Odontomachus chicomendesi, França & Fernandes & Bravo, 2024

França, Eder Cleyton Barbosa, Fernandes, Itanna Oliveira & Bravo, John Edwin Lattke, 2024, Looking at upside-down ants: Taxonomic revision of the Neotropical species of Odontomachus Latreille, 1804 (Hymenoptera: Formicidae: Ponerinae), Zootaxa 5502 (1), pp. 1-166 : 87-131

Odontomachus cornutus Stitz, 1933 ( Figures 11-A , 79 and 80 )

Odontomachus cornutus Stitz, 1933: 67 (☿); Kugler, 1980: 227 (♀ and ♂).

Type specimens: ECUADOR: Bucay, 31.v.-5.xi.1905 (no collector’s name); Type-depository: Hamburg Museum (Bolton, 2023). Holotype destroyed in World War II ( Brown, 1976).

Etymology: From the Latin word cornutus = having horns; probably referring to the acute ocular prominences.

Diagnosis: (☿) Vertex rectangular wider than long, smooth and shining. Ocular prominences acute projected latero-anteriorly forming spines. Central disc of pronotum forming concentric circles. Petiole pedunculate, node conical; in dorsal view, longitudinal carina continuous to petiolar spine. (♀) Vertex, ocular prominences and petiole as in worker (Kluger, 1980). (♂) Not examined, see Kluger (1980) for description.

Worker

Additional material examined: EL 0.4, HL 4.1, HoW 3.4, HvW 2.4, HW 2.5, ML 2.4, SL 4.2, PnW 1.6, WL 4.9, FL 4.8, PTH 1.5, PTL 0.9, PTW 0.5, GL 3.6, TL 15.9. CI 0.60, MI 0.58, PtI 1.66, SI 1.02, VI 0.96 (n= 1).

Head. Vertex rectangular wider than long in frontal view; smooth and shining, regular longitudinal striae diverging posteriorly limited to frontal area. Posterior margin concave medially; occipital carina not forming nuchal collar. Cephalic constriction conspicuous. Frontal lobes smooth and shining with inconspicuous striae posteriorly. Ocular bridges posteriorly forming angle. Eye relatively large (EL> 0.4); rounded, posterior margin rounded. Ocular prominence acute, directed latero-anteriorly forming spine. Clypeus smooth and shining on median portion. Malar area wrinkle anteriorly in latero-oblique view, smooth and shining posteriorly.

Mandible: Slightly longer than anterior head portion. External margin straight. Masticatory margin with denticles that serially increase in size apically. Palp formula 4,4.

Antenna: Scape surpassing occipital corner by more than pedicel and first flagellomere length.

Mesosoma: Pronotum in lateral view triangular, dorsal margin convex; neck well developed anteriorly in dorsal view; lateral margin rounded with obtuse angle posteriorly; central disc of pronotum with regular striae forming concentric circles, lateral striae also concentric. Propleuron with micro-sculpturing giving coriaceous aspect but not imbricated on external margin. Mesonotum slightly longer than wider; with regular transverse striae. Mesopleuron with vestigial carina on anteroventral portion not forming projection; mostly smooth and shining, anterior portion with fine irregular transverse striae. Mesometanotal sulcus well-marked, scrobiculate; curved anteriorly. Metanotum as long as half-length of mesonotum; with regular fine transverse striae; strongly curved anteriorly. Metapleural gland bulla smooth and shining. Propodeum in lateral view straight; posterior margin length about 1/4 of dorsal length. Metasternum with metasternal process open, cuticle border fine forming lateral projections; with regular transverse striae.

Petiole: Pedunculate; peduncle shorter than petiolar node. Conical node, almost cylindrical, anterior margin in lateral view slightly convex, posterior margin more convex than anterior. Anterior margin with longitudinal carina elevating posteriorly, continuous with petiolar spine. Spine robust, short, straight, tip slightly curved posteriorly, as long as node half-length. Entire surface smooth and shining, with at most inconspicuous longitudinal striae on lateral face on ventral portion. Subpetiolar process longer than wider; tip acute, concave posteriorly; posterior portion not concave.

Gaster: First tergum almost as high as long; anterior and dorsal margin meeting in a convex surface not forming an angle. dorsal face of all terga smooth and shining.

Color and size. Body nut-brown. Leg, antenna and mesopleuron slightly lighter. Large size ants.

Pilosity. Dense short pubescence on body surface, separated from each other less than its length. Pubescence on gaster denser and slightly longer. Short, erect hairs on pronotum and covering gastral terga, shorter than procoxa width.

Natural history: Little is known about this species. Kugler (1980) found a nest in soft deep soil under leaflitter in a montane forest at Chocó, Colombia. He could not find any larvae, even after digging down to 30 cm. The author suspected that it could be a nest fragment, but considering the depth of the nest of O. chelifer which might reach 1.34 m ( Guimarães et al. 2018), another relatively large size species, the nest of O. cornutus is probably deeper.

According to Jimenéz (2008), O. cornutus also occurs at Río Calima, Valle del Cauca, Colombia suggesting a wider distribution, but still restricted to South America’s northwest portion ( Fig. 80 ).

Comments: Odontomachus cornutus is the only species of the genus with acute ocular prominences, making it easily recognizable. Its shining vertex is wider than long, while in O. hastatus it is posteriorly narrow and relatively elongated. Even though the vertex of O. mormo is relatively elongate, the petiolar node of both species are very contrasting: in O. cornutus the peduncle does not have blunt anterior projections, which are present in O. mormo ; O. cornutus has the entire body with widespread relatively long and suberect pubescence. Such pubescence is absent in O. mormo .

Material examined: COLOMBIA: Chocó: 10 Km SW S. [San] José del Palmar (2☿) .

Odontomachus cupreus França, Fernandes & Lattke new species urn:lsid:zoobank.org:act:DC040086-174B-475C-8ED7-A5534310F624

( Figures 25-B , 27-A , 32-B , 78 , 81 and 82 )

Type specimens: Holotype. BRAZIL: São Paulo: São José do Rio Preto , UNESP-Jardim, Amorim L. col., 19.X.1977, 1510, Coll. Diniz, DZUP591299 1☿ [ DZUP].

Paratypes. Same data as holotype. DZUP 591300 1☿, DZUP 591301 1☿, DZUP 549106 1☿, DZUP 591894 1☿ [ DZUP]/ DZUP 591895 1☿, DZUP 591896 1☿ [ MZSUP]/ HYM-INPA 034967 1☿, HYM-INPA 034968 1☿ [ INPA] .

Etymology: The specific epithet cupreus (from the Latin word cupreum = copper, meaning “of copper”) is a Latin singular masculine adjective in the nominative case and refers to the reddish-brown color of individuals of this species resembling copper.

Diagnosis: (☿) Propleuron with fine transverse striae. Dorsal posterior portion of procoxa micro-sculptured.

Conical petiolar node with regular transverse striae strongly punctate. Mesosoma and pedicel usually red copperish. (♀) Propleuron, petiole and body color as in worker. (♂) Unknown.

Worker

Holotype measurements: EL 0.4, HL 2.5, HoW 1.6, HvW 1.4, HW 1.6, ML 1.2, SL 1.9, PnW 1, WL 2.5, FL 2.0, PTH 1.0, PTL 0.5, PtW 0.3, GL 2.2, TL 8.6. CI 0.72, MI 0.54, PtI 2.0, SI 0.8, VI 0.87.

EL 0.3-0.4, HL 1.8-2.4, HoW 1.4-1.9, HvW 1.2-1.9, HW 1.3-2.1, ML 1.0-1.3, SL 1.7-2.1, PnW 0.8-1.1, WL 2.2-2.9, FL 1.9-2.4, PTH 0.8-1.1, PTL 0.4-0.6, PTW 0.3-0.5, GL 1.7-3.2, TL 7.1-10.0. CI 0.70-0.95, MI 0.50-0.60, PtI 1.83-2.75, SI 0.82-0.95, VI 0.82-0.94 (n= 30).

Head. Vertex trapezoidal, wider than long in frontal view; with regular longitudinal striae diverging posteriorly. Posterior margin concave medially; occipital carina not forming nuchal collar. Cephalic constriction conspicuous. Frontal lobes striated. Ocular bridges posteriorly forming arc. Eye relatively large (EL> 0.4); rounded, posterior margin rounded. Clypeus with irregular longitudinal striae on median portion. Malar area wrinkly anteriorly in latero-oblique view, posterior portion smooth and shining.

Mandible. Almost as long as anterior head portion. External margin slightly convex. Masticatory margin with denticles that serially increase in size apically. Palp formula 4,3.

Antenna. Scape surpassing occipital corner about pedicel half-length or less.

Mesosoma. Pronotum in lateral view rounded, dorsal margin convex; neck well developed anteriorly in dorsal view; lateral margin rounded; central disc of pronotum with regular longitudinal striae, lateral striae forming concentric circles. Propleuron with fine transverse striae. Mesonotum larger than wide; with regular transverse striae. Mesopleuron with anterior portion with regular transverse striae curved dorsal-anteriorly; median portion smooth and shining; posterior portion with regular transverse striae; cuticle border present on anteroventral portion forming callosity. Mesometanotal sulcus well-marked, scrobiculate, slightly curved anteriorly. Metanotum as long as notopropodeal sulcus; with irregular transverse or longitudinal striae extending to anterior propodeum portion. Metapleural gland bulla with transverse striae effacing posteriorly. Propodeum in lateral view almost straight, with two shallow impressions; posterior margin length about 1/3 of dorsal length. Metasternum with metasternal process close and bilobate; regular transverse striae.

Petiole. Sessile. Conical node, anterior margin in lateral view continuous with petiolar spine. Spine short, acute, straight or curved posteriorly. Regular transverse striae punctate covering all surface, coarser on ventral portion. Subpetiolar process wider than long, keel-shaped; tip acute; posterior portion concave.

Gaster. First tergum almost higher than long; anterior and dorsal margin meeting in an obtuse angle; dorsal face convex; dorsal face of all terga imbricate.

Color and size. Head reddish brown. Mesosoma and pedicel red copperish. Gaster dark brown. Legs and antennae yellowish. Relatively medium size ants.

Pilosity. Hook-shaped, dense, conspicuous and short pubescence spread over the body, separated each other about its half-length. Short hairs on anterior portion of procoxa smaller than coxae width. Pronotum with two or three pairs of curved erect hairs almost long than maximum procoxa width. Short subdecumbent hairs on posterior 3/4 of gastral terga directed posteriorly, as long as procoxa hairs.

Gyne

Non-type measurements: EL 0.4-0.5, HL 2.1-2.3, HoW 1.7-1.9, HvW 1.4-1.5, HW 1.6-1.8, ML 1.2-1.3, SL 1.8-2.0, PnW 1.1-1.3, WL 2.8-3.1, FL 2.1-2.2, FWL 5.0-5.5, PTH 1.1-1.3, PTL 0.5, PtW 0.5-0.6, GL 2.4-3.2, TL 8.5-9.9. CI 0.76-0.78, MI 0.56-0.57, PtI 2.20-2.60, SI 0.85-0.90, VI 0.83-0.93 (n= 4).

Mesosoma. Pronotum slightly longer than high in lateral view, dorsal margin slightly concave; lateral margins converging anteriorly forming a neck in dorsal view; pronotal dorsum with regular transverse striae curved anteriorly and forming arcs directed posteriorly on lateral face. Subalar area conspicuous with pronotum. Mesoscutum anterior and dorsal face forming single convexity in lateral view; with regular longitudinal striae in dorsal view; striae between parapsidial line and parascutal area oblique, directed medially. Parapsidial line curved, well-marked. Parascutal area smooth and shining. Notopropodeal sulcus scrobiculate. Axilla with longitudinal inconspicuous striae anteriorly. Mesoscutellum smooth and shining with vestigial longitudinal carina. Metascutellum smooth and shining with longitudinal carina.

Wing. Color pale yellow, translucid. Forewing ML1 slightly curved. RsL1 straight to slightly curved; as long as Rs+M. RSL2-3 slightly sinuous. ML2 short, about half-length of Rs+M. Rs+M and ML2 forming single slightly curved line, not forming angle with RsL2-3. Vein cu-a not reaching M+Cu ML1 angle by less than its half-length. Hindwing 1rs-m vein slightly curved. 7 to 9 hamuli.

Petiole. Conical node, anterior face straight. Irregular transverse striae coarser than worker.

Pilosity. Pubescence of mesosoma strongly curved, dense separated each other by less than its length. Pubescence on mesoscutellum and metascutellum longer and denser than pronotum pubescence, about half-length longer. Pronotum with three or four pairs of short hairs, shorter than pronotum height.

Natural history: Odontomachus cupreus has been found nesting in the soil, under rotten trunks, and inside the nests of Cornitermes cumulans (Kollar, 1832) . There are records of specimens collected in hollow trunks of Caesalpinia pluviosa DC. ( Santos et al. 2021) . In this study, the authors identified the specimens as O. brunneus , but their vouchers were checked and identified as O. cupreus (INPA-HYM 033677 and Voucher 6010). They observed that workers responded to help signs made by other workers stuck on adhesive tape. The stuck workers displayed stridulation behavior while the free workers were aggressive against the adhesive tape.

This species can be collected using pitfalls and leaf litter sifters. Odontomachus cupreus lives in forested areas, gallery forests, meadows, pastures and urban areas. It is widely distributed in Brazil’s Cerrado and in the Atlantic Rain Forest ( Fig. 78 ). Considering the number of specimens examined and the range of O. cupreus , we suspect that the specimens from Amazonas were probably mislabeled.

Comments: Odontomachus cupreus specimens were usually identified as O. bauri due to its petiolar striation. However, the striae of O. bauri are coarser and are not punctate, as in O. cupreus . The procoxa of most specimens of O. cupreus have fine basal striae on their posterior portion and the propleuron has fine transverse striae. Some populations, especially in Minas Gerais and São Paulo, have effaced striae on the procoxa and propleuron. They can be distinguished from O. bauri by the petiolar sculpturing and the reddish brown and dark brown almost black body color of O. bauri . Using just body color is not recommended to distinguish both (see O. bauri comments). We also often found specimens of O. cupreus identified as O. brunneus . In this case, the conspicuous and reticulate node can distinguish O. cupreus from the almost or completely smooth and shining node of O. brunneus .

In a taxonomic revision of the Neartic species, Macgown et al. (2014) based on male morphology and the comments of Deyrup & Cover (2004), proposed that O. brunneus records from South America, are actually O. ruginodis . We found no specimens of O. brunneus from South America. However, O. cupreus is very similar to O. ruginodis , but O. ruginodis never has striae on the propleuron and procoxa. In the Fernandes et al. (2021) phylogeny, specimens identified as O. brunneus (Vouchers I58, I60 and I61) from the Brazilian Cerrado were examined and identified as O. cupreus . If we supposed that O. cupreus is a variation of O. ruginodis from South America, it would be paraphyletic to O. chicomendesi sp. nov. and O. brunneus . The distribution of O. brunneus and O. ruginodis restricted to Central America corroborates the hypothesis of O. cupreus as a species ( Fig. 78 ).

Material examined: 112☿, 6♀. BRAZIL: Amazonas: Humaitá (14☿) . Bahia: Salvador (1☿) . Distrito Federal: Brasília (1☿) . Goiás: Jataí (11☿, 1♀) ; Mineiros (22☿); RPPN Serra do Tombador (1☿) ; Serranópolis (4☿). Mato Grosso: Cuiabá (2☿) ; Sto A. [Santo Antônio] do Levenger (2☿) ; São Luiz de Cáceres (1☿) . Mato Grosso do Sul: [ Amambaí ] (1☿) ; Paranaíba (2☿); Porto Murtinho (7☿). Minas Gerais: Arassuahy [Araçuaí] (4☿) ; Boa Esperança (4☿); Rio Pardo de Minas (1☿) ; Ritápolis (1☿); Santana do Riacho (1☿) . Paraná: Jaguariaíva (1☿) ; P. E. [Parque Estadual] do Cerrado (2☿, 1♀) ; Ponta Grossa (2☿). São Paulo: Lins (13☿) ; Mirassol (5☿, 4♀); Mogi-Guaçu (1☿); São José do Rio Preto (3☿) ; Tanabi (1☿). Sergipe: São Cristóvão (1☿) .

PARAGUAY: [Caaguazú]: Pastoreo [Doctor Juan Manuel Frutos] (2☿). [no further data] Ptº 14 de Mayo; R Nabilecche [Río Nabileque] (1☿) .

Odontomachus davidsoni Hoenle, Lattke & Donoso, 2020 ( Figures 9-B , 10-B and 80 )

Odontomachus davidsoni Hoenle et al. 2020: 86 (☿ and ♀).

