC). •6w., 2s., 1q.; Réserve Spéciale Kalambatritra, Ampanihy; -23.4635, 46.4631; alt. 1270 m; 9 Feb 2009; B.L. Fisher et al. leg.; CASENT0150522, CASENT0150523, CASENT0153214, CASENT0153217, CASENT0217989, CASENT0846595, CASENT0846596, CASENT0846597 (CASC).

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.

Description

Major workers. Measurements (N = 10): HL: 1.18–1.34 (1.25); HW: 1.15–1.31 (1.23); SL: 0.61–0.69 (0.64); EL: 0.13–0.17 (0.15); WL: 0.93–1.05 (0.99); PSL: 0.23–0.28 (0.26); MTL: 0.58–0.66 (0.61); PNW: 0.5–0.6 (0.53); PTW: 0.13–0.17 (0.16); PPW: 0.45–0.58 (0.51); CI: 97.4–101.3 (99.0); SI: 48.7–55.8 (52.1); PSLI: 18.9–21.9 (20.8); PPI: 27.5–33.7 (30.7); PNI: 40.9–45.0 (43.0); MTI: 47.6–52.2 (49.7). Head. In full-face view cordate, widened posteriorly (Fig. 35B). In lateral view oval; ventral and dorsal faces strongly convex; inner hypostomal teeth visible. Sides of the head with sparse, long, suberect pilosity; whole head with moderately dense, long, suberect to erect 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. Occipital lobes shiny, with sparse and thick rugoreticulation; genae shiny, with moderately dense to dense, fine rugoreticulation; frons and malar area with sparse, thick, and longitudinal rugae, interspaces smooth or with fine rugulae. Centre of clypeus smooth and shiny, lateral sides with longitudinal rugae; median notch present, narrow, and shallow; median longitudinal carina present but sometimes indistinct; lateral longitudinal carinae absent. Scape, when laid back, reaching slightly beyond the midlength of head; pilosity decumbent to erect (Fig. 35B, D). Inner hypostomal teeth distinct, closely spaced, triangular, with rounded apex and moderately narrow base; outer hypostomal teeth distinct, slightly lower than inner hypostomal teeth, lobe-like, with base wide and tops directed slightly inward (Fig. 84P). Mesosoma. In lateral view, promesonotum short, angular, and moderately high, posterior mesonotum with tubercle-like projection, dropping steeply to propodeum; promesonotal groove absent; metanotal groove absent; propodeal spines long, massive basally, with acute apex; humeral area laterally weakly produced (Fig. 35D). Surface shiny, with thick and dense rugoreticulation, sometimes sculpture weakening on dorsum. Pilosity moderately sparse, long, and erect (Fig. 35D, F). Petiole. Shagreened; peduncle relatively long, without horizontal lobes on its basal part; node low, triangular, with rounded apex, in rear view node slightly convex; pilosity moderately sparse and erect (Fig. 35D, F). Postpetiole. Shagreened; in dorsal view sides with acute, triangular, moderately long projections; pilosity long, moderately sparse, and erect (Fig. 35D, F). Petiole. First gastral tergite shagreened, at least on its basal part; pilosity moderately sparse, long, and erect (Fig. 35D, F). Colour. Head and mesosoma reddish brown to brown; dorsum of mesosoma dark brown; gaster brown; legs yellow to brown (Fig. 35D, F).

Figure 35. 

Pheidole ferruginea sp. nov., full-face view (A), profile (C), and dorsal view (E) of paratypeminor worker (CASENT0494994) and full-face view (B), profile (D), and dorsal view (F) of holotype major worker (CASENT0494997).

Minor workers. Measurements (N = 10): HL: 0.52–0.57 (0.55); HW: 0.44–0.48 (0.46); SL: 0.57–0.63 (0.6); EL: 0.11–0.13 (0.12); WL: 0.67–0.72 (0.7); PSL: 0.15–0.18 (0.16); MTL: 0.44–0.49 (0.46); PNW: 0.31–0.35 (0.33); PTW: 0.07–0.09 (0.08); PPW: 0.13–0.14 (0.13); CI: 80.9–86.2 (83.9); SI: 124.6–134.8 (130.0); PSLI: 27.5–31.4 (29.3); PPI: 57.4–67.9 (63.1); PNI: 69.6–74.8 (71.7); MTI: 96.9–102.3 (100.9). Head. Occipital margin straight or indistinctly concave; occipital carina absent (Fig. 35A). Pilosity sparse, long, suberect to erect. Whole head foveolate, sometimes with additional longitudinal rugae on frons. Clypeus foveolate; median longitudinal carina absent; two lateral longitudinal carinae absent. Scape, when laid back, surpassing posterior head margin by two-fifths of its length; pilosity suberect to erect (Fig. 35A, C). Mesosoma. In lateral view, promesonotum box-like; promesonotal groove absent; metanotal groove absent; propodeal spines very long, massive basally, with acute apex (Fig. 35C). Whole surface foveolate, sometimes with additional rugae on mesosomal dorsum; sometimes katepisternum with smooth notch. Pilosity sparse, long, and erect (Fig. 35C, E). Petiole. Peduncle relatively long and thin; node low, globular; with few long, erect setae (Fig. 35C, E). Postpetiole. Short, low, and convex; with few long, erect setae at the anterior edge (Fig. 35C, E). Petiole. Pilosity sparse and erect (Fig. 35C, E). Colour. Unicolourous, yellow to brown (Fig. 35C, E).

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.

Pheidole rugocephala sp. nov.

Figs 36A–F, 85U, 88E

Type material

Holotype. Madagascar. •1 major worker; Toamasina; 6.9 km NE Ambanizana, Ambohitsitondroina; -15.58506, 50.00952; alt. 825 m; 2 Dec 1993; B.L. Fisher leg.; BLF00976, CASENT0923224 (CASC). Paratypes. Madagascar. •1w., 1q.; same data as for holotype; CASENT0198470, CASENT0872239 (CASC).

Other material

Madagascar. –Antsiranana: •1q.; 9.2 km WSW Befingotra, Rés. Anjanaharibe-Sud; -14.75, 49.46667; alt. 1200 m; 9 Nov 1994; B.L. Fisher leg. CASENT0196913 (CASC). •1s.; Masoala National Park; -15.3014, 50.22776; alt. 280 m; 7 Mar 2014; B.L. Fisher et al. leg.; CASENT0377718 (CASC). •1s.; Masoala National Park; -15.32331, 50.30751; alt. 60 m; 10 Mar 2014; B.L. Fisher et al. leg.; CASENT0376820 (CASC). –Toamasina: •1w., 1s.; Montagne d’Anjanaharibe, 18.0 km 21°NNE Ambinanitelo; -15.18833, 49.615; alt. 470 m; 8 Mar 2003; B.L. Fisher et al. leg.; CASENT0037624, CASENT0037758 (CASC). •1s.; Montagne d’Anjanaharibe, 19.5 km 27°NNE Ambinanitelo; -15.17833, 49.635; alt. 1100 m; 12 Mar 2003; B.L. Fisher et al. leg.; CASENT0038365 (CASC). •4s.; Parc National Mananara-Nord, 7.1 km 261° Antanambe; -16.455, 49.7875; alt. 225 m; 14 Nov 2005; B.L. Fisher et al. leg.; CASENT0068924, CASENT0069459, CASENT0069581, CASENT0069598 (CASC). •1s.; Reserve Betampona, Camp Vohitsivalana, 37.1 km 338° Toamasina; -17.88667, 49.2025; alt. 520 m; 1 Dec 2005; B.L. Fisher et al. leg.; CASENT0069316 (CASC).

Diagnosis

Head in full-face view cordate, widened posteriorly; sides of the head with moderately dense, long, suberect to erect pilosity; antennal scrobes present, strongly developed, well delimited and forming distinct dorsal concavity beneath frontal carina; scrobe surface foveolate; delimited ventrally and posteriorly by carinulae; propodeal spines long (PSL: 0.13–0.2 (0.15)); first gastral tergite shagreened on its basal part. Minor workers. Head foveolate; propodeal spines long (PSL: 0.09); promesonotum box-like; mesosoma foveolate; katepisternum with smooth notch.

Description

Major workers. Measurements (N = 10): HL: 0.86–1.18 (0.97); HW: 0.87–1.12 (0.93); SL: 0.42–0.52 (0.45); EL: 0.12–0.14 (0.13); WL: 0.72–0.99 (0.82); PSL: 0.13–0.2 (0.15); MTL: 0.43–0.51 (0.45); PNW: 0.47–0.62 (0.53); PTW: 0.13–0.19 (0.14); PPW: 0.35–0.53 (0.41); CI: 95.3–101.3 (97.5); SI: 46.0–50.9 (48.2); PSLI: 14.3–17.0 (15.7); PPI: 30.9–36.9 (33.6); PNI: 53.7–59.9 (56.4); MTI: 45.5–52.3 (48.5). Head. In full-face view cordate, widened posteriorly (Fig. 36B). In lateral view oval; ventral face convex; dorsal face relatively flat; inner hypostomal teeth visible. Sides of the head with moderately dense, long, suberect to erect pilosity; whole head with moderately dense, long, suberect to erect pilosity. Antennal scrobes strongly developed, well delimited and forming distinct dorsal concavity beneath frontal carina, scrobe surface foveolate, delimited ventrally and posteriorly by carinulae. Occipital lobes shiny, with sparse and thick rugae, sculpture weakening posteriorly; frons and malar area with thick, sparse, longitudinal or irregular rugae, interspaces smooth; genae with dense and fine rugulae. Centre of clypeus smooth and shiny, lateral sides with longitudinal rugae; median notch present, narrow, and shallow; median longitudinal carina absent; lateral longitudinal carinae present. Scape, when laid back, reaching slightly beyond the midlength of head; pilosity decumbent to erect (Fig. 36B, D). Inner hypostomal teeth distinct, closely spaced, triangular, with rounded apex pointed inward; outer hypostomal teeth distinct, slightly higher than inner hypostomal teeth, lobe-like, with base wide and acute tops directed outward; inner and outer teeth closely spaced and connected by concavity (Fig. 85U). Mesosoma. In lateral view, promesonotum short, angular, and moderately high, posterior mesonotum with tubercle-like projection, dropping steeply to propodeum; promesonotal groove absent; metanotal groove absent; propodeal spines long, with narrow base and acute apex; humeral area laterally weakly produced (Fig. 36D). Surface shiny, with sparse rugoreticulation, dorsum with weaker sculpture and sometimes smooth patches. Pilosity moderately sparse, long, and erect (Fig. 36D, F). Petiole. Finely foveolate; peduncle relatively long, with horizontal, triangular lobes on its basal part; node high, triangular, with rounded apex, in rear view node dorsoventrally depressed; pilosity moderately sparse and erect (Fig. 36D, F). Postpetiole. Finely foveolate; short; in dorsal view sides with acute, very wide, and long triangular projections; pilosity long, moderately dense and erect (Fig. 36D, F). Petiole. First gastral tergite shagreened on its basal part; pilosity moderately dense, very long and erect (Fig. 36D, F). Colour. Head and mesosoma reddish brown to brown; gaster brown; legs yellow to brown (Fig. 36D, F).

Figure 36. 

Pheidole rugocephala sp. nov., full-face view (A), profile (C), and dorsal view (E) of paratype minor worker (CASENT0198470) and full-face view (B), profile (D), and dorsal view (F) of holotype major worker (CASENT0923224).

