Tetragona perangulata ( Cockerell, 1917 )

Nogueira, David Silva, Oliveira, Favízia Freitas De & Oliveira, Marcio Luiz De, 2022, Revision of the Tetragona clavipes (Fabricius, 1804) species-group (Hymenoptera: Apidae: Meliponini), Zootaxa 5119 (1), pp. 1-64 : 45-51

Tetragona perangulata ( Cockerell, 1917)

Figures 7B , 8B , 9B , 10B , 11B , 12 , 15B , 16 , 17 .

Trigona perangulata Cockerell, 1917: 125 [original description]; Lutz & Cockerell, 1920: 500 [geographic records, citation]; Wille, 1983: 60 [citation]; Ayala, 1999: 5 [type examined];

Trigona clavipes ; Salt, 1929: 443, 444 [uncertain identity, geographic records, nesting site, nest characteristics, behavior, habitat]; Nates Parra, 1983: 157 [uncertain identity, partim, geographic record, citation];

Trigona clavipes perangulata ; Schwarz, 1934: 5, 14 [key, geographic record]; Trigona (Tetragona) perangulata ; Schwarz, 1938: 472, 474 [taxonomic note, geographic records]; Wille & Michener, 1973: 14, 25, 35, 48, 59, 71, 83, 210–213, 265 [nesting site, nest entrance, figure 109, behavior]; Villa & Vergara, 1982: 150, 157, 158 [geographic records, nesting site]; Nates Parra, 1983: 157 [uncertain identity, geographic record]; Nates Parra & Cepeda, 1983: 69, 73 [behavior]; Vergara et al., 1986: 183 [partim, geographic record]; Michener, 1990: 125 [list]; Roubik, 1992: 503, 508, 515, 523 [key, geographic range, type examined, nesting site, nest entrance, behavior, figure]; Griswold et al., 1995: 684, 690 [nesting site, nest entrance, nest characteristics, list]; Smith-Pardo, 2003: 338 [list];

Trigona (Tetragona) clavipes perangulata ; Michener, 1954: 169 [synonymy of Trigona perangulata and Trigona clavipes subspecies perangulata , geographic records];

Trigona (Tetragona) clavipes ; Parra, 1984: 283 [uncertain identity, geographic record]; Nates-Parra, 2001: 240, 247 [partim];

Tetragona perangulata ; Moure, 1971: 48, 50 [taxonomic notes]; Camargo, 1988: 364 [list, geographic record]; Jarau & Barth, 2008: 274 [list, nesting site, comparative notes, geographic records, taxonomic characters, nest entrance]; Aguilar et al., 2013: 117 [geographic records]; Camargo & Pedro, 2013 [catalog]; Grüter, 2020: 71 [citation]; Nogueira et al., 2020: 67, 68 [taxonomic characters].

Diagnosis: worker ( Table 2 ): paraocular maculation reaching a superior line of the clypeus in comparison with T. clavipes , T. quadrangula and T. korotaii sp. nov., with the widest portion of the paraocular maculation on the level of the upper portion of the epistomal suture, narrowing obliquely upwards towards compound eyes with the narrowest portion of the maculation reaching or surpassing the level of the medium tangent of the antennal socket. Malar area black. Metatibia as in T. clavipes , with at least one black or dark brown maculation on wider region (except in callows specimens, which may be more yellowish). Black terga with contrasting yellow apical third and a broad yellow longitudinal band wider than the flagellum. Male: Sixth sternum in ventral view with a median recess strongly thin and elongated in the shape of an inverted V with a rounded bottom ( Fig. 7B ); with abundant and elongated setae, longer than the width of the median recess. Seventh sternum in ventral view with paraboloid lateral structure; with a median projection less wide than its distance to the lateral structure; median projection straight with curved and pointed apex. Eighth sternum elongated in ventral view, about 2x its width and with a median Y-shaped structure. Valve about 2x the length of the basal bulb. Gonocoxite in ventral view wider than long, with rounded apex. Long setae at the apex of the gonostylus, longer than the width of the gonostylus ( Figs. 7–11B ).

Description: male from Colón ( Panama) ( Fig. 7–11B , 17 ).