Type specimens: Paratypes. ECUADOR: Esmeraldas, Res [erva] Río Canandé , 0.5252 -79.2079, 300m, 20.V.2018, P. Hoenle & A. Agosti leg. [col.], forest selected logged AB_3 plot 8, BOLD ID: ODECU002-19, DZUP548820 1☿ / AB_4 plot 8 BOLD ID: ODECU001-19, DZUP548819 1☿ [ DZUP] .

Etymology: In honor to Carleton Davidson (1963–2001), founder of Clyde’s Restaurant Group who had a lifelong interest in Ecuador environments (Hoenlle et al. 2020).

Diagnosis: (☿) Rectangular vertex narrow posteriorly, with regular longitudinal striae oblique posteriorly. Central disc of pronotum with transverse striae curved anteriorly, external striae forming concentric ellipses. Petiole pedunculate. Dorsum of gastral terga smooth and shining. Body not covered with long pubescence. (♀) Vertex, petiole and pubescence as in worker. (♂) Unknown.

Worker and gyne descriptions: See Hoenle et al. (2020).

Natural history: Odontomachus davidsoni still is known only from its type-locality ( Fig. 81 ) ( Hoenle et al. 2020). This species is known to lives in hollow twigs and has been observed only foraging on the vegetation, with the highest worker activity between 8–11 pm. One nest with 18 workers and a delate queen was collected. The nest entrance was found under a bromeliad and had a tunnel of 40 cm length leading to a chamber. Gynes were collected with light traps. According to Hoenle et al. (2020) it may be threatened and more studies about its biology and distribution are necessary.

Comments: Odontomachus davidsoni is very similar to O. hastatus due to their slender body, orangish to reddish brown color, and elongate vertex ( Fig. 6-A ). Both species can be separated by the vertex striae present in O. davidsoni , but absent in O. hastatus ; the pronotum striae form concentric circles in O. davidsoni , but in O. hastatus they are transverse with an effaced central portion. The metasternal process is also diagnostic, forming a pair of blunt projections in O. davidsoni and in O. hastatus forms an arc with a middle notch.

Gynes have the same differences as the workers, except for the pronotum striae that are transverse in both species. The posterior wings of O. davidsoni have a spurious longitudinal vein in the basal cell between the RsL and ML veins, but this vein is absent in O. hastatus .

Odontomachus dubius França, Fernandes & Lattke new species urn:lsid:zoobank.org:act:420AA277-B46F-498B-9BC4-0A6300CC9A73 ( Figures 19-B , 67 and 83 )

Type specimens: Holotype. COLOMBIA: Cauca, Popayán, Est. [Estación] La Rejoya , 2.5168 -76.5937, station, R. Feitosa col., [20-25.xi.2019], RMF1010, ICN107291 1☿, [ ICN-MHN] . Paratypes: Same data as holotype, 1☿ DZUP591907 [ DZUP] and HYM-INPA 034969 1☿ [ INPA] . VENEZUELA: Monagas, Pq. [Parque] Nc. [Nacional] El Guácharo , 1150m, 29.x.1986, Cancello & Brandão [col.], geral 1☿ MZSP91973 , 2☿ MZSP91974 , 2☿ MZSP91975 [ MZSP] .

Etymology: The specific epithet dubius (from the Latin word dubius = doubtful, dubious, uncertain) is a Latin singular masculine adjective in the nominative case and refers to the resemblance with O. chelifer which can cause doubts about its identity.

Diagnosis: (☿) Appendages dark brown slightly lighter than the body color. Rectangular vertex almost as long as wide striated longitudinally. Central disc of pronotum with transverse striae. Metasternum forming a pair of triangular lobes projected posteriorly almost long than wide. Petiole sessile, angulate node. Dorsum of first gaster tergum with fine transverse striae. (♀) Unknown. (♂) Unknown.

Worker

Holotype measurements: EL 0.6, HL 3.9, HoW 2.6, HvW 2, HW 2.2, ML 2.3, SL 3.7, PnW 1.5, WL 4.7, FL 4.4, PTH 1.6, PTL 0.9, PTW 0.55, GL 4.5, TL 16.3. CI 0.56, MI 0.58, PtI 1.77, SI 0.94, VI 0.90.

Head: Vertex rectangular to trapezoidal in frontal view, length larger than its half-width; with regular longitudinal striae diverging posteriorly. Posterior margin concave medially; occipital carina not forming nuchal collar. Cephalic constriction conspicuous. Frontal lobes striated. Ocular bridges posteriorly forming arc. Eye relatively large (EL> 0.4); completely rounded. Clypeus with wrinkle micro-sculpturing on median portion. Malar area in latero-oblique view, with fine regular transverse to oblique striae on anterior portion extending to occipital corners.

Mandible: Almost as long as anterior head portion. External margin straight to slightly convex. Masticatory margin with denticles that serially increase in size apically. Palp formula 4,3.

Antenna: Scape surpassing occipital corner by more than pedicel length.

Mesosoma: Pronotum in lateral view triangular, dorsal margin convex; in dorsal view, neck well developed anteriorly; lateral margin rounded with obtuse angle posteriorly; central disc of pronotum with regular transverse striae, lateral striae might form concentric ellipses curved anteriorly. Propleuron with fine oblique striae coarser on lateral portion. Mesonotum with regular transverse striae. Mesopleuron with rounded carina on anteroventral portion; mostly smooth and shining, anterior portion with regular transverse striae curved dorsal-anteriorly. Mesometanotal sulcus well-marked, scrobiculate. Metanotum with regular transverse striae; lateral margins in dorsal view slightly concave. Metapleural gland bulla smooth and shining at most with weakly fine striae on central portion. Propodeum in lateral view slightly concave on middle length; posterior margin length about 1/3 of dorsal length. Metasternum with metasternal process cuticle border expanded like lamellae laterally and forming bilobate triangular blunt projections; regular transverse striae anteriorly.

Petiole: Sessile. Angulate node, anterior margin in lateral view meeting dorsal face in an obtuse angle, anterior margin varying in length. Spine robust, slightly curved posteriorly. Oblique striae on lateral face extending posteriorly limited to ventral portion. Subpetiolar process longer than wider; tip rectangular to blunt; posterior portion not concave.

Gaster: First tergum longer than higher; anterior and dorsal margin meeting in a slightly convex surface not forming an angle. First and second terga dorsal face with regular transverse striae slightly curved anteriorly. Third and fourth terga slightly striate anteriorly, striae gradually fading posteriorly.

Color and size: Body dark brown, head slightly darker. Legs and antennae slightly lighter than body color. Medium to large size ants.

Pilosity: Scattered pubescence spread over the body separated each other by about its length. Relatively short hairs on anterior portion of procoxa smaller than coxae width; short hairs on ventral portion of gaster. Long suberect hairs on gastral terga directed posteriorly.

Natural history: The holotype nest series was found under a rock in an open field (Feitosa, 2023 personal communication). The paratypes from the Parque Nacional El Guácharo were collected in the soil next to the El Guácharo cave entrance, Colombia (Brandão, 2023 personal communication). Considering that lots of specimens of O. chelifer have their metasternum hidden by glue, some of them may be O. dubius sp. nov., and consequently, it has a wider distribution than observed ( Fig. 67 ).

Comments: Specimens of O. dubius sp nov. were found in collections identified as O. chelifer due to their similarities. The only stable diagnostic character for O. dubius sp. nov. that can separate it from O. chelifer is the metasternal process forming a pair of triangular lobes with a lamellate aspect laterally. A specimen from an unknown Colombian locality in the Borgmeier collection (MZUSP) was found with the metasternal lobes more separated than usual, but with the triangular lobes and lamellate aspect. Some specimens of O. chelifer were found in the Colombian Andes region, suggesting that both species might be sympatric in other areas.

Material examined: 62☿ COLOMBIA: [no further data] Casalma , Ap. Monte (1☿) . VENEZUELA: Cojedes: 10km W Chaparral (1☿). Miranda: Altos de Pipe (60☿) .

Odontomachus erythrocephalus Emery, 1889 ( Figures 24-A , 44-A , 58 , 84 , 85 and 86 )

Odontomachus haematodes r. erythrocephalus Emery, 1890: 44 (☿); Forel, 1914: 9 (♀); status as species by Dalla Torre, 1893: 50; subspecies of Odontomachus haematodus by Forel, 1899: 21; Wheeler & Wheeler, 1952: 650 (larvae); status as species by Kempf, 1972: 7, 170.

Type specimens: Syntype [Images]. [ COSTA RICA: Puntarenas], [Puerto] Jiménez [A. Alfaro col.], ANTWEB CASENT0904006 1☿ [ MSNG] .

Etymology: Combination of the Greek words erythros = red + cephalo = head; referring to the reddish head.

Diagnosis: (☿) Head red to yellowish contrasting to the body dark brown almost black. Petiole sessile, campanulate, with fine, irregular transverse striae, almost smooth and shining. Body color dark brown, almost black. (♀) Head and body color, petiole shape and striae as in worker. (♂) Body brown, propodeum and petiole dark brown almost black, gaster light brown. Mesoscutum with oblique fine rugae and punctures. Metascutellum with longitudinal keel. Petiole with fine rugae becoming ascendent posteriorly.

Worker

Syntype measurements: EL 0.41, HL 2.68, HoW 2.18, HvW 1.78, HW 1.97, ML 1.28, SL 2.59, PnW 1.3, WL 3.55, FL 1.92 (femur inclinate), PTH 1.29 (subpetiolar process covered), PTL 0.69, PtW 0.53, GL 2.82, TL 11.02. CI 0.73, MI 0.47, SI 0.96, VI 0.90.

Additional material examined: EL 0.4-0.5, HL 2.5-2.9, HoW 1.9-2.3, HvW 1.4-1.9, HW 1.6-2.1, ML 1.4-1.7, SL 2.4-2.8, PnW 1.1-1.3, WL 3.1-3.7, FL 2.6-3.0, PTH 1.2-1.4, PTL 0.6-0.7, PTW 0.4-0.5, GL 2.8-3.5, TL 10.7- 12.3. CI 0.64-0.75, MI 0.55-0.62, PtI 2.0-2.33, SI 0.92-1.0, VI 0.87-0.90 (n= 7).

Head. Vertex trapezoidal, wider than long in frontal view; with regular longitudinal striae diverging posteriorly. Posterior margin concave medially; occipital carina not forming nuchal collar. Cephalic constriction conspicuous. Frontal lobes striated. Ocular bridges posteriorly forming arc. Eye relatively large (EL> 0.4); rounded, posterior margin rounded. Clypeus with fine regular longitudinal striae. Malar area smooth and shining in latero-oblique view.

Mandible. Almost as long as anterior head portion. External margin convex. Masticatory margin with denticles that serially increase in size apically. Palp formula 4,3.

Antenna. Scape surpassing occipital corner about pedicel half-length or less.

Mesosoma. Pronotum in lateral view rounded, dorsal margin convex; neck well developed anteriorly in dorsal view; lateral margin rounded; central disc of pronotum with regular longitudinal striae that might form concentric triangles, lateral striae forming concentric circles. Propleuron with fine regular transverse striae effaced on central portion. Mesonotum longer than wide. Mesopleuron with rounded carina on anteroventral portion; mostly smooth and shining, anterior portion with regular transverse striae curved dorsal-anteriorly. Mesometanotal sulcus well-marked, scrobiculate. Metanotum as long as notopropodeal sulcus; wrinkle sculptured. Metapleural gland bulla with transverse striae effacing posteriorly. Propodeum in lateral view straight; posterior margin length about 1/3 of dorsal length. Metasternum with metasternal process open, cuticle border bilobate separated on middle line by longitudinal sulcus; smooth and shining with inconspicuous micro-sculpturing.

Petiole. Sessile. Campanulate node, anterior margin in lateral view usually more convex than posterior. Spine acute, straight or curved posteriorly. Fine, irregular transverse striae on anterior and lateral face, coarser on ventral portion, extending posteriorly limited to ventral portion. Subpetiolar process wider than long keel-shaped; tip acute; posterior portion concave.

Gaster. First tergum almost higher than long; anterior and dorsal margin meeting in an obtuse angle; dorsal face convex. dorsal face of all terga slightly imbricate, almost smooth and shining.

Color and size. Head red, yellowish or orangish. Body dark brown almost black. Legs and antennae yellowish. Medium small ants.

Pilosity. Scattered, conspicuous and short pubescence spread over the body; head and mandible pilosity dense separated each other about their length. Pubescence on mesosoma, petiole and gaster separated each other by more than its length. Pubescence on petiole limited to anterior face. Short hairs on anterior portion of procoxa smaller than coxae width. Pronotum with three or four pairs of long, curved erect hairs longer than maximum procoxa width. Long erect hairs on posterior 3/4 of gastral terga directed posteriorly, as long as pronotum hairs.

Gyne

Non-type measurements: EL 0.5, HL 2.7-2.8, HoW 2.2-2.3, HvW 1.8-1.9, HW 2.0-2.1, ML 1.5-1.6, SL 2.5, PnW 1.5, WL 3.3-3.7, FL 2.8-2.9, FWL 7.6, PTH 1.7, PTL 0.7, PtW 0.7, GL 4.4-5.1, TL 13.0-13.5. CI 0.74-0.75, MI 0.55-0.57, PtI 2.42, SI 0.89-0.92, VI 0.90 (n= 2).

Mesosoma. Pronotum slightly longer than high in lateral view, dorsal margin concave separating anterior from posterior portion; lateral margins converging anteriorly forming a neck in dorsal view; pronotal dorsum with regular transverse and curved striae forming arcs directed posteriorly on lateral face. Subalar area conspicuous with pronotum; with fine regular striae. Mesoscutum anterior and dorsal margin forming single convexity in lateral view; with regular longitudinal striae in dorsal view; striae between parapsidial line and parascutal area oblique, directed medially. Parapsidial line curved, well-marked. Parascutal area smooth and shining. Notopropodeal sulcus smooth and shining, with at most two vestigial carina that delimits three pits. Axilla with inconspicuous and fine longitudinal striae anteriorly, smooth and shining posteriorly. Mesoscutellum smooth and shining. Metascutellum smooth and shining with vestigial longitudinal carina.

Wing. Color pale yellow, translucid. Forewing ML1 slightly curved. RsL1 straight; as long as Rs+M. RSL2-3 slightly curved. ML2 slightly shorter than Rs+M. Rs+M and ML2 forming single slightly curved line, not forming angle with RsL2-3. Vein cu-a not reaching M+Cu ML1 angle almost by its half-length. Hindwing 1rs-m vein slightly curved. 8 hamuli.

Petiole. Conical node, anterior face straight to slightly concave, posterior face convex. Petiolar spine long, robust and curved posteriorly.

Pilosity. Pubescence on mesosoma dense separated each other by less than its length. Pubescence on mesoscutellum and metascutellum longer and less dense than pronotum pubescence, about half-length longer. Pronotum with two pairs of short hairs, shorter than procoxa width.

Male

Non-type measurements: EL 0.65, HL 1.0, OW 0.15, OES 0.24, IOW 0.27, WL 3.0, FWL 5.9, PTH 1.0, PTL 0.6, PTW 0.5, GL 2.1, TL 6.1. PtI 1.66 (n= 1).

Head. Maxillary palp equal 4. Labial palps broken. Lateral ocelli surpassing posterior margin in frontal view; posterior margin between lateral ocelli straight, not reaching median ocellus.

Mesosoma. Pronotal dorsum with inconspicuous micro sculpture, almost smooth and shining; with fine longitudinal rugae on ventral portion in lateral view. Propleuron with inconspicuous micro sculpture. Mesoscutum anteriorly with oblique rugulae; dorsal surface rugulose-punctate, with median longitudinal rugae, rugulae becoming oblique laterally. Axilla with fine longitudinal striae and punctuation with reticulate aspect. Mesoscutellum convex; with rugulae on anterior and lateral portion gradually becoming smooth and shining posteriorly, median longitudinal rugae coarser. Metascutellum with sparse effaced rugae, surface between them smooth and shining with longitudinal carina well-developed forming keel. Anepisternum anteriorly with oblique fine rugae, ventral portion imbricate gradually becoming smooth and shining dorsally. Katepisternum imbricate with fine longitudinal rugae on posteroventrally. Metapleuron densely rugose. Metepisternum with rugulae with dense micro sculpture between them. Metapleuron and propodeum separated by longitudinal sulcus scrobiculate. Propodeal dorsum with coarse rugae; dorsal and posterior face delimited by coarser rugae; posterior face with coarse rugae.

Wing. Translucid. Hamuli varying from 8 or 9.

Petiole. Sessile; dorsal portion acute; with rugulae on ventral portion becoming oblique, coarser and sparser posteriorly, surface between them punctate. Subpetiolar process acute and curved posteriorly. Petiolar sternum concave posteriorly.

Color and size. Head, prothorax, mesothorax legs and antenna brown. Propodeum and petiole dark brown almost black. Gaster light brown. Relatively small size.