Minor workers. Measurements (N = 1): HL: 0.48; HW: 0.43; SL: 0.42; EL: 0.09; WL: 0.6; PSL: 0.09; MTL: 0.35; PNW: 0.3; PTW: 0.08; PPW: 0.14; CI: 89.4; SI: 99.1; PSLI: 18.6; PPI: 58.6; PNI: 69.2; MTI: 82.2. Head. Occipital margin straight or indistinctly concave; occipital carina absent (Fig. 36A). Pilosity sparse, long, suberect to erect. Whole head foveolate. Clypeus with median longitudinal carina absent; two lateral longitudinal carinae absent. Scape, when laid back, surpassing posterior head margin by one-fifth of its length; pilosity suberect to erect (Fig. 36A, C). Mesosoma. In lateral view, promesonotum box-like; promesonotal groove present; metanotal groove present; propodeal spines moderately long, triangular, with acute apex (Fig. 36C). Whole surface foveolate, only katepisternum with smooth notch. Pilosity sparse, long and erect (Fig. 36C, E). Petiole. Shiny; peduncle foveolate, short and thin; node finely foveolate, globular; with few long, erect setae (Fig. 36C, E). Postpetiole. Finely foveolate with smooth centre; short, low and convex; with few long, erect setae at the anterior edge (Fig. 36C, E). Petiole. With few long, erect setae (Fig. 36C, E). Colour. Unicolourous, yellow (Fig. 36C, E).

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.

Comments

This species is most similar to P. ferruginea sp. nov. Major workers. Pheidole rugocephala sp. nov. can be distinguished from P. ferruginea sp. nov. by foveolate surface of antennal scrobes; lack of longitudinal rugae on frons, inner hypostomal teeth pointed inward, presence of smooth surface on promesonotum, and presence of horizontal lobes on the basal part of petiolar peduncle. Minor workers. Pheidole rugocephala sp. nov. can be distinguished from P. ferruginea sp. nov. by short and triangular propodeal spines, never foveolate clypeus, and presence of promesonotal and metanotal grooves.

Pheidole manantenensis sp. nov.

Figs 37A–F, 85F, 87J

Type material

Holotype. Madagascar. •1 major worker; Antsiranana; Parc National de Marojejy, Manantenina River, 27.6 km 35°NE Andapa, 9.6 km 327°NNW Manantenina; -14.435, 49.76; alt. 775 m; 12 Dec 2005; Fisher et al. leg.; BLF13478, CASENT0068456 (CASC). Paratypes. Madagascar. •1w., 1q.; same data as for holotype; CASENT0068457, CASENT0923218 (CASC).

Other material

Madagascar. –Antsiranana: •2w.; Makirovana Forest; -14.104, 50.03574; alt. 225 m; 4 May 2011; B.L. Fisher et al. leg.; CASENT0230806, CASENT0230809 (CASC). •1w.; Makirovana Forest; -14.16044, 49.95216; alt. 550 m; 1 May 2011; B.L. Fisher et al. leg.; CASENT0231430 (CASC). •4w.; Makirovana Forest; -14.16666, 49.95; alt. 715 m; 1 May 2011; B.L. Fisher et al. leg.; CASENT0243103, CASENT0243118, CASENT0243120, CASENT0243129 (CASC). –Toamasina: •1w., 1s.; Montagne d’Akirindro 7.6 km 341°NNW Ambinanitelo; -15.28833, 49.54833; alt. 600 m; 17 Mar 2003; B.L. Fisher et al. leg.; CASENT0038921, CASENT0039038 (CASC).

Diagnosis

Head in full-face view cordate, widened posteriorly; sides of the head with moderately dense, short, suberect to erect pilosity; antennal scrobes present, weakly impressed, and indistinctly delimited ventrally and posteriorly by carinulae; scrobe surface foveolate, with indistinct, thick, moderately sparse, irregular rugae; propodeal spines long (PSL: 0.2–0.26 (0.24)); gaster shagreened; inner hypostomal teeth distinct, high, narrow, and triangular, closely spaced, with rounded apex; outer hypostomal teeth approximately as high as inner hypostomal teeth, but thinner and with wider base, lobe-like. Minor workers. Head and mesosoma foveolate; propodeal spines long (PSL: 0.14–0.17 (0.15)); promesonotum low, convex, short, with posterior declivity smoothly declining towards propodeum.

Description

Major workers. Measurements (N = 10): HL: 0.96–1.13 (1.07); HW: 0.95–1.15 (1.07); SL: 0.52–0.58 (0.55); EL: 0.1–0.12 (0.12); WL: 0.79–0.94 (0.88); PSL: 0.2–0.26 (0.24); MTL: 0.49–0.57 (0.52); PNW: 0.45–0.54 (0.51); PTW: 0.12–0.15 (0.14); PPW: 0.33–0.44 (0.4); CI: 98.2–102.5 (100.5); SI: 48.3–54.7 (51.1); PSLI: 20.0–24.5 (22.1); PPI: 31.9–38.2 (34.8); PNI: 45.6–48.9 (47.1); MTI: 45.6–51.3 (48.2). Head. In full-face view cordate, slightly longer than wide, anterior of eyes slightly convex, posterior of eyes convex, occipital margins of lobes convex (Fig. 37B). 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, short, suberect to erect pilosity; whole head with moderately dense, long, suberect to erect pilosity. Antennal scrobes impressed and delimited ventrally and posteriorly by carinulae; scrobe surface foveolate, with indistinct, thick, moderately sparse, irregular rugae. Occipital lobes shiny, foveolate, with additional thick, sparse, irregular rugae, rugae weakening posteriorly; frons with moderately sparse, thick, longitudinal rugae, on the posterior part rugae longitudinal to irregular, interspaces superficially foveolate; genae foveolate, with additional, indistinct, and irregular rugulae; malar area with thick, sparse, longitudinal rugae, interspaces smooth to indistinctly rugulose. Centre of clypeus shiny and with indistinct, short rugulae, lateral sides with longitudinal rugae; median notch present, moderately wide, 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. 37B, D). Inner hypostomal teeth distinct, high, narrow, and triangular, closely spaced, with rounded apex; outer hypostomal teeth approximately as high as inner hypostomal teeth, but thinner and with wider base, lobe-like (Fig. 85F). Mesosoma. In lateral view, promesonotum low and arched, dorsal mesonotum slightly concave, posterior mesonotum relatively steep, with small tubercle-like projections; promesonotal groove absent; metanotal groove absent; propodeal spines long, triangular, thin, with sharp apex and relatively narrow base; humeral area laterally weakly produced (Fig. 37D). Surface foveolate, with additional sparse and indistinct rugoreticulation; dorsoventral propodeum with fading sculpture. Pilosity sparse, long, erect (Fig. 37D, F). Petiole. Shiny; peduncle moderately long, foveolate, with distinct, short horizontal lobes on its basal part; node relatively high, triangular, with rounded apex, in rear view node slightly convex; pilosity moderately sparse and erect (Fig. 37D, F). Postpetiole. Shiny and foveolate; in dorsal view sides with moderately long, wide, acute, and triangular projections; pilosity moderately long, and erect (Fig. 37D, F). Petiole. Shiny and shagreened; pilosity moderately sparse, long, and erect (Fig. 37D, F). Colour. Brown to dark brown; legs yellow; malar area brighter that the rest of the head (Fig. 37D, F).

Figure 37. 

Pheidole manantenensis sp. nov., full-face view (A), profile (C), and dorsal view (E) of paratype minor worker (CASENT0923218) and full-face view (B), profile (D), and dorsal view (F) of holotype major worker (CASENT0068456).

Minor workers. Measurements (N = 10): HL: 0.47–0.51 (0.49); HW: 0.41–0.46 (0.44); SL: 0.45–0.54 (0.49); EL: 0.08–0.11 (0.09); WL: 0.58–0.64 (0.62); PSL: 0.14–0.17 (0.15); MTL: 0.35–0.38 (0.37); PNW: 0.28–0.32 (0.31); PTW: 0.06–0.08 (0.07); PPW: 0.1–0.13 (0.11); CI: 85.2–93.0 (88.9); SI: 105.1–119.7 (111.8); PSLI: 29.2–34.3 (31.5); PPI: 60.0–73.2 (65.8); PNI: 67.5–74.1 (70.7); MTI: 79.8–88.6 (84.0). Head. Occipital margin straight or indistinctly concave; occipital carina indistinct (Fig. 37A). Pilosity moderately dense, long, suberect. Head foveolate; genae with fading sculpture. Clypeus 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 erect (Fig. 37A, C). Mesosoma. In lateral view, promesonotum low, convex, short, with posterior declivity smoothly declining towards propodeum; promesonotal groove absent; metanotal groove present; propodeal spines long, triangular, with acute apex and narrow base (Fig. 37C). Sculpture foveolate. Pilosity moderately sparse, short, and erect (Fig. 37C, E). Petiole. Peduncle moderately short and thin; with few short, erect setae (Fig. 37C, E). Postpetiole. Short, low, and convex; with few short, erect setae (Fig. 37C, E). Petiole. With sparse, erect pilosity (Fig. 37C, E). Colour. Dark yellow (Fig. 37C, E).

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.

Pheidole vohemarensis sp. nov.

Figs 38A–F, 85AA, 88K

Type material

Holotype. Madagascar. •1 major worker; Antsiranana; Ambondrobe, 41.1 km 175° Vohemar; -13.71533, 50.10167; alt. 10 m; 1 Dec 2004; Fisher et al. leg.; BLF11264, CASENT0107946 (CASC). Paratypes. Madagascar. •2w., 1q.; same data as for holotype; CASENT0107947, CASENT0217986, CASENT0872084 (CASC).

Other material

Madagascar. –Antsiranana: •1s.; Binara Forest; -13.26207, 49.60505; alt. 692 m; 20 Oct 2013; B.L. Fisher et al. leg.; CASENT0353324 (CASC). •2w., 1s.; Forêt d’Analabe, 30.0 km 72°ENE Daraina; -13.08333, 49.90833; alt. 30 m; 27 Nov 2003; B.L. Fisher et al. leg.; CASENT0041430, CASENT0041449, CASENT0048846 (CASC). •1w., 2s.; Forêt de Bekaraoka, 6.8 km 60°ENE Daraina; -13.16667, 49.71; alt. 150 m; 7 Dec 2003; B.L. Fisher et al. leg.; CASENT0044312, CASENT0044422, CASENT0044438 (CASC). •2w., 2s.; Forêt de Binara, 7.5 km 230°SW Daraina; -13.255, 49.61667; alt. 375 m; 1 Dec 2003; B.L. Fisher et al. leg.; CASENT0041832, CASENT0041833, CASENT0041890, CASENT0041895 (CASC). •1s.; Parc National de Marojejy, Manantenina River, 27.6 km 35°NE Andapa, 9.6 km 327°NNW Manantenina; -14.435, 49.76; alt. 775 m; 15 Nov 2003; B.L. Fisher et al. leg.; CASENT0045241 (CASC). •1w.; Rés. Analamerana, 28.4 km 99° Anivorano-Nord; -12.74667, 49.49483; alt. 60 m; 5 Dec 2004; B.L. Fisher et al. leg.; CASENT0054163 (CASC).

Diagnosis

Head, in full-face view, cordate, widened posteriorly; sides of the head with dense, long, suberect to erect pilosity; antennal scrobes present, impressed, and indistinctly delimited ventrally and posteriorly by carinulae; scrobe surface foveolate, with distinct, thick, moderately sparse, longitudinal rugae; propodeal spines long (PSL: 0.19–0.23 (0.21)); gaster shagreened; inner hypostomal teeth distinct, high, narrow, and triangular, closely spaced, with rounded apex; outer hypostomal teeth slightly lower and thinner than inner hypostomal teeth, lobe-like. Minor workers. Head foveolate, genae with smooth notch; propodeal spines long (PSL: 0.12–0.14 (0.13)); promesonotum low, slightly convex, short, with posterior declivity relatively steep; mesosoma foveolate, katepisternum and mesonotum with smooth notches.

