Cylindromyrmex

Cylindromyrmex is a genus of mostly arboreal-nesting termite hunters, rarely encountered but distributed throughout New World tropics, including the Antilles. (Borowiec 2016)

Identification
Borowiec (2016) - Worker With a combination of large eyes, conspicuously costate or striate sculpture, torulo-posttorular complex horizontal and concealing antennal sockets, two pectinate spurs on mid and hind tibiae, and simple pretarsal claws, the workers of Cylindromyrmex can be readily distinguished from all other dorylines. The only other genus with large eyes, conspicuously sulcate sculpture, and two tibial spurs is Chrysapace, but it has fully exposed antennal sockets, possesses toothed pretarsal claws, and occurs only in the Old World. The extinct Procerapachys, which can also have sulcate sculpturing, has a single pectinate spur on each mid and hind tibiae.

Male The males of Cylindromyrmex are also easily differentiated from all other genera by two tibial spurs, simple pretarsal claws, no transverse groove on the mesopleuron, and well-developed wing venation with costal (C) vein present in fore wing, two submarginal cells and marginal cell closed. The only other genus with two tibial spurs and similar venation is the Old World genus Chrysapace, but it has a transverse groove on the mesopleuron and pretarsal claws armed with a tooth. Putative males of the extinct Procerapachys have only one spur on each mid and hind tibiae.

Distribution
Borowiec (2016) - Cylindromyrmex is an exclusively Neotropical lineage with ten extant species and three extinct species known from Dominican amber (De Andrade 1998a). Its distribution extends from the state of Veracruz, Mexico to Rio Grande do Sul in southern Brazil (De Andrade 1998a, Quiroz-Robledo 2003). Known from Cuba and Hispaniola, Cylindromyrmex darlingtoni is also the only member of the Dorylinae endemic in the Antilles. Cylindromyrmex whymperi has been apparently introduced and established in Galapagos Islands (De Andrade 1998a).

Fossils
Fossils are known from:.

Biology
Borowiec (2016) - Members of this lineage have been reported to be termite predators (De Andrade 1998a). Some authors described Cylindromyrmex as termite inquilines based on records of workers from termite nests (Wheeler 1936, Overal and Bandeira 1985). It seems possible, however, that these specimens represent raiding foragers of arboreal nesting ants, as complete nest series containing brood and reproductives are so far known apparently only from wood (Fernández and Escobar 1997, De Andrade 1998a, Mariano et al. 2004, Philip Ward pers. comm.). A colony of Cylindromyrmex whymperi has been recently found in Peru and studied in captivity by Josh Richards, an ant keeper from Lima, Peru. He has observed that these ants readily pursue and sting termites, which are brought to the nest paralyzed but apparently not dead. When outnumbered in a confrontation, Cylindromyrmex workers first sting as many termites as possible before attempting to carry some of them back to the nest (Josh Richards pers. comm.). Gobin et al. (2001) described a novel type of gland between sternites VI and VII in Cylindromyrmex whymperi and demonstrated that this species employs mass recruitment to termite prey. Morgan et al. (2008) chemically analyzed Dufour’s gland secretions of the same species. Three species of Cylindromyrmex (Cylindromyrmex brasiliensis, Cylindromyrmex brevitarsus and Cylindromyrmex longiceps) have been reported occurring in sympatry, collected in Malaise traps in a single locality in Bahia, Brazil. The flying males and gynes were present in samples from the end of August to beginning of December, with at least one of the samples containing all the three species (Delabie and Reis 2000). All known queens of Cylindromyrmex are winged and brood production is apparently synchronized (Mariano et al. 2004, Josh Richards pers. comm.).

Brown (1975) - The species of Cylindromyrmex nest in cavities in sound or rotten wood, under bark, in hollow stems of standing or fallen plants such as the castor bean, and in similar situations. The small-eyed species are evidently more cryptic in habits than are large-eyed forms such as C. striatus. The workers and even nests of several species have been found in termite galleries, and males and queens of Cylindromyrmex parallelus collected in a log on Barro Colorado Island, Panama Canal Zone, by A. E. Emerson, are pinned with workers and sexual forms of a termite in the MCZ. Consequently, the old assumption that the genus is termitotherous is probably correct. Still, it would be useful to have some detailed observations on the feeding habits.