Pilosity. Head, mesosoma, petiole and gaster with dense appressed pubescence. Pronotum with two pairs of thin short erect hairs. Thin and suberect short hairs on posterior portion of gastral terga.

Natural history: Little is known about this species that occurs from Andean regions at Ecuador, Colombia, and Venezuela (Lattke pers. obs.) to Central America ( Fig. 58 ). The mandible closing movement can reach velocities of 64m / s- 1 ( Spagna et al. 2008) and it has been observed carrying seeds of Siparuna aspera (Ruiz & Pav.) A. DC. ( Siparunaceae ) and Passiflora ligularis A. Juss. ( Passifloraceae ) in treeless pasture areas ( Escobar et al. 2007). Three pyrazines were found in its mandibular gland extractions ( Xu et al. 2018). It can be found in undisturbed areas, city parks, and even gardens in suburban areas. Nests can be found in accumulations of leaf litter at the base of trees or under rotten logs or within them. One nest found within a rotten log in the Venezuelan state of Táchira contained hundreds of workers that aggressively poured out of the nest upon being uncovered (Lattke, pers. obs.)

Comments: Among haematodus group species, O. erythrocephalus stands out because of the red to yellowish head that contrasts with the dark brown almost black body. Not one specimen was found misidentified in collections, indicating the usefulness of this diagnostic character. The campanulate node with coarse striae closely resembles the O. bauri petiole. Even so, the head color is enough to recognize O. erythrocephalus . The main variations are worker size, which varies from relatively small to medium size, and the pronotal striae. As in O. bauri , the pronotal striae might be longitudinal with concentric arcs or form concentric triangles. The hamuli numbers were different on the wings of the examined male, with eight hamuli on the left hind wing and nine on the right.

Material examined: 46☿, 7♀, 1♂. COLOMBIA: Antioquia: Cornoná (2☿) ; Medellín (7☿. 1♀). Cauca: Calibio (1♀) ; Cerro las Tres Cruces (4☿. 2♀). [Cundinamarca:] Sasaima (2☿) . [Valle del Cauca:] Cali (3☿. 1♀) . COSTA RICA: Alajuela: 10km N Quesada (2☿) ; Río Toro Amarillo. vicinity of Guapiles (2☿) . Cartago: La Fuente (2☿) ; Turrialba (2☿). Heredia: Estación Biológica La Selva (2☿) ; [no further data] (2☿. 1♀) . Limón: Bristol (1☿. 1♀) . ECUADOR: Pichincha: Puerto Quito (1☿) . PANAMA: Cocle: El Valle de Antón (5☿. 1♂) . Panama: Parque Nacional Altos de Campana (1☿) ; [Parque Nacional Soberanía] (1☿) . Panama Oeste: Barro Colorado Island (6☿) ; Parque Nacional Altos de Campana (1☿) .

Odontomachus haematodus ( Linnaeus, 1758) ( Figures 21-B , 26-A , 31-B , 35-B , 39-B , 46-A , 87 , 88 , 89 and 90 )

Formica haematoda ( Linnaeus, 1758) : 582 (♀); combination in Myrmecia by Fabricius, 1804: 425; combination in Odontomachus by Latreille, 1804: 179; Mayr, 1867: 47 (☿); André, 1887: 290 (♂); Emery, 1899: 5 (larvae).

Formica maxillosa Geer, 1773: 601 (♀); junior synonym of Formica haematoda by Retzius, 1783: 75.

Odontomachus hirsutiusculus Smith, 1858: 78 (☿); junior synonym of Odontomachus haematodus by Roger, 1861: 24; subspecies of Odontomachus insularis by Dalla Torre, 1893: 51; subspecies of Odontomachus haematodus by Forel, 1893a: 354; Forel, 1909: 252 (♀); junior synonym of Odontomachus haematodus by Brown, 1976: 104.

Odontomachus haematoda var. brunneipes Menozzi, 1935: 191 (☿); junior synonym of Odontomachus haematodus by Brown, 1976: 104.

Odontomachus haematoda var. pallipes Crawley, 1916: 368 (☿); junior synonym of Odontomachus haematodus by Brown, 1976: 104.

Type specimens: Syntype [ Images ]. Odontomachus haematoda var. brunneipes Menozzi, 1935 ; [ Guyana: Bambooncamp , Haimaracamp e Canister falls], [Beccari, N. Gigliolli, G. R. & Ignesti, U. cols.], Pav. N. 11, ANTWEB CASENT0904007 1☿ [ MSNG] .

Holotype [ Images ]. Odontomachus haematoda var. pallipes Crawley, 1916 ; BRITISH GUIANA [ GUYANA]: Onderneeming,G.E.Bodkin[col.],1916-68, 20.4.15 [20.iv.1915], BMNH (E)1013976,ANTWEB CASENT0900582 1☿ [ BMNH].

Syntype [Images]. Odontomachus hirsutiusculus Smith, 1858 ; BRAZIL: [Pará], Santarém, [H. W. Bates col.], Smith, 54 63, BMNH (E) 1013975, CASENT0900581 1☿ [ BMNH] .

Etymology: From Greek word haematodes = bloody, blood-red. In the original description Linnaeus mentions only the mandible being red (maxillis porrectis rubris), hence, the name probably refers to this character. An alternative interpretation is the combination of the haemato + Greek verb odax = biting, probably referring to the unique mandible shape of Odontomachus that could cause bleeding.

Diagnosis: (☿) Metasternal process forming a pair of acute spines. Petiole sessile, conical with acute spine; petiolar striae fine, irregular and inconspicuous. Gaster pubescence usually very dense, separated from each other by less than 1/3 of its length. (♀) metasternal process, petiolar striae and gaster pubescence as in worker. (♂) Mesosoma with light brown spots on parapsidial lines. Propodeum dark brown with coarse rugae. Metasternal process as in worker.

Worker

Non-type measurements: EL 0.30-0.50, HL 1.50-2.80, HoW 1.50-2.70, HvW 1.20-1.60, HW 1.40-2.00, ML 1.20- 1.60, SL 2.00-2.70, PnW 0.80-1.20, WL 2.70-3.40, FL 2.20-2.90, PTH 0.90-1.40, PTL 0.50-0.60, PTW 0.30-0.50, GL 2.20-3.60, TL 8.50-11.80. CI 0.61-0.73, MI 0.53-0.63, PtI 1.80-2.80, SI 0.92-1.05, VI 0.80-0.88 (n= 30).

Head. Vertex trapezoidal, wider than long in frontal view; with regular longitudinal striae. Posterior margin concave medially; occipital carina not forming nuchal collar. Cephalic constriction conspicuous. Frontal lobes striated. Ocular bridges posteriorly forming arc. Eye relatively large (EL> 0.4); rounded, posterior margin rounded. Clypeus smooth and shining with inconspicuous piligerous punctuation on median portion. Malar area with inconspicuous irregular striae in latero-oblique view, posterior portion smooth and shining.

Mandible. Almost as long as anterior head portion. External margin slightly convex. Masticatory margin with denticles that serially increase in size apically. Palp formula 4,3.

Antenna. Scape long, surpassing occipital corner about pedicel half-length or less.

Mesosoma. Pronotum in lateral view rounded, dorsal margin convex; neck well developed anteriorly in dorsal view; lateral margin rounded; central disc of pronotum with regular longitudinal striae forming different shapes, lateral striae forming concentric circles or arcs directed anteriorly. Propleuron smooth and shining; regular fine oblique striae restricted to anterior portion gradually fading posteriorly. Mesonotum larger than wide; with regular transverse striae slightly curved anteriorly. Mesopleuron with rounded carina on anteroventral portion; mostly smooth and shining, anterior portion with regular transverse striae curved dorsal-anteriorly. Mesometanotal sulcus well-marked, scrobiculate. Metanotum vestigial. Metapleural gland bulla with transverse striae effacing posteriorly. Propodeum in lateral view straight; posterior margin length about 1/3 of dorsal length. Metasternum with metasternal process close, cuticle border forming pair of acute spines directed ventrally; inconspicuous sculpturing, apparently smooth and shining.

Petiole. Sessile. Conical node, anterior margin in lateral view usually continuous with petiolar spine. Spine acute, straight or curved posteriorly. Fine, irregular and transverse striae limited to ventral portion, not extending to posterior face. Subpetiolar process wider than long keel-shaped; tip acute; posterior portion concave.

Gaster. First tergum almost higher than long; anterior and dorsal margin meeting in an obtuse angle; dorsal face convex. dorsal face of all terga imbricate.

Color and size. Body dark brown almost black to light brown. Legs and antennae brown to yellowish. Relatively medium size ants.

Pilosity. Hook-shaped, scattered, conspicuous and short pubescence spread over the body, separated each other about their length. Short hairs on anterior portion of procoxa smaller than coxae width. Pronotum with three or four pairs of long, curved erect hairs longer than maximum procoxa width. Long erect hairs on posterior 2/3 of gaster tergum directed posteriorly, as long as pronotum hairs.

Gyne

Non-type measurements: EL 0.40-0.60, HL 2.20-2.60, HoW 1.80-2.30, HvW 1.40-1.70, HW 1.60-1.90, ML 1.20- 1.50, SL 2.10-2.60, PnW 1.10-1.40, WL 3.10-3.60, FL 2.10-2.70, FWL 6.50-7.50, PTH 1.20-1.60, PTL 0.50-0.70, PTW 0.50-0.70, GL 3.00-4.20, TL 9.90-12.30. CI 0.72-0.75, MI 0.5-0.63, PtI 2.0-2.6, SI 0.87-1.0, VI 0.84-0.89 (n= 11).

Mesosoma. Pronotum slightly longer than high in lateral view, dorsal margin slightly concave; in dorsal view, lateral margins constricted anteriorly and parallels forming a neck; with regular transverse striae forming arcs directed posteriorly on lateral face. Subalar area conspicuous with pronotum. Mesoscutum anterior and dorsal margin forming single convexity in lateral view; with median sulcus anteriorly in dorsal view; regular longitudinal striae; striae between parapsidial line and parascutal area oblique, directed medially. Parapsidial line curved, well-marked. Parascutal area smooth and shining. Notopropodeal sulcus scrobiculate. Axilla smooth and shining posteriorly. Mesoscutellum smooth and shining. Metascutellum smooth and shining with longitudinal carina.

Wing. Color varying from light and pale brown translucid to whitish translucid with iridescent shining. Forewing ML1 slightly curved. RsL1 straight to slightly curved; as long as Rs+M. RSL2-3 slightly curved. ML2 short, about half-length of Rs+M. Rs+M and ML2 forming single slightly curved line, not forming angle with RsL2-3. Vein cu-a not reaching M+Cu ML1 angle by less than its half-length. Hindwing 1rs-m vein slightly curved. 7 to 10 hamuli.

Petiole. Conical node, anterior face straight to slightly convex.

Pilosity. Pubescence on mesosoma strongly curved, dense separated each other less than its length. Pubescence on mesoscutellum and metascutellum longer and denser than pronotum pubescence, about half-length longer. Pronotum with three or four pairs of short hairs, shorter than pronotum height.

Male

Non-type measurements: EL 0.65-0.68, HL 0.90-1.00, OW 0.15-0.18, OES 0.21-0.27, IOW 0.24-0.27, WL 2.60- 2.70, FWL 5.20-5.40, PTH 0.80-1.10, PTL 0.50, PTW 0.40-0.50, GL 1.90-3.00, TL 5.60-6.50. PtI 1.60-2.20 (n= 5).

Head. Palp formula 4,3. Lateral ocelli partially to completely surpassing posterior margin; posterior margin between lateral ocelli concave, not reaching median ocellus.

Mesosoma. Pronotal dorsum with inconspicuous micro sculpture of striate aspect; with oblique fine irregular striae, slightly coarser on posterior portion in lateral view. Propleuron with micro sculpture. Mesoscutum anteriorly with fine transverse striae, longitudinal sulcus standing out from striae; dorsal surface with oblique striae diverging posteriorly towards parapsidial lines or mesoscutum completely micro sculptured with reticulate aspect. Axilla, mesoscutellum, metascutellum and mesopleuron with dense micro sculpture of reticulate aspect. Mesoscutellum convex.Metascutellum with longitudinal carina. Mesopleuron with micro sculpture. Metapleuron and metepisternum with rugulae and dense micro sculpture between them. Metapleuron and propodeum separated by longitudinal sulcus scrobiculate anterior to spiracle. Propodeal dorsum with coarse rugae; dorsal and posterior face delimited by coarser rugae; posterior face with coarse rugae or rugulae.

Wing. Whitish translucid. Hamuli varying from 7 to 9. Pterostigma nut brown contrasting to wing surface.

Petiole. Sessile; dorsal portion acute; with micro sculpture.

Color and size. Head, prothorax, mesothorax and legs and antenna pale yellow. Light brown spots on ocelli region and parapsidial lines. Propodeum dark brown. Petiole brown. Gaster light brown, terga with posterior portion yellowish. Relatively small size.

Pilosity. Head, mesosoma, petiole and gaster with dense appressed pubescence. Pronotum with two thin erect hairs on posterior portion. Thin and erect short hairs on posterior portion of gaster segments I and II, hairs on segments III, IV and V amost two times larger.

Natural history: It is an opportunistic species nesting from the soil strata to epiphytes ( Blüthgen et al. 2000; Gibernau et al. 2007), abandoned ant gardens ( Orivel et al. 1998), and even in houses between cracks in the wall or under plant pots with flowers ( Delabie et al. 1995). The nests found in Aechmea aquilega (Salisb.) Griseb. ( Bromeliaceae ) can be polydomous and built between the leaves, sprouts, or under the tree bark that hosts the epiphyte ( Talaga et al. 2015). The colonies are monogynous and might number about 200-500 individuals ( O’Fallon et al. 2016). However, considering its opportunistic nesting habit, the colonies might be facultative polygynous.

There are observations of O. haematodus preying on semi-aquatic tadpoles Thoropa taophora (Miranda-Ribeiro, 1923) ( Cycloramphidae ). A single snap of the jaws is enough to subdue the tadpole that tries to escape by throwing itself from the rocks where it lives into the water next to it ( Facure & Giaretta, 2009).

Cuticular chemical profiles are described by Smith et al. (2016) studying colonies from Alabama, U.S.A. Touchard et al. (2015) found 150 linear short peptides in the venom of O. haematodus . The authors also observed that the peptide composition does not vary among workers from the same colony and the farther the colonies are, the larger is the variation in the composition of the peptides.

Odontomachus haematodus is widely distributed in the Neotropical Region ( Fig. 90 ) and considered exotic in the Nearctic ( Macgown et al. 2014), African, Indomalaysia, and Australasia regions (see comments section). According to Brown (1976), it is more frequent in forested areas than in dry areas like the Brazilian Cerrado. But we also studied several specimens from these areas and large cities like Rio de Janeiro and São Paulo based on older museum specimens.

Comments: Odontomachus haematodus can be easily misidentified with other widely distributed species (e.g., O. bauri , O. cupreus sp. nov., and O. chicomendesi sp. nov.) due to the imbricate gastral terga. The petiole of O. haematodus has the anterior face straight and slightly inclined posteriorly with fine transverse striae that are continuous to the petiolar spine. Some specimens of O. bauri also display this petiolar morphology, especially in the northeast of Brazil. However, the striae of O. bauri are always coarser than those of O. haematodus , especially on the ventral portion of the lateral face and the posterior face. Brown (1976) mentions that the metasternum of O. haematodus with its bispinous process is diagnostic and contrasts with the bilobate metasternal process of O. bauri . Even though it may be difficult to observe, especially in badly mounted specimens, this feature is reliable and stable in all of the specimens examined. The size and body color of both species are very variable and cannot be used for identification. Comparing O. haematodus mesopleuron without projections to the mesopleural acute projection present in O. chicomendesi sp. nov. is enough to separate them. Odontomachus cupreus sp. nov. can be recognized by its coarse striate-punctate node and reddish body color. However, as in O. bauri , body color alone is not diagnostic since some small individuals of young colonies of O. haematodus also can have brown reddish body color. All these features also can be applied to identifying the gynes. Males are also very similar to those of O. bauri due to the bicolored yellowish and brown body, and the metasternal process forming a pair of spines.

Workers and gynes display a wide variation of body color from light brown with yellowish appendages to dark brown almost black with light brown or ferruginous legs. The gastral terga can be strongly imbricate, with a coriaceous aspect to weakly imbricate and almost smooth and shining. There is no geographical pattern to this variation but smaller specimens tend to be lighter and have weakly imbrication sculpture on the gastral terga.