Description

Major workers. Measurements (N = 10): HL: 1.03–1.11 (1.08); HW: 0.99–1.1 (1.05); SL: 0.49–0.57 (0.52); EL: 0.12–0.14 (0.13); WL: 0.83–0.91 (0.87); PSL: 0.19–0.23 (0.21); MTL: 0.48–0.54 (0.51); PNW: 0.48–0.57 (0.53); PTW: 0.13–0.17 (0.15); PPW: 0.4–0.51 (0.47); CI: 95.1–99.3 (98.0); SI: 47.1–53.1 (49.7); PSLI: 18.2–20.8 (19.1); PPI: 29.1–36.1 (33.1); PNI: 45.9–52.9 (49.9); MTI: 44.8–50.8 (48.8). Head. In full-face view cordate, slightly longer than wide, anterior of eyes slightly convex, posterior of eyes convex, occipital margins of lobes convex (Fig. 38B). In lateral view sub-oval; ventral and dorsal faces convex; dorsal face not depressed posteriorly; inner hypostomal teeth visible. Sides of the head with dense, long, suberect to erect pilosity; whole head with dense, short, suberect to erect pilosity. Antennal scrobes impressed and indistinctly delimited ventrally and posteriorly by carinulae; scrobe surface foveolate, with distinct, thick, moderately sparse, longitudinal rugae. Occipital lobes shiny, with thick, sparse, irregular rugae, rugae weakening posteriorly, interspaces with fine and dense rugulae; frons with moderately sparse, thick, longitudinal rugae, on the posterior part rugae longitudinal to irregular, interspaces with fine and dense rugulae, sometimes rugulae fading on the central part of frons; genae with dense, moderately thick, and irregular rugulae, interspaces with dense and fine rugulae; malar area with thick, sparse, longitudinal rugae, interspaces smooth to indistinctly rugulose. Centre of clypeus shiny and with indistinct, short rugulae, lateral sides with longitudinal rugae; median notch present, moderately wide 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. 38B, D). Inner hypostomal teeth distinct, high, narrow, and triangular, closely spaced, with rounded apex; outer hypostomal teeth slightly lower and thinner than inner hypostomal teeth, lobe-like (Fig. 85AA). Mesosoma. In lateral view, promesonotum relatively low and arched, posterior mesonotum relatively steep, without tubercle-like projections; promesonotal groove absent; metanotal groove absent; propodeal spines long, triangular, thin, with sharp apex and moderately narrow base; humeral area laterally weakly produced (Fig. 38D). Surface foveolate, with additional sparse to dense, moderately thick rugoreticulation; dorsoventral propodeum with fading sculpture. Pilosity moderately sparse, long, erect (Fig. 38D, F). Petiole. Shiny; peduncle moderately long, foveolate, with very distinct, wide, horizontal lobes on its basal part; node relatively high, thin, with rounded apex, in rear view node with deep dorsoventral concavity; pilosity long and erect (Fig. 38D, F). Postpetiole. Shiny and foveolate; in dorsal view sides with long, wide, acute, and triangular projections; pilosity long and erect (Fig. 38D, F). Petiole. Shiny and shagreened; pilosity dense, long, and erect (Fig. 38D, F). Colour. Bright brown to dark brown, sometimes head with brighter colouration; legs yellow (Fig. 38D, F).

Figure 38. 

Pheidole vohamarensis sp. nov., full-face view (A), profile (C), and dorsal view (E) of paratype minor worker (CASENT0107947) and full-face view (B), profile (D), and dorsal view (F) of holotype major worker (CASENT0107946).

Minor workers. Measurements (N = 10): HL: 0.44–0.53 (0.49); HW: 0.4–0.45 (0.43); SL: 0.47–0.51 (0.5); EL: 0.08–0.11 (0.1); WL: 0.58–0.63 (0.6); PSL: 0.12–0.14 (0.13); MTL: 0.35–0.4 (0.37); PNW: 0.28–0.33 (0.3); PTW: 0.07–0.09 (0.08); PPW: 0.1–0.13 (0.11); CI: 82.6–94.3 (86.7); SI: 110.2–122.0 (115.8); PSLI: 23.4–30.9 (26.5); PPI: 60.4–82.9 (71.7); PNI: 67.2–75.3 (71.0); MTI: 81.0–90.9 (86.7). Head. Occipital margin straight or indistinctly concave; occipital carina indistinct (Fig. 38A). Pilosity moderately dense, long, suberect. Head foveolate; genae with smooth notch. Clypeus 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 erect (Fig. 38A, C). Mesosoma. In lateral view, promesonotum low, slightly convex, short, with posterior declivity relatively steep; promesonotal groove absent; metanotal groove present; propodeal spines long, triangular, with acute apex and narrow base (Fig. 38C). Sculpture foveolate; katepisternum and mesonotum with smooth notches. Pilosity moderately sparse, short, and erect (Fig. 38C, E). Petiole. Peduncle moderately short and thin; with few short, erect setae (Fig. 38C, E). Postpetiole. Short, low, and convex; with few short, erect setae (Fig. 38C, E). Petiole. With moderately sparse, erect pilosity (Fig. 38C, E). Colour. Yellow (Fig. 38C, E).

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, triangular; 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.

Pheidole longipilosa sp. nov.

Figs 39A–F, 84X, 87B

Type material

Holotype. Madagascar. •1 major worker; Toliara; Forêt Classée d’Analavelona, 29.2 km 343°NNW Mahaboboka; -22.675, 44.19; alt. 1100 m; 18 Feb 2003; B.L. Fisher et al. leg.; CASENT0498298 (CASC). Paratypes. Madagascar. •9w.; same data as for holotype; CASENT0235043, CASENT0498295CASENT0498297, CASENT0872154CASENT0872158 (CASC).

Other material

Madagascar. –Fianarantsoa: •1w.; Parc National d’Isalo, Sahanafa River, 29.2 km 351°N Ranohira; -22.31333, 45.29167; alt. 500 m; 10 Feb 2003; Fisher et al. leg.; CASENT0031767 (CASC). –Toliara: •7w., 4s., 3q.; Forêt Classée d’Analavelona, 29.2 km 343°NNW Mahaboboka; -22.675, 44.19; alt. 1100 m; 18 Feb 2003; Fisher et al. leg.; CASENT0031011, CASENT0031099, CASENT0049501, CASENT0073507, CASENT0235043, CASENT0496936, CASENT0496937, CASENT0496938 (CASC). •1w., 5s.; Forêt Classée d’Analavelona, 29.4 km 343°NNW Mahaboboka; -22.675, 44.18667; alt. 1050 m; 21 Feb 2003; Fisher et al. leg.; CASENT0023935, CASENT0023938, CASENT0023943, CASENT0023946, CASENT0023947, CASENT0024032 (CASC).

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.

Description

Major workers. Measurements (N = 10): HL: 1.05–1.11 (1.09); HW: 0.98–1.04 (1.0); SL: 0.46–0.52 (0.48); EL: 0.13–0.16 (0.13); WL: 0.84–0.97 (0.88); PSL: 0.14–0.17 (0.15); MTL: 0.47–0.5 (0.49); PNW: 0.52–0.58 (0.54); PTW: 0.12–0.14 (0.13); PPW: 0.35–0.4 (0.37); CI: 90.0–93.9 (91.7); SI: 46.5–51.8 (48.1); PSLI: 12.5–15.4 (13.7); PPI: 31.9–39.0 (34.9); PNI: 53.1–55.5 (54.3); MTI: 45.8 –50.9 (48.8). Head. In full-face view longer than wide, anterior of eyes slightly convex, posterior of eyes convex (Fig. 39B). In lateral view sub-oval; ventral and dorsal faces convex; dorsal face not depressed posteriorly; inner hypostomal teeth visible. Sides of the head with dense, very long, erect pilosity; whole head with moderately dense, long, suberect to erect pilosity. Antennal scrobes indistinct and not delimited by carinulae; scrobe surface shiny, with thick, longitudinal, and long rugae; interspaces distinctly rugulose. Occipital lobes shiny, with thick, irregular rugae, interspaces with distinct, irregular rugulae not fading posteriorly; frons with moderately dense, thick, longitudinal rugae, interspaces smooth, with indistinct rugulae weakening anteriorly; genae shiny, with dense and thin, irregular rugulae, central part with smooth notch; malar area with thin, longitudinal, dense rugae, interspaces with distinct rugulae. Centre of clypeus shiny and smooth, lateral sides with longitudinal rugulae; median notch present, narrow, and moderately deep; median longitudinal carina present, indistinct; lateral longitudinal carinae present. Scape, when laid back, reaching the midlength of head; pilosity erect (Fig. 39B, D). 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 (Fig. 84X). Mesosoma. In lateral view, promesonotum relatively high and arched, posterior mesonotum relatively steep, with tubercle-like projections; promesonotal groove absent; metanotal groove absent; propodeal spines small to moderately long, triangular, with sharp apex and wide base; humeral area laterally weakly produced (Fig. 39D). Surface shiny, promesonotum foveolate with additional indistinct, sparse rugoreticulations, sometimes sculpture fading on dorsal surface and lower parts of lateral sides; katepisternum rugoreticulate, with smooth notch; anepisternum and lateral sides of propodeum with distinct rugoreticulation; dorsoventral side of propodeum smooth or with indistinct sculpture. Pilosity moderately dense, very long, and erect (Fig. 39D, F). Petiole. Shiny; peduncle short, finely foveolate, with indistinct horizontal lobes on its basal part; node with fading sculpture, relatively low, triangular, with rounded apex, in rear view node slightly convex; pilosity long and erect (Fig. 39D, F). Postpetiole. Shiny, finely shagreened, with dorsum at least partially smooth; in dorsal view sides with moderately short, acute, and triangular projections; pilosity long, long and erect (Fig. 39D, F). Petiole. Shiny and smooth, only basal part of first gastral tergite finely shagreened; pilosity dense, long, and erect (Fig. 39D, F). Colour. Reddish brown to bright brown (Fig. 39D, F).

Figure 39. 

Pheidole longipilosa sp. nov., full-face view (A), profile (C), and dorsal view (E) of paratype minor worker (CASENT0235043) and full-face view (B), profile (D), and dorsal view (F) of holotype major worker (CASENT0498298).

Minor workers. Measurements (N = 10): HL: 0.49–0.55 (0.51); HW: 0.44–0.47 (0.45); SL: 0.46–0.47 (0.46); EL: 0.09–0.11 (0.1); WL: 0.57–0.6 (0.58); PSL: 0.09–0.11 (0.1); MTL: 0.33–0.36 (0.34); PNW: 0.29–0.32 (0.31); PTW: 0.06–0.08 (0.07); PPW: 0.1–0.12 (0.11); CI: 85.7–90.4 (88.6); SI: 100.2–105.6 (103.0); PSLI: 18.1–21.8 (20.0); PPI: 56.5–64.5 (60.8); PNI: 65.9–70.1 (68.2); MTI: 73.5–80.1 (76.1). Head. Occipital margin straight or indistinctly concave; occipital carina absent (Fig. 39A). Pilosity moderately sparse, moderately long, suberect to erect. Head shiny, foveolate, with additional short, longitudinal, thick rugae on frons; genae with fading sculpture and smooth notch. Clypeus smooth, with basal part finely rugulose; median longitudinal carina present; two lateral longitudinal carinae indistinct. Scape, when laid back, surpassing the posterior head margin by one-fifth of its length; pilosity suberect to erect (Fig. 39A, C). Mesosoma. In lateral view, promesonotum low, convex, short, with posterior declivity smoothly declining towards propodeum; promesonotal groove absent; metanotal groove absent; propodeal spines short, triangular, with acute apex (Fig. 39C). Sculpture foveolate; katepisternum smooth; anepisternum and mesonotum with fading sculpture and sometimes smooth. Pilosity moderately sparse, long, and erect (Fig. 39C, E). Petiole. Peduncle short and thin with ventral face slightly convex; with few short, erect setae (Fig. 39C, E). Postpetiole. Short, low, and convex; with few short, erect setae (Fig. 39C, E). Petiole. With moderately sparse, erect pilosity (Fig. 39C, E). Colour. Yellow (Fig. 39C, E).