De Andrade (1998) - The nesting place and feeding habits of Cylindromyrmex are still fragmentarily known. Wheeler (1936) listed Cylindromyrmex brasiliensis and williamsi (=Cylindromyrmex whymperi) as termite inquilines. Overal & Bandera (1985) equally supposed that specimens of Cylindromyrmex striatus collected in a termite nest should be termite inquilines. Their statement is partly contradicted by their report in the same paper of striatus workers attacking Nasutitermes surinamensis in laboratory.

Nomenclature

 *  CYLINDROMYRMEX [Cerapachyinae: Cylindromyrmecini]
 * Cylindromyrmex Mayr, 1870b: 967. Type-species: Cylindromyrmex striatus, by monotypy.
 * Cylindromyrmex subgenus of Cerapachys: Forel, 1892l: 243.
 * Cylindromyrmex revived status as genus: Dalla Torre, 1893: 16.
 * Cylindromyrmex senior synonym of Holcoponera Cameron (junior homonym): Forel, 1892f: 256.
 * Cylindromyrmex senior synonym of Hypocylindromyrmex, Metacylindromyrmex: Brown, 1975: 36.
 * HOLCOPONERA [junior homonym, junior synonym of Cylindromyrmex]
 * Holcoponera Cameron, 1891: 92. Type-species: Holcoponera whymperi, by monotypy. [Unresolved junior homonym of Holcoponera Mayr, 1887, above.]
 * Holcoponera Cameron junior synonym of Cylindromyrmex: Forel, 1892f: 256.
 * HYPOCYLINDROMYRMEX [junior synonym of Cylindromyrmex]
 * Hypocylindromyrmex Wheeler, W.M. 1924a: 106 [as subgenus of Cylindromyrmex]. Type-species: Cylindromyrmex longiceps, by original designation.
 * Hypocylindromyrmex junior synonym of Cylindromyrmex: Brown, 1975: 36.
 * METACYLINDROMYRMEX [junior synonym of Cylindromyrmex]
 * Metacylindromyrmex Wheeler, W.M. 1924a: 106 [as subgenus of Cylindromyrmex]. Type-species: Cylindromyrmex godmani, by original designation.
 * Metacylindromyrmex junior synonym of Cylindromyrmex: Kempf, 1972a: 91; Brown, 1975: 36.

Borowiec (2016) - Cylindromyrmex has three generic synonyms: Holcoponera Cameron, Hypocylindromyrmex Wheeler, and Metacylindromyrmex Wheeler. Cameron’s Holcoponera has been considered a synonym since the end of 19th century (Forel 1892a), and the two other names were introduced as subgenera by Wheeler (1924a) but have not been used as valid since Brown’s (1975) work on the ‘Cerapachyinae’. De Andrade (1998a) revised, illustrated, and keyed all the species of Cylindromyrmex, subsequently adding new records and a second fossil taxon from Dominican amber (De Andrade 2001).

Cylindromyrmex is the sister genus to Acanthostichus (Brady et al. 2006, Brady et al. 2014, Borowiec, in prep.). A morphology-based internal phylogeny is also available, inferred by De Andrade (1998a).