According to Macgown et al. 2014, O. haematodus is an exotic species in the Southeast of the United States of America and it was observed occurring in Alabama by O’Fallon et al. (2016). Macgown et al. (2014) stated that the lack of records of this species on the Nearctic region corroborates their hypothesis. Wheeler (1900) describes the larvae of O. haematodus with specimens collected in Texas, but probably, these specimens were misidentified. Also, the name Odontomachus haematodus was used several times to describe subspecies and variations, and some of them are currently valid as species ( Brown, 1976). These facts associated with the lack of records in collections corroborate the hypothesis of O. haematodus being an exotic species in the Nearctic region.

The easternmost record of O. haematodus in the Neotropical region is from Trindade Island ( Fig. 90 ), located 1,116 Km east from Brazil’s coast (IBGE, 2021). Their opportunistic biology and the commercial activities can explain this wide distribution, including the Nearctic invasion. According to the AntMaps records, this species also is exotic in Africa, Indo-Malaysia, and Australasia. These records are probably misidentifications of other species with a sessile and conical petiolar node. In Africa, the records are probably of Odontomachus troglodytes Santschi, 1914 . Both species have slender and acute petiolar spine and the metasternal process forms a pair of spines ( Fisher & Smith, 2008). According to Brown (1976), O. troglodytes has a shorter scape length, and coarser petiolar striation when compared to O. haematodus ; and the males are entirely yellowish. Even so, he mentions that some specimens are difficult to separate. The Indo-Malaysian records probably would be specimens of Odontomachus monticola Emery, 1892 , whose metasternal process resembles the pair of spines of O. haematodus ( Yoshimura et al. 2007) . Odontomachus monticola can be recognized by the pronotum striae forming concentric circles and the almost cylindrical petiolar node with straight anterior and posterior faces that slightly converge apically. Also, its palp formula is 4,4. Lastly, in the Australasia region, the species that most resembles O. haematodus is Odontomachus simillimus Smith, 1858 . According to Brown (1976) the striation of workers and queens are very similar, but the metasternal process is low and round.

Material examined: 255☿, 24♀ and 31♂. ARGENTINA : Tucumán: Racó (2☿) . BOLIVIA: [Cochabamba]: Chimoré (1☿) . BRAZIL: Acre: Mâncio Lima (2☿) ; Rio Branco (2☿) ; Tarauacá (1☿) ; [Porto Acre] (1☿) . Amapá: Cutias (2☿) ; Serra do Navio (1☿) . Amazonas : [Borba] (1☿) ; Coari (1☿) ; Humaitá (1☿) ; Itacoatiara (1☿) ; Jari (9☿, 1♀) ; Manacapuru (3☿) ; Manaus (11☿, 3♂) ; Novo Airão (1☿) ; Presidente Figueiredo (2☿, 1♀) ; Tapauá (1☿) . Bahia: Camacã (1☿) ; Guarupá-Mirim (3☿) ; Ilhéus (5☿) ; Itacaré (2☿) ; Itaquara (1☿) ; Porto Seguro (2☿) ; Travessão (1☿) ; [no further data] (1☿) . Ceará: Guaramiranga (2☿) ; Ubajara (1☿) . Espírito Santo: Anchieta (1☿) ; Cachoeiras de Itapemirim (1☿) ; FLONA do Rio Preto (1♀) ; Guaçuí (1☿, 1♀) ; Guiriri (1☿) ; Ilha da Trindade (1☿) ; Linhares (3☿) ; Pinheiros (1☿) ; Santa Teresa (1☿) ; São Mateus (1☿) ; Soorotema (2☿, 1♀) . Goiás: Niquelândia (4☿) . Maranhão: Igarapé Gurupi-Uma , Aldeia Araçu 50km E de Canindé (3☿) . Itapecuru-Mirim (1☿) . Mato Grosso: Chapada dos Guimarães (4☿) ; Poxoréu (4☿) ; Santo Antônio do Leverger (1☿) . Mato Grosso do Sul: Aquidauana (1☿) ; Bonito (2☿) ; Corumbá (18☿, 1♀) ; Cuiabá (1♀) ; Jardim (1☿) ; Terenos (2☿) . Minas Gerais: Boa Esperança (3☿) ; Bom Despacho (1☿) ; Itutinga (2☿) ; Nanuque (4☿) ; Parque Estadual do Rio Doce (1☿) ; Viçosa (2☿) . Pará : Belém (5☿) ; Belterra (2☿) ; Benevides (2☿) ; [ Castanhal ] (1☿) ; Marabá (3☿) ; Oriximiná (2☿, 2♀) ; Paragominas (6☿, 1♀) ; Parauapebas (1☿) ; [ Santarém ] (5☿) ; Santo Antônio do Tauá (2☿) ; Terra Santa (4☿) ; [ Without location] (3☿) . Paraíba: João Pessoa (2☿) . Paraná: Antonina (3☿, 26♂) ; Jaguariaiva (7☿, 3♀) ; Tibagi (5☿) ; Parque Estadual Palmito , 2.8km SE Paranaguá (3☿, 1♀) . Pernambuco: Ilha de Itamaracá (2☿) ; [ Moreno ] (1☿) ; Recife (2☿, 1♀) . Piauí: BR 316, Km 371 (3☿) ; Rio Urucuí [Uruçuí]- Preto (1☿) ; São João do Piauí (1☿, 1♂) . Rio de Janeiro: Belford-Roxo (1☿, 1♀) ; Itatiaia (1☿) ; Rio de Janeiro (2☿) ; Quissamã (1☿) ; Seropédica (1☿) . Rondônia: Cacoal (2☿) ; Guajará-Mirim (1☿, 1♀) ; Ouro Preto do Oeste (2☿) ; Porto Velho (3☿) . Roraima: Caracaraí (5☿, 2♀) . Sergipe: Areia Branca (1☿, 1♀) ; São Cristóvão (1☿) ; Parque Nacional do Itabaiana (1☿) . São Paulo: Sete Barras (1☿) . Tocantins: Araguacema (7☿) ; Araguaína (1☿) . ECUADOR: [Morona-Santiago]: Los Tayon [Tayos] (1☿) . Napo: Limoncocha (3☿, 1♂) . FRENCH GUIANA: Cayenne: Roura: Montagne dês Chevaux (3☿) . Saint-Laurent-du-Maroni: Saül: Bélvédére de Saül (2☿) . GUIANA: [Cuyuni-Mazaruni]: Kartabo (2☿) . [ Without location] Courantyne R [river] (1☿) . PERU: Cusco: Estación Biológica Villa Carmen (1☿) ; Kosñipata (7☿) . Madre de Dios: Puerto Maldonado (3☿) . SURINAME: Maroujine : (3☿, 1♀) . [Saramacca]: La Poulle [Poule] (9☿, 2♀) . Dirkshoope (4☿, 1♀) . Vank (7☿) . UNITED STATES: West Indies [Saint Thomas Islands]: [no further data] (1☿) .

Odontomachus hastatus ( Fabricius, 1804) ( Figures 8-A , 9-A , 10-A , 38-A , 80 , 91 , 92 and 93 )

Myrmecia hastata Fabricius, 1804: 426 (☿); combination in Odontomachus by Illiger, 1807: 194. Wheeler & Wheeler 1971: 1212 (larvae).

Odontomachus maxillaris Smith, 1858: 77 (☿ and ♀); junior synonym of Odontomachus hastatus by ( Roger, 1863): 22; status as species by Mayr, 1863: 437; junior synonym of Odontomachus hastatus by Dalla Torre, 1893: 51.

Type specimens: Syntype [Images]. Myrmecia hastata Fabricius, 1804 ; AMERICA MERIDIONALE [ SOUTH AMERICA]: Rio de Janeiro, Coll. Roger, 19040, [Dom. Smidt col?], GBIF-D/FoCol 1059, FOCOL1059 1☿ [ ZMHB] .

Syntype [Images]. Odontomachus maxillaris Smith, 1858 ; [ BRAZIL:Amazonas]. Tefé [?]. Ega, 56 69, BMNH (E) 1013978, CASENT0900583 1♀ [ BMNH] .

Etymology: From the Latin word hasta = spear; probably referring to the slender body, and the pedunculate petiolar node with acute spine, resembling to a spear.

Diagnosis: (☿) Rectangular vertex narrow posteriorly, smooth and shining. Central disc of pronotum smooth and shining or with inconspicuous transverse striae. Petiole pedunculate. Dorsum of gastral terga smooth and shining. Body covered with long sparse pubescence with length about half-length of hairs on gaster. (♀) Vertex, petiole, gaster and pubescence as in worker. (♂) Body bicolored, yellowish-brown areas contrasting with dark brown. Mesoscutum with transverse to oblique rugae, with light brown “Y shaped” area contrasting with pale yellow background. Petiole pedunculate.

Worker

Syntype measurements: EL 0.45, HL 2.98, HoW 1.95, HvW 0.93, HW 1.47, ML 1.74, SL 3.35, PnW 1.16, WL 4.59, FL 3.20, PTH 1.59, PTL 1.04, PTW 0.49, GL 3.68, TL 14.03. CI 0.49, MI 0.58, PtI 1.52, SI 1.12, VI 0.63.

Additional material examined: EL 0.40-0.60, HL 2.80-3.60, HoW 1.90-2.40, HvW 1.00-1.50, HW 1.20-1.80, ML 2.00-2.50, SL 3.10-4.20, PnW 1.10-1.40, WL 4.40-5.60, FL 3.8-4.9, PTH 1.20-2.00, PTL 1.00-1.40, PTW

0.50-0.60, GL 3.00-4.30, TL 13.60-17.20. CI 0.36-0.56, MI 0.63-0.78, PtI 1.09-1.72, SI 0.96-1.31, VI 0.66-1.25 (n= 30).

Head. Vertex rectangular narrow posteriorly in frontal view, posterior margin about half-width of anterior wide; smooth and shining, regular longitudinal striae diverging posteriorly limited to frontal area. Posterior margin straight, slightly concave medially; occipital carina posteriorly forming nuchal collar. Cephalic constriction not well-marked. Frontal lobes with inconspicuous striae. Ocular bridges posteriorly forming angle. Eye relatively large (EL> 0.4); rounded, posterior margin rounded. Clypeus smooth and shining. Malar area smooth and shining in latero-oblique view.

Mandible. As long as anterior head portion or more. External margin straight. Masticatory margin with acute denticles that serially increase in size apically. Apical teeth long and acutes. Palp formula 4,4.

Antenna. Scape surpassing occipital corner by pedicel and first flagellomere length.

Mesosoma. Pronotum in lateral view triangular, dorsal margin straight to slightly convex; lateral margins straight and converging anteriorly in dorsal view, cervical area continuous to the pronotum, with obtuse angle posteriorly; central disc of pronotum smooth and shining, lateral striae regular and transverse. Propleuron smooth and shining. Mesonotum longer than wide; with regular transverse striae. Mesothorax constricted in lateral view. Mesopleuron without rounded carina on anteroventral portion; mostly smooth and shining, anterior portion with regular transverse striae curved dorsal-anteriorly. Mesometanotal sulcus vestigial, smooth and shining. Metanotum with regular or irregular transverse striae; posteriorly constricted on laterals around metanotal spiracles. Metapleural gland bulla with inconspicuous transverse striae finer than metapleural striae. Propodeum in lateral view straight; posterior margin length about 1/4 of dorsal length. Metasternum with metasternal process close, cuticle border forming an arch with shallow notch medially sometimes slightly elevate forming two low blunt projections; with transverse striae.

Petiole. Pedunculate. Conical node, anterior margin in lateral view continuous with petiolar spine acute and straight. Oblique to longitudinal inconspicuous striae on lateral face limited to ventral portion. Subpetiolar process longer than wider; tip acute; posterior portion not concave.

Gaster. First tergum longer than high; anterior and dorsal margin forming a single convex surface. dorsal face of all terga smooth and shining.

Color and size. Mesosoma reddish brown to ferruginous, head and gaster lighter and orangish. Legs and antennae yellowish. Large and slender ants.

Pilosity. Scattered pubescence spread over the head and mesosoma. Head pubescence short, length about scape width, subdecumbent directed anteriorly. Mesosoma pubescence, except propodeum, erect as long or more than head pubescence. Propodeum pubescence dense, curved and decumbent. Gastral pubescence dense. Hairs on gaster thin, erect and dense mixed with pubescence.

Gyne

Non-type measurements: EL 0.6, HL 3.2-4.0, HoW 2.4-2.8, HvW 1.3-.17, HW 1.5-2.2, ML 2.2-2.6, SL 3.6-4.4, PnW 1.1-1.8, WL 4.9-6.1, FL 4.0-5.0, FWL 9.0-9.6, PTH 1.9-2.4, PTL 1.2-1.4, PtW 0.7-0.8, GL 4.2-5.3, TL 16.3- 19.3. CI 0.44-0.56, MI 0.64-0.75, PtI 1.42-1.71, SI 1,08-1,18, VI 0.76-0.86 (n= 5).

Mesosoma. Pronotum longer than high in lateral view, dorsal margin straight; lateral margins straight and converging anteriorly in dorsal view, cervical area continuous to pronotum; pronotal dorsum with regular transverse striae effacing on lateral posterior portion. Subalar area separated from pronotum by sulcus slightly impressed; with effaced striae. Mesoscutum anterior and dorsal margins meeting in convex surface, form an almost rounded right angle; longer than wider in dorsal view; with regular longitudinal striae; striae between parapsidial line and parascutal area oblique, directed medially. Parapsidial line curved, well-marked. Parascutal area smooth and shining. Notopropodeal sulcus as striated as mesoscutum.Axilla with regular longitudinal striae. Mesoscutellum with regular longitudinal striae anteriorly. Metascutellum smooth and shining.

Wing. Color pale brown translucid. Forewing ML1 curved. RsL1 slightly curved; as long as Rs+M. RSL2- 3 sinuous. ML2 as long as Rs+M. RS+M and ML2 meeting with RSL 2-3 in an obtuse angle giving discal cell pentagonal shape. Vein cu-a not reaching M+Cu ML1 angle by less than 1/3 of its length. Hindwing 1rs-m vein slightly curved. 12 or 13 hamuli.

Petiole. As in worker.

Pilosity. Pubescence on mesosoma as long as first flagellomere, dense separated each other less than its length.

Male

Non-type measurements: EL 0.77-0.89, HL 1.08-1.10, OW 0.18-0.24, OES 0.24-0.27, IOW 0.27-0.31, WL 3.40- 3.70, FWL 6.30-6.60, PTH 0.80-1.10, PTL 0.90, PTW 0.50, GL 2.80-3.00, TL 7.38-7.8. PtI 0.88-1.22 (n= 3).

Head. Palp formula 5,3. Lateral ocelli surpassing posterior margin; posterior margin between lateral ocelli concave, reaching median ocellus.

Mesosoma. Pronotal dorsum with irregular coarse rugae next to anterior margin and fine irregular transverse striae posteriorly; with oblique to longitudinal irregular striae with central portion effaced in lateral view. Propleuron with effaced micro sculpture. Mesoscutum dorsal surface with transverse striae anteriorly gradually becoming rugulae diverging latero-posteriorly, coarser irregular longitudinal rugae on median line, discontinuous anteriorly. Axilla and mesoscutellum with irregular longitudinal striae gradually becoming regular posteriorly. Mesoscutellum convex. Metascutellum with inconspicuous irregular oblique striae converging posteriorly, longitudinal carina on anterior portion. Anepisternum and katepisternum with band of micro sculptured area. Metapleuron with coarse transverse irregular striae. Metepisternum with coarse striae on dorsal portion effacing ventrally. Metapleuron and propodeum separated by longitudinal shallow sulcus extending from metepisternum to propodeal spiracle, and by sharp irregular longitudinal carina posterior to it. Propodeal dorsum with irregular longitudinal striae sparse on anterior portion; dorsal and posterior face not delimited by coarser striae or rugae; posterior face with transverse to oblique regular striae extending to lateral faces and metapleuron.

Wing. Light brown translucid. Hamuli varying from 10 to 12. Pterostigma brown to nut brown, contrasting with wing surface.

Petiole. Pedunculate; dorsal portion acute; with irregular longitudinal striae on ventral portion becoming smooth and shining dorsally. Subpetiolar process acute. Sternum not concave posteriorly.

Color and size. Head light brown with ocelli area brown. Prothorax and mesothorax brown; mesoscutum with light brown spots on median anterior portion and parapsidial lines area. Propodeum and petiole dark brown. Gaster light brown. Relatively large size.

Pilosity. Head, mesosoma and petiole with dense decumbent pubescence. Dorsal portion of petiole and gaster with shorter and less dense pubescence. Pronotum with series of thin erect hairs on posterior portion, one pair larger than the others. Thin and erect short hairs on posterior portion of gaster segments I and II, hairs on segments III, IV and V almost two times longer.