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.

This group contains two sympatric species: P. annemariae and P. marieannae sp. nov. Both species are known from northern part of the evergreen rainforest biome. The distribution of Pheidole annemariae spreads between Toamasina and Andapa, while P. marieannae is known from lowlands between Antalaha and Vohemar.

Key to the P. annemariae group

1 Major workers. Genae with smooth notch, inner and outer hypostomal teeth connected by indistinct concavity, posterior mesonotum with distinct teeth-like projections, propodeal spines very long (Fig. 40A, E). Minor workers. Mesonotal spines distinct, small, and triangular, katepisternum, anepisternum, and mesonotum never entirely smooth (Fig. 40B) P. annemariae Forel
Major workers. Genae never with smooth notch, inner and outer hypostomal teeth not connected by concavity, posterior mesonotum with tubercle-like projections, propodeal spines moderately long (Fig. 40C, F). Minor workers. Mesonotal spines indistinct, katepisternum, anepisternum, and mesonotum entirely smooth (Fig. 40D) P. marieannae sp. nov.
Figure 40. 

Pheidole annemariae Forel, head and profile of major worker (A), hypostomal teeth (E), profile of minor worker (B). Pheidole marieannae sp. nov., head of major worker (C), hypostomal teeth (F), profile of minor worker (D).

Pheidole annemariae Forel, 1918

Figs 41A–F, 84E, 86E

Type material

Pheidole annemariae Forel, 1918: 152 (s.w.q.). Lectotype [designated here]: major worker (middle specimen, CASENT0101688): Madagascar, Toamasina, Ilôt Prune, coll. Friederichs (MHNG) [examined]. Paralectotypes: 2 major workers (CASENT0810541, top and bottom specimens, the same pin as lectotype) (MHNG) [examined], 3 minor workers (CASENT0101829) (MHNG) [examined], 1 minor worker (CASENT0923206) (MHNG) [examined]: the same data as lectotype.

Other material

Madagascar. –Antsiranana: •9w., 6s., 2q.; Ambondrobe, 41.1 km 175° Vohemar; -13.71533, 50.10167; alt. 10 m; 29 Nov 2004; B.L. Fisher et al. leg.; CASENT0056072, CASENT0056515, CASENT0056530, CASENT0056545, CASENT0056546, CASENT0056667, CASENT0056671, CASENT0107936, CASENT0109695, CASENT0109742, CASENT0110543, CASENT0110544 (CASC). •1w.; Binara Forest; -13.26392, 49.59919; alt. 1065 m; 18 Oct 2013; B.L. Fisher et al. leg.; CASENT0369879 (CASC). •3w., 1s., 1q.; Cap Est, Forêt d’Andranoanala; -15.24644, 50.46538; alt. 15 m; 16 Mar 2014; B.L. Fisher et al. leg.; CASENT0376154, CASENT0376165, CASENT0376166 (CASC). •7w., 3s.; Forêt de Binara, 9.1 km 233°SW Daraina; -13.26333, 49.60333; alt. 800 m; 5 Dec 2003; B.L. Fisher et al. leg.; CASENT0076461, CASENT0076462, CASENT0076463, CASENT0043234 (CASC). •4w., 1q.; Makirovana Forest; -14.17066, 49.95409; alt. 415 m; 28 Apr 2011; B.L. Fisher et al. leg.; CASENT0231267, CASENT0231268, CASENT0236175 (CASC). •4w.; Makirovana Forest; -14.16666, 49.95; alt. 715 m; 2 May 2011; B.L. Fisher et al. leg.; CASENT0231155, CASENT0231282 (CASC). •1w.; Masoala, Cap Est, Forêt d’Andranoanala; -15.26158, 50.4758; alt. 15 m; 15 Mar 2014; B.L. Fisher et al. leg.; CASENT0377896 (CASC). •9w., 3s.; Parc National de Marojejy, Manantenina River, 28.0 km 38°NE Andapa, 8.2 km 333°NNW Manantenina; -14.43667, 49.775; alt. 450 m; 12 Nov 2003; B.L. Fisher et al. leg.; CASENT0077287CASENT0077290 (CASC). –Toamasina: •1w., 1s.; Nosy Mangabe, 7.43 km S Maroantsetra; -15.4973, 49.76223; alt. 3 m; 25 Jul 2007; B.L. Fisher et al. leg.; CASENT0129807 (CASC). •5w.; P.N. Masoala, 40 km 154°SSE Maroantsetra; -15.72667, 49.95667; alt. 150 m; 14 Oct 2001; Dejean et al. leg.; CASENT0004830, CASENT0004831, CASENT0004970CASENT0004972 (CASC). •2w.; Tampolo, Masoala Peninsula, 40.4 km 154°SSE Maroantsetra; -15.73, 49.96; alt. 30 m; 28 Nov 2001; B.L. Fisher et al. leg.; CASENT0418227, CASENT0418228 (CASC).

Diagnosis

Head in full-face view rectangular, slightly longer than wide, anterior of eyes slightly convex, posterior of eyes convex; sides of the head with moderately dense, long, erect pilosity; antennal scrobes present, impressed and not delimited; scrobe surface rugo-foveolate, with distinct, thin, moderately sparse, longitudinal rugae; propodeal spines long (PSL: 0.21–0.24 (0.22)); gaster finely shagreened; inner hypostomal teeth distinct, moderately high, closely spaced, triangular, with rounded apex and moderately narrow base; outer hypostomal teeth approximately the same size as inner hypostomal teeth, triangular; inner and outer teeth closely spaced and connected by indistinct concavity. Minor workers. Head foveolate, genae smooth; propodeal spines long (PSL: 0.15–0.18 (0.16)); promesonotum low, slightly convex, short, with posterior declivity relatively steep; mesonotal spines present; mesosoma foveolate, katepisternum and mesonotum with smooth notches.

Redescription

Major workers. Measurements (N = 10): HL: 1.16–1.3 (1.21); HW: 1.06–1.19 (1.11); SL: 0.5–0.52 (0.51); EL: 0.13–0.16 (0.14); WL: 0.87–1.02 (0.93); PSL: 0.21–0.24 (0.22); MTL: 0.51–0.56 (0.53); PNW: 0.56–0.64 (0.59); PTW: 0.13–0.16 (0.15); PPW: 0.38–0.47 (0.43); CI: 90.7–93.9 (91.8); SI: 43.5–47.6 (46.2); PSLI: 17.6–20.1 (18.6); PPI: 31.1–38.1 (33.8); PNI: 51.2–55.2 (53.0); MTI: 44.7–50.0 (47.6). Head. In full-face view rectangular, slightly longer than wide, anterior of eyes slightly convex, posterior of eyes convex, occipital margins of lobes slightly convex (Fig. 41B). 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 visible. In lateral view weakly impressed on vertex. 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 rugo-foveolate, with distinct, thin, moderately sparse, longitudinal rugae. Occipital lobes shiny, with thick, sparse, irregular rugae, interspaces with fine rugulae fading posteriorly; frons with moderately sparse, thick, longitudinal, and sometimes interrupted rugae, interspaces variable, smooth to finely rugoreticulate, most often rugoreticulation fading on the central part of frons; genae with dense, fine and irregular rugulae, area behind eyes with smooth notch; malar area with thick, sparse, longitudinal rugae, interspaces rugoreticulate. Centre of clypeus shiny and sometimes with indistinct, short rugulae, lateral sides with sparse and longitudinal rugae; median notch present, moderately wide and shallow; median longitudinal carina present; lateral longitudinal carinae present. Scape, when laid back, reaching the midlength of head; pilosity suberect to erect (Fig. 41B, D). Inner hypostomal teeth distinct, moderately high, closely spaced, triangular, with rounded apex and moderately narrow base; outer hypostomal teeth approximately the same size as inner hypostomal teeth, triangular; inner and outer teeth closely spaced and connected by indistinct concavity (Fig. 84E). Mesosoma. In lateral view, promesonotum low and arched, dorsal mesonotum finely concave, posterior mesonotum relatively steep, with distinct teeth-like projections; promesonotal groove absent; metanotal groove absent; propodeal spines long, triangular, with sharp apex and moderately wide base; humeral area laterally weakly produced (Fig. 41D). Surface variable; promesonotum finely foveolate to rugo-foveolate, foveolae or rugo-foveolae always fading on lateral sides of pronotum and promesonotal dorsum, sometimes dorsal surface with smooth notch on its centre, additional indistinct, sparse, and vertical rugulae cover promesonotal dorsum and sometimes also its lateral sides; anepisternum and katepisternum rugo-foveolate, katepisternum with smooth notch on its central part; propodeum rugo-foveolate, its dorsoventral surface and lower part of lateral sides with fading sculpture. Pilosity moderately dense, long, erect (Fig. 41D, F). Petiole. Shiny; peduncle moderately long, foveolate, with indistinct horizontal lobes on its basal part; node relatively high, moderately thick, with rounded apex, in rear view node with shallow dorsoventral concavity or straight; pilosity moderately sparse and erect (Fig. 41D, F). Postpetiole. Shiny and foveolate, its dorsum with fading sculpture; in dorsal view sides with moderately long, wide, acute, and triangular projections; pilosity long and erect (Fig. 41D, F). Petiole. Shiny and finely shagreened; pilosity moderately dense, long, and erect (Fig. 41D, F). Colour. Brown to brownish black; legs yellowish brown (Fig. 41D, F).

Figure 41. 

Pheidole annemariae Forel, full-face view (A), profile (C), and dorsal view (E) of minor worker (CASENT0077287) and full-face view (B), profile (D), and dorsal view (F) of major worker (CASENT0077290).

Minor workers. Measurements (N = 10): HL: 0.52–0.57 (0.54); HW: 0.48–0.54 (0.5); SL: 0.47–0.51 (0.48); EL: 0.11–0.12 (0.11); WL: 0.62–0.67 (0.64); PSL: 0.15–0.18 (0.16); MTL: 0.36–0.41 (0.39); PNW: 0.32–0.35 (0.34); PTW: 0.06–0.08 (0.07); PPW: 0.12–0.16 (0.13); CI: 89.3–94.6 (92.4); SI: 92.1–102.3 (97.6); PSLI: 29.3–31.9 (30.7); PPI: 50.0–60.5 (55.8); PNI: 65.1–70.4 (67.5); MTI: 73.5–80.3 (77.8). Head. Occipital margin straight or indistinctly concave; occipital carina indistinct (Fig. 41A). Pilosity moderately dense, long, suberect. Head foveolate; frons sometimes with fading sculpture; genae smooth. Clypeus finely foveolate with additional short, longitudinal rugulae; 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. 41A, C). Mesosoma. In lateral view, promesonotum low, slightly convex, short, with posterior declivity relatively steep; mesonotal spines present, small and triangular, promesonotal groove absent; metanotal groove absent; propodeal spines long, triangular, with acute apex and narrow base (Fig. 41C). Sculpture foveolate; katepisternum and mesonotum with smooth notches. Pilosity sparse, short, and erect (Fig. 41C, E). Petiole. Peduncle relatively short and thin; with few short, erect setae (Fig. 41C, E). Postpetiole. Short, low and convex; with few short, erect setae (Fig. 41C, E). Petiole. With sparse, erect pilosity (Fig. 41C, E). Colour. Yellow (Fig. 41C, E).