Worker
Borowiec (2016) - Head: Antennae with 12 segments. Apical antennal segment moderately enlarged, broader than and about equal in length to two preceding segments combined. Clypeus without cuticular apron. Lateroclypeal teeth absent. Parafrontal ridges absent. Torulo-posttorular complex horizontal. Antennal scrobes present. Labrum with median notch or concavity. Proximal face of stipes projecting beyond inner margin of sclerite, concealing prementum when mouthparts fully closed. Maxillary palps 3- or 2-segmented. Labial palps 2-segmented. Mandibles triangular, with teeth or edentate. Eyes present, always composed of more than 5 ommatidia and usually more than 20 ommatidia. Ocelli present or absent. Head capsule with differentiated vertical posterior surface above occipital foramen. Ventrolateral margins of head without lamella or ridge extending towards mandibles and beyond carina surrounding occipital foramen. Posterior head corners dorsolaterally immarginate. Carina surrounding occipital foramen ventrally present. Mesosoma: Pronotal flange separated from collar by distinct ridge or not. Promesonotal connection with suture completely fused. Pronotomesopleural suture visible, unfused up to notal surface. Mesometapleural groove deeply impressed, conspicuous. Transverse groove dividing mesopleuron absent. Pleural endophragmal pit concavity present. Mesosoma dorsolaterally immarginate. Metanotal depression or groove on mesosoma absent. Propodeal spiracle situated low on sclerite. Propodeal declivity with distinct dorsal edge or margin and rectangular in posterior view. Metapleural gland bulla visible or not visible through cuticle. Propodeal lobes present, well developed. Metasoma: Petiole anterodorsally marginate, dorsolaterally immarginate, dorsolaterally marginate, and laterally above spiracle marginate. Helcium in relation to tergosternal suture placed at posttergite and supraaxial. Prora forming a V-shaped protrusion. Spiracle openings of abdominal segments IV–VI circular. Abdominal segment III anterodorsally immarginate and dorsolaterally immarginate. Abdominal segment III more than half size of succeeding segment IV, which is weakly constricted at presegmental portion (uninodal waist). Girdling constriction of segment IV present, i.e. pre-and postsclerites distinct. Cinctus of abdominal segment IV gutter-like, not sculptured. Abdominal segment IV not conspicuously largest segment. Abdominal tergite IV not folding over sternite, and anterior portions of sternite and tergite equally well visible in lateral view. Girdling constriction between pre- and posttergites of abdominal segments V and VI absent. Girdling constriction between pre- and poststernites of abdominal segments V and VI present or absent. Pygidium large, with impressed medial field and armed with modified setae. Hypopygium unarmed. Legs: Mid tibia with two pectinate spurs. Hind tibia with two pectinate spurs. Hind basitarsus not widening distally, circular in cross-section. Posterior flange of hind coxa not produced as raised lamella. Metatibial gland present as oval patch of whitish cuticle. Metabasitarsal gland absent. Hind pretarsal claws simple. Polymorphism: Monomorphic.

Brown (1975) - Essentially monomorphic but varying considerably in size, even in uninidal series. Color reddish brown to black or piceous; appendages yellow to black, or dark with the tibiae extensively marked with yellow or ivory. Integument thick and hard.

Head longer than broad, and deep dorsoventrally; sides parallel, with blunt posterior corners and concave posterior border. Clypeus short, crowded by the antennal insertions and the frontal carinae, which approach or reach the anterior margin of the head; in the latter case, the median section of the clypeus is vertical. Posteromedian section of clypeus extending posteriad between the nearly approximate frontal lobes as a deeply sunken groove that broadens very slightly behind to encompass the tiny frontal area, which is not or barely distinguishable. Frontal carinae subparallel behind, where they arise more or less near the cranial midlength, curving mesad anteriorly to form a rounded plate or twin blunt points at or above the main part of the clypeus. The carinal lobes are extended laterad in the form of broad laminae roofing the medial sides of broad antennal scrobes that extend forward from the level of the eyes to the antennal insertions. Behind median groove, space between carinae is broad, flat, or gently convex. Ocelli present or absent. Antennae very short, broad, and flattened, 12-merous; scape only about 2-3 times as long as broad, and not, or just barely, reaching the eye; funiculi gradually broadened toward apices; a vague club formed of the 3 apical segments, or no club distinguishable; apical segment as long as, or longer than, the 2 preceding segments taken together. Mandibles subtriangular, thick and slightly downcurved; apical and basal borders distinct and meeting at an angle; apical border with 4-10 low teeth or crenulations, sometimes virtually edentate, but with a more or less acute apex. Maxillary palpi with 2 segments, labial palpi with 2 or 3 segments; mouthparts of Cylindromyrmex striatus described by Gotwald 1969:43, 47, pI. 31. Compound eyes situated near or behind cranial midlength, ranging from small and flat, with 20 or less indistinct facets, to large and moderately convex, taking up nearly 1/3 length of the sides of the head and having 500 or more facets.