Natural history: Odontomachus hastatus is an arboreal species with nests built under epiphytal agglomerates or their roots. There is a record of ant gardens and their nests in epiphytes ( Table 1) and nests built in the litter formed in young palm leaf axilla ( Gibernau et al. 2007). The nests under bromeliad roots occur from 1.5 to 4.6 height meters, and the colonies display facultative polygyny varying from 35 to 536 individuals (291.2 ± 163) ( Camargo & Oliveira, 2012). Up to 12 queens have been observed in a single colony and the number of queens is related to the size of the root mass where the nest is built ( Oliveira et al. 2011). Also, worker numbers are positively related to the queen number.

Oliveira et al. (2011) describe the queen dominance and the agonistic behavior in polygynous colonies. In the monogynous colonies, both the queen and workers interact similarly and workers offer trophic eggs to the queen. Worker foraging starts at about 17:30h with the highest activity close to 20:00h and finishing between 6:00h to 8:00h ( Camargo & Oliveira, 2012)

Odontomachus hastatus is a keystone species in epiphytic ecosystems. It can exert top-down effects on the detritivore aquatic and terrestrial communities associated with the bromeliads by decreasing their density and changing their composition ( Gonçalves et al. 2017). It also helps plant growth where their nests are built by improving nitrogen availability as a consequence of their predation activities ( Gonçalves et al. 2017). This species is a host of Agamomermir costaricensis Poinar et al. 2006 (Nematoda: Mermithidae ) and there is a record of parasitism by a species of Kapala Cameron, 1884 ( Hymenoptera : Eucharitidae ) ( Poinar et al. 2006). According to Santos et al. 2010, its karyotype is 2n= 44 (4SM+2ST+38T).

Family Genus/Species Reference Anthurium gracile (Rudge) Lindl. *

Philodendron insigne Schott

Araceae Gibernau et al. 2017 Philodendron linnaei Kunth

Philodendron squamiferum Poepp.

Aechmea mertensii (G. Mey.) Schult. & Schult. f. *

Cyclanthus sp. Gibernau et al. 2017 Gusmania sp.

Bromeliaceae

Quesnelia arvensis (Vell.) Mez

Camargo & Oliveira, 2012 Vriesea procera (Mart. Ex Schult. & Schult. f.) Wittm.

Vriesea splendens (Brongn.) Lem. Gibernau et al. 2017 Clusiaceae Clusia sp. Gibernau et al. 2017 Codonanthe calcarata (Miq.) Hanst.

Gesneriaceae Gibernau et al. 2017 Codonanthe crassifolia (Focke) C. V. Morton *

Piperaceae Peperomia macrostachya (Vahl) A. Dietr. Gibernau et al. 2017

* Indicates species with ant gardens.

Although O. hastatus is widely distributed in humid Neotropical areas, there are relatively few specimens in collections, when compared to the other widely distributed species. Its arboreal habit and nocturnal activity probably explain the low number of specimens and its apparent patchy distribution ( Fig. 80 ). Considering that epiphytes are more frequent and diverse in wet tropical forests ( Zotz, 2016), these areas can offer more resources for O. hastatus to nesting, explaining the relatively few records in the Brazilian Cerrado and Caatinga.

Comments: Among the Neotropical species, O. hastatus is very similar to O. davidsoni . Both have an elongated and narrow vertex, mandibles longer than half-length of the head with acute apical teeth, and the pronotum is not constricted anteriorly, gradually forming a neck. Odontomachus hastatus can be quickly recognized by the vertex, smooth and shining, the pronotum with transverse striae sometimes smooth and shining on the central portion; whereas O. davidsoni has the vertex striate and the pronotum forming concentrical striae on central portion.

The main variations are the level of sculpturing of some segments. The central portion of the pronotum can be completely smooth and shining to transversely striate slightly effaced. The central portion of the mesopleuron and the lateral faces of the petiole also vary from smooth and shining to slightly effaced striated. These variations are not stable in the populations and in the series.

Material examined: 64☿, 9♀, 4♂. BRAZIL: Amazonas: 60km north from Manaus (1☿) ; BR 174, Km 42 (1☿) ; Benjamin Constant (3☿) ; Barcelos (1☿) ; Presidente Figueiredo (3☿) ; Manaus (10☿) . Bahia: Camacã (2☿, 1♀) . Pará: Cachimbo , (1♂) ; Jari (1☿) . Pernambuco: Moreno (1☿) . Rio de Janeiro: Angra dos Reis, Jiboia Island (1☿) . Roraima: Caracaraí (1☿) . Sergipe: Aracaju (1☿) . São Paulo: Alcatrazes Islands (3☿) ; Caraguatatuba (11☿, 4♀, 3♂) ; Cubatão (1☿) ; Ubatuba (4☿, 1♀) . COSTA RICA: [ Locality not found] La Salvadora (1☿) . ECUADOR: Napo: Limoncocha (1☿, 1♀) . FRENCH GUIANA: Saint-Laurent-du-Maroni: Maripassoula : [Without location] (1☿) . Régina: Réserve Naturelle Nationale des Nouragues (1♀) . Roura: Montagne dès Chevaux (6☿) . Sinnamary: Paracou Research Station (1☿) . GUYANA [ SURINAME]: Brokopondo: Sara [ Creek River ] (2☿) . PANAMA: Los Espaveses: Bosque Protector San Lorenzo (1☿) . PANAMA: Coclé: El Coopé (1☿) . PERU: Loreto: Iquitos (1♀) . Madre de Dios: Puerto Maldonado (4☿) . VENEZUELA: Bolívar: Cañon Del Diablo (1☿) .

Odontomachus insularis Guérin-Méneville, 1844 ( Figures 23-B , 36-A , 60 , 94 , 95 and 96 )

Odontomachus insularis Guérin-Méneville, 1844: 423 (☿ and ♂); subspecies of Odontomachus haematodus by Emery, 1890: 44; Forel, 1897: 298 (♀); status as species by Dalla Torre, 1893: 51; subspecies of Odontomachus haematodus by Forel, 1899: 20; status as species by Taylor & Wilson, 1961: 142; subspecies of Odontomachus haematodus by Smith, 1967: 348; status as species and lectotype designation by Brown. 1976: 104. 135.

Odontomachus haematodus insularis var. pallens Wheeler, 1905: 82 . 122 (☿, ♀ and ♂); junior synonym of Odontomachus insularis by Brown. 1976: 104; as unavailable name by Bolton, 1995: 296.

Odontomachus haematodus insularis var. wheeleri Mann, 1920: 405 (☿); junior synonym of Odontomachus insularis by Brown. 1976: 104; declared as unavailable name by Bolton, 1995: 298.

Type specimens: Paralectotypes. Odontomachus insularis ; CUBA: [no further data]. ic. R. a. (type), GBIF-D/ FoCol 0683, FOCOL0683 1♀ / GBIF-D/ FoCol 0684, FOCOL0684-1 1☿ / GBIF-D/ FoCol 0684, FOCOL0684-2 1☿ [ ZSM] .

Paralectotypes. Odontomachus haematodus insularis var. pallens ; BAHAMAS: Nassau, May-June 1904, W.M. Wheeler, Coll. COTYPE No. A.M.N.H., USNMENT 00531664 1☿ [ NMNH] .

Etymology: From the Latin word insularis = of/from islands; referring to the type-locality of this species.

Diagnosis: (☿) Scape reaching occipital corner but not surpassing it. Mandible denticles acute. Central disc of pronotum with striae forming concentric triangles. Petiolar node conical, anterior face continuous to petiolar spine needle-like strongly curved posteriorly, smooth and shining. Body light brown to yellowish, gaster dark brown. (♀) Mesosoma and petiole light brown to yellowish. Petiole in lateral view with long curved spine. (♂) Body pale yellow with propodeum brown. Mesoscutum with central portion effaced and micro-sculpturing becoming coarser laterally. Petiole with fine rugae on ventral portion becoming ascendent, coarser and sparse on posterior portion.

Worker

Syntype measurements: FOCOL0684-2 was not measured due to the bad conservation status. EL 0.31, HL 2.7, HoW 2.13, HvW 1.77, HW 1.97, ML 1.38, SL 2.33, PnW 1.19, WL 3.33, FL 2.41, PTH 1.23, PTL 0.51, PTW 0.56, GL 2.89, TL 10.81. CI 0.72, MI 0.51, PtI 2.41, SI 0.86, VI 0.89.

Non-type measurements: EL 0.3-0.4, HL 2.3-2.6, HoW 1.8-2.1, HvW 1.4-1.8, HW 1.6-1.9, ML 1.2-1.3, SL 2.0-2.3, PnW 1.0-1.2, WL 2.7-3.0, FL 2.2-2.5, PTH 1.1-1.2, PTL 0.5, PTW 0.4-0.5, GL 2.6-3.1, TL 9.5-10.0. CI 0.69-0.73, MI 0.50-0.52, PtI 2.20-2.40, SI 0.84-0.91, VI 0.87-0.94 (n= 4).

Head. Vertex trapezoidal, wider than long in frontal view; with regular longitudinal striae. Posterior margin concave medially; occipital carina not forming nuchal collar. Cephalic constriction conspicuous. Frontal lobes striated. Ocular bridges posteriorly forming arc. Eye relatively large (EL> 0.4), rounded, posterior margin rounded. Clypeus with inconspicuous longitudinal striae on median portion. Malar area with longitudinal sulcus in latero-oblique view, anterior portion with fine irregular transverse striae, posterior portion smooth and shining.

Mandible. Almost as long as anterior head portion. External margin convex. Masticatory margin with denticles that serially increase in size apically. Apical teeth acute. Palp formula 4,3.

Antenna. Scape reaching occipital corner but not surpassing it.

Mesosoma. Pronotum in lateral view rounded, dorsal margin convex; neck well developed anteriorly in dorsal view; lateral margin rounded; central disc of pronotum with regular longitudinal striae forming acute angle anteriorly, external striae forming concentric triangles, striae becoming irregular posteriorly. Propleuron smooth and shining with effaced transverse striae on external margin. Mesonotum larger than wide; with regular transverse striae. Mesopleuron with vestigial carina on anteroventral portion; mostly smooth and shining, anterior portion with regular transverse striae. Mesometanotal sulcus well-marked, scrobiculate; curved anteriorly. Metanotum as long as mesonotum half-length; with fine regular transverse striae. Metapleural gland bulla with transverse striae effacing posteriorly. Propodeum in lateral view slightly concave posteriorly, posterior margin length about 1/3 of dorsal length. Metasternum with metasternal process open, cuticle border bilobate separated on middle line by longitudinal sulcus or completely separated forming lateral cuticle borders; regular transverse fine striae.

Petiole. Sessile. Conical node, anterior margin in lateral view strongly convex on dorsal portion, continuous with petiolar spine. Spine long, acute, curved posteriorly, as long as petiolar node half-length. Smooth and shining, with at most inconspicuous transverse striae on ventral portion of lateral face. Subpetiolar process wider than long keel-shaped; tip acute; posterior portion concave.

Gaster. First tergum almost higher than long; anterior and dorsal margin meeting in an obtuse angle; dorsal face convex. dorsal face of all terga smooth and shining.

Color and size. Body light brown to yellowish. Gaster dark brown. Legs and antennae pale yellow. Relatively small ants.

Pilosity. Hook-shaped, scattered, conspicuous and short pubescence spread over the body, separated each other about their length. Pubescence on mandible and gaster sparser than body surface. Pubescence on petiole limited to anterior face extending to lateral faces. Short hairs on anterior portion of procoxa, pronotum and gastral terga, smaller than maximum procoxa width.

Gyne

Syntype measurements: EL 0.39, HL 2.4, How 1.79, HvW 1.42, HW 1.59, ML 1.08, SL 1.85, PnW 1.23, WL 2.98, FL 1.79, femur inclinated, not possible to measure, FWL 4.61, PtW 0.52, PTH, PTL and GL no measured due to forewing covering the petiole and first gaster tergum. CI 0.74, MI 0.50, SI 0.86, VI 0.89.

Non-type measurements: EL 0.4, HL 2.4, HoW 2.0, HvW 1.6, HW 1.8, ML 1.3, SL 2.0, PnW 1.2, WL 3.1, FL 2.3, FWL 6.2, PTH 1.4, PTL 0.5, PtW 0.5, GL 3.0, TL 10.3. CI 0.75, MI 0.54, PtI 2.8, SI 0.83, VI 0.88 (n= 1).

Mesosoma. Pronotum slightly longer than high in lateral view, dorsal margin slightly concave; lateral margins converging anteriorly forming a neck in dorsal view; with regular transverse striae forming arcs directed posteriorly on lateral face. Subalar area conspicuous with pronotum. Mesoscutum anterior and dorsal margin forming single convexity in lateral view; with regular longitudinal striae in dorsal view; striae between parapsidial line and parascutal area oblique, directed medially. Parapsidial line curved, well-marked. Parascutal area smooth and shining. Notopropodeal sulcus scrobiculate. Axilla with inconspicuous and fine longitudinal striae. Mesoscutellum smooth and shining. Metascutellum smooth and shining with vestigial longitudinal carina.

Wing. Color pale brown, translucid. Forewing ML1 curved. RsL1 straight; as long as Rs+M. RSL2-3 slightly curved. ML2 short, about half-length of Rs+M. Rs+M and ML2 forming single slightly curved line, not forming angle with RsL2-3. Vein cu-a not reaching M+Cu ML1 angle by less than its half-length. Hindwing 1rs-m vein slightly curved. 8 hamuli.

Petiole. Conical node, anterior and posterior face straights. Petiolar spine needle-like, long and curved posteriorly.

Pilosity. Pubescence on mesosoma strongly curved, dense separated each other less than its length. Pubescence on mesoscutellum and metascutellum longer and less dense than pronotum pubescence, about half-length longer. Pronotum with two pairs of short hairs, shorter than procoxa width.

Male

Non-type measurements: EL 0.62, HL 0.83, OW 0.16, OES 0.21, IOW 0.18, WL 2.4, FWL 4.4, PTH 0.8, PTL 0.5, PTW 0.4, GL 1.9, TL 5.13. PtI 1.6 (n= 1).

Pilosity. Head, mesosoma, petiole and gaster with dense appressed pubescence. Pronotum without hairs. Thin and erect short hairs on posterior portion of gastral terga IV and V.

Head. Lateral ocelli partially surpassing posterior margin; posterior margin between lateral ocelli straight, not reaching median ocellus.

Mesosoma. Pronotal dorsum with inconspicuous micro sculpture; with fine, irregular longitudinal striae, striae on dorsal portion curved dorsally in lateral view. Propleuron with inconspicuous micro sculpture. Mesoscutum anteriorly with inconspicuous punctuations; dorsal surface with fine median longitudinal carina anteriorly; central portion with inconspicuous micro sculpture becoming coarser laterally. Axilla with fine punctuation with reticulate aspect. Mesoscutellum convex; with inconspicuous micro sculpture on central portion. Metascutellum with dense micro sculpture of reticulate aspect, with longitudinal carina. Mesopleuron mostly smooth and shining. Metapleuron with rugae. Metepisternum with rugulae with dense micro sculpture between them. Metapleuron and propodeum separated by longitudinal sulcus scrobiculate. Propodeal dorsum with coarse rugae; dorsal and posterior face delimited by coarser ruga; posterior face with coarse rugae.

Wing. Translucid. Hamuli equal 8. Pterostigma brown, contrasting with wing surface.

Petiole. Sessile. In lateral view dorsal portion acute; with fine rugae on ventral portion becoming ascendent. coarser and sparse on posterior portion. Subpetiolar process acute and curved posteriorly. Petiolar sternum concave posteriorly.

Color and size. Head, prothorax, mesothorax legs and antenna pale yellow. Lateral portion of mesoscutum and axilla brown. Propodeum and metapleuron dark brown. Petiole and gaster brown. Relatively small size.

Natural history: Wheeler & Wheeler (1952) mention that Clausen (1941) found larvae of O. insularis hosting two species of parasitoids, Kapala terminalis Ashmed, 1892 and Kapala sp. ( Hymenoptera : Eucharitidae ). The planidia of K. terminalis are usually found attached close to the head on the esophagus of the larvae. The parasitoid provokes shortness of pupal legs that never attain more than half the full length.

It is restricted to Central America in Cuba ( Fig. 60 ), and according to Brown (1976) it also occurs in the Bahamas.

Comments: Odontomachus insularis is similar to O. biolleyi due to its small yellowish body. They can be separated by the well-developed petiolar spine in O. insularis , but short in O. biolleyi . The scape length in O. insularis does not surpass the occipital corner, but it does in O. biolleyi .

Males have a pale-yellow body with a dark brown and rugose propodeum, similar to the males of O. haematodus and O. bauri . However, the male of O. insularis can be separated due to the rugulose mesoscutum and petiolar node. The sculpturing on the dorsal mesoscutal face is coarser in O. insularis than in either O. haematodus or O. bauri .

The Lectotype designated by Brown (1976) is deposited in MHNH. In the ZSM site (SNSB, 2023) the type specimens are mentioned as “ holotypes ”, but they should be considered paralectotypes (Article 73.2.2. ICZN).