Biology

The species was collected between 3–1343 m in elevation, in rainforest, littoral rainforest, montane rainforest, and occasionally beach vegetation on sandy soil. Nests were located in rotten logs and branches, and once in canopy.

Comments

Pheidole annemariae is most similar to P. marieannae sp. nov. Majors workers. It differs from P. marieannae sp. nov. in presence of smooth notch on genae, inner and outer teeth closely spaced and connected by indistinct concavity, posterior mesonotum with distinct teeth-like projections, and longer propodeal spines. Minor workers. It differs from P. marieannae sp. nov. in presence of small but distinct mesonotal spines and katepisternum, anepisternum, and mesonotum never entirely smooth.

Pheidole marieannae sp. nov.

Figs 42A–F, 85H, 87L

Type material

Holotype. Madagascar. •1 major worker; Antsiranana; Sava Region: Parc National de Marojejy, near Manantenina River; -14.43677, 49.77541; alt. 475 m; 5 Feb 2018; Fisher et al. leg.; BLF40635, CASENT0923219 (CASC). Paratypes. Madagascar. •2w., 1q.; same data as for holotype; CASENT0808177, CASENT0808178, CASENT0872220 (CASC).

Other material

Madagascar. –Antsiranana: •6w., 6s.; Ambondrobe, 41.1km 175° Vohemar; -13.71533, 50.10167; alt. 10 m; 29 Nov 2004; B.L. Fisher et al. leg.; CASENT0056064, CASENT0056072, CASENT0056523, CASENT0056661, CASENT0056664, CASENT0109696, CASENT0109743, CASENT0110541, CASENT0110669 (CASC). •10w., 5s., 1m.; Forêt Ambanitaza, 26.1 km 347° Antalaha; -14.67933, 50.18367; alt. 240 m; 26 Nov 2004; B.L. Fisher et al. leg.; CASENT0054895, CASENT0054958, CASENT0054962, CASENT0055564, CASENT0057274, CASENT0059041, CASENT0109525, CASENT0109533, CASENT0109546, CASENT0109546, CASENT0109547 (CASC). •5w., 2q.; Makirovana Forest; -14.104, 50.03574; alt. 225 m; 4 May 2011; B.L. Fisher et al. leg.; CASENT0212371, CASENT0212374, CASENT0212508, CASENT0230797 (CASC). •4w.; Makirovana Forest; -14.17066, 49.95409; alt. 415 m; 29 Apr 2011; B.L. Fisher et al. leg.; CASENT0231309, CASENT0243330, CASENT0243346 (CASC). •9w.; Makirovana Forest; -14.16044, 49.95216; alt. 550 m; 1 May 2011; B.L. Fisher et al. leg.; CASENT0212460, CASENT0230823, CASENT0231428, CASENT0231431, CASENT0243617, CASENT0243658, CASENT0245034 (CASC). •7w., 6s.; Parc National de Marojejy, Manantenina River, 28.0 km 38°NE Andapa, 8.2 km 333°NNW Manantenina; -14.43667, 49.775; alt. 450 m; 12 Nov 2003; B.L. Fisher et al. leg.; CASENT0045856, CASENT0045977, CASENT0045997, CASENT0046003, CASENT0046078, CASENT0046082, CASENT0048630, CASENT0077344, CASENT0077345 (CASC). •6w., 1s., 1q., 1m.; Sava Region: Parc National de Marojejy, near Manantenina River; -14.43677, 49.77541; alt. 475 m; 5 Feb 2018; B.L. Fisher et al. leg.; CASENT0808180, CASENT0808236, CASENT0808251, CASENT0808252, CASENT0825304, CASENT0825339, CASENT0826675 (CASC).

Diagnosis

Head in full-face view rectangular, slightly longer than wide, anterior of eyes slightly convex, posterior of eyes convex; sides of the head with moderately dense, long, erect pilosity; antennal scrobes present, slightly impressed, and not delimited; scrobe surface foveolate, with distinct, thin, moderately dense, longitudinal rugae; propodeal spines moderately long (PSL: 0.16–0.21 (0.19)); gaster finely shagreened; inner hypostomal teeth distinct, moderately high, closely spaced, triangular, with rounded apex directed outward and moderately wide base; outer hypostomal teeth smaller and narrower than inner hypostomal teeth, triangular with moderately narrow base. Minor workers. Head foveolate, genae smooth; propodeal spines long (PSL: 0.11–0.19 (0.13)); promesonotum low, slightly convex, short, with posterior declivity steep; mesonotal spines present but indistinct; mesosoma foveolate, katepisternum, anepisternum, and mesonotum smooth.

Description

Major workers. Measurements (N = 10): HL: 1.03–1.14 (1.07); HW: 0.91–1.02 (0.96); SL: 0.41–0.5 (0.44); EL: 0.11–0.13 (0.12); WL: 0.72–0.91 (0.78); PSL: 0.16–0.21 (0.19); MTL: 0.42–0.54 (0.45); PNW: 0.48–0.59 (0.53); PTW: 0.12–0.16 (0.14); PPW: 0.36–0.46 (0.41); CI: 86.2–92.6 (89.2); SI: 42.8–49.7 (46.1); PSLI: 15.1–19.4 (17.3); PPI: 31.5–36.7 (34.7); PNI: 51.2–60.2 (55.7); MTI: 43.9–53.3 (47.2). Head. In full-face view rectangular, slightly longer than wide, anterior of eyes slightly convex, posterior of eyes convex, occipital margins of lobes convex (Fig. 42B). 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 visible. In lateral view weakly impressed on vertex. Sides of the head with moderately dense, long, erect pilosity; whole head with moderately dense, short, suberect to erect pilosity. Antennal scrobes shallow and not delimited by carinulae; scrobe surface foveolate, with distinct, thin, moderately dense, longitudinal rugae. Occipital lobes shiny, with thick, sparse, irregular rugae, interspaces with fine rugulae fading posteriorly; frons with moderately dense, thick, longitudinal, and sometimes interrupted rugae, interspaces finely rugulose, most often rugulae fading on the central part of frons; genae with dense, irregular rugulae, area behind eyes with slightly fading sculpture; malar area with thick, sparse, longitudinal rugae, interspaces rugoreticulate. Centre of clypeus shiny and smooth, lateral sides with sparse and longitudinal rugae; median notch present, moderately wide, and shallow; median longitudinal carina present; lateral longitudinal carinae present. Scape, when laid back, reaching the midlength of head; pilosity suberect to erect (Fig. 42B, D). Inner hypostomal teeth distinct, moderately high, closely spaced, triangular, with rounded apex directed outward and moderately wide base; outer hypostomal teeth smaller and narrower than inner hypostomal teeth, triangular with moderately narrow base (Fig. 85H). Mesosoma. In lateral view, promesonotum relatively low and arched, dorsal mesonotum finely concave, posterior mesonotum relatively steep, with tubercle-like projections; promesonotal groove absent; metanotal groove absent; propodeal spines long, triangular, with sharp apex and moderately wide base; humeral area laterally weakly produced (Fig. 42D). Surface variable; promesonotum finely rugo-foveolate, rugo-foveolae always fading on lateral sides of pronotum and promesonotal dorsum; katepisternum and mesonotum smooth; anepisternum rugulae; propodeum rugoreticulate, its dorsoventral surface with fading sculpture. Pilosity moderately sparse, long, erect (Fig. 42D, F). Petiole. Shiny; peduncle relatively short, foveolate, with indistinct horizontal lobes on its basal part; node relatively high, moderately thick, with rounded apex, in rear view node indistinctly convex; pilosity moderately sparse and erect (Fig. 42D, F). Postpetiole. Shiny and shagreened; in dorsal view sides with moderately long, wide, acute, and triangular projections; pilosity moderately sparse and erect (Fig. 42D, F). Petiole. Shiny and finely shagreened; pilosity moderately dense, long, and erect (Fig. 42D, F). Colour. Brown to dark brown; legs yellowish brown (Fig. 42D, F).

Figure 42. 

Pheidole marieannae sp. nov., full-face view (A), profile (C), and dorsal view (E) of paratype minor worker (CASENT0808178) and full-face view (B), profile (D), and dorsal view (F) of holotype major worker (CASENT0923219).

Minor workers. Measurements (N = 10): HL: 0.41–0.49 (0.46); HW: 0.39–0.46 (0.42); SL: 0.39–0.47 (0.43); EL: 0.09–0.11 (0.1); WL: 0.47–0.58 (0.54); PSL: 0.11–0.19 (0.13); MTL: 0.28–0.37 (0.32); PNW: 0.25–0.31 (0.29); PTW: 0.06–0.07 (0.06); PPW: 0.1–0.13 (0.11); CI: 87.6–95.1 (92.2); SI: 97.1–114.1 (101.8); PSLI: 24.6–38.4 (28.3); PPI: 50.0–64.2 (56.2); PNI: 62.6–73.7 (67.6); MTI: 71.4–83.7 (76.4). Head. Occipital margin straight or indistinctly concave; occipital carina indistinct (Fig. 42A). Pilosity moderately dense, short, suberect. Head foveolate; genae smooth. Clypeus with median longitudinal carina absent; two lateral longitudinal carinae absent. Scape, when laid back, reaching the posterior head margin; pilosity suberect to erect (Fig. 42A, C). Mesosoma. In lateral view, promesonotum low, slightly convex, short, with posterior declivity steep; mesonotal spines present but indistinct; promesonotal groove absent; metanotal groove indistinct; propodeal spines long, triangular, with acute apex and narrow base (Fig. 42C). Sculpture foveolate; katepisternum, anepisternum, and mesonotum smooth. Pilosity sparse, moderately long, and erect (Fig. 42C, E). Petiole. Peduncle moderately short and thin with ventral face slightly convex; with few short, erect setae (Fig. 42C, E). Postpetiole. Short, low, and convex; with few short, erect setae (Fig. 42C, E). Petiole. With sparse and erect pilosity (Fig. 42C, E). Colour. Yellow (Fig. 42C, E).

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 sub-oval 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 one- to 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.

Key to the P. makaensis group

1 Major workers. Frons with thick, sparse, and interrupted rugae, interspaces almost entirely smooth, outer hypostomal teeth lobe-like, distinctly bigger than inner hypostomal teeth (Fig. 43A, J). Minor workers. Pilosity on the whole body dense and very long (Fig. 43F) P. makaensis sp. nov.
Major workers. Frons with thick, dense to moderately dense, longitudinal or interrupted rugae, interspaces rugo-foveolate, foveolate or with rugulae, outer hypostomal teeth never lobe-like, approximately as big as inner hypostomal teeth or slightly higher (Fig. 43B–E, K–N). Minor workers. Pilosity on the whole body moderately sparse and never very long (Fig. 43G–I) 2
2 Major workers. Rugae on frons fading posteriorly and never connected with rugae on the occipital lobes, genae smooth, inner hypostomal teeth directed inward (Figs 43B, K, 44A). Minor workers. Genae smooth, katepisternum with smooth notch (Fig. 43G) P. fitarata sp. nov.2
Major workers. Rugae on frons not fading posteriorly and connected with rugae on the occipital lobes, genae never entirely smooth, inner hypostomal teeth never directed inward (Figs 43D, E, 44C, D). Minor workers. Genae with fading foveolae or only with smooth notch, katepisternum never smooth (Fig. 44H, I) 3
3 Major workers. Body dark yellow, promesonotum high and arched, frons with interspaces never rugo-foveolae, outer hypostomal teeth dentate and directed outward (Figs 43E, N, 44D). Minor workers. Scape reaching posterior margin of head, and head with additional longitudinal rugae on frons and malar area, propodeal spines never directed upward (Figs 43I, 44G) P. ehazoara sp. nov.
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 (Figs 43D, M, 44C). Minor workers. Scape surpassing the posterior head margin by one-fifth of its length, and head without additional longitudinal rugae on frons and malae area, propodeal spines directed upward (Figs 43H, 44F) P. avaratra sp. nov.
Figure 43. 

