Lateral ocellus separated from the eye by two times its diameter. Compound eyes with bristles. Clypeus with acute apex, lateral margins parallel in part upper and rectangular lateral lobes. Short malar space. Occiput with carina. Labial palpus with four segments. Proepisternum with reduced lateral carina. Pronotal carina limited to a small length in the center of the pronotum and acute. Pronotum with lateral fovea. Mesoscutum with reduced posterolateral lamella. Dorsal groove of the mesoepisternum present and transverse to the sclerite. Metasomal tergum I, in dorsal view, with lateral margins diverging gradually from the base to apex.

Considering the Gymnopolybia description (currently Agelaia) from Stelopolybia species (currently Angiopolybia) by Ducke (1914), these genera are not well-defined by the proposed characters. Based on this, some morphological characters that had been proposed to separate Angiopolybia from Agelaia such as stelocyttarous nest (with envelope) (Ducke 1914), very weak or absent occipital carina (Richards 1943), and flat metapleural basalar area (Richards 1978; Andena et al. 2007) were not considered here because they are not exclusive to Angiopolybia, since they are present in few species of Agelaia. Moreover, the scutellum with line or depression (Richards 1943; Andena et al. 2007) was not considered because they are not exclusive to Agelaia, since some Angiopolybia species present the line, despite it is inconspicuous. Richards (1943) commented that the characters determined by him, isolated, had no effect on separating the taxa.

Based on previous and this study, A. pallens and A. zischkai are closer, as are A. paraensis and A. obidensis. Angiopolybia pallens and A. zischkai are distinguished by rounded gena and propodeum with posterior submedian translucent mark inserted in a depression, whereas the other species have angled gena and submedian translucent mark of the propodeum not inserted in a depression. We documented other differences in the step 1 of the identification key for the genus (see below).

Bolivia, Brazil, Colombia, Costa Rica, Ecuador, French Guiana, Guyana, Panama, Peru, Suriname, Trinidad and Tobago, Venezuela.

Cayenne, French Guiana.

Anterior wing of 7–8.5 mm; eyes with medium-sized and sparse bristles; rounded gena; pronotal lamella low on the anterior margin, one fifth of the height of antennal socket; pronotal lobe developed in the anterior lateral region, below the pronotal fovea; defined and deep pronotal fovea; axillary fossa with anterior margin directed to the posterior region; posterior submedian translucent mark of the propodeum inserted in a round depression; basal metapleural area with parallel upper and lower margins.

Size. (1) Head 1.27 mm long, 2 mm high, and 2.27 mm wide; (2) mesosoma 3.50 mm long, anterior wing 8.11 mm long, and posterior wing 5.27 mm long; (3) metasoma 5.53 mm long. Head. (1) Lateral ocelli with 0.15 mm and median ocellus with 0.17 mm of diameter, not inserted in a declivity of the vertex, and the lateral ocelli separated from the eyes for twice its diameter. (2) Compound eyes with medium-sized and sparse bristles. (3) Frons with interantennal space about twice the height of antennal socket. Anterior tentorial fovea closer to the antennal socket than to the eye. The central region of the frons with long bristles. (4) Antennal socket 0.22 mm high. (5) Clypeus as high as wide, contact with eyes for a distance greater than twice the height of antennal socket, and lateral lobe touching the eye. Medium bristles in the basal half and long bristles in the apical half. (6) Gena with half of the width of the eye at the level of the ocular sinus. Mesosoma. (1) Anterior lamella of pronotum with a height of one fifth of the height of antennal socket. Pronotal fovea with a circular shape, deep and with slight anterior prominence. (2) Mesoscutum convex and as long as wide. (3) Tegula 1.7 times longer than wide. (4) Axillary fossa with anterior margin directing to the posterior region. (5) Propodeum with translucent posterior submedian mark, anterior to the propodeal valve, inserted in a round depression. Propodeal valve complete and expanded, median region with half of the height of antennal socket. (6) Anterior wing with prestigma 1.4 times longer than wide. (7) Posterior wing with eight hamuli. Metasoma. (1) Metasomal tergum I two times longer than broad. Tergum with angulation in the posterior third, in lateral view. (2) Metasomal tergum II 0.82 times longer than broad. Color. Brown in general. Yellow: lateral of the apex, lateral of the front, interantennal region, disc of the clypeus, mandibles, gena, antennal segments: scape, dorsal of FL6 (flagellomere 6) and 7, FL8–10, band contouring the posterior margin of the pronotum, tegula, thin bands in the posterior margins of the metasomal terga I, II and III, and metasomal sterna II and III. Yellowish brown: central longitudinal and lateral region of the mesoscutum, tibiae and tarsi, femora median and posterior, metasomal tergum I, anterior half of metasomal tergum II, metasomal sterna I–IV. Black: FL1–5, ventral of FL 6 and 7, metasomal terga 4–6 and metasomal sterna 5 and 6. Wings with hyaline cells, pterostigma and venation in general yellowish-brown, except brown in the veins C, Sc+R, M+Cu and M.