Coloration: integument predominantly yellowish-brown. With yellow paraocular maculation with the widest portion on the level of the upper portion of the epistomal suture, narrowing obliquely upwards towards compound eyes, with the narrowest portion of the maculation reaching the level of the medium tangent of the antennal socket. Clypeus and labrum yellow. Radicle yellowish-brown. Supraclypeal triangle yellow. Gena and malar area black. Mandible yellow with black condyles. Scape yellow. Pedicel yellowish-brown. Flagellum brown with yellowishbrown in ventral view of F2 to F11. Pronotal lobe and metepisternum yellow. Propodeum yellow circled in black and with longitudinal black line. Mesepisternum black with anterior yellow maculation. Epimeron black. Scutellum and axilla yellow. Proleg, mesoleg and metaleg yellowish-brown with tarsomeres brown. Metatibia with posterior line and apical fourth dark brown. Tegula yellowish-brown. Pterostigma and wing venation dark brown. Wing membranes slightly smoky. T1 brown with lateral black maculation. T2 and T3 with apical half yellowish brown and basal half dark brown with longitudinal band yellowish brown. T4 with basal third dark brown and apical 2/3 yellowish-brown. T5 dark brown with apical 2/3 yellowish brown. T6 and T7 yellowish brown. Sterna yellowish brown.

Punctation: piliferous punctures thin and sparse, except in the face very dense; integument predominantly smooth, polished and shiny between punctures, propodeal triangle impunctate, smooth, polished and shiny.

Pubescence: frons with yellow simple hairs (0.13 mm) and yellow branched hairs (0.05 mm). Some hairs behind genal area with curved apex (0.20 mm). Malar area with very short setae (0.01 mm). Scape with sparse yellow simple setae (0.05 mm). Vertex with yellow long simple and branched setae (0.28 mm). Supra clypeal area with erect simple setae (0.13). Clypeus in lateral view with yellow erect slightly curved simple setae (0.14 mm). Labrum with yellow simple setae (0.16 mm). Coxae with yellow simple hairs: procoxa (0.19 mm), mesocoxa (0.11 mm) and metacoxa (0.23 mm). Trochanters with yellow simple hairs: protrochanter (0.12 mm), mesotrochanter (0.12 mm) and metatrochanter (0.20 mm). Basal area of profemur with yellow simple hairs (0.13 mm), apical area with erect hairs (0.04 mm). Basal area of mesofemur with yellow simple hairs (0.19 mm), apical area with short erect hairs (0.08 mm). Metafemur with erect hairs on basal area (0.09 mm) and apical area (0.09 mm). Protibia (0.19 mm), mesotibia (0.22 mm) and metatibia (0.15 mm) with yellow simple setae on posterior border. Basitarsi with yellow simple setae: probasitarsus (0.19 mm), mesobasitarsus (0.15 mm) and metabasitarsus (0.18 mm). Mesoscutum with dark brown erect simple setae (0.19 mm) and yellow short branched hairs (0.06 mm). Mesoscutellum with dark brown simple hairs (0.32 mm). Propodeal triangle glabrous; sides of propodeum with yellow simple hairs (0.22 mm), and branched hairs (0.11 mm). Mesepisternum with yellow simple hairs (0.22 mm) and branched hairs (0.08 mm). Metepisternum with yellow simple hairs (0.71 mm) and branched hairs (0.05 mm). T1 glabrous with very short setae on basal third (0.01 mm). T2 with basal half glabrous and apical half with yellow simple erect setae (0.04 mm). T3 and T4 with yellowish-brown simple setae (0.06 mm and 0.0. 5 mm, respectively). T5 with simple (0.07 mm) and branched (0.05 mm) setae on central area and simple setae on sides (0.17 mm). T6 with simple (0.05 mm) and branched (0.03 mm) setae on central area and simple setae on sides (0.10 mm). T7 with yellowish-brown simple setae (0.04 mm). Sterna with yellow simple hairs (0.07 mm: 0.13 mm: 0.16 mm: 0.14 mm: 0.25 mm = length of S1, S2, S3, S4 and S5, respectively); some hairs on S3 and S4 with curved apex.

Structures: body length 6.29 mm. Head 1.34x wider than long (1.75 mm: 2.35 mm, length and width, respectively). Length of compound eye 3.46x breadth (1.49 mm: 0.43 mm, length and width of the compound eye, respectively). Inner orbits concave, converging downwards. Maximum interorbital distance less than the length of the eyes (1.43 mm: 1.49 mm). Flagellum diameter 4.2x malar area (0.17 mm: 0.04 mm). Clypeus slightly less than half its width (0.52 mm: 0.9 mm = length and width of clypeus respectively). Clypeus-ocellar distance 1.03 mm. Interalveolar distance slightly less than the alveolorbital distance (0.19 mm: 0.27 mm = interalveolar distance and alveolorbital distance, respectively). Interocellar distance greater than ocellorbital distance (0.45 mm: 0.33 mm). Scape length 3.2x its width (0.69 mm: 0.21 mm). Flagellum wider than the pedicel (0.17 mm: 0.16 mm). Mandible with two teeth, the first shorter and further back than the second. Spine of protibial strigillae (0.06 mm). Mesotibial spur 0.30 mm long. Metatibia with apical paraboloid border; length of metatibia 3.2x its width (2.18 mm: 0.67 mm). Metabasitarsus 2x longer than broad (0.74 mm: 0.36 mm). Intertegular distance 1.45 mm. With 5 hamuli on posterior wing.