Pheidole makaensis sp. nov., head and profile of major worker (A), hypostomal teeth (J), profile of minor worker (F). Pheidole fitarata sp. nov., head and profile of major worker (B), hypostomal teeth (K), profile of minor worker (G). Pheidole rugofitarata sp. nov., head and profile of major worker (C), hypostomal teeth (L). Pheidole avaratra sp. nov., head and profile of major worker (D), hypostomal teeth (M), profile of minor worker (H). Pheidole ehazoara sp. nov., head and profile of major worker (E), hypostomal teeth (N), profile of minor worker (I).

Figure 44. 

Pheidole fitarata sp. nov., profile of major worker (A), head of minor worker (E). Pheidole rugofitarata sp. nov., profile of major worker (B). Pheidole avaratra sp. nov., profile of major worker (C), head of minor worker (F). Pheidole ehazoara sp. nov., profile of major worker (D), head of minor worker (G).

Pheidole makaensis sp. nov.

Figs 45A–F, 85D, 87H

Type material

Holotype. Madagascar. •1 major worker; Toliara; Makay Mts.; -21.25864, 45.16412; alt. 500 m; 8 Dec 2010; Fisher et al. leg.; BLF25762, CASENT0205745 (CASC). Paratype. Madagascar. •1w.; same data as for holotype; CASENT0923188 (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.

Description

Major workers. Measurements (N = 1): HL: 1.24; HW: 1.03; SL: 0.46; EL: 0.14; WL: 0.84; PSL: 0.16; MTL: 0.54; PNW: 0.51; PTW: 0.14; PPW: 0.34; CI: 83.5; SI: 44.2; PSLI: 13.3; PPI: 40.4; PNI: 49.0; MTI: 51.9. Head. In full-face view longitudinal, longer than wide, anterior of eyes straight, posterior of eyes slightly convex, occipital margin of lobes straight, inclining towards centre (Fig. 45B). In lateral view sub-oval; ventral and dorsal faces convex; dorsal face finely depressed posteriorly, forming indistinct transverse depression between frons and occipital lobes; inner hypostomal teeth visible. Sides of the head with moderately dense, very long, erect pilosity; whole head with moderately dense, long, suberect to erect pilosity. Antennal scrobes indistinct and not delimited by carinulae; scrobe surface shiny, with thick, sparse, longitudinal rugae; interspaces smooth and finely foveolate on the posterior part. Occipital lobes shiny, with thick, sparse, short, irregular, and slightly arcuate rugae, interspaces smooth; frons with moderately sparse, thick, longitudinal, interrupted rugae, interspaces smooth and indistinctly rugulae on the posterior part; genae shiny and smooth; malar area with thick, sparse, longitudinal rugae, interspaces smooth. Centre of clypeus shiny and smooth, lateral sides with longitudinal rugae; median notch present, narrow, and moderately deep; median longitudinal carina absent; lateral longitudinal carinae absent. Scape, when laid back, not reaching the midlength of head; pilosity erect (Fig. 45B, D). 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 (Fig. 85D). Mesosoma. In lateral view, promesonotum relatively low and arched, dorsal mesonotum slightly concave, posterior mesonotum steep, with tubercle-like projections; promesonotal groove absent; metanotal groove absent; propodeal spines short, triangular, with sharp apex and relatively wide base; humeral area laterally weakly produced (Fig. 45D). Surface foveolate with additional indistinct and thin rugulae; dorsal and lateral surfaces of pronotum with fading sculpture and sometimes with smooth notches; katepisternum and anepisternum with sculpture reduced and smooth notches. Pilosity moderately dense, very long, and erect (Fig. 45D, F). Petiole. Shiny; peduncle moderately long, finely foveolate, with distinct, short, horizontal lobes on its basal part; node with fading sculpture, relatively low, triangular, with rounded apex, in rear view node slightly convex; pilosity moderately sparse and erect (Fig. 45D, F). Postpetiole. Shiny and finely shagreened, dorsum with fading sculpture; in dorsal view sides with very short, acute, and triangular projections; pilosity long, dense, and erect (Fig. 45D, F). Petiole. Shiny and finely shagreened; pilosity dense, very long, and erect (Fig. 45D, F). Colour. Reddish brown to brown; lateral sides of mesosoma, and sometimes malar area and lower frons, yellowish brown; legs yellow to yellowish brown (Fig. 45D, F).

Figure 45. 

Pheidole makaensis sp. nov., full-face view (A), profile (C), and dorsal view (E) of paratype minor worker (CASENT0923188) and full-face view (B), profile (D), and dorsal view (F) of holotype major worker (CASENT0205745).

Minor workers. Measurements (N = 1): HL: 0.48; HW: 0.41; SL: 0.44; EL: 0.1; WL: 0.55; PSL: 0.07; MTL: 0.35; PNW: 0.28; PTW: 0.07; PPW: 0.11; CI: 86.8; SI: 107.0; PSLI: 15.5; PPI: 64.6; PNI: 67.8; MTI: 84.3. Head. Occipital margin straight or indistinctly concave; occipital carina absent (Fig. 45A). Pilosity dense, very long, erect. 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. 45A, C). Mesosoma. In lateral view, promesonotum low, convex, short, with posterior declivity smoothly declining towards propodeum; promesonotal groove absent; metanotal groove indistinct; propodeal spines small, triangular, with acute apex (Fig. 45C). Sculpture foveolate; anepisternum, katepisternum, and mesonotum with fading sculpture. Pilosity moderately sparse, long, and erect (Fig. 45C, E). Petiole. Peduncle short and thin with ventral face slightly convex; node moderately low, globular, and small; with few short, erect setae (Fig. 45C, E). Postpetiole. Short, low, and slightly convex; with few short, erect setae (Fig. 45C, E). Petiole. With moderately sparse, very long, erect pilosity (Fig. 45C, E). Colour. Yellow (Fig. 45C, E).

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.

Pheidole fitarata sp. nov.

Figs 46A–F, 84S, 86S

Type material

Holotype. Madagascar. •1 major worker; Antsiranana; Ampasindava, Forêt d’Ambilanivy, 3.9 km 181°S Ambaliha; -13.79861, 48.16167; alt. 600 m; 4 Mar 2001; Fisher et al. leg.; BLF03293, CASENT0420028 (CASC). Paratypes. Madagascar. •4w., 1s.; same data as for holotype; CASENT0420024CASENT0420027, CASENT0872215CASENT0872217 (CASC).

Other material

Madagascar. –Antsiranana: •1s.; Ambondrobe, 41.1km 175° Vohemar; -13.71533, 50.10167; alt. 10 m; 29 Nov 2004; B.L. Fisher leg.; CASENT0056078 (CASC). •1w., 4s.; Ampasindava, Forêt d’Ambilanivy, 3.9 km 181°S Ambaliha; -13.79861, 48.16167; alt. 600 m; 4 Mar 2001; B.L. Fisher et al. leg.; CASENT0406700, CASENT0420217, CASENT0420226, CASENT0464109, CASENT0464294 (CASC). •2s.; Forêt d’ Andavakoera, 21.4 km 75°ENE Ambilobe; 4.6 km 356°N Betsiaka; -13.11833, 49.23; alt. 425 m; 15 Dec 2003; B.L. Fisher leg.; CASENT0044135, CASENT0044209 (CASC). •1s.; Montagne des Français, 7.2 km 142°SE Antsiranana (=Diego Suarez); -12.32278, 49.33817; alt. 180 m; 22 Feb 2001; B.L. Fisher et al. leg.; CASENT0461044 (CASC). •1s.; Nosy Be, Réserve Naturelle Intégrale de Lokobe, 6.3 km 112°ESE Hellville; -13.41933, 48.33117; alt. 30 m; 19 Mar 2001; B.L. Fisher et al. leg.; CASENT0462968 (CASC). •1s.; R.S. Manongarivo, 12.8 km 228°SW Antanambao; -13.97667, 48.42333; alt. 780 m; 11 Oct 1998; B.L. Fisher leg.; CASENT0198865 (CASC). •3s.; Réserve Spéciale d’Ambre, 3.5 km 235°SW Sakaramy; -12.46889, 49.24217; alt. 325 m; 26 Jan 2001; B.L. Fisher et al. leg.; CASENT0406576, CASENT0423829, CASENT0423892 (CASC). •1w., 1s.; Réserve Spéciale de l’Ankarana, 13.6 km 192°SSW Anivorano Nord; -12.86361, 49.22583; alt. 210 m; 16 Feb 2001; B.L. Fisher et al. leg.; CASENT0420164, CASENT0420181 (CASC). •7s., 1q.; Réserve Spéciale de l’Ankarana, 22.9 km 224°SW Anivorano Nord; -12.90889, 49.10983; alt. 80 m; 10 Feb 2001; B.L. Fisher et al. leg.; CASENT0420207, CASENT0439168, CASENT0439173, CASENT0439183, CASENT0439197, CASENT0439313, CASENT0439315, CASENT0008034 (CASC).

Diagnosis

Major workers. Head in full-face view elongated; sides of the head with moderately dense, long, erect pilosity; occipital lobes smooth to indistinctly foveolate, with thick, sparse, short, transverse, and sometimes arcuate rugae; 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. 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 relatively convex; mesosoma foveolate, katepisternum with smooth notch; propodeal spines short, triangular.

Description

Major workers. Measurements (N = 10): HL: 1.13–1.43 (1.29); HW: 0.91–1.12 (1.07); SL: 0.43–0.53 (0.49); EL: 0.12–0.17 (0.14); WL: 0.79–1.02 (0.92); PSL: 0.16–0.21 (0.18); MTL: 0.51–0.59 (0.55); PNW: 0.45–0.64 (0.56); PTW: 0.11–0.19 (0.16); PPW: 0.29–0.47 (0.41); CI: 77.9–88.4 (82.7); SI: 43.5–49.0 (46.5); PSLI: 12.7–15.3 (14.0); PPI: 34.8–42.9 (38.2); PNI: 49.1–56.7 (52.8); MTI: 48.4 –56.1 (52.0). Head. In full-face view longitudinal, longer than wide, anterior of eyes straight, posterior of eyes slightly convex, occipital margin of lobes straight, inclining toward centre (Fig. 46B). In lateral view sub-rectangular; ventral and dorsal faces finely convex; dorsal face finely depressed posteriorly, forming 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).

Figure 46. 

Pheidole fitarata sp. nov., full-face view (A), profile (C), and dorsal view (E) of paratype minor worker (CASENT0420025) and full-face view (B), profile (D), and dorsal view (F) of holotype major worker (CASENT0420028).

Minor workers. Measurements (N = 6): HL: 0.45–0.47 (0.46); HW: 0.39–0.41 (0.4); SL: 0.44–0.46 (0.45); EL: 0.07–0.1 (0.08); WL: 0.52–0.56 (0.55); PSL: 0.08–0.09 (0.08); MTL: 0.33–0.36 (0.35); PNW: 0.26–0.28 (0.27); PTW: 0.06–0.06 (0.06); PPW: 0.09–0.1 (0.1); CI: 85.9–89.5 (87.7); SI: 111.0–113.6 (111.6); PSLI: 17.3–20.4 (18.5); PPI: 61.0–65.2 (62.5); PNI: 66.4–68.8 (67.8); MTI: 82.8–89.4 (86.7). Head. Occipital margin straight or indistinctly concave; occipital carina absent (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). Mesosoma. In lateral view, promesonotum low, convex, short, with posterior declivity relatively convex; promesonotal groove absent; metanotal groove indistinct; propodeal spines short, triangular, with acute apex (Fig. 46C). Sculpture foveolate; katepisternum with smooth notch. Pilosity sparse, long, and erect (Fig. 46C, E). Petiole. Peduncle short and thin with ventral face slightly convex; with few short, erect setae (Fig. 46C, E). Postpetiole. Short, low, and convex; sometimes with few short, erect setae (Fig. 46C, E). Petiole. With moderately sparse, erect pilosity (Fig. 46C, E). Colour. Yellow (Fig. 46C, E).

