Revision of the Tetragona clavipes (Fabricius, 1804) species-group (Hymenoptera: Apidae: Meliponini) Author Nogueira, David Silva 0000-0003-2045-0191 dsnogueira90@gmail.com Author Oliveira, Favízia Freitas De 0000-0003-4366-5005 favizia.freitas@ufba.br Author Oliveira, Marcio Luiz De 0000-0002-3950-1086 mlolivei@inpa.gov.br text Zootaxa 2022 2022-03-23 5119 1 1 64 http://dx.doi.org/10.11646/zootaxa.5119.1.1 journal article 151368 10.11646/zootaxa.5119.1.1 eb69b1e4-7442-4727-a42c-426584a1a846 1175-5326 6378560 F5425B3C-1877-4C3F-AE76-0CE37998437D Tetragona clavipes species-group Diagnosis (worker): clypeus yellow, with or without brown maculations, and sometimes divided by a longitudinal brown line. Basal triangle of the clypeus, when present, from dark brown to black. Paraocular area with yellow maculation, not extending above middle tangent of the antennal socket (except in T. perangulata ). Forewing with apical half of membrane hyaline (with whitish microtrichia) and basal half slightly yellow (with yellowish-brown microtrichia). Setae on the sixth tergum about 1.5x–2x or more the length of the setae of fourth tergum, more abundant than in fifth tergum (except in T. korotaii sp. nov. ). T2–T6 black with contrasting yellow band on apical third. TABLE 2. Comparative diagnosis of workers of species-group Tetragona clavipes ( Fabricius, 1804 ) .
Characteristics T. clavipes T. quadrangula T. perangulata T. mourei sp. nov. T. korotaii sp. nov.
Paraocular maculation below the epistomal suture X X X
Paraocular maculation in line to the epistomal suture X X
Paraocular maculation smoky superiorly X
Malar area black X X X X
Malar area yellow X
Metatibia with a black or dark brown area X X X
Metatibia completely yellow or yellowish-brown X X
T2 with a longitudinal yellow band wider than the diameter of the f lagellum X
T6 with branched hairs and simple setae mixed X X X X
T6 with only branched hairs X
FIGURE 1. Worker of Tetragona clavipes ( Fabricius, 1804 ) from Manacapuru (Amazonas, Brazil ). 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 2. Male of Tetragona clavipes ( Fabricius, 1804 ) from Bom Jardim de Goiás ( Goiás , Brazil ). 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 3. Physogastric queen of Tetragona clavipes ( Fabricius, 1804 ) from Ribeirão Preto ( São Paulo , Brazil ). 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 4. Heads of Tetragona clavipes ( Fabricius, 1804 ) in frontal view. A: from Atalaia do Norte (Amazonas, Brazil ). B: from Careiro (Amazonas, Brazil ). C: from Amajari ( Roraima , Brazil ). D: from Aripuanã ( Mato Grosso , Brazil ). E: from Acrelândia ( Acre , Brazil ). F: from Casanare ( Colombia ). G: from Campo Novo ( Rondônia , Brazil ). H: from Panguana ( Huánuco , Peru ). I: from Manacapuru (Amazonas, Brazil ). Scales: 0.5 mm . FIGURE 5. Lectotype of Centris clavipes Fabricius, 1804 . A: Right metatibia (A), metasoma in dorsal view (B), body in left lateral view (C), head in frontal view (D), body in right lateral view (E), labels (F). Scales: 0.5 mm (A, B and D) and 1 mm (C and E). FIGURE 6. Nest entrances of Tetragona clavipes ( Fabricius,1804 ) . Iranduba, Amazonas, Brazil (A). Cacoal, Rondônia , Brazil (B). Viçosa, Minas Gerais , Brazil (C). São Paulo , São Paulo , Brazil (D). Coeroeni, Suriname (E). Fonts: David Nogueira (A, C, D), Odair Diogo da Silva (B), Ayrton Vollet-Neto (E). 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 . TABLE 3. Nucleotide differences (%) between Tetragona and Ptilotrigona species found from the comparison of the sequences available in the genbank (https://www.ncbi.nlm.nih.gov/genbank/) provided by Rasmussen & Cameron (2010) . Bold = 16S; Italic = 28S. Differences greater than 8.63% were disregarded (-); *Northern Brazil; **Southeastern Brazil.
T. clavipes * T. clavipes ** T. dissecta T. perangulata T. ziegleri T. goettei T. atahualpa P. lurida P. pereneae
T. clavipes * 0 0 0 0 0.13 0.13 0.21 1.02 0.02
T. clavipes ** 1.89 0 0.39 0.52 0.52 0.52 0.27 1.42 0.55
T. dissecta 0.19 2.08 0 0.13 0.13 0.25 0.27 1.02 0.15
T. perangulata 2.46 3.78 2.65 0 0.26 0.26 0.27 1.15 0.28
T. ziegleri 5.68 6.43 5.87 5.49 0 0.37 0.13 0.86 0.99
T. goettei 4.56 5.31 4.75 4.18 5.89 0 0.38 1.11 0.23
T. atahualpa 8.63 8.1 - 8.29 7.94 8.33 0 1.02 0.02
P. lurida - - - - - - - 0 0.12
P. pereneae - - - - - - - 0.19 0
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 13. Worker of Tetragona quadrangula ( Lepeletier, 1836 ) from Teresina de Goiás ( Goiás , Brazil ). 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 14. Male of Tetragona quadrangula ( Lepeletier, 1836 ) from Lagoa da Confusão ( Tocantins , Brazil ). Metatibia (A), head in frontal view (B), metasoma in dorsal view (C) and body in lateral view (D). Scales: 0.5 mm (A, B and C) and 1 mm (D). 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). FIGURE 18. Holotype of Tetragona mourei sp. nov. from Bom Jesus do Piauí ( Piauí , Brazil ) 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 19. Paratype male of Tetragona mourei sp. nov. from Lizarda ( Maranhão , Brazil ). 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 20. Holotype of Tetragona korotaii sp. nov. from Coeroeni ( Suriname ). 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). Key of T. clavipes species-group species (workers) 1. Hairs or setae inconspicuous and sparse on T2–T5 ( Fig. 20C ), being longer on the apical area of the terga; central area of T6 only with short branched hairs, smaller than the diameter of the flagellum; mandibular condyle yellowish brown ( Fig. 20B ); malar area and contour of the clypeus yellow ( Fig. 20B ); only known from Suriname .......... Tetragona korotaii sp. nov. - Hairs or setae conspicuous and abundant on T2–T5 ( Fig. 13C ); central area of T6 with elongated simple setae, larger than the diameter of the flagellum; mandibular condyle black ( Fig. 13B ); malar area and contour of the clypeus black ( Fig. 13B ); Neotropical region and beyond.......................................................................... 2 2(1). Paraocular maculation blurry (smoky) yellow ( Fig. 18B ) or not superiorly ( Figs. 16B ), from the vicinity of the epistomal suture obliquely upwards towards compound eye in line or below the inferior tangent of the antennal socket.................. 3 - Paraocular maculation yellow from the vicinity of tentorial pit obliquely upwards towards compound eye in line or below the inferior tangent of the antennal socket.................................................................... 4 3(2). Paraocular maculation yellow, from the upper portion of the epistomal suture to compound eye to the middle tangent of the antennal socket ( Fig. 16B ); T2 with a yellow longitudinal central line wider or equal to the diameter of the flagellum ( Fig. 16C ); T3 with a thin longitudinal central yellow line; metatibia yellow with brown or black areas; found in Central America, Colombia and Ecuador .............................................................. Tetragona perangulata - Paraocular maculation smoky yellow superiorly, from the upper portion of the epistomal suture to compound eye to the inferior tangent of the antennal socket ( Fig. 18B ); T2 without yellow longitudinal central line, or with a yellow line less wide than the diameter of the flagellum ( Fig. 18C ); T3 without a thin longitudinal central yellow line; metatibia completely yellow; found in Brazilian Cerrado................................................................ Tetragona mourei sp. nov. 4(2). Metafemur, metatibia and metabasitarsus yellow ( Fig. 13A, D ); nest entrance usually in the form of a narrow vertical gap with landing area made of rigid cerumen around ( Fig. 15A ); found in eastern and southeastern Amazon, central and southeastern Brazil ............................................................................ Tetragona quadrangula - Metafemur, metatibia and metabasitarsus brown with at least one black marking ( Fig. 1A, D ); nest entrance very variable, usually in form of slit in tree trunk without projections ( Fig. 6A, B, D ); with wide distribution from the eastern Andes to Uruguay ................................................................................................................................................................................. Tetragona clavipes