Variations (workers): some specimens from Colombia and Ecuador have yellowish metatibia and metabasitarsus. In addition, these specimens also present the paraocular maculation in upper part, which differentiates them from T. quadrangula and T. mourei sp. nov.. These specimens are likely to be callows bees.

Geographic distribution ( Fig. 12 ): Colombia (Antioquia, Boyacá, Caldas, Cauca, Cesar, Córdoba, Cundinamarca, Huila, Magdalena, Santander, Tolima, Valle del Cauca ( Camargo & Pedro, 2013)) , Costa Rica (Cartago, Guanacaste ( Camargo & Pedro, 2013) Puntarenas, San José) , Ecuador (Esmeraldas, Imbabura, Napo) , Panama ( Barro Colorado , Colón, Chiriquí, Darién ( Camargo & Pedro, 2013)) .

Biological notes: like other species of the clavipes species-group, T. perangulata has strong defensive behavior. The nest entrance was described by Wille & Michener (1973) and Roubik (1992) as being elongated, black and ovoid, in the shape of a funnel projected out of the tree with brittle resin. Wille & Michener (1973) reported a nest in an anthropized area, in the trunk of a Ficus tree, seven meters from the ground, and commented that the nest entrance can be easily confused with various types of fungi in humid areas ( Fig. 15B ).