Material examined: 4☿, 1♀, 1♂. CUBA: [Ciudad de La Havana]: Havana (4☿, 1♂) . [Mayabeque]: Jaruco (1♀) .

Odontomachus laticeps Roger, 1861 ( Figures 20-A , 34-A , 40-B , 97 , 98 , 99 and 100 )

Odontomachus haematodus var. laticeps Roger, 1861: 25 (☿ in text); status as species by Dalla Torre 1893: 51; subspecies of Odontomachus haematodus by Emery, 1894: 50; status as species by Brown, 1976: 104.

Odontomachus haematodes r. striativentris Emery, 1890: 44 (☿); status as species by Dalla Torre 1893: 1893; subspecies of

Odontomachus haematodus by Forel, 1899: 21; status as species by Kempf, 1972: 172; junior synonym of Odontomachus laticeps by Brown, 1976: 104.

Type specimens: Syntype [ Images ]. Odontomachus haematodus r. striativentris . [ COSTA RICA: Alajuela]. Jiménez col., ANTWEB CASENT0904008 1☿ [ MSNG] .

Etymology: Combination of the Latin words latus = wide, broad + ceps = head; probably referring to the relative wide vertex compared to the O. haematodus .

Diagnosis: (☿ and ♀) Propleuron with regular transverse fine striae, striae on posterior portion forming concentric arcs. Ventral portion of pronotum in lateral view with long hairs, almost as long as the procoxa hairs and curved anteriorly. Propleura with long erect hairs, about as long as procoxa hairs. First and second gastral terga longitudinally striated to smooth and shining. (♂) Body yellowish brown. Mesoscutum and mesoscutellum in dorsal view with inconspicuous micro sculpture. Propodeum in dorsal view with longitudinal regular striae diverging posteriorly forming concentric circles on the posterior face. Petiole higher than longer.

Worker

Syntype measurements: EL 0.48, HL 3.37, HoW 2.59, HvW 2.3, HW 2.49, ML 1.55, SL 2.93, PnW 1.46, WL 4.13, FL 2.79, PTH 1.52, PTL 0.63, PtW 0.59, GL 3.26, TL 12.49. CI 0.73, MI 0.45, PtI 2.41, SI 0.86, VI 0.92.

Non-type measurements: EL 0.5-0.4, HL 2.7-3.5, HoW 2.1-2.6, HvW 1.7-2.3, HW 1.9-2.5, ML 1.5-2.0, SL 2.4-3.0, PnW 1.2-1.5, WL 3.0-3.9, FL 2.7-3.3, PTH 1.3-1.5, PTL 0.5-0.6, PtW 0.5-0.6, GL 3.1-3.5, TL 10.8-13.5. CI 0.70-0.91, MI 0.56-0.70, PtI 2.0-2.66, SI 0.86-1.16, VI 0.86-0.90 (n= 4).

Head. Vertex trapezoidal, wider than long in frontal view; with regular longitudinal striae diverging and effacing posteriorly, reaching posterior margin. Posterior margin concave medially; occipital carina not forming nuchal collar. Cephalic constriction conspicuous. Frontal lobes striated. Ocular bridges posteriorly forming arc. Eye relatively large (EL> 0.4); rounded, posterior margin rounded. Clypeus with regular fine longitudinal striae on median portion. Malar area with inconspicuous irregular transverse striae in latero-oblique view, posterior portion smooth and shining.

Mandible. Almost as long as anterior head portion. External margin convex, base almost straight. Masticatory margin with blunt denticle series gradually increasing in size. Palp formula 4,3.

Antenna. Scape surpassing occipital corner about pedicel half-length.

Mesosoma. Pronotum in lateral view rounded, dorsal margin slightly convex; neck well developed anteriorly in dorsal view; lateral margin rounded; central disc of pronotum with regular longitudinal striae, lateral striae forming concentric arcs directed anteriorly or circles. Propleuron with regular transverse fine striae, striae on posterior portion forming concentric arcs. Mesonotum larger than wide, with regular transverse striae. Mesopleuron with rounded carina on anteroventral portion; mostly smooth and shining, anterior portion with fine regular transverse striae curved dorsal-anteriorly. Notopropodeal sulcus scrobiculate, slightly curved anteriorly. Metanotum as long as notopropodeal sulcus with fine irregular transverse striae. Metapleural gland bulla with inconspicuous, fine, transverse striae on dorsal portion. Propodeum in lateral view with two weak impressions; posterior margin length about 1/3 of dorsal length; striae on anterior portion usually longitudinal forming concentric ellipses in dorsal view. Metasternum with metasternal process open, cuticle border forming lateral projections; fine regular transverse striae on anterior portion.

Petiole. Sessile. Conical node, resembling an isosceles triangle, anterior margin in lateral view straight continuous with petiolar spine. Spine short or long, robust, curved posteriorly. Regular fine transverse and punctate striae covering all surface, except by posterior face. Subpetiolar process wider than long keel-shaped; tip acute; posterior portion not concave.

Gaster. First tergum almost higher than long; anterior and dorsal margin meeting in an obtuse angle; dorsal face convex. dorsal face of first second and third terga smooth varying from with fine regular longitudinal striae finely punctate to smooth and shining.

Color and size. Body dark brown to reddish brown. Gaster darker than body. Legs and antennae usually lighter than body. Relatively medium ants.

Pilosity. Scattered, decumbent, conspicuous and short pubescence spread over the body, separated each other about their length. Pubescence on mandible and petiole sparser than body surface. Long pubescence strongly curved directed anteriorly on posterior ventral portion of pronotum, about double length of body pubescence. Pubescence on petiole limited to anterior face extending to lateral faces. Short hairs on anterior portion of procoxa and propleuron, smaller than procoxa width. Pronotum with three or four pairs of long, curved erect hairs longer than maximum procoxa width.

Gyne

Non-type measurements: EL 0.5-0.4, HL 2.4-3., HoW 1.9-2.4, HvW 1.9-2.0, HW 2.1-2.2, ML 1.7-1.8, SL 2.6-2.8, PnW 1.6, WL 3.7-3.9, FL 2.6-3.0, FWL 7.7, PTH 1.4-1.6, PTL 0.7-0.6, PtW 0.7-0.8, GL 3.7-4.7, TL 12.9-13.6. CI 0.70-0.91, MI 0.56-0.70, PtI 2.0-2.6, SI 0.86-1.16, VI 0.86-0.90 (n= 4).

Mesosoma. Pronotum slightly longer than high in lateral view, dorsal margin slightly concave; lateral margins converging anteriorly forming a neck in dorsal view; with regular transverse curved striae forming concentric arcs directed anteriorly, central striae might be longitudinal, striae forming acute arcs directed dorsally in lateral view. Subalar area conspicuous with pronotum; with irregular transverse striae. Mesoscutum anterior and dorsal margin forming single convexity in lateral view; with or without median sulcus anteriorly in dorsal view; regular longitudinal striae; striae between parapsidial line and parascutal area longitudinal. Parapsidial line curved, well-marked. Parascutal area smooth and shining. Notopropodeal sulcus scrobiculate. Axilla with regular longitudinal striae. Mesoscutellum longitudinal striae effaced with longitudinal carina. Metascutellum smooth and shining with longitudinal carina.

Wing. Color light brown, translucid. Pterostigma darker than wing surface. Forewing ML1 curved. RsL1 straight; as long as Rs+M. RSL2-3 slightly sinuous. Rs+M and ML2 forming single curved line with RsL2-3; Rs+M longer than ML2; Vein cu-a not reaching M+Cu ML1 angle by less than its half-length. Hindwing 1rs-m vein slightly curved. 9 hamuli.

Petiole. Conical node, anterior face straight to slightly concave, posterior face convex on dorsal portion. Petiolar spine coarser than worker’s spine.

Pilosity. Pubescence on mesosoma dense separated each other less than its length. Pubescence on mesoscutellum less dense. Pronotum with two or three pairs of short hairs, as long as procoxa width.

Male

Non-type measurements: EL 0.68, HL 0.99, OW 0.15, OES 0.24, IOW 0.27, WL 2.9, FWL 5.5, PTH 0.9, PTL 0.50, PtW 0.5, GL 2.7, TL 6.59. PtI 1.8 (n= 1).

Head. Palp formula 5,3. Lateral ocelli partially surpassing posterior margin; posterior margin between lateral ocelli concave, not reaching median ocellus.

Mesosoma. Pronotal dorsum with micro sculpture; with micro sculpture effaced on dorsal portion and short fine striae posteriorly in lateral view. Mesoscutum anteriorly with irregular transverse striae; dorsal surface with regular oblique striae diverging laterally, striae on posterior portion converging medially. Axilla with fine striae and micro sculpture between them. Mesoscutellum slightly concave; with inconspicuous striae anteriorly and inconspicuous micro sculpture. Metascutellum smooth and shining, with sharp longitudinal carina.Anepisternum and katepisternum with micro sculpture on central portion gradually becoming sooth and shining. Metapleuron with fine irregular longitudinal striae, coarser anteriorly. Metepisternum with fine irregular longitudinal striae on dorsal portion and micro sculpture of reticulate aspect. Metapleuron and propodeum separated by longitudinal carina extending from metepisternum to propodeal spiracle and sharp irregular longitudinal carina posteriorly to it. Propodeal dorsum with oblique striae diverging and meeting in transverse striae on posterior face forming concentric circles; dorsal and posterior face of propodeum not delimitated by coarser striae; posterior face with irregular transverse striae curved ventrally.

Wing. Light brown translucid. Hamuli equal 8. Pterostigma brown, contrasting with wing surface.

Petiole. Sessile; dorsal margin acute; with micro sculpture on ventral portion gradually becoming smooth and shining dorsally. Subpetiolar process acute and curved posteriorly. Sternum concave posteriorly.

Color and size. Entire body yellowish brown with propodeum slightly darker. Relatively medium size.

Pilosity. Head, mesosoma and gaster with dense appressed pubescence. Petiolar pubescence less dense. Pronotum with three pairs of thin erect hairs on posterior portion, two of them larger. Thin and erect short hairs on posterior portion of gaster segments I and II, hairs on segments III, IV and V slightly almost two times larger. Gaster sterna IV and V with several long hairs.

Natural history: Horvitz (1981), studied ant interactions with seeds of Calathea G. Mey., 1818 ( Marantaceae ), observed on two occasions workers feeding larvae with bits of arils. The workers of O. laticeps removed the aril from 78% of the seeds in his experiment and carried them to the nest. There is a record of it in a Chamaedorea radicalis Mart. ( Araceae ) palm plantation foraging on the litter and low vegetation ( Lara-Villalón et al. 2015), and in pine forests ( Brown, 1976). It occurs in Central America ( Fig. 100 ) and according to Brown (1976), this species occurs mostly in lowland forested areas up to 1,070 m above sea level.

Comments: Odontomachus laticeps can easily be confused with at least another five Odontomachus species, due to its lack of exclusive diagnostic characters, and the remarkable variation in the gastral tergal sculpture. The massive head, the slightly angulate pronotum and the smooth and shining gastral terga make it resemble O. biumbonatus and O. opaciventris . Odontomachus scalptus differs from them by the head being relatively small, the pronotum less angulate, almost completely convex in lateral view, and the petiolar node conical, which is cylindrical in O. biumbonatus and O. opaciventris . The conical petiolar node, resembling an isosceles triangle in lateral view, can also occur in other widely distributed and abundant species such as O. haematodus and O. bauri . The sympatric O. ruginodis may have similar sculpturing on the node, but the relative massive head, the dark body, and the long pilosity of the pronotum and procoxa are enough to distinguish O. laticeps . Furthermore, the long pilosity on the posterior ventral portion on the lateral face pronotum and propleuron present in O. laticeps are enough to separate it from them.

Brown (1978) mentions specimens of O. laticeps from Bolivia collected by Mann that were smaller than usual and have striate-punctate sculpturing on the gastral terga. We examined individuals of O. meinerti corresponding to this from Peru and Ecuador, so the specimens mentioned by Brown (1978) probably correspond to O. meinerti . The gastral tergal sculpturing is usually diagnostic for Odontomachus species, and at first sight, two forms can be distinguished from populations of O. laticeps : one with smooth and shining gastral terga and another with longitudinal striae. Brown (1976) observed intermediary forms, also observed in this study, confirming the variation of this character. According to him, populations from upland areas usually are dark-brown with a smooth and shining gastral terga, and specimens from wet lowlands tend to be reddish-brown with striated terga. Even so, it is possible to find exceptions in both extremes of this variation ( Brown, 1976). Unfortunately, it was not possible to examine as many specimens as he did to confirm its hypothesis.

The males are similar to O. chelifer due to the medium to large size and the propodeal striation forming concentric circles on the posterior face. Odontomachus laticeps is yellowish-brown in color and has inconspicuous sculpturing on the mesoscutum and mesoscutellum that contrasts with the darker brown and conspicuous sculpturing of O. chelifer .

Material examined: 33☿, 2♀, 1♂. BELIZE: Cayo: Augustine (2☿) . Orange Walk: Rio Bravo Pres. (1☿) . COSTA RICA: Chiapas: Izapa (3☿) . Heredia: La Selva (2☿) ; [ Sarapiquí ], 22km N Volcan Barba (1☿) ; Pto [Puerto] Viejo (1☿) . [Limón]: R[io] Toro Amarillo Vic [inity] Guapiles (1☿) . [San José]: San José (4☿) . [no further data] (1☿) . GUATEMALA: [El Petén]: Cave 4 min S Flores (1☿) . Izabal: Morales , 5km NW (1☿) . MÉXICO: [Guanajuato]: Cueva San Ignacio, “Acahuizotia” (2♀) . Jalisco: Casimiro , Castillo (5☿) . Puebla: Cuetzalan (1☿, 1♂) . Tamaulipas: Sa. de Guatemala, Rancho del Cielo (2☿) . Veracruz: Park Cañon 2 min W of Fortin, H [High] Way 150 (1☿) ; Km 17 on Fortin-Huatusco road bridge 4 Aug (3☿) ; Reserva [de la Biosfera de] los Tuxtlas (1☿) . PANAMA: Canal Zone (1☿) . VENEZUELA: Aragua: Parque Nacional Henri Pittier Grande (1☿) .

Odontomachus meinerti Forel, 1905 ( Figures 18-A , 33-B , 35-A , 43-B , 100 , 101 , 102 and 103 )

Odontomachus haematodus r. meinerti Forel, 1905: 156 (☿); junior synonym of Odontomachus minutus by Brown, 1976: 104; status as species and senior synonym by Bolton, 1995: 296.

Odontomachus haematoda minuta Luederwaldt, 1918: 36 (☿); status as species by Kempf, 1972: 7; junior synonym of Odontomachus meinerti by Bolton, 1995: 296.

Odontomachus haematodus dulcis Mann, 1922: 18 (☿); junior synonym of Odontomachus minutus by Brown, 1976: 102; junior synonym of Odontomachus meinerti by Bolton, 1995: 296.

Type specimens: Syntype [Images]. Odontomachus haematoda dulcis Mann, 1922 ; HONDURAS: Progresso, W. M. Mann [col.], [ii-iii.1920], Cotype N °24445, U.S. N.M., USNM ENT 00533104 1☿ [ NMNH] .

Syntype [ Images ]. Odontomachus haematoda minutus . Luederwaldt. 1918; BRÉSIL[ BRAZIL]: Prov. Matto[Mato] Grosso, [Conceição de Itanhaém], P. Germain [col.], 1886, ANTWEB CASENT0904009 1☿ [ MSNG] .

Etymology: In honor to Frederik Vilhelm August Meinert (1833-1912), a Danish arachnologist who collected the type specimens of this species.

Diagnosis: (☿) Propleuron smooth and shining. Petiolar node conical resembling an isosceles triangle, almost entirely smooth and shining; petiolar spine short, robust and curved posteriorly. First and second gastral terga varying from punctuate, anteriorly punctuate gradually forming longitudinal striae posteriorly or completely striate-punctate longitudinally (♀) Petiole and gastral terga as in worker. (♂) Entire body pale yellow. Mesoscutellum in antero-dorsal view with dorsal margin rounded. Posterior face of propodeum with fine longitudinal to oblique irregular striae with micro sculpture between them. Petiolar node with dorsal portion blunt.

Worker

Non-type measurements: EL 0.20-0.40, HL 1.60-2.50, HoW 1.20-2.60, HvW 0.90-1.50, HW 1.10-1.70, ML 0.90- 1.50, SL 1.50-2.30, PnW 0.70-1.10, WL 2.00-3.20, FL 0.90-2.40, PTH 0.60-1.10, PTL 0.30-0.50, PTW 0.30-0.50, GL 1.20-3.00, TL 6.30-10.40. CI 0.70-0.72, MI 0.55-0.64, PtI 2.0-2.66, SI 0.9-1.12, VI 0.85-0.93 (n= 67).