Trunk elongate, boxlike, with subparallel, vertical sides and a gently convex dorsum (sometimes nearly flat); dorsal sutures represented at most by a feeble, promesonotal, arched line and a weakly indicated metanotal groove with a median pit, but often these sutures are obliterated. Lateral sutures reduced to an inverted U- or A-shaped system outlining the mesopleuron. Pronotum not transversely marginate in front, humeral angles rounded; propodeal declivity distinct and flat or nearly flat, but not, or bluntly, margined laterally and above. Propodeal spiracle situated below the middle axis of the trunk, round, oval, or elliptical, opening directed dorsad and usually slightly caudad. Metapleural gland opening a horizontal slit very near the bottom edge of the trunk (below a rather prominent bulla); a horizontal groove extends forward along the lower side of the trunk to the vicinity of the mesometanotal suture.

Petiolar node subcuboidal, usually a little longer than high and about as broad as long, a little broader behind than in front, sides slightly convex and vertical (receding ventrad); subpetiolar process stout and bluntly angular in front, subsiding concavely behind. Postpetiole much wider than petiole, wider than long, and almost as wide as succeeding (first gastric) segment, from which it is separated by a wide pretergital belt belonging to the latter; constriction between these two segments is marked, especially in side view. Stridulatory file present and extremely fine on middle of first gastric segment pretergite but not readily visible unless gaster is flexed.

In C. striatus, Gotwald (1969: 126) found the tergum and sternum of the postpetiole (true abdominal somite III) to be fused, while in the succeeding segment, which I call the “first gastric” in this paper, the tergum and sternum are connected only by membrane, as are those of the segments following. The first gastric segment is larger than the postpetiole and the second and third gastric segments, but not markedly so, and these segments are well developed, mobile, and extend free. Apical (VII true abdominal) somite well developed, tapered caudad, the tergum flattened (obliquely truncate) and margined along the sides with subreclinate spinules that appear to be short, stout setae in raised sockets, arranged on each side in a more or less regular single row. The pygidium itself ends in a pair of blunt, flattened teeth lying just above the sting, which is well developed and tends to be curved in a sword shape (laterally compressed) and usually is extruded part way.

Legs stout, moderate to short in length; femora dorsoventrally incrassate and anteroposteriorly compressed, their flexor surfaces with a long, deep sulcus to receive the curved flexor surface of the tibia when the leg is folded. Tibiae more or less incrassate, especially in the assumed cryptic-foraging small-eyed species; apical spurs long, broadly pectinate, especially on foreleg; middle and hind legs each with an extra, small, pectinate outer spur next to the large one.

Metatarsus, especially that of middle leg, extremely variable in proportions: long and slender in C. striatus, much shorter and broadened apically in the small-eyed species, with a semicirclet of 4-5 stout, spinelike setae on the outer apical edge, best developed also in the small-eyed species, where the middle legs may serve (along with the pygidium) as "pushers" in helping the ant through narrow cracks or passages in soil or rotten wood, perhaps through defensive walls being raised by termite prey.

“Pusher legs” are also found in the termite predators of tribe Acanthostichini, in genera Centromyrmex and “Wadeura” (Pachycondyla) of tribe Ponerini, in Cryptopone (prey unknown) of the same tribe, and in the termitotherous myrmicine genus Metapone; in Melissotarsus, similar middle legs bear glands that may have a very special use in marking trails along the substrate above the ant's body as it moves along (see Delage-Darchen, 1972, Insectes Sociaux 19:213 If.).

Other tarsal segments also with 2-5 spinelike setae at apices. Claws simple, thickened basally.