Examined material: workers (47): COLOMBIA: 1 (ZSM), Süda[?] America [?], M.W. / M.W., 1858., 82 / Tr. Quadrangula [sic], 1900 Friese det. / Trigona clavipes F., E. Clément det. [probably from Colômbia]; Antioquia: 1 (UNAL), 22626 / Co Ant. R. Cler [?], Bgve [?] de la Vuedei [?], I. Arvila [?], 10-III-07 / [verse] LABUN 015045; 3 (UFV), COLÔMBIA, Antioquia, Medellin; 1 (UNB), UNB 028998 / COLOMBIA, Antioquia, Santa Fé, 06°32.09’N e 75°49.833’W, vii.2008, A. Aguiar col.; Boyacá: 1 (UNAL), 8207 / ICN-MHN HY-2955 / I / 69, Pto. Boyacá, Vereda El Pescado, Restreyo-Mejíz [?] / [verse] ICN 011224; 2 (RPSP), COLOMBIA, 1976, GUIOMAR leg.; 1 (RPSP), idem except: Y; 1 (RPSP), idem except: Tetragona clavipes cf. perangulata Cockerell, Camargo Det. Camargo ; Caldas: 1 (ZSM), La Dorada, Kauda., coll. [?] Prinz. Therese / La Dorada, -Kauda / Sammlung I.K.H, Prinzessin Therese von Bayern / Trigona clavipes F., E. Clément det.; 1 (ZSM), La Dorada, Kauda., coll. [?] Prinz. Therese / Sammlung I.K.H, Prinzessin Therese von Bayern / Trigona clavipes F., E. Clément det.; Cauca: 1 (UNAL), 13392 / ANTIOQUIA-BETULIA, CERCA AL RIO CAUCA, EM PASTIZAL Y COMPOSITAE, COL: FRANCO-O. CEPEDA, 29-XII-84 / [verse] LABUN 004622; Cesar: 1 (UNAL), 20391 / COL. Cesar Pueblo Bello 1085, msnm 11-05-08 Col:, M. Cepeda; Córdoba: 1 (UNAL), 2345 / MAGDALENA, EL CAMPANO, 1300 M, COL: R OSPINA [?], 26-vi-87 / [verse] LABUN 004126; Cundinamarca: 1 (UNAL), 8225 / ICN-MHN HY-2973 / Pandi, Cund., em Maracuyá, Col. G. Nates, 12.III.77 / [verse] ICN 011242; Huila: 1 (UNAL), 29407 / COLOMBIA, Huila, Rivera., N 2°46’98.9”W 75°16’3.24”, 648m. Colector Rose Trindade, Captura: rede entomológica / Tetragona sp. , D. Guevara 2017 / [verse] Tetragona ; Magdalena: 1 (UNAL), 22235 / COL. Magdalena. Cienaga Siberia, (Agua Linda) 21/09/2010, Col: J. Medina / [verse] LABUN 014945; 1 (UNAL), 2366 / Parque Tayrona, Col: Villa, Nov-6, 76 / [verse] LABUN 004132; 1 (UNAL), 21073 / 13 / Cienaga Magdalena, SNSM voa [?] Scin [?] Jainer [?], 1400 msnm, Col. D. Obiegor [?] / [verse] LABUN 015743; Santander: 1 (UNAL), 21845 / COL. Sant. Lebrija. Sn. Nicolás 21-11-09 Col: A Rodriguez / [verse] LABUN 015742; 1 (UNAL), 21294 / Colombia. Santander., Floridablanca. Casiano. Finca las Brisas, 1160m 2octu2004, Col: G. Nates & A. Parra / [verse] LABUN 015035; Tolima: 1 (UNAL), 13316 / TOL. ICONONZO. AL VUELO, COL: NATES, 20.III.78 / [verse] LABUN 004546; COSTA RICA: Cartago: 1 (ZMUC), Collection, Sehild-Burgdorf, Costa Rica, Turrialba / Coll. Wüstnei; Puntarenas: 2 (RPSP), COSTA RICA, (S.) Puntarenas, Prov., Gromaco, 34km. SE. of Potrero Grande, on Rio Coto Brus. 21 July 1963, 1000 ft. (C. D. Michener & W. Kerfoot) / T. (Tetragona) perangulata Ckll. , det.A. Wille 1972; 1 (RPSP), EL DOMINICAL, P. PUNTARENAS, COSTA RICA, 6 JULIO 1960 / T. (Tetragona) clavipes perangulata Cokerell, Det. A. Wille ; 1 (RPSP), EL DOMINICAL, P. PUNTARENAS, COSTA RICA, 6 JULIO 1960, A WILLE / T. (Tetragona) clavipes perangulata Cokerell, Det. A. Wille ; San José: 1 (RPSP), El Rey., San José; Costa Rica., 2-9-1961, Col. A. WILLE / Trigona (Tetragona) perangulata Cockerell, Det. A. Wille ; ECUADOR: Esmeraldas: 1 (QCAZ), ECUADOR ESMERALDAS, Kumanii lodge 41m, 78°55’15”W 00°45’14”N, 12–15APR2005 K. Vivanco, EX: Barrido de red / QCAZ I 259375; 1 (QCAZ), Ecuador IV-83, ROCAFUERTE ESMERALDAS SDM., Legit: EF. BRLONES[?] / CAPTURADA COM PANAL[?] / QCAZ I 26932; 1 (RPSP), ECUADOR, ESMERALDAS, PITZARA.III.85, LEG. G. ONORE / Ry “MARACUYÁ” / Tetragona sp. n., Det. Camargo 1984 / T. perangulata (Cock. 1917) , Det. Camargo, 2000; Imbabura: 1 (QCAZ), Ecuador LITA, IMBABURA, XI- 86, Legit: F.RODRIGUEZ / QCAZ I 27238; 1 (QCAZ), idem except: QCAZ I 27249; 1 (QCAZ), Ecuador Lita, IMBABURA, XI-86, Legit: X.Permíno / QCAZ I 27044; 1 (QCAZ), ECUADOR, Imbabura —Lita, 710 m. s.n.ni, I-XI-86, Legít: Fco Jarrin / QCAZ I 27039; 1 (QCAZ), idem except: QCAZ I 27045 / Tetragona sp 2 ; 1 (QCAZ), Ecuador, Imbabura-Lita, 730m, XI-1986, Legit: R. Ramires / QCAZ I 26935; 1 (QCAZ), Ecuador, Imbabura- Lita, XI-86 100 mts, Legit: L. Torres / QCAZ I 26934; 1 (QCAZ), Ecuador, Imbabura-Lita, XI-86 600m, Legit: L. Torres / QCAZ I 27329; Napo: 1 (QCAZ), Ecuador, NAPO, Arquidona, XII-86, Legit: M. Ferro / QCAZ I 26871; 1 (QCAZ), idem except: QCAZ I 26872; PANAMÁ: Barro Colorado: 1 (RPSP), Barro Colorado I, Canal Zone, Panama, 17.III.1956 No., C.W. & M.E. Rettenmeyer / T. (Tetragona) perangulata Ckll. , det. A. Wille 1972; Colón: 1 (STRI), PANAMA: Colón Prov., Colon, km San Lor., 20 Sep. 1979, D. Roubik coll. # 26; Chiriquí: 1 (NHM), Bugaba, Panama., Champion. / Trigona (Tetragona) clavipes var. perangulata Cockerell, Det. H.F.Schwarz / NHMUK 010812306; 1 (NHM), idem except: NHMUK 010812307.