Size. (1) Head 1.01 mm long, 1.83 mm high, and 2.09 mm wide; (2) mesosoma 2.87 mm long, anterior wing 7.1 mm long, and posterior wing 4.5 mm long; (3) metasoma 6.08 mm long. Head. (1) Lateral ocelli with 0.17 mm and median ocellus with 0.18 mm of diameter. (2) Compound eyes with small-sized and sparse bristles. (3) Frons with interantennal space with 1.85 times the height of antennal socket. Anterior tentorial fovea closer to the eye that to the antennal socket. (4) Antennal socket with 0.21 mm high. (5) Clypeus as high as wide and apex less acute than in the female. Pubescence stronger than in the female. (6) Gena with half of the width of the eye at the level of the ocular sinus. Mesosoma. (1) Mesoscutum 0.9 times longer than wide. (2) Propodeum with translucent posterior submedian mark, anterior to the propodeal valve, inserted in a not round depression. (3) Anterior wing with prestigma 1.6 times longer than wide. Metasoma. (1) Metasomal tergum I 1.9 times longer than broad. (2) Metasomal tergum II 0.8 times longer than wide. Genitalia (Fig. 6a–e). Paramere 1.6 mm long and 0.6 mm wide; parameral spine with one fifth of the paramere, curved upwards and with small-sized and sparse bristles; lobe with a rounded apex and slightly curved downwards. Aedeagus 1.2 mm long; enlarged valve with a small emargination in the tip; apical portion 0.42 mm long and straight, ventral margin with denticles directed for the anterior region; denticulation decreasing in size from the apex to the base and more sclerotized than the rest of the apical portion; small-sized bristles with alveolar base, closer in the lower half and sparse in the upper half; median expansion without denticles and with acute apex; lateral apodeme not flattened dorsoventrally at the apex; basal apodeme arched to the venter. Digitus 2.5 times longer than wide; apical process not curved in the region of the upper half and with bristles of alveolar base small and sparse; rounded anteroventral lobe with a strip of black scale-like bristles crossing it obliquely at the base of the digitus; bristles absent in the basal articulation. Cuspis approximately 0.48 mm long, with five black scale-like bristles on the lateral lobe, and small bristles with alveolar base and close throughout the area of the cuspis, except sparse in the central region and on the ventral margin.

Anterior wing between 7–8.5 mm in length. Anterior margin of the pronotum (below the fovea) more curved. Pronotum, in lateral view, with frontal region more projected forward. Coloration varied between populations from black with yellow marks [like A. pallens dark morph (sensu Richards, 1978)] to yellowish brown.

The nest of A. pallens was initially described by Möbius (1856) for Polybia ampullaria (junior synonym of the taxon), and later described by Wenzel (1998) as a nest with a flask-shaped envelope and with a long downward entrance with its hole horizontally; pedicel initially single and later being able to be multiple; flexible card with long fibers, usually yellow or amber; and adjacent combs, suspended or fused from each other and without contact with the envelope. Additionally, the following nest variation was found: built on an irregular substrate (thin branch with several leaves), connected to the substrate by a central pedicel (thick) and several support pedicels (fines); 11 combs overlapping and also connected by a central pedicel and multiple support, with the third and fourth combs with the largest circumferences and combs decreasing in circumference towards the ends, and without contact with the envelope; hexagonal cells of the combs with diameter of 2.5 mm; single envelope with long fibers arranged longitudinally, circumference gradually decreasing towards the entrance and without entrance of tubular shape.

