Cephalotes hamulus

Identification
Species included in the clade: Cephalotes argentiventris, Cephalotes auricomus, Cephalotes flavigaster, Cephalotes hamulus, Cephalotes resinae, Cephalotes taino, Cephalotes unimaculatus, Cephalotes vinosus.

Distribution
This taxon was described from Dominican Republic. It is also found in Haiti.

Nomenclature

 *  hamulus. Cryptocerus hamulus Roger, 1863a: 209 (w.) DOMINICAN REPUBLIC. Kempf, 1951: 149 (m.); De Andrade & Baroni Urbani, 1999: 86 (q.). Combination in Cryptocerus (Paracryptocerus): Emery, 1924d: 307; in Hypocryptocerus: Wheeler, W.M. 1936b: 201; in Zacryptocerus: Brandão, 1991: 384; in Cephalotes: De Andrade & Baroni Urbani, 1999: 84. Junior synonym of haemorrhoidalis: Emery, 1924d: 307; Menozzi & Russo, 1930: 162. Revived from synonymy as subspecies of haemorrhoidalis: Wheeler, W.M. 1936b: 201. Senior synonym of haytianus: Wheeler, W.M. 1936b: 201; Kempf, 1951: 149. Revived status as species and senior synonym of affinis: De Andrade & Baroni Urbani, 1999: 84.
 * haytianus. Cryptocerus hamulus var. haytianus Forel, 1901e: 337 (w.) HAITI. Junior synonym of haemorrhoidalis: Wheeler, W.M. & Mann, 1914: 38; of hamulus: Kempf, 1951: 149.
 * affinis. Hypocryptocerus haemorrhoidalis subsp. affinis Wheeler, W.M. 1936b: 202 (w.) HAITI. Combination in Zacryptocerus: Brandão, 1991: 384. Junior synonym of hamulus: De Andrade & Baroni Urbani, 1999: 85. See also: Kempf, 1951: 147.

Description
Cryptocerus hamulus ROGER, 1 863: 209. Worker. Original description. Type locality: Santo Domingo. Type material: one syntype worker labelled " Cryptc. hamulus, S. Domingo, Type" in the MCSN, examined. Additional syntype workers presumably in the "Zoologisches Museum an der Humboldt-Universitat", Berlin. Cryptocerus haemorrhoidalis (LATREILLE), WHEELER & MANN, 1 9 14: 38, fig. 16 (nee LATREILLE, 1 802 b). Worker, male. Misidentification proposed by WHEELER, 1 936 b: 20 1. 85 Cryptocerus hamulus var. haytianus (sic) FOREL, 1 90 1 b: 3 37. Worker. Original description. Type locality: Haiti. Type material: 2 probably syntype workers labelled "Cr. h. haytianus, Haiti" in MHNG (examined). Synonymy by WHEELER, 1 93 6 b: 20 1. Hypocryptocerus haemorrhoidalis hamulus (ROGER), WHEELER, 1936 b : 2 0 1. First combination in Hypocryptocerus. Hypocryptocerus haemorrhoidalis hamulus (ROGER), KEMPF, 1 9 5 1 : 149. Worker and male. Hypocryptocerus haemorrhoidalis affinis WHEELER, 1 93 6 b: 202. Worker. Original description. Type locality: Poste Terre Rouge (Haiti). Type material: 2 syntype workers in the MCZC (KEMPF, 1 9 5 1 : 147), examined. N e w s y n o n y m. ?Zacryptocerus haemorrhoidalis, KUGLER, 1 978: 474, figs. 1 98-1 99. Worker, sting. Zacryptocerus haemorrhoidalis hamulus (ROGER), BRANDAO, 1 99 1 : 384. First combination in Zacryptocerus. D i a g n o s i s. A member of the hamulus clade characterised by the following apomorphies: completely black body with longitudinal rugosities on the anterior fourth of the first gastral tergite and thin, short, appressed hairs on the gaster. W 0 r k e r (Fig. 26). - Vertexal angles round, with crenulate margin; vertexal margin gently concave medially. Vertex with a median pair of small denticles and a trace of transverse carina. Frontal carinae reaching the middle of the eyes posteriorly, their sides expanded, not completely covering the cheeks. Cheeks posteriorly marginate. Anterior clypeal border concave in the middle and bearing two small lateral teeth. Eyes gently convex. Mandibles without lateral carinae, robust, and directed ventrally; the masticatory margin with a developed apical followed by a smaller preapical tooth and by 0-4 minute denticles or by an edentate margin. Mesosoma robust, slightly convex in side