Head. In frontal view, vertex trapezoidal, wider than long; with regular longitudinal striae diverging and effacing posteriorly, reaching posterior margin. Posterior margin concave medially; occipital carina not forming nuchal collar. Cephalic constriction conspicuous. Frontal lobes striated. Ocular bridges posteriorly forming arc. Eye relatively large (EL> 0.4); rounded, posterior margin rounded. Clypeus smooth and shining with inconspicuous piligerous punctuation on median portion. Malar area smooth and shining in latero-oblique view.

Mandible. Almost as long as anterior head portion. External margin convex, base almost straight. Masticatory margin with denticles that serially increase in size apically. Palp formula 4,3.

Antenna. Scape surpassing occipital corner about pedicel half-length or less.

Mesosoma. Pronotum in lateral view rounded, dorsal margin convex; in dorsal view, neck well developed anteriorly; lateral margin rounded; central disc of pronotum with regular longitudinal striae, lateral striae forming concentric longitudinal ellipses or arcs directed anteriorly. Propleuron smooth and shining; regular fine oblique striae restricted to anterior portion gradually fading posteriorly. Mesonotum larger than wide, with regular transverse striae slightly curved anteriorly. Mesopleuron with rounded carina on anteroventral portion; mostly smooth and shining, anterior portion with inconspicuous regular transverse striae curved dorsal-anteriorly. Mesometanotal sulcus well-marked, scrobiculate. Metanotum vestigial. Metapleural gland bulla with inconspicuous, fine, transverse striae on dorsal portion. Propodeum in lateral view with two weak impressions; posterior margin length about 1/3 of dorsal length. Metasternum with metasternal process open, cuticle border forming straight lateral projections; fine regular transverse striae on anterior portion.

Petiole. Sessile. Conical node, resembling an isosceles triangle, anterior margin in lateral view straight continuous with petiolar spine. Petiolar spine short, robust, curved posteriorly. Sculpturing inconspicuous limited to ventral portion. Subpetiolar process wider than long keel-shaped; tip acute; posterior portion concave.

Gaster. First tergum almost higher than long; anterior and dorsal margin meeting in an obtuse angle; dorsal face convex. dorsal face of first and second terga punctate anteriorly gradually forming longitudinal striae on posterior portion; third and fourth terga punctate.

Color and size. Body yellow pale to nut brown. rarely dark brown. Legs and antennae yellowish. Relatively small to medium size ants.

Pilosity. Scattered, decumbent, conspicuous and short pubescence spread over the body, separated each other about their length. Pubescence on mandible and petiole sparser than body surface. Pubescence on petiole limited to anterior face extending to lateral faces. Short hairs on anterior portion of procoxa smaller than coxae width. Pronotum with three or four pairs of long, curved erect hairs longer than procoxa width. Long erect hairs on gastral terga directed posteriorly, as long as pronotum hairs.

Gyne

Non-type measurements: EL 0.40, HL 1.90-2.30, HoW 1.60-1.80, HvW 1.20-1.40, HW 1.40-1.60, ML 1.10-1.40, SL 1.60-2.20, PnW 1.00-1.20, WL 2.60-3.20, FL 1.80-2.30, FWL 5.90, PTH 1.10-1.20, PTL 0.40-0.5, PTW 0.50- 0.60, GL 2.40-3.20, TL 8.60-10.40. CI 0.68-0.76, MI 0.54-0.63, PtI 2.22-2.75, SI 0.84-0.95, VI 0.80-0.87 (n= 10).

Mesosoma. Pronotum slightly longer than high in lateral view, dorsal margin slightly concave; lateral margins converging anteriorly forming a neck in dorsal view; pronotal dorsum with regular transverse striae gradually becoming irregular posteriorly, striae completely longitudinal in lateral view. Subalar area conspicuous with pronotum; with irregular longitudinal striae. Mesoscutum anterior and dorsal margin forming single convexity in lateral view; with or without median sulcus anteriorly in dorsal view; regular longitudinal striae; striae between parapsidial line and parascutal area longitudinal. Parapsidial line curved, well-marked. Parascutal area smooth and shining. Notopropodeal sulcus scrobiculate. Axilla with regular longitudinal striae effacing or not posteriorly. Mesoscutellum varying from densely punctate to smooth and shining.Metascutellum sculpture as like mesoscutellum, with or without longitudinal carina.

Wing. Color pale yellow, translucid. ML1 curved. RsL1 straight; as long as Rs+M. RSL2-3 curved. Rs+M and ML2 forming obtuse angle with RsL2-3; Rs+M longer or as long as ML2; Vein cu-a not reaching M+Cu ML1 angle by almost its half-length. Forewing 1rs-m vein slightly curved. 7 hamuli.

Petiole. Conical node. Anterior face straight to slightly concave. Posterior face convex on dorsal portion.

Pilosity. Pubescence on mesosoma strongly curved, dense separated each other less than its length. Pubescence on mesoscutellum less dense. Pronotum with five pairs of long hairs, as long as procoxa width.

Male

Non-type measurements: EL 0.50-0.60, HL 0.80-0.90, OW 0.12-0.18, OES 0.21-0.27, IOW 0.21-0.27, WL 2.20- 2.50, FWL 4.30-5.00, PTH 0.60-0.70, PTL 0.40-0.50, PTW 0.30-0.70, GL 2.00-2.40, TL 5.30-5.80. PtI 1.40-1.7 (n= 5).

Head. Palp formula 4,3. Lateral ocelli partially surpassing posterior margin; posterior margin between lateral ocelli concave, not reaching median ocellus.

Mesosoma. Pronotal dorsum with micro sculpture of striate aspect; with micro sculpture of striate aspect effaced anteriorly in lateral view. Propleuron with inconspicuous micro sculpture. Mesoscutum, axilla and mesoscutellum with effaced micro sculpture. Mesoscutum with longitudinal carina. Mesoscutellum convex, dorsal margin slightly straight on median portion. Anepisterno smooth and shining or with inconspicuous micro sculpture on antero-dorsal portion. Metapleuron with rugulae. Metepisternum with longitudinal irregular striae with micro sculpture between them. Metapleuron and propodeum separated by longitudinal sulcus, sometimes rugae on propodeum and metapleuron crossing it posteriorly. Propodeal dorsum with fine irregular striae diverging posteriorly; dorsal and posterior face delimited by coarser irregular striae; posterior face with fine longitudinal to oblique irregular striae with micro sculpture between them.

Wing. Whitish or grayish translucid. Hamuli equal to 7. Pterostigma dark brown, contrasting with wing surface.

Petiole. Sessile; in lateral view dorsal portion rounded; imbricate gradually becoming smooth and shining dorsally. Subpetiolar process subquadrate. Sternum concave posteriorly.

Color and size. Entire body yellowish brown. Relatively small size.

Pilosity. Head, mesosoma, petiole and gaster with dense appressed pubescence. Pronotum with three or four thin erect hairs on posterior portion. Thin and erect short hairs on posterior portion of gaster segments I and II, hairs on segments III, IV and V almost two times larger.

Natural history: Despite the abundance of O. meinerti specimens in collections, relatively little is known about its biology. The species nests in rotten branches on the ground in forests and coffee plantations ( De La Mora & Philpott, 2010), or under rotten trunks ( Brown, 1976). The larvae have three instars and pre-pupae, and the male larvae are larger than the workers ( Fox et al. 2017). New colonies in artificial conditions can produce male eggs after 46 days ( Brandão 1983). There are records of pupae parasitized by species of Kapala ( De la Mora et al. 2015; De La Mora & Philpott, 2010). Its karyotype is 2n= 44 (2K= 4SM+6ST+34T) ( Santos et al. 2010).

Odontomachus meinerti is widely distributed from southern Brazil to southern Mexico in forest areas ( Fig. 100 ). According to Brown (1976) it is common in gallery forests in Cerrado areas and also occurs in Paraguay and it probably occurs in Bolivia and Guyana. There is a record of it in Colombia ( Rodriguez et al. 2008), though no voucher from this study was examined.Amongst the other widely distributed species (i.e., O. bauri , O. haematodus , and O. chelifer ), it has not yet been recorded from any islands. Its nesting habit in rotten branches would facilitate dispersal process to islands next to the continent. However, the establishment of the colonies would be difficulted due to other species that also live in litter or on the ground. Studies about the biology and the dynamic of the colonies are necessary to understand its distribution.

Comments: Odontomachus meinerti is usually confused with O. laticeps due to similarities in the node shape and the gastral terga sculpturing. Odontomachus meinerti lacks the conspicuous fine striae and the punctures on the petiolar node that O. laticeps has, and O. meinerti is usually smaller than O. laticeps . Odontomachus scalptus also has striate-punctate gastral terga, but it has longitudinal striae on the mesonotum, while O. meinerti has transverse striae.

Odontomachus meinerti is the Neotropical species with the wider morphological variation, regarding size and sculpturing. Workers can be quite small, almost like O. spissus , with areolate to punctuate gastral terga and yellowish to light brown body color. Odontomachus meinerti has eyes relatively big, vertex rectangular and striate, and the petiolar node conical; while O. spissus has eyes relatively small, vertex rectangular and smooth and shinning, and the petiolar node scale-like. The extreme opposite of variation are specimens with median body size, longitudinally striate-punctate gastral terga, dark brown body, and lighter colored appendages. In collections, the smaller yellowish specimens with areolate gaster terga were usually identified as O. minutus and the larger ones as O. meinerti . Even with this contrasting variation, specimens with intermediate morphology can be found in the same locality or even in the same series.

Gynes and males have relatively stable morphology compared to the workers. Gynes present areolate-punctuate sculpture and yellowish bodies are rare, and specimens with anteriorly punctuate and posteriorly striate gastral terga are the most frequent. Males usually have the mesoscutum, mesoscutellum, and metascutellum weakly sculptured to smooth and shining.

Although O. meinerti has the typical morphology of the haematodus group, according to Fernandes et al. (2021), it is closely related to O. laticeps ( Fig. 1 ), placed between the O. chelifer + O. affinis clade and other members of the haematodus group. Observing the vouchers from this study (Vouchers I28, I44, I55, I68, I86, and I89) it was possible to confirm their identity as O. meinerti . In the original description of Odontomachus haematodus r. meinerti, Forel (1904) does not mention how many specimens he studied. Brown (1976) did not examine type specimens and mentioned that a type of O. meinerti was examined for him by Dr. Børge Petersen. No types were found in the MHNG. A single specimen in alcohol labeled as a type was found in ZMUC.

Material examined: 485☿, 27♀ and 10♂. BRAZIL: Acre: Mâncio Lima (9☿, 3♀) ; Rio Branco (4☿) . Amapá: Serra do Navio (3☿, 2♀) . Amazonas: Manaus (16☿, 1♀) . Bahia: Andaraí (3☿) ; Camacã (1☿); Guarupá-Mirim (1♂); Ilhéus (2☿); Itacaré (1♂); Maracás (1☿); Porto Seguro (1♂). Ceará: Chapada de Ipiapaba (2☿) ; Crato (2☿); Guaramiranga (2☿); Maranguape (2☿); Meruoca (2☿); Ubajara (1☿). Espírito Santo: FLONA do Rio Preto (1☿) ; Linhares (3☿); Soretama (1☿). Goiás: Cavalcante (5☿) ; Jataí (122☿, 8♀); Montividiu (8☿); Niquelândia (2☿); Serranópolis (14☿, 1♀). Mato Grosso: Canarana (1☿, 1♀) . Mato Grosso do Sul: Bonito (4☿) ; Chapada dos Guimarães (3☿); Rondonópolis (3☿). Minas Gerais: 6km W from Cristália (1☿) ; Belo Horizonte (1☿); Betim (13☿, 1♀); Estrela do Sul (1☿) ; Ipaba (19☿); Monte Camelo (1☿); Nova Lima (2☿); Poço Fundo (1♀); Santana do Riacho (1☿) ; São Tiago (1♂); Timóteo (1♀); Uberlândia (1☿); Viçosa (19☿, 3♀); Vista Alegre (1☿). Pará: Belterra (6☿) ; Jari (2☿); Melgaço (1☿); Oriximiná (16☿); Paragominas (7☿, 1♀); Terra Santa (2♂). Paraná: Antonina (19☿, 1♀, 1♂) ; Foz do Iguaçu (4☿) ; Guaíra (2☿); Guarapuava (1☿); Ilha do Mel (1☿) ; Morretes (5☿); Parque Estadual do Palmito (1♀) ; Tibagi (4☿); Toledo (1☿); Turneiras do Oeste (10☿, 1♀) . Pernambuco: Recife (7☿) . Rio de Janeiro: Angra dos Reis, Ilha Grande (3☿) ; Itatiaia (3☿). Rondônia: Ouro Preto do Oeste (2☿) ; Porto Velho (19☿, 1♀).

Roraima: Amajari (3☿); Caracaraí (2☿). Santa Catarina: Ibirama (1☿); Florianópolis (5☿). São Paulo: Bálsamo (1☿); Biritiba Mirim (1☿); Botucatu (4☿); Ipeúna (2☿); Itirapina (2☿); Mirassol (3☿); Mogi das Cruzes (5☿); Monções (1☿); Morro Aprazível (3☿); Piracicaia (9☿); Rio Claro (2☿); Rio Grande da Serra (1☿); Salesópolis (4☿). Sergipe: Malhador (1☿); Nossa Senhora das Dores (1☿); São Cristóvão (3☿). BELIZE: [Cayo]: [St. Herman’s Blue Hole National Park], Caves Branch (3☿). COSTA RICA : Cartago: [no further data] (2☿). [Puntarenas]: Osa Peninsula (1☿). ECUADOR : Zamora-Chinchipe: Reserva Copalinga (1☿) . Morona-Santiago: Los Tayos (5☿). FRENCH GUIANA : Cayenne : Régina : Nouragues Natural Reserve (3☿, 3♂). Saint-Laurent-du-Maroni: Saül: Bélvedére de Saül (1☿). GUATEMALA : Izabal: 5km NW Morales (1☿). HONDURAS : Atlántida: Tela (1☿). MEXICO :

Chiapas: Tuxtla Chico (1☿) ; [Ocosingo] (1☿) . Vera Cruz: Reserva los Tuxtlas (3☿) . PANAMA: Chiriqui: 20.4km N of Sán Felix (1☿) . PERU: Cusco: Estación Biológica Villa Carmen (9☿) . Madre de Dios: Paquitza (2☿) ; Puerto Maldonado (2☿) . VENEZUELA: Bolívar: Cerro Los Pijiguaos (1☿) . Falcón: Cueva Zumbador , 36km W Chichiriviche (1☿) ; Finca Rita , 36.7km W Chinchiriviche (1☿) . Tachira: Pregonero (1☿) .

Odontomachus mormo Brown, 1976 ( Figures 9-C , 11-B , 60 , and 104)

Odontomachus mormo ( Brown, 1976) : 161 (☿).

Type specimens: Holotype [Images]. ECUADOR: Prov[ince] Guayas, 3km Sw Bucay, July 24’73 W Brown [col], mt – base for, MCZ Holotype 34795 1☿ [ MCZ] .

Paratype. ECUADOR: Prov [ince] Pichinca, 3 km E Tandapi, 1300 m., no. B-303, 1975, in wet ravine, [Peck], S. & Peck, J. col., MZSP 77724 1☿ [ MZSUP] .

Etymology: From the Greek word mormo = fear, frighten; a female spirit in the Greek folklore who was invoked by mothers or nurses to frighten children to keep them from misbehaving. The name probably refers to the notorious and potentially frightening size of this species, at the time, the largest described Odontomachus .

Diagnosis: (☿) Vertex trapezoidal, almost long than wide, smooth and shining. In dorsal view, central disc of pronotum with regular transverse striae; external striae forming concentric ellipses curved anteriorly. Petiole pedunculate, node angulate with blunt projection anteriorly. Body surface smooth and shining with inconspicuous short pubescence. (♀) Vertex, petiole and body surface as in worker. (♂) Unknown.

Worker

Paratype measurements: EL 0.65, HL 4.39, HoW 2.96, HvW 2.13, HW 2.43, ML 2.8, SL 4.76, PnW 1.27, WL 4.39, FL 4.18, PTH 1.43, PTL 0.84, PtW 0.42, GL 3.42, TL 15.84. CI 0.55, MI 0.63, PtI 1,70, SI 1,08, VI 0,87.

Non-type measurements: EL 0.6-0.7, HL 4.3-4.6, HoW 2.8-2.9, HvW 2.0-2.4, ML 2.7-2.9, SL 4.7-4.9, PnW 1.6-1.8, WL 5.9-6.3, FL 5.4-5.9, PTH 2.0-2.1, PTL 1.1-1.2, PTW 0.5-0.6, GL 4.4-4.7, TL 18.5-19.5. CI 0.51-0.53, MI 0.60-0.65, PtI 1.72-1.90, SI 1.04-1.12, VI 0.87-0.91 (n= 6).