Angiopolybia pallens is distinguished by the presence of a lobe in the lateroanterior region of the pronotum, absent in A. zischkai, and metapleural basalar area with parallel upper and lower margins, diverging in A. zischkai. Some A. pallens specimens can be confused with A. zischkai because it can resemble the typical form of A. zischkai, of color darker.

Despite the geographic disjunction by thousand kilometers in the distribution of A. pallens between the Amazon and Atlantic Forest (Carvalho et al. 2014, 2021), considering the morphological analysis of specimens collected in both biomes, we verified that the differences, when found, are very subtle and follow the variation found between different populations. Considering the populations from the Atlantic Forest, the color varied from yellow with brown marks to completely brown, while among the populations from the Amazon Forest it varied from black with yellow marks to yellowish brown, with some forms existing in both biomes. The occipital carina present in A. pallens, despite weak or not, found in this study not reported by Richards (1978) and Andena et al. (2007). They treated the occipital carina absent for the taxon. The prestigma longer than wide, also, was not cited by Richards (1978) and Andena et al. (2007). However, this character was found by Silveira and Carpenter (1995) for some specimens, and here we also found the two forms, prestigma about as long as wide and longer than wide. The redescription of A. pallens was made based on the lectotype of Polybia infernalis de Saussure, 1854 (junior synonym of A. pallens). The information about the male specimen described is: BRA, Roraima, Amajarí, Serra do Tepequém, SESC Tepequém. 1–16.i.2016 / Malaise grande, J.A. Rafael, F.F. Xavier Filho col.

♀, TYPE / MUSEUM PARIS, Amérique, Leprieur 1834 / Polybia infernalis Sauss., Type. / 289694 / Polybia infernalis Sauss., Lectotype ♀, Richards, 1971 (MNHN, Paris), record MNHN, Paris EY25588 (Fig. 1a, 1b, 1c). Polybia infernalis is a junior synonym of Rhopalidia pallens Lepeletier, 1836, but here we used it to the redescription of the species. Type specimen analyzed by images.

We examined 469 females and 32 males for A. pallens; see supplementary material S1.

Bolivia: Beni, Cochabamba; Brazil: Acre, Alagoas (new record), Amapá, Amazonas, Bahia, Ceará, Espírito Santo, Maranhão, Mato Grosso, Pará, Pernambuco, Piauí, Rio de Janeiro, Rondônia, Roraima, Santa Catarina, São Paulo Sergipe; Colombia: Meta, Caquetá, Nariño, Vaupés, Putumayo, Amazonas; Ecuador: Esmereldas, Napo, Pichincha; French Guiana; Guyana; Panama; Peru: Loreto, San Martin, Huánuco, Pasco, Junín, Cuzco; Suriname; Trinidad and Tobago; Venezuela: Monagas (Fig. 10a).

Zumbi, Ecuador.

Anterior wing of 8–9.5 mm; eyes with medium-sized and sparse bristles; rounded gena; pronotal lamella low on the anterior margin, one fifth of the height of antennal socket; pronotal lobe not developed in the lateral anterior region, below of the pronotal fovea; defined pronotal fovea, but little deep; axillary fossa with anterior margin directed to the posterior region; posterior submedian translucent mark of the propodeum inserted in a round depression; metapleural basalar area with divergent upper and lower margins.