view. Scapular angles bearing a pair of small denticles. Anterior pronotal border variably marked by a carina. Humeral angles with a pair of obtuse or stout spines directed anteriorly. Pronotal sides straight, ending in a small tooth visible in dorsal view. Promesonotal suture impressed laterally, morc superficial in the middle of the dorsum. Mesonotal teeth small, obtuse or round or absent. Propodeal suture deeply impressed dorsally. Propodeum with differentiated basal and declivous faces. Basal face slightly more than 1 /3 broader than long, with convex or obtuse sides anteriorly, straightening posteriorly and with or without a small denticle; declivous face as long as the basal face, with posteriorly slightly converging sides. Propodeal spines subequal to the basal face of the propodeum, thick and diverging. Petiole ca. 1 12 broader than long, its anterior face truncate and slightly concave; its sides converging posteriorly, bearing a pair of anterior denticles and a pair of pointed teeth medially, both variable in size. Postpetiole broader than and as long as the petiole; postpetiolar spines broad and variably pointed, inserted anteriorly and curved backwards. Gaster with a membranaceous anterior crest, broad in most specimens, and reaching the first gastral stigma at maximum. Legs. Fore coxae without a broad lobe or angle anteriorly. Hind femora slightly inflated in tl:e middle; mid and hind basitarsi laterally compressed; their sides narrowing distally in side view. Sculpture. Head minutely reticulate; vertexal area covered by broad impressions or irregular foveae gradually shallower towards the cheeks, sparser and small anteriorly. Frontal carinae reticulate and faintly rugulose on their sides. Ventral part of the head laterally covered by thick, regular striae. Hypostomal area deeply reticulate. Cheeks reticulate and rugulose on their posterior half. Mesosoma, petiole and postpetiole covered by dense, longitudinal, irregular rugae, and by superficial, minute reticulation. The striae more regular on the pleurae. Declivous face of the propodeum partially or entirely covered by superficial to thick, longitudinal rugae, and reticulation variably impressed. First gastral tergite reticulate, with longitudinal rugosities on its anterior fourth and on its ventral marginal area. Corresponding sternite with superficial longitudinal rugosities on the sides, the rest with impressed reticulation and opaque, or faintly reticulate and slightly shining in the middle, and with sparse piligerous foveae. 86 Coxae with sparse, transversal or longitudinal rugosities and reticulate. Femora deeply reticulate and with sparse, thin, irregular, transversal rugae variably impressed; dorsally with a pair of distal, short, carinae more impressed on the mid and hind legs. Outer face of tibiae and basi tarsi with longitudinal, slightly anastomosing rugosities, otherwise reticulate. Pilosity. Body with the following types of hairs: ( 1 ) truncate, erect to suberect, long on the vertexal area, on the mesosoma and on the pedicel, shorter on the anterior part of the head, on the gaster and legs; (2) thick and appressed, long on the vertexal area, on the mesosoma and pedicel; (3) thick, short, sparse, appressed on the anterior half of the head, thinner on the gaster and on the legs; (4) thin, very long and pointed on the sternites. Colour. Black. Frontal carinae, scapes and tarsomeres dark orange to light brown. Vertexal area, mesosoma and pedicel with some golden reflexes. Measurements (in mm) and indices: TL 6. 1 6-7. 1 6; HL 1 .46-1 .68; HW 1 .68-1 .96; EL 0.44-0.52; PW 1 .52-1 .80; PeW 0.72-0.86; PpW 0.96-1 .08; HBaL 0.68-0.76; HBaW 0. 