Male (2): PANAMA: Colón: 2 ( STRI) , ♂ / PANAMA: Colón Prov. Puerto Pilon 25 km NE, 22 Ago 1984, D. Roubik No .

Figures

FIGURE 7. Sixth sternum of Tetragona Lepeletier & Serville, 1828 species in ventral view: T. clavipes (Fabricius, 1804) (A), T. perangulata (Cockerell, 1917) (B), T. quadrangula (Lepeletier, 1836) (C), T. mourei sp. nov. (D), T. essequiboensis (Schwarz, 1940) (E), T. handlirschii (Friese, 1900) (F), T. truncata Moure, 1971 (G), T. goettei (Friese, 1900) (H), T. dorsalis (Smith, 1854) (I) and T. ziegleri (Friese, 1900) (J). Scale: 0.5 mm.

FIGURE 8. Seventh sternum of Tetragona Lepeletier & Serville, 1828 species in ventral view and apex of the median projection in lateral view: T. clavipes (Fabricius, 1804) (A), T. perangulata (Cockerell, 1917) (B), T. quadrangula (Lepeletier, 1836) (C), T. mourei sp. nov. (D), T. essequiboensis (Schwarz, 1940) (E), T. handlirschii (Friese, 1900) (F), T. truncata Moure, 1971 (G), T. goettei (Friese, 1900) (H), T. dorsalis (Smith, 1854) (I) and T. ziegleri (Friese, 1900) (J). Scale: 0.5 mm.

FIGURE 9. Eighth sternum of Tetragona Lepeletier & Serville, 1828 species in ventral view: T. clavipes (Fabricius, 1804) (A), T. perangulata (Cockerell, 1917) (B), T. quadrangula (Lepeletier, 1836) (C), T. mourei sp. nov. (D), T. essequiboensis (Schwarz, 1940) (E), T. handlirschii (Friese, 1900) (F), T. truncata Moure, 1971 (G), T. goettei (Friese, 1900) (H), T. dorsalis (Smith, 1854) (I) and T. ziegleri (Friese, 1900) (J). Scale: 0.5 mm.

FIGURE 10. Genital capsule of Tetragona Lepeletier & Serville, 1828 species in dorsal and ventral view: T. clavipes (Fabricius, 1804) (A), T. perangulata (Cockerell, 1917) (B), T. quadrangula (Lepeletier, 1836) (C), T. mourei sp. nov. (D), T. essequiboensis (Schwarz, 1940) (E), T. handlirschii (Friese, 1900) (F), T. truncata Moure, 1971 (G), T. goettei (Friese, 1900) (H), T. dorsalis (Smith, 1854) (I) and T. ziegleri (Friese, 1900) (J). Scale: 0.5 mm.

FIGURE 11. Valve and basal bulb of Tetragona Lepeletier & Serville, 1828 species in lateral view: T. clavipes (Fabricius, 1804) (A), T. perangulata (Cockerell, 1917) (B), T. quadrangula (Lepeletier, 1836) (C), T. mourei sp. nov. (D), T. essequiboensis (Schwarz, 1940) (E), T. handlirschii (Friese, 1900) (F), T. truncata Moure, 1971 (G), T. goettei (Friese, 1900) (H), T. dorsalis (Smith, 1854) (I) and T. ziegleri (Friese, 1900) (J). Scale: 0.5 mm.

FIGURE 12. Records of the occurrence of clavipes species-group of Tetragona Lepeletier & Serville, 1828. Scale in kilometers.

FIGURE 15. Nest entrance of Tetragona quadrangula (Lepeletier, 1836) from Goiás, Brazil (A). Draw of nest entrance of Tetragona perangulata (Cockerell, 1917) in top view with an arrow indicating the entrance (B). Fonts: Adriano Gonçalves (A) and David Nogueira (B).

FIGURE 16. Worker of Tetragona perangulata (Cockerell, 1917) from Puntarenas (Costa Rica). Metatibia (A), head in frontal view (B), metasoma in dorsal view (C) and body in lateral view. Scales: 0.5 mm (A, B and C) and 1 mm (D).

FIGURE 17. Male of Tetragona perangulata (Cockerell, 1917) from Colón (Panama). Metatibia (A), head in frontal view (B), metasoma in dorsal view (C) and body in lateral view. Scales: 0.5 mm (A, B and C) and 1 mm (D).