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.

Pheidole rugofitarata sp. nov.

Figs 47A–C, 85V, 88F

Type material

Holotype. Madagascar. •1 major worker; Mahajanga; Parc National Tsingy de Bemaraha, 2.5 km 62°ENE Bekopaka, Ankidrodroa River; -19.13222, 44.81467; alt. 100 m; 11 Nov 2001; Fisher et al. leg.; BLF04341, CASENT0078206 (CASC). Paratype. Madagascar. •1s.; same data as for holotype; CASENT0078208 (CASC).

Other material

Madagascar. – Mahajanga: •2s.; Forêt Ambohimanga, 26.1 km 314° Mampikony; -15.96267, 47.43817; alt. 250 m; 13 Dec 2004; Fisher et al. leg.; CASENT0054800, CASENT0055004 (CASC). •2s.; Forêt de Tsimembo, 11.0 km 346°NNW Soatana; -18.99528, 44.4435; alt. 50 m; 21 Nov 2001; Fisher et al. leg.; CASENT0483554, CASENT0483640 (CASC). •1s.; Parc National d’Ankarafantsika, Ampijoroa Station Forestière, 5.4 km 331°NW Andranofasika; -16.29889, 46.813; alt. 70 m; 26 Mar 2001; Fisher et al. leg.; CASENT0468979 (CASC). •3s.; Parc National d’Ankarafantsika, Forêt de Tsimaloto, 18.3 km 46°NE de Tsaramandroso; -16.22806, 47.14361; alt. 135 m; 2 Apr 2001; Fisher et al. leg.; CASENT0431979, CASENT0432158, CASENT0432186 (CASC). •1s.; Parc National de Baie de Baly, 12.4 km 337°NNW Soalala; -16.01, 45.265; alt. 10 m; 26 Nov 2002; Fisher et al. leg.; CASENT0025410 (CASC). •5s.; Parc National de Namoroka, 16.9 km 317°NW Vilanandro; -16.40667, 45.31; alt. 100 m; 12 Nov 2002; Fisher et al. leg.; CASENT0038862, CASENT0023652, CASENT0023665, CASENT0023667, CASENT0493106 (CASC). •2s.; Parc National de Namoroka, 9.8 km 300°WNW Vilanandro; -16.46667, 45.35; alt. 140 m; 4 Nov 2002; Fisher et al. leg.; CASENT0032768, CASENT0492055 (CASC). •4s.; Parc National Tsingy de Bemaraha, 3.4 km 93°E Bekopaka, Tombeau Vazimba; -19.14194, 44.828; alt. 50 m; 6 Nov 2001; Fisher et al. leg.; CASENT0078127, CASENT0477444, CASENT0477453, CASENT0477482 (CASC). •4s.; Réserve d’Ankoririka, 10.6 km 13°NE de Tsaramandroso; -16.26722, 47.04861; alt. 210 m; 9 Apr 2001; Fisher et al. leg.; CASENT0471030, CASENT0471209, CASENT0484762 (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.

Description

Major workers. Measurements (N = 10): HL: 1.29–1.41 (1.36); HW: 1.05–1.17 (1.12); SL: 0.48–0.53 (0.5); EL: 0.14–0.17 (0.15); WL: 0.95–1.02 (0.99); PSL: 0.17–0.21 (0.18); MTL: 0.54–0.57 (0.55); PNW: 0.58–0.68 (0.62); PTW: 0.14–0.17 (0.16); PPW: 0.38–0.47 (0.43); CI: 81.2–85.8 (82.9); SI: 41.8–49.0 (44.3); PSLI: 12.2–15.2 (13.5); PPI: 34.0–42.1 (37.3); PNI: 53.1–59.0 (55.3); MTI: 46.9 –52.5 (49.1). Head. In full-face view longitudinal, longer than wide, anterior of eyes straight, posterior of eyes slightly convex, occipital margin of lobes straight, inclining toward centre (Fig. 47C). 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, long, suberect to erect pilosity. Antennal scrobes indistinct and not delimited by carinulae; scrobe surface shiny, with moderately dense, longitudinal rugae, interspaces superficially foveolate. Occipital lobes shiny, indistinctly foveolate, with thick, sparse, short, transverse and sometimes arcuate rugae; frons with moderately dense, thick, longitudinal rugae, interspaces smooth and indistinctly foveolate on the posterior part; genae shiny, with dense, thin, longitudinal rugulae and smooth notch in the centre; 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 midlength of head; pilosity suberect to erect (Fig. 47B, C). Inner hypostomal teeth distinct, high, closely spaced, triangular, with rounded apex; outer hypostomal teeth wider and higher than inner hypostomal teeth, lobe-like (Fig. 85V). 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. 47B). Surface finely foveolate with additional irregular, thin, and sparse rugoreticulation; dorsal surface of promesonotum with fading foveolae and thin, sparse, and transverse rugulae; katepisternum with reduced sculpture. Pilosity moderately dense, very long and erect (Fig. 47A, B). 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 long and erect (Fig. 47A, B). Postpetiole. Shiny and finely shagreened, dorsum with fading sculpture; in dorsal view sides with moderately short, acute, and triangular projections; pilosity long and erect (Fig. 47A, B). Petiole. Shiny and finely shagreened; pilosity dense, long, and erect (Fig. 47A, B). Colour. Reddish brown to brown, lateral sides of mesosoma and legs yellowish brown (Fig. 47A, B).

Figure 47. 

Pheidole rugofitarata sp. nov., dorsal view (A), profile (B), and full-face view (C) of holotype major worker (CASENT0078206).

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.

Comments

Major workers. Pheidole rugofitarata sp. nov. is most similar to P. fitarata sp. nov. and differs from it in presence of distinct and never-interrupted longitudinal rugae on frons and antennal scrobes, rugae on frons connected with rugae on occipital lobes, never entirely smooth genae, and lobe-like outer hypostomal teeth.

Pheidole avaratra sp. nov.

Figs 48A–F, 84F, 86F

Type material

Holotype. Madagascar. •1 major worker; Antsiranana; Forêt Ambato, 26.6 km 33° Ambanja; -13.4645, 48.55167; alt. 150 m; 10 Dec 2004; Fisher et al. leg.; BLF11593, CASENT0107719 (CASC). Paratype. Madagascar. •1w.; same data as for holotype; CASENT0923184 (CASC).

Other material

Madagascar. – Antsiranana: •1w., 4s.; Forêt Ambato, 26.6 km 33° Ambanja; -13.4645, 48.55167; alt. 150 m; 8 Dec 2004; Fisher et al. leg.; CASENT0056638, CASENT0056639, CASENT0056704, CASENT0056727, CASENT0107724 (CASC). •1w., 12s.; Nosy Be, Réserve Naturelle Intégrale de Lokobe, 6.3 km 112°ESE Hellville; -13.41933, 48.33117; alt. 30 m; 19 Mar 2001; Fisher et al. leg.; CASENT0403256, CASENT0403258, CASENT0462862, CASENT0463005, CASENT0463136, CASENT0466195, CASENT0466206, CASENT0466207, CASENT0466209, CASENT0466222, CASENT0466243, CASENT0466247, CASENT0466295 (CASC). •2w., 2s.; Nosy Faly; -13.3624, 48.49101; alt. 15 m; 26 Feb 2013; Fisher et al. leg.; CASENT0303115, CASENT0303182 (CASC). •1w., 1s.; Réserve Spéciale d’Ambre, 3.5 km 235°SW Sakaramy; -12.46889, 49.24217; alt. 325 m; 26 Jan 2001; Fisher et al. leg.; CASENT0406637, CASENT0406688 (CASC). •1w., 2s.; Réserve Spéciale de l’Ankarana, 13.6 km 192°SSW Anivorano Nord; -12.86361, 49.22583; alt. 210 m; 16 Feb 2001; Fisher et al. leg.; CASENT0411220, CASENT0440802, CASENT0440818 (CASC).

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.

Description

Major workers. Measurements (N = 10): HL: 0.92–1.2 (1.0); HW: 0.71–1.02 (0.8); SL: 0.368–0.502 (0.4); EL: 0.11–0.16 (0.14); WL: 0.71–0.91 (0.99); PSL: 0.12–0.18 (0.13); MTL: 0.41–0.54 (0.44); PNW: 0.4–0.56 (0.45); PTW: 0.1–0.15 (0.12); PPW: 0.25–0.38 (0.3); CI: 78.0–87.9 (82.9); SI: 46.5–51.5 (49.2); PSLI: 11.8–15.2 (13.3); PPI: 35.0–46.2 (38.9); PNI: 51.2–58.9 (55.5); MTI: 52.2 –57.3 (53.8). Head. In full-face view longitudinal, longer than wide, anterior of eyes straight, posterior of eyes slightly convex, occipital margin of lobes straight, inclining toward centre (Fig. 48B). In lateral view sub-oval; ventral and dorsal faces convex; dorsal face not depressed posteriorly; inner hypostomal teeth visible. Sides of the head with moderately dense, long, erect pilosity; whole head with dense, moderately long, suberect to erect pilosity. Antennal scrobes indistinct and not delimited by carinulae; scrobe surface shiny, with thick, moderately sparse, longitudinal rugae; interspaces superficially rugo-foveolae. Occipital lobes shiny, with thick, sparse, short, irregular, and arcuate rugae, interspaces superficially rugulose; frons with moderately sparse, thick, longitudinal rugae, interspaces smooth and indistinctly rugo-foveolate on the posterior part; genae shiny, with dense and fine rugulae, and smooth notch behind eyes; malar area with thick, sparse, longitudinal rugae, interspaces superficially rugulae. Centre of clypeus shiny and smooth, lateral sides with longitudinal rugae; median notch present, narrow, and moderately deep; median longitudinal carina absent; lateral longitudinal carinae present. Scape, when laid back, not reaching the midlength of head; pilosity suberect to erect (Fig. 48B, D). 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 (Fig. 84F). Mesosoma. In lateral view, promesonotum relatively low and arched, dorsal mesonotum slightly concave, posterior mesonotum steep, with tubercle-like projections; promesonotal groove absent; metanotal groove absent; propodeal spines small to moderately long, triangular, with sharp apex and relatively wide base; humeral area laterally weakly produced (Fig. 48D). Surface foveolate, dorsal surface of promesonotum with fading sculpture and sometimes with additional transverse, thin rugulae. Pilosity moderately sparse, very long, and erect (Fig. 48D, F). Petiole. Shiny; peduncle moderately long, finely foveolate, with indistinct horizontal lobes on its basal part; node with fading sculpture, relatively high, triangular, with rounded apex, in rear view node slightly convex; pilosity moderately sparse and erect (Fig. 48D, F). Postpetiole. Shiny and finely shagreened, dorsum with fading sculpture; in dorsal view sides with very short, acute, and triangular projections; pilosity moderately long, sparse, and erect (Fig. 48D, F). Petiole. Shiny and shagreened; pilosity dense, moderately long, and erect (Fig. 48D, F). Colour. Brown to dark brown; lateral sides of mesosoma, malar area, and lower frons yellowish brown; legs yellow to yellowish brown (Fig. 48D, F).