1 6-0.20; CI 1 1 4.6-1 1 6.6; PI 1 08.8-1 1 1 .3; PPeI 200.0-233.3; PPpI 1 58.3-1 75.0; HBaI 23 .5-26.3. G y n e (previously undescribed, tentative specific attribution). - Differing from auricomus in the following characters: head less convex, vertexal angles convex and broader. Vertex with a longer pair of median denticles and with the transversal carina more impressed. Mesosoma. Humeral angles with a pair of longer anterior teeth. Pronotal crest strongly marked by a row of higher, pointed denticles. Petiole with the anterior face separate from the posterior one by a visible carina interrupted medially. Postpetiole and gaster missing. Legs. Fore coxae without a broad lobe or angle anteriorly. Hind femora slightly inflated in t􀄸e middle; mid and hind basitarsi laterally compressed; their sides narrowing distally in side VIew. Sculpture. Vertexal area superficially reticulate, irregularly foveolate-rugulose, the foveae more regular and small anteriorly. Ventral side of the head punctate and covered by thick, regular striae. Cheeks reticulate and irregularly rugose. Mesosoma and petiole reticulate and covered by thick, irregular foveae and rugosities, the rugosities more longitudinally oriented on the mesonotum, scutellum, propodeum and petiole. Lower mesopleurae and metapleurae reticulate and with thin rugosities, the rugosities more regular and sparser on the metapleurae. Declivous face of the propodeum reticulate and with short rugosities on the middle only: longitudinal superiorly and transversal inferiorly. Legs reticulate; outer face of the tibiae with thin, irregular, longitudinal rugosities. Pilosity. Body with the following types of hairs: (1 ) truncate, erect to sub erect, long on the vertexal area, on the mesosoma and on the petiole, shorter on the anterior part of the head and on the legs; (2) thick and decumbent, long on the vertexal area, on the mesosoma and petiole, shorter on the anterior half of the head, dense on the middle of the basal face of the propodeum; (3) thin, short, sparse, appressed on the legs. Colour. Black. Frontal carinae and antennae dark orange-ferrugineous. Body with silvery reflexes given by the pilosity. Measurements (in mm) and indices: TL (gaster missing in the unique specimen; length of the gaster measured on the complete print left in the glue) - 9. 1 8 ; HL 1 .96; HW 2. 1 2; EL 0.56; PW 2.08; PeW 0.76; HBaL 0.76; HBa W 0.22; CI 1 08.2; PI 1 0 1 .9; PPe! 2 73.7; HBaI 28.9. Male (description of KEMPF, 1 9 5 1 ). - "Head, thorax, coxae, peduncle fuscous brown. Legs, tips of mandibles, scape and gaster yellowish brown. Funiculus light brown. Mandibles with a distinct apical and preapical tooth; the following teeth vestigial. Head transverse; the median head length shorter than the interocular width. Frontal carinae divergent, fading out above the eyes. Antennal scrobe obsolescent. Eyes large, their maximum diameter sub equal to half of the median head length. Occiput evenly rounded transversely without forming distinct lateral angles. Integument finely reticulate-punctate. Transverse rugae between the upper border of the eyes and the front. Scape about half as long as the second funicular segment. Sides of pronotum diverging behind, with a distinct scapular tooth. A vestigial transverse carina across the pronotum. Laterotergite excavated beneath just above the fore coxae. Scutum and scutellum considerably convex. Mayrian furrows deeply impressed. Upper mesopleura strongly convex. Epinotum transverse, the posterior corners o f the basal face dentate. Integument finely reticulate-punctate, irregularly rugulose. Petiole about as long as wide, with a vestigial tooth on each side. Postpetiole flat above, subquadrate, with a tubercular tooth on each side. Gaster finely reticulate, subfulgid. The genitalia have