Size. (1) Head 1.03 mm long, 2.09 mm high, and 2.24 mm wide; (2) mesosoma 3.54 mm long, anterior wing 9.42 mm long, and posterior wing 6.10 mm long; (3) metasoma 5.9 mm long. Head. (1) Lateral ocelli with 0.16 mm and median ocellus with 0.18 mm of diameter, not inserted in a declivity of the vertex and the lateral ocelli separated from the eyes for twice its diameter. (2) Compound eyes with medium-sized and sparse bristles. (3) Frons with interantennal space with 1.75 times the height of antennal socket. Anterior tentorial fovea closer to the antennal socket than to the eye. Central region of the frons with long bristles. (4) Antennal socket 0.24 mm high. (5) Clypeus 0.9 times higher than wide, contact with eyes for a distance greater than twice the height of antennal socket, and lateral lobe touching the eye. Long bristles all over the clypeus and very long bristles on the apical margin. (6) Gena with half of the width of the eye at the level of the ocular sinus. Mesosoma. (1) Anterior lamella of pronotum with height of one fifth of the height of antennal socket. Pronotal fovea with ellipsoid shape, little deep and with slight anterior prominence. (2) Mesoscutum subconvex and as long as wide. (3) Tegula 1.5 times longer than wide. (4) Axillary fossa with anterior margin directing to the posterior region. (5) Propodeum with translucent posterior submedian mark, anterior to the propodeal valve, inserted in a round depression. Propodeal valve complete and expanded, median region with two thirds of the height of antennal socket. (6) Anterior wing with prestigma as long as wide. (7) Posterior wing with eight hamuli. Metasoma: (1) Metasomal tergum I 2.1 times longer than wide. Tergum with angulation in the posterior third, in lateral view. (2) Metasomal tergum II 0,82 times longer than broad. Color. Dark brown in general. Yellow: lateral of the vertex, lateral of the frons, clypeus (but dark brown disc), interantennal region, mandibles, malar space, gena, slender band contouring the posterior margin of the pronotum, outer margins of the tegula, wide spot at along the anterior margin of the scrobal furrow and along the dorsal groove of the mesoepisternum, anterior half of the metanotum, submedian longitudinal band on the propodeum, lateral margin of the propodeum, upper region of the metapleural basalar area. Yellowish brown: FL7–10 of the antenna, anterior and median coxae, trochanters, femora, tibiae and tarsi (but with dorsal brown spots). Black: ocellar area, FL1–6 of the antenna, mesoscutum and metasoma. Wings with hyaline cells, except yellow in the costal, medial, submarginal I and marginal; and yellowish-brown venation, except brown in the veins C, Sc+R, M+Cu, M and in the beginning of the Cu.

Unknown.

We found a specimen with yellow color and black marks, and the abdomen, apparently, wider in dorsal view.

Not described, but Valverde et al. (2019) in the identification key of social wasps from Costa Rica commented that the nest envelope resembles an inverted flask.

Angiopolybia zischkai resembles A. pallens, but it is distinguished by the pronotum without a developed lobe in the lateroanterior region, which is present in A. pallens; metapleural basalar area with divergent upper and lower margins, which are parallel in A. pallens; and pronotal fovea with translucent mark of elliptical shape, which is circular in A. pallens.

One specimen designated as holotype of Angiopolybia brunnescens (Fig. 7h) but not described by Richards (specimen deposited in the NHM) is an A. zischkai specimen with more yellowish coloration, similar to the color of some A. pallens. The two specimens with the labels: Paratype / PERU: 1.609, Maracapata [Marcapata] (Perú) / Stelopolybia infernalis Ducke rev.11. [1911] (1 ♂, MZUSP), and 1.609, Maracapata [Marcapata] (Perú) / Stelopolybia infernalis Ducke rev.11. [1911] / A. pallens (Lep.) f. zischkai, Rich. 4 ♀ (1 ♀, MZUSP), that are two of the paratypes of A. zischkai, are specimens of A. pallens with coloration resemble to the holotype of A. zischkai. Although Gomes et al. (2018, 2020) reported A. zischkai samples from the Brazilian states Rondônia and Acre, we did not find A. zischkai for these regions.

♀, Holotype / Zumbi, Rio Zamora, 700M, Ecuador / XI.2.41, D.B.Laddey / Angiopolybia pallens ssp., zischkai Rich., ♀ Holotype / AMNH_IZC 00332335 (AMNH, New York) (Fig. 3). Type specimen analyzed by images.

Paratype: Paratype / PERU: Dept. Huanuco, Divisoria, 7.viii.1949, J.M.Schuncke., B.M.1952-645 / Angiopolybia zischkai Rich., ♀, paratype (1 ♀, NHM).