De Andrade (1998) - Monomorphic but variable in size. Head longer than broad with slightly convex, subparallel or parallel sides. Clypeus short. Frontal carinae parallel or subparallel diverging posteriorly. Ocelli present or reduced to an impressed pit. Compound eyes placed on the middle or slightly behind the mid line of the head and with a variable number of ommatidia (16-500). Antennae 12-jointed. Funicular joints 8-10 with spine-like seta on the proximal border; last joint with similar spine-like seta but almost on its all surface. Scapes reaching or slightly surpassing the anterior border of the eyes. Funiculi thickening from the base to the apex. Mandibles subtriangular, dorsally flat or convex. Masticatory margin of the mandibles with 4-14 irregular denticles or edentate. Apex of the mandibles with pointed apical tooth. Palpal formula 2.2 or 2.3. Mesosoma elongate. Cylindric, with parallel sides and weakly convex dorsally. Promesonotal and propodeal sutures absent, simply marled by a pit or superficially impressed. Promesopleural suture superficially or deeply impressed. Meso-metapleural suture superficially impressed. Humeral angle, round. Propodeum with basal and declivous faces distinct separated or not by a margin. Propodeal spiracle round or oval and placed at mid height in lateral view. Petiole subcylindric, as long as broad, longer than broad, or shorter than broad. Petiolar sides subparallel and often diverging posteriorly. Ventral petiolar process small or large, subtriangular, subtruncate, or subround. Postpetiole (abdominal segment III or gastral segment I) broader than petiole, broader than long, and as broad as the first gastric segment (abdominal segment IV or gastral segment II). Postpetiolar sternite antero-medially without or with a variably marked triangular “lip.” Pygidium obliquely or perpendicularly truncate: apex of pygidium with or without a notch. Sides of pygidium surrounded by a set of many irregularly distributed denticle, within 2-4 larger denticles above the sting, or with a row of denticles enlarging apically. Sting developed, curved upwards and with flat sides. Legs incrassate or slender. Femora with a concavity of variable deepness to receive the tibiae. All tibiae with a large, pectinate spur. Mid and hind tibiae with an additional, smaller, pectinate spur close to the large one. Basitarsi of the three pairs of legs of variable length and with 3-7 spine-like setae on the outer apical edge. First, second and third tarsomeres with similar spines. Fourth (apical) tarsomeres of variable length. Pretarsal claws thicker proximally than distally and with a small denticle or an angle on the proximal part. Head, mesosoma and petiole covered by longitudinal striae of variable thickness. Postpetiole smooth or striate. First. second and third gastric tergites smooth and variably reticulate-punctate or longitudinally striated. Remaining gastric tergites, sternites and pygidium smooth and/or reticulate-punctate. Legs smooth to superficially punctate: some species with hind or hind and mid coxae longitudinally striated. Body with pointed hairs of different size and variably distributed, generally denser on the gaster. Colour dark ferrugincous to black. Legs concolour with or lighter than the body. Some species with yellowish tibiae.

Queen
De Andrade (1998) - Very similar to the worker but differing from it in the following characters. Size slightly or much larger than the worker. Ocelli and compound eyes larger. Fore wing with well marked veins and pterostigma. Rsf5 connected eith RI. Mf2 and r-m medially interrupted. Mf4 and CuA I variably pigmented. Hind wing with R+Sc, M+CuA and A pigmented. Distal veins faintly pigmented. CuA and IA more pigmented than Rs and M. In some species the wings have violaceous relflexes. Dorsum of the mesonotum with or without striae on the sides. Mesopleurae striate or not on the anterior part. Scutellum smooth or with variably impressed longitudinal striae.