been lost. Pile (sic) fulvous and erect, abundant and conspicuous on head and thorax; sparser and appressed on gaster and legs. Wings slightly infumated; veins brunneous. Fore wing with a closed and appendiculate marginal cell, and a transverse cubital vein. Submarginal cell short. Transverse median vein about half as long as the second abscissa of the median vein." Note: the sole known male of this species in the MCZC lacks the last 6 gastral segments and the genitalia as already mentioned by KEMPF ( 1 95 1 ). Measurements (in mm) and indices: TL 5 .62 (excluding gastral segments 2-8 missing in the only available specimen); HL 0.94; HW 1 . 1 6; EL 0.46; PW 1 1 .08; PeW 0.48; Pp W 0.52; HBaL 0.85; HBa W 0 . 1 1 ; CI 123.4; PI 1 07.4; PPeI 225.0; PPpI 207.7; HBal 12.9. M a t e r i a 1 e x a m i n e d . Haiti: no further locality, 1 gyne [MNHN]; idem, 2 workers (probable syntypes of hamulus haytianus) [MHNG]; Diquini, 1 worker, W. M. MANN [NHMB]; Mts. N of Jacmel, 1 0 workers, W. M. MANN [MCZC, MZSP, NHMB]; Manneville, 2 workers, W. M. MANN [MCZC]; Port au Prince, 4 workers and 1 male, W. M. MANN [MCZC, NHMB]; Port au Prince, 1 worker, 06.XII. 1 9 0 1, G. KELTEL [MHNG]; unintelligible locality, 1 worker, [MHNG]; Porte Terre Rouge, 2000', 05.X. 1 934, 2 workers (syntypes of affinis) [MCZC]. Dominican Republic: S. Domingo (no further data, syntype of hamulus), 1 worker, [MCSN]; Provo La Vega, 1 2 km NE Jarabacoa, 550 m, 02.X. 1 988, pine forest, 2 workers, M. A. IVIE, T. K. PIIILIPS & K. A. JOHNSON [USNM]. D i s c u s s i o n. - C. hamulus is recognisable by its completely black body with the anteri or fourth of the first gastral tergite completely or partially covered by longitudinal rugosities. It results as outgroup of the sister species resinae and auricomus. The three species share the pilosity completely or partially hiding the sculpture. This and a number of other taxa of the hamulus clade were formerly regarded as subspecies of C. haemorrhoidalis, a species which can not be identified with certainty at present (see chapter 5 .7, Cephalotes incertae sedis). Due to the close sympatry and apparent lack of transitional forms between most of these nominal taxa, in this paper, we consider a number of them worthy of specific rank. This is the case with hamulus. We named the whole clade after this species since it is the oldest described taxon in the clade whi<;:h corresponds to a still identifiable species (due to the availability of type material). C. haemorrhoidalis affinis WHEELER, described on two workers only (examined) has been separated on minute sculptural and morphological differences of the propodeum and of the pedicel which appear to be variable and inconsistent within the material available for the present study. Here it has been considered as a synonym of hamulus as KEMPF ( 1 9 5 1 ) already suspected. We tentatively attribute a gyne from Haiti (MNHN) without postpetiole and gaster to hamulus for the following reasons: ( 1 ) there are three species of the hamulus clade known from Haiti, i. e. hamulus, vinosus and unimaculatus; (2) gynes are systematically larger than workers, (3) the measurements of the Haitian gyne are larger than those of hamulus workers but within the range of vinosus and unimaculatus workers as it results from the following recapitulation:

hamulus vznosus unimaculatus Haitian worker worker worker gyne HW (mm) 1 .68-1 .96 1 .92-2. 1 2 1 .68-2 . 1 6 2 . 1 2 E L (mm) 0.44-0.52 0.54-0.56 0.48-0.55 0.56 PeW (mm) 0.72-0.86 0.84-0.88 0.64-1 .00 0.76 The most visible character distinguishing hamulus from vinosus and unimaculatus workers, however, is the coloration of the gaster; the Haitian gyne is without gaster. D i s t r i b u t i o n : Hispaniola.