Head. Vertex rectangular almost long than wide in frontal view; smooth and shining, regular longitudinal striae diverging posteriorly limited to frontal area. Posterior margin concave medially; occipital carina not forming nuchal collar. Cephalic constriction conspicuous. Frontal lobes Smooth and shining. Ocular bridges posteriorly forming arc. Eye relatively large (EL> 0.4); rounded, posterior margin rounded. Clypeus smooth and shining on median portion. Malar area smooth and shining in latero-oblique view.

Mandible. Almost as long as anterior head portion. External margin straight to slightly convex. Masticatory margin with denticles that serially increase in size apically. Palp formula 4,4.

Antenna. Scape long, surpassing occipital corner by more than pedicel and first flagellomere length.

Mesosoma. Pronotum in lateral view triangular, dorsal margin convex; neck well developed anteriorly in dorsal view; lateral margin rounded with obtuse angle posteriorly; central disc of pronotum with regular transverse striae slightly curved anteriorly, lateral striae forming concentric ellipses curved anteriorly. Propleuron smooth and shining. Mesonotum slightly wider than long, almost quadrate, with regular transverse striae. Mesopleuron with vestigial carina on anteroventral portion; mostly smooth and shining, anterior portion with regular transverse striae, posterior ventral portion forming triangular lobe anterior to middle coxa insertion. Mesometanotal sulcus well-marked, smooth and shining. Metanotum as long as half-length of mesonotum; with regular transverse striae. Metapleural gland bulla smooth and shining. Propodeum in lateral view slightly concave; posterior margin length about 1/4 of dorsal length. Metasternum with metasternal process open cuticle border fine medially separated; regular transverse striae curved posteriorly.

Petiole. Pedunculate. Angulate node, anterior margin in lateral view meeting dorsal margin in an almost straight angle with blunt projection; anterior margin short as high as subpetiolar process; dorsal margin slightly concave. Spine robust, long, slightly curved posteriorly, as long as dorsal margin half-length. Entire surface smooth and shining, with at most inconspicuous longitudinal to oblique striae on lateral face on ventral portion. Subpetiolar process longer than wider; tip rectangular to blunt; posterior portion not concave.

Gaster. First tergum longer than higher; anterior and dorsal margin forming a single convex surface; dorsal face of all terga smooth and shining.

Color and size. Body nut-brown. Gaster slightly darker, terga gradually becoming light brown posteriorly. Leg and antenna yellowish. Large size ants.

Pilosity: Inconspicuous short pubescence on body surface, separated each other more than 1.5x its length. Pubescence on metapleural gland bulla slightly denser than metapleuron. Short, subdecumbent hairs on gastral terga, except by first terga, shorter than procoxa width.

Natural history: The holotype and one paratype were collected in a ravine in a wet forest, and the second paratype was found in leaf a litter sample using Berlese funnel sample ( Brown, 1976). They were found respectively at 1,300 m and 680 m altitude. It occurs in Ecuador ( Fig. 60 ), and according to Rodriguez (2008), it also occurs in Colombia in areas from 515–1,500 m altitude.

Comments: Odontomachus mormo can be quickly recognized due to its unique petiole shape. Among the Neotropical species, only O. hastatus and O. davidsoni have pedunculate petioles and rounded ocular prominences, but the petiole of O. mormo has a blunt projection in the anterior region Also, the posterior portion of the vertex is almost as wide as the anterior portion, and the nuchal carina does not protrude posteriorly.

The only evident variation is the extent of the pronotum concentric striae on the dorsum. The petiolar spine can be slightly curved to almost straight as usual for the genus.

Material examined: 6☿. ECUADOR: Esmeraldas: Reserva biológica Bilsa (5☿). Reserva Río Canandé (1☿) .

Figures

FIGURE 1. Phylogenetic relationships of the Neotropical species of Odontomachus adapted from Fernandes et al. (2021). Colors representing the species group, yellow (hastatus group) and blue (haematodus group).

FIGURE 6. Variation of worker morphology of Odontomachus. In cephalic frontal view A, O. hastatus, mandibles elongated, vertex trapezoidal and long; B, O. chelifer mandibles short, vertex rectangular and short; C, O. bauri mandibles short and vertex trapezoidal. Pronotum in lateral view D, O. hastatus, triangular; E, O. biumbonatus angular; F, O. bauri, rounded. Petiole in lateral view G, O. hastatus, pedunculate and anteriorly elongated; H, O. chelifer sessile and angular; I, O. biumbonatus sessile and cylindrical; J, O. bauri sessile and conical. Gaster in lateral view K, O. hastatus, anterior and dorsal margins continuous forming a single convexity; L, O. chelifer and M, O. bauri, anterior margin straight forming blunt angle with dorsal convex margin. Dash lines and gray area representing shape of each structure. Scale bar = 1mm.

FIGURE 8. Petiole of workers in lateral view. A, pedunculate (O. hastatus); B, sessile (O. affinis). Dash lines highlighting anterior portion of petioles.

FIGURE 9. Vertexes of Odontomachus workers in frontal view. A, smooth and shining, nuchal collar present (O. hastatus); B, striate, nuchal collar present (O. davidsoni); C, smooth and shining, nuchal collar absent (O. mormo). Dash lines highlighting occipital carina.

FIGURE 10. Pronotum of Odontomachus workers in dorsal view. A, central disk smooth and shining (O. hastatus); B, central disk with conspicuous striae (O. davidsoni). Arrows pointing to the central portion of pronotum.

FIGURE 11. Ocular prominences of Odontomachus workers in frontal view. A, acute and projecting anteriorly (O. cornutus). Source: MZSP, imaged by: Mônica Ulysséa; B, rounded and not projecting anteriorly (O. mormo). Dash lines highlighting ocular prominence shapes.

FIGURE 18. Anterior face of petiolar nodes of Odontomachus workers in lateral view. A, bluntly angular, vertical anterior margin of petiolar node in lateral view longer than the half-length of petiolar sloping margin (O. chelifer); B, bluntly angular, vertical anterior margin of petiolar node in lateral view shorter than 1/3 length of petiolar sloping margin (O. bradleyi); C, conical (O. meinerti). Dash lines highlighting anterior face shape.

FIGURE 19. Metaesternal process of Odontomachus workers in ventral view, middle coxae on superior portion and hind coxae on inferior portion. A, metasternal process forming a pair of lateral cuticular borders widely separated each other (O. chelifer); B, metaesternal process forming a pair of triangular lobes almost as long as wide (O. dubius sp. nov.).

FIGURE 20. Pronotum of Odontomachus workers in lateral view. A, posterior portion of ventral margin of pronotum in lateral view with anteriorly curved long hairs, almost as long as the anterior procoxal hairs and propleuron with long erect hairs (O.laticeps); B, posterior portion of ventral margin of pronotum in lateral view without long hairs, propleuron with short decumbent hairs (O. bauri). Vertical arrows indicating the hairs on propleuron and horizontal arrows indicating the hairs on ventral portion of pronotum.

FIGURE 21. Petiolar node shapes of Odontomachus workers in lateral view. A, node cylindrical (O. xeta sp. nov.); B, node conical (O. haematodus). Dash lines highlighting anterior and posterior face shapes.

FIGURE 23. Odontomachus workers in lateral view. A, O. biolleyi (CASENT0907425, imaged by: Ziv Lieberman, Source: AntWeb.org. B, O. insularis. Arrows pointing to the petiole node spines.

FIGURE 24. Body color pattern of Odontomachus workers in lateral view. A, head ferruginous, strongly contrasting with the dark brown body (O. erythrocephalus); B, head and body dark brown almost black (O. bauri).

FIGURE 25. Mesopleuron of Odontomachus workers in lateral view. A, mesopleuron with anteroventral portion forming acute projection directed ventrally of (O. chicomendesi sp. nov.); B, mesopleuron without anteroventral portion forming acute projection directed ventrally of (O. cupreus sp. nov.). Dash lines highlighting the mesopleuron shape.

FIGURE 26. Metaesternal process of Odontomachus workers in ventral view, middle coxae on superior portion and hind coxae on inferior portion. A, metasternal process forming pair of spines (O. haematodus); B, metaesternal process not forming pair of spines of (O. bauri). Dash lines highlighting the metasternal process shapes.

FIGURE 27. Propleuron of Odontomachus workers in ventral view. A, propleuron with fine transverse striae (O. cupreus sp. nov.); B, propleuron smooth and shining with coarse lateral striae (O. ruginodis).

FIGURE 31. Propleuron of Odontomachus gynes in ventral view. A, propleuron with conspicuous striae (O. scalptus); B, propleuron smooth and shining with lateral striae restricted anteriorly of (O. haematodus).

FIGURE 32. Pronotum of Odontomachus gynes in lateral view. A, dorsal face anteriorly concave forming obtuse angle posteriorly (O. xeta sp. nov.); B, anteriorly straight becoming convex posteriorly (O. cupreus sp. nov.). Dash lines highlighting the pronotum shapes.

FIGURE 33. Pronotal striae of Odontomachus gynes in dorsal view. A, dorsum with striae strongly curved (O. scalptus); B, dorsum with striae dorsum with striae weakly curved (O. meinerti). White squares with lines representing the striation pattern of pronotum in dorsal view.

FIGURE 34. Metascutellum striae of Odontomachus gynes in dorsal view. A, with longitudinal striae weakly impressed (O. laticeps); B, smooth and shining (O. scalptus); C, with longitudinal striae well-marked converging posteriorly (O. caelatus). Arrows pointing to the mesoscutellum.

FIGURE 35. Axillae sculpturing of Odontomachus gynes in dorsal view. A, axilla with longitudinal striae (O. meinerti); B, axilla smooth and shining (O. haematodus). Arrows pointing to the axillae.

FIGURE 36. Petiolar nodes of Odontomachus gynes in lateral view. A, node smooth and shining (O. insularis); B, node with coarse striae (O. bauri).

FIGURE 38. Petiolar nodes of Odontomachus males in lateral view. A, pedunculate (O. hastatus); B, sessile with anterior face strongly convex (O. chelifer); C, sessile with anterior face slightly convex (O. affinis). Dash lines highlighting the shape of anterior face of petiole node.

FIGURE 39. Mesoscutellum shapes of Odontomachus males in antero-dorsal view. A, concave (O. chelifer); B, convex (O. haematodus). Dash lines highlighting the mesoscutellum shape.

FIGURE 40. Propodeum of Odontomachus males in posterior view. A, dorsal and posterior face separated by coarse striae, without concentrical striae (O. affinis); B, dorsal and posterior face not separated by coarse striae, with concentrical striae (O. laticeps). Arrows pointing to the propodeum striae patterns.

FIGURE 43. Propodeum and posterior face of petiole node of Odontomachus males.A, propodeum coarsely rugose and petiole node with apex acute and ascending striae (O. erythrocephalus); B, Propodeum with striae forming concentrical longitudinal arcs and transversal striae on lateral portion, petiole node with blunt apex without ascending striae (O. meinerti). Arrows pointing to the petiole node sculpturing and dash lines highlighting the petiole node apex shapes.

FIGURE 44. Mesoscutum sculpturing of Odontomachus males in dorsal view.A, with conspicuous oblique and irregular striae and posterior portion with coarse longitudinal striae (O. erythrocephalus); B, central portion with effaced striae and posterior portion without coarse longitudinal striae (O. insularis). Arrows pointing to the posterior portion of the mesoscuta.

FIGURE 46. Metasternal process of Odontomachus males in ventral view.A, forming a pair of spines (O. haematodus); B, with pair of lateral cuticle borders (O. bauri). Dash lines highlighting the metasternal process shapes.

FIGURE 58. Map of distribution of O. erythrocephalus and O. bauri. Triangles representing the type-localities.

FIGURE 60. Map of distribution of O. spissus, O. biolleyi, O. insularis and O. mormo. Triangles representing the typelocalities.

FIGURE 67. Map of distribution of O. chelifer, O. bradleyi and O. dubius sp. nov. Triangles representing the type-localities.

FIGURE 78. Map of distribution of O. cupreus sp. nov., O. ruginodis and O. chicomendesi sp. nov. Triangles representing the type-localities.

FIGURE 79. Odontomachus cornutus worker. A, head in frontal view; B, full lateral view; C, full lateral view. Source: MZSP, imaged by: Mônica Ulysséa.

FIGURE 80. Map of distribution of O. hastatus, O. cornutus and O. davidsoni. Triangles representing the type-localities.

FIGURE 81. Odontomachus cupreus holotype. A, head in frontal view; B, full lateral view; C, metasternum in ventral view with middle coxae at superior portion and hind coxae at inferior portion; D, mesosoma in dorsal view; E, petiole in lateral view; F, gaster in dorsal view.

FIGURE 82. Odontomachus cupreus gyne. A, head in frontal view; B, mesosoma in dorsal view; C, full lateral view.

FIGURE 83. Odontomachus dubius holotype.A, head in frontal view; B, full lateral view; C, metasternum in ventral view with middle coxae at superior portion and hind coxae at inferior portion; D, mesosoma in dorsal view; E, petiole in lateral view; F, gaster in dorsal view.

FIGURE 84. Odontomachus erythrocephalus worker. A, head in frontal view; B, full lateral view; C, metasternum in ventral view with middle coxae at superior portion and hind coxae at inferior portion; D, mesosoma in dorsal view; E, petiole in lateral view; F, gaster in dorsal view.

FIGURE 85. Odontomachus erythrocephalus gyne. A, head in frontal view; B, mesosoma in dorsal view; C, full lateral view.

FIGURE 86. Odontomachus erythrocephalus male. A, head in frontal view; B, mesosoma in dorsal view; C, propodeum in posterior view; D, full lateral view.

FIGURE 87. Odontomachus haematodus worker. A, head in frontal view; B, full lateral view; C, metasternum in ventral view with middle coxae at superior portion and hind coxae at inferior portion; D, mesosoma in dorsal view; E, petiole in lateral view; F, gaster in dorsal view.

FIGURE 88. Odontomachus haematodus gyne. A, head in frontal view; B, mesosoma in dorsal view; C, full lateral view.

FIGURE 89. Odontomachus haematodus male.A, head in frontal view; B, mesosoma in dorsal view; C, propodeum in posterior view; D, full lateral view.

FIGURE 90. Map of distribution of O. haematodus. Triangle representing the type-locality.

FIGURE 91. Odontomachus hastatus worker.A, head in frontal view; B, full lateral view; C, metasternum in ventral view with middle coxae at superior portion and hind coxae at inferior portion; D, mesosoma in dorsal view; E, petiole in lateral view; F, gaster in dorsal view.

FIGURE 92. Odontomachus hastatus gyne. A, head in frontal view; B, mesosoma in dorsal view; C, full lateral view.

FIGURE 93. Odontomachus hastatus male. A, head in frontal view; B, mesosoma in dorsal view; C, propodeum in posterior view; D, full lateral view.

FIGURE 94. Odontomachus insularis worker.A, head in frontal view; B, full lateral view; C, metasternum in ventral view with middle coxae at superior portion and hind coxae at inferior portion; D, mesosoma in dorsal view; E, petiole in lateral view; F, gaster in dorsal view.

FIGURE 95. Odontomachus insularis gyne. A, head in frontal view; B, mesosoma in dorsal view; C, full lateral view.

FIGURE 96. Odontomachus insularis male. A, head in frontal view; B, mesosoma in dorsal view; C, propodeum in posterior view; D, full lateral view.

FIGURE 97. Odontomachus laticeps worker.A, head in frontal view; B, full lateral view; C, metasternum in ventral view with middle coxae at superior portion and hind coxae at inferior portion; D, mesosoma in dorsal view; E, petiole in lateral view; F, gaster in dorsal view.

FIGURE 98. Odontomachus laticeps gyne. A, head in frontal view; B, mesosoma in dorsal view; C, full lateral view.

FIGURE 99. Odontomachus laticeps male. A, head in frontal view; B, mesosoma in dorsal view; C, propodeum in posterior view; D, full lateral view.

FIGURE 100. Map of distribution of O. meinerti and O. laticeps. Triangles representing the type-localities.

FIGURE 101. Odontomachus meinerti worker. A, head in frontal view; B, full lateral view; C, metasternum in ventral view with middle coxae at superior portion and hind coxae at inferior portion; D, mesosoma in dorsal view; E, petiole in lateral view; F, gaster in dorsal view.

FIGURE 102. Odontomachus meinerti gyne. A, head in frontal view; B, mesosoma in dorsal view; C, full lateral view.

FIGURE 103. Odontomachus meinerti male. A, head in frontal view; B, mesosoma in dorsal view; C, propodeum in posterior view; D, full lateral view.