Male
Borowiec (2016) - Head: Antennae with 13 segments. Clypeus without cuticular apron. Parafrontal ridges absent. Torulo-posttorular complex vertical. Maxillary palps 2-segment ed. Labial palps 3- or 2-segmented. Mandibles triangular, edentate. Ventrolateral margins of head without lamella or ridge extending towards mandibles and beyond carina surrounding occipital foramen. Carina surrounding occipital foramen ventrally present. Mesosoma: Pronotal flange separated from collar by distinct ridge or not separated. Notauli absent or present. Transverse groove dividing mesopleuron absent. Propodeal declivity with distinct dorsal edge or margin. Metapleural gland opening present. Propodeal lobes present. Metasoma: Petiole anterodorsally marginate, dorsolaterally immarginate, and laterally above spiracle marginate. Helcium in relation to tergosternal suture placed at posttergite and supraaxial. Prora forming a U-shaped margin with median ridge. Spiracle openings of abdominal segments IV–VI circular. Abdominal segment III more than half size of succeeding segment IV; latter weakly constricted at presegmental portion (uninodal waist). Girdling constriction of segment IV present, i.e. pre- and postsclerites distinct. Cinctus of abdominal segment IV gutter-like, not sculptured or cross-ribbed. Girdling constriction between pre- and postsclerites of abdominal segments V and VI absent. Abdominal segment IV not conspicuously largest segment. Abdominal sternite VII simple. Abdominal sternite IX distally armed with two spines, with lateral apodemes about as long as medial apodeme, directed anteriorly (towards head). Genitalia: Cupula long relative to rest of genital capsule and shorter ventrally than dorsally. Basimere broadly fused to telomere, with sulcus discernable at junction, and ventrally with left and right arms abutting. Telomere gradually tapering toward apex. Volsella laterally flattened, at apex with dorsal lobe and hooked ventrally. Penisvalva laterally compressed, rounded at apex. Legs: Mid tibia with two pectinate spurs. Hind tibia with two pectinate spurs. Posterior flange of hind coxa not produced as raised lamella. Metatibial gland absent. Metabasitarsal glands absent. Hind pretarsal claws simple. Wings: Tegula present, broad, demiovate in shape. Vein C in fore wing present. Pterostigma broad. Abscissa R·f3 present, running toward distal wing margin and enclosing cell with Rs·f5. Abscissae Rs·f2–3 present, connecting with Rs+M&M·f2. Cross-vein 2r-rs present, differentiated from Rs·f4 by presence of Rs·f2–3. Abscissae Rs·f4–5 differentiated into Rs·f4 and Rs·f5 by 2rs-m. Abscissa M·f2 in fore wing contiguous with Rs+M. Abscissa M·f4 in fore wing present, reaching wing margin. Cross-vein 1m-cu in fore wing present. Cross-vein cu-a in fore wing present, arising from M+Cu and proximal to M·f1. Vein Cu in fore wing present, with both branches Cu1 and Cu2. Vein A in fore wing with abscissae A·f1 and A·f2 present. Vein C in hind wing absent. Vein R in hind wing present, extending past Sc+R but not reaching distal wing margin. Vein Sc+R in hind wing present. Abscissa Rs·f1 in hind wing present, shorter than 1rs-m. Abscissa Rs·f2 in hind wing present, not reaching wing margin. Cross-vein 1rs-m in hind wing fused with M·f1. Vein M+Cu in hind wing present. Abscissa M·f1 in hind wing present. Abscissa M·f2 in hind wing present. Cross-vein cu-a in hind wing present. Vein Cu in hind wing present. Vein A in hind wing with abscissae A·f1 and A·f2 present.

Brown (1975) - Similar in size to the conspecific worker, but body more slender; head shorter, subglobular, with very large convex eyes taking up more or less the anterior half of the sides of the head. Antennae long and slender, 13-merous, with very short, cylindrical scape, even shorter, cup-shaped pedicel, and remaining flagellar segments long and subcylindrical.

Frontal lobes short, raised, separated in front, but fused behind. Clypeus with a broad, sloping anterior part and a narrow posterior part between frontal lobes. Mandibles substantial, narrow-subtriangular, with curved, edentate (cultrate) apical margins and an acute, incurved apical tooth. Mandibles crossing over each other at full closure, leaving no anteclypeal space. Palpi segmented 2,2 or 2,3.

Trunk compact and subcylindrical, pterothorax only gently convex and slightly raised above truncal outline. Notauli present (C. parallelus) but incomplete behind, not meeting as a Y or V. Metanotum small, transversely elliptical. Propodeum long, declivity nearly perpendicular and with a very strong margin and a vertical median septum or carina.

Wings with complete ponerine venation, but Mf2 and r-m have wide breaks, apparently at the crossings of a fold line; radial cell fairly short, closed apically at or very near the margin. Pterostigma thick, convex, heavily pigmented. Hind wing with distal free abscissae ending near mid-wing; r-m weak and broken; anal lobe lacking.

Legs short and only moderately robust; anterior and middle coxae inserted far apart. Middle and hind tibiae each with 2 pectinate apical spurs, one larger than the other. Tarsal claws simple but with thickened, sometimes angular bases.

Petiole much like that of worker; anterior face steep and marginate. Subpetiolar process present. Postpetiole only a little wider than petiole and not much narrower than gastric segments I-IV, which are subequal among themselves in length and width; postpetiole separated from gaster by a distinct constriction. Pygidium bluntly rounded; hypopygium ending in paired, long posterior spines and a short median tooth (in C. parallelus). Genital capsule complete; parameres tend to be divided into basal and distal pieces (gonocoxites and gonostyli?).

Integument coarsely striate as in worker and queen, but the pterothorax (including pleura) are largely smooth and shining, with numerous, fine punctures; postpetiole variably sculptured; striate, longitudinally rugulose, or partly reticulate; gaster shining, very finely reticulate, and with scattered punctures.

Pilosity fine, short, predominantly decumbent. Color black, legs and antennae often prevailingly yellow or light brown.

De Andrade (1998) - Size variable, generally smaller, but in some species as large as or larger than the gyne. Head shorter than broad, as long as broad, or longer than broad. Vertex convex. Frontal carinae developed but never completely hiding the antennal socket. Sides of the frontal carinae subparallel, or broad anteriorly and converging posteriorly, or strongly broad anteriorly and touching each other posteriorly. Antennae 13-segmented, varying from 1/3 to 1/2 of the maximum body length. Ocelli large. Compound eyes very large, slightly longer than 1/2 or the head length and largely on the anterior half of the head sides. Scapes very short. First funicular joint less than or about 1/2 of the length of the second one: second and last two apical joints thinner than joints 3-10. Mandibles slender, edentate except for a visible apical pointed tooth. Mesosoma robust. Pronotum with subparallel or diverging sides. Mesonotum and scutellum gently convex. Pair Mayrian furrows impressed or not. Parapsidal furrows variably impressed. Propodeum with the sides converging posteriorly. Basal face of the propodeum separate from the declivous one by a well marked carina. Petiole cylindric, as long as or longer than broad. Anterior face of the petiole truncate and separate from the dorsal one by a marked carina. Subpetiolar process variable in size, subtriangular or subtruncate. Postpetiole broader than the petiole. Postpetiolar sides diverging posteriorly or gently convex. First gastric segment broader than the postpetiole. Second gastric segment as broad as or slightly narrower the first segment, rarely broader than the first segment. Remaining gastric segments narrowing posteriorly. Legs long and slender. Head with deep punctures or piligerous foveae sometimes separated by irregular or regular striae. Mesooma pro- and rnesopleurae smooth and with punctures or piligerous foveae of variable size. Propodeum and metapleurae with thick, longnudrnal. rugosities, sometimes irregular. Petiole and postpetiole smooth or with irregular longitudinal rugosities, very superficial on the postpetiole. Gaster and legs smooth and variably punctate. Body with pointed hairs denser than in the female castes. Sometimes the posterior part of the head, pronotum, gaster and legs with dense pilosity of variable size. Wings as in Fig. 5, similar to the one of the gyne. Colour brown to black. Legs concolour with or lighter than the body. Some species with yellowish tibiae.

Larvae
Not described. Cocoons absent.