Figure 48. 

Pheidole avaratra sp. nov., full-face view (A), profile (C), and dorsal view (E) of paratype minor worker (CASENT0923184) and full-face view (B), profile (D), and dorsal view (F) of holotype major worker (CASENT0107719).

Minor workers. Measurements (N = 5): HL: 0.43–0.48 (0.46); HW: 0.39–0.44 (0.42); SL: 0.38–0.47 (0.44); EL: 0.09–0.11 (0.1); WL: 0.5–0.58 (0.55); PSL: 0.08–0.1 (0.93); MTL: 0.29–0.37 (0.35); PNW: 0.27–0.31 (0.29); PTW: 0.05–0.08 (0.07); PPW: 0.1–0.11 (0.11); CI: 87.6–94.8 (90.4); SI: 96.4–110.8 (105.5); PSLI: 17.3–22.0 (20.0); PPI: 46.4–70.3 (61.9); PNI: 67.4–72.8 (69.9); MTI: 74.6–87.6 (83.7). Head. Occipital margin straight or indistinctly concave; occipital carina absent (Fig. 48A). Pilosity moderately dense, short, suberect. Head shiny, foveolate; genae with smooth notch. Clypeus smooth or finely foveolate; 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. 48A, C). Mesosoma. In lateral view, promesonotum low, convex, short, with posterior declivity smoothly declining towards propodeum; promesonotal groove absent; metanotal groove indistinct; propodeal spines short, triangular, with acute apex (Fig. 48C). Sculpture foveolate. Pilosity sparse, long, and erect (Fig. 48C, E). Petiole. Peduncle moderately short and thin with ventral face slightly convex; with few short, erect setae (Fig. 48C, E). Postpetiole. Short, low, and convex; with few short, erect setae (Fig. 48C, E). Petiole. With moderately sparse, erect pilosity (Fig. 48C, E). Colour. Yellow (Fig. 48C, E).

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 avaratra sp. nov. differs from P. fitarata sp. nov. in shorter and higher promesonotum and katepisternum entirely foveolate.

Pheidole ehazoara sp. nov.

Figs 49A–F, 84N, 86N

Type material

Holotype. Madagascar. •1 major worker; Toliara; Ehazoara Canyon, 26 km E Betioky; -23.68333, 44.63333; alt. 175 m; 27 Apr 1997; Fisher et al. leg.; BLF01518, CASENT0872059, middle specimen (CASC). Paratype. Madagascar. •2w., 2s.; same data as for holotype; CASENT0198555, CASENT0923186, CASENT0872218, CASENT0872219 (CASC).

Diagnosis

Major workers. Head in full-face view elongated; sides of the head with dense, very long, erect pilosity; occipital lobes with thick, sparse, short, irregular, and arcuate rugae, interspaces indistinctly rugulose; inner hypostomal teeth distinct, narrow, and triangular, closely spaced, with narrow and rounded apex directed outward; outer hypostomal teeth bigger and wider than inner hypostomal teeth, dentate. Minor workers. Head foveolate, foveolate, with additional longitudinal rugae on frons and malar area; genae with fading sculpture; scape, when laid back, reaching head margin; promesonotum low, convex, short, with posterior declivity smoothly declining towards propodeum; mesosoma foveolate; propodeal spines short, triangular.

Description

Major workers. Measurements (N = 3): HL: 1.07–1.1 (1.08); HW: 0.85–0.88 (0.87); SL: 0.41–0.44 (0.42); EL: 0.14–0.15 (0.14); WL: 0.82–0.86 (0.84); PSL: 0.13–0.14 (0.14); MTL: 0.44–0.46 (0.45); PNW: 0.49–0.51 (0.5); PTW: 0.15–0.16 (0.16); PPW: 0.38–0.42 (0.4); CI: 81.5–83.6 (82.3); SI: 41.8–49.0 (45.9); PSLI: 13.5–15.0 (14.09); PPI: 34.8–37.3 (36.2); PNI: 54.9–55.7 (55.4); MTI: 47.2–52.5 (50.4). Head. In full-face view longitudinal, longer than wide, anterior of eyes straight, posterior of eyes slightly convex, occipital margin of lobes straight, inclining towards centre (Fig. 49B). In lateral view sub-oval; ventral and dorsal faces convex; dorsal face finely depressed posteriorly, forming indistinct transverse depression between frons and occipital lobes; inner hypostomal teeth visible. Sides of the head with dense, very long, erect pilosity; whole head with dense, moderately long, suberect to erect pilosity. Antennal scrobes indistinct and not delimited by carinulae; scrobe surface shiny, with thick, moderately sparse, longitudinal rugae; interspaces rugulose. Occipital lobes shiny, with thick, sparse, short, irregular, and arcuate rugae, interspaces indistinctly rugulose; frons with moderately sparse, thick, longitudinal rugae, on the anterior part of frons interspaces with fine, short, and longitudinal rugulae, on the posterior part of frons interspaces with fine and irregular rugulae; genae shiny, with fine, dense and longitudinal rugulae, and behind the eyes smooth; malar area with thick, sparse, longitudinal rugae, interspaces smooth to indistinctly rugulose. Centre of clypeus shiny and smooth, lateral sides with longitudinal rugae; median notch indistinct; median longitudinal carina absent; lateral longitudinal carinae absent. Scape, when laid back, not reaching the midlength of head; pilosity erect (Fig. 49B, D). Inner hypostomal teeth distinct, narrow, and triangular, closely spaced, with narrow and rounded apex directed outward; outer hypostomal teeth bigger and wider than inner hypostomal teeth, dentate (Fig. 84N). Mesosoma. In lateral view, promesonotum relatively low and arched, posterior mesonotum relatively convex, with small, tubercle-like projections; promesonotal groove absent; metanotal groove indistinct; propodeal spines short, triangular, with sharp apex and very wide base; humeral area laterally weakly produced (Fig. 49D). Surface foveolate; promesonotum with additional indistinct and thin, transverse rugulae; its dorsal surface with fading sculpture; propodeum with additional longitudinal rugae, its dorsoventral side with fading sculpture. Pilosity moderately dense, long, suberect to erect (Fig. 49D, F). Petiole. Shiny; peduncle moderately short, finely foveolate, with distinct, short, horizontal lobes on its basal part; node with fading sculpture, relatively high, triangular, with rounded apex, in rear view node slightly convex; pilosity long and erect (Fig. 49D, F). Postpetiole. Shiny and finely shagreened, dorsum with fading sculpture; in dorsal view sides with moderately short, very wide, acute, and triangular projections; pilosity long and erect (Fig. 49D, F). Petiole. Shiny and finely shagreened on the basal part of first gastral tergite; pilosity dense, long, and erect (Fig. 49D, F). Colour. Dark yellow; lower part of malar area brown; gaster yellowish brown to brown (Fig. 49D, F).

Figure 49. 

Pheidole ehazoara sp. nov., full-face view (A), profile (C), and dorsal view (E) of paratype minor worker (CASENT0923186) and full-face view (B), profile (D), and dorsal view (F) of holotype major worker (CASENT0872059).

Minor workers. Measurements (N = 2): HL: 0.52–0.51; HW: 0.47–0.46; SL: 0.42–0.4; EL: 0.12–0.1; WL: 0.6–0.59; PSL: 0.1–0.09; MTL: 0.37–0.34; PNW: 0.33–0.32; PTW: 0.09–0.09; PPW: 0.17–0.15; CI: 83.6–81.5; SI: 41.8–49.0; PSLI: 15.0–13.9; PPI: 36.3–34.8; PNI: 54.9–55.7; MTI: 47.2–52.5. Head. Occipital margin straight or indistinctly concave; occipital carina indistinct (Fig. 49A). Pilosity moderately dense, short, suberect. Head shiny, foveolate, with additional longitudinal rugae on frons and malar area; genae with fading sculpture. Clypeus with fine, short, and longitudinal rugulae; median longitudinal carina absent; two lateral longitudinal carinae absent. Scape, when laid back, reaching head margin; pilosity suberect to erect (Fig. 49A, C). Mesosoma. In lateral view, promesonotum low, convex, short, with posterior declivity smoothly declining towards propodeum; promesonotal groove absent; metanotal groove present; propodeal spines small, triangular, with acute apex (Fig. 49C). Sculpture foveolate. Pilosity moderately sparse, moderately short, and erect (Fig. 49C, E). Petiole. Shiny; peduncle finely foveolate; short and thin; node moderately low, globular, and small; with few short, erect setae (Fig. 49C, E). Postpetiole. Moderately long, low, and slightly convex; with few short, erect setae (Fig. 49C, E). Petiole. With moderately sparse, erect pilosity (Fig. 49C, E). Colour. Dark yellow (Fig. 49C, E).

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.

Key to the Pheidole curvistriata group

1 Major workers. Frons with sparse, thick, longitudinal rugae with smooth to indistinctly rugulose interspaces; mesosoma with rugulae and sometimes indistinct foveolae, outer hypostomal teeth higher than inner hypostomal teeth but very narrow (Fig. 50A, I, E). Minor workers. Rugae on upper part of frons distinct and never curved outward, mesonotum never smooth (Fig. 51A, E, I) P. curvistriata sp. nov.
Major workers. Frons with longitudinal rugae denser, with interspaces smooth to distinctly rugo-foveolate or foveolate; mesosoma foveolate and sometimes with indistinct rugulae, outer hypostomal teeth higher than inner hypostomal teeth but wide (Fig. 50B–D, F–H, J–L). Minor workers. Rugae on upper part of frons indistinct or distinct and curved outward, mesonotum smooth (Fig. 51B–D, F–H, J–L) 2
2 Major workers. Frons and mesosoma with interspaces finely rugo-foveolate (Fig. 50C, K). Minor workers. Rugae on upper part of frons curved outward, mesosoma always with thick rugae, propodeal spines longer (Fig. 51C, G, K) P. makirovana sp. nov.
Major workers. Frons and mesosoma with fine foveolae (Fig. 50B, D, J, L). Minor workers. Rugae on upper part of frons never curved outward, propodeal spines shorter, mesosoma never with thick rugae (Fig. 51B, D, F, H, J, L) 3
3 Major workers. Sides of head with sparse and short pilosity, inner hypostomal teeth very low (Fig. 50D, H). Minor workers. Clypeus foveolate to rugoreticulate, rugae on head very indistinct (Fig. 51D) P. mantadia sp. nov.
Major workers. Sides of head with dense and long pilosity, inner hypostomal teeth higher (Fig. 50B, F). Minor workers. Clypeus smooth, rugae on head distinct but thin (Fig. 51B) P. moramanaensis sp. nov.
Figure 50. 

Major workers. Pheidole curvistriata sp. nov., head (A), profile (I), hypostomal teeth (E). Pheidole moramanaensis sp. nov., head (B), profile (J), hypostomal teeth (F). Pheidole makirovana sp. nov., head (C), profile (K), hypostomal teeth (G). Pheidole mantadia sp. nov., head (D), profile (L), hypostomal teeth (H).

Figure 51. 

Minor workers. Pheidole curvistriata sp. nov., head (A), profile (E), dorsal view of mesosoma (I). Pheidole moramanaensis sp. nov., head (B), profile (F), dorsal view of mesosoma (J). Pheidole makirovana sp. nov., head (C), profile (G), dorsal view of mesosoma (K). Pheidole mantadia sp. nov., head (D), profile (H), dorsal view of mesosoma (L).

Pheidole curvistriata sp. nov.

Figs 52A–F, 84L, 86L

Type material

Holotype. Madagascar. •1 major worker; Fianarantsoa; Parc National de Ranomafana, Sahamalaotra River, 6.6 km 310°NW Ranomafana; -21.23667, 47.39667; alt.