We examined three females of A. zischkai; see supplementary material S1.

Bolivia: Cochabamba; Colombia: Amazonas, Cundinamarca, Magdalena, Putumayo, Valle del Cauca (new record); Costa Rica: Heredia; Ecuador: Orellhana, Zamora-Chinchipe; Panama: Colón; Peru: Cuzco, Huánuco, Junín, Loreto, Pasco, Ucayali (Fig. 10b).

Óbidos, Pará, Brazil.

Anterior wing of 12–14 mm; eyes with very small-sized and sparse bristles; angulate gena with enlarged lower and upper region; pronotal lamella very elevated along the anterior margin, one third of the height of antennal socket; axillary fossa with anterior margin directed to the anterior region; posterior submedian translucent mark not inserted in a depression.

Size. (1) Head 1.77 mm long, 3.13 mm high, and 3.56 mm wide; (2) mesosoma 5.69 mm long, anterior wing 13.38 mm long, and posterior wing 8.55 mm long; (3) metasoma 8.57 mm long. Head. (1) Lateral ocelli with 0.26 mm and median ocellus with 0.28 mm of diameter, inserted in a declivity of the vertex, and the lateral ocelli separated from the eyes for 1.7 times its diameter. (2) Compound eyes with very small-sized and sparse bristles. (3) Frons with interantennal space with 1.7 times the height of antennal socket. Anterior tentorial fovea closer to the antennal socket than to the eye. Central region of the frons with very long bristles. (4) Antennal socket 0.34 mm high. (5) Clypeus as high as wide, contact with eyes for a distance of approximately the height of antennal socket and lateral lobe not touching the eye. Clypeus with long bristles, but with very long bristles on the apical margin. (6) Gena wider than half of the width of the eye at the level of the ocular sinus. Mesosoma. (1) Anterior lamella of pronotum with height of one third of the height of antennal socket. Pronotal fovea with ellipsoid shape, shallow and without anterior prominence. (2) Mesoscutum convex and as long as wide. (3) Tegula 1.5 times longer than wide. (4) Axillary fossa with anterior margin directing to the anterior region. (5) Propodeum with translucent posterior submedian mark, anterior to the propodeal valve, not inserted in a depression. Propodeal valve complete and expanded, median region with two third of the height of antennal socket. (6) Anterior wing with prestigma as long as wide. (7) Posterior wing with 11 hamuli. Metasoma. (1) Metasomal tergum I 1.7 times longer than broad. Tergum with angulation in the posterior third, in lateral view. (2) Metasomal tergum II 0.8 times longer than broad and without a row of very long bristles on the posterior margin. Color. Brown in general. Yellow: median longitudinal band and lateral of the frons, interantennal elevation, band surrounding the disc of the clypeus, mandibles, lower quarter of the gena, malar space, band contouring the posterior margin of the pronotum, tegula, spot anterior to the scrobal furrow of the mesepisternum, longitudinal submedian band and thin lateral band in the mesoscutum, axilla, anterior half of the scutellum, metanotum, submedian band in the propodeum, margin anterior to the propodeal valve, upper region of the metapleural basalar area, apex of the coxae, femora, tibiae and tarsi, bands along the posterior margins of the metasomal terga I–III and sternum II. Yellowish-brown: vertex and gena. Black: metasomal terga II–VI and metasomal sterna II–VI. Reddish-brown: inferior margin of the clypeus and the mandibular teeth. Wings with yellowish-brown in the cells and venation, except reddish-brown in the veins C, Sc+R, M+Cu, M and in the beginning of the Cu.

Size. (1) Head 1.6 mm long, 2.7 mm high, and 3.2 mm wide; (2) mesosoma 5.2 mm long, anterior wing 12.4 mm long, and posterior wing 8.10 mm long; (3) metasoma 8.3 mm long. Head. (1) Lateral ocelli with 0.24 mm and median ocellus with 0.27 mm of diameter. (2) Frons with interantennal space with 1.5 times the height of antennal socket. Anterior tentorial fovea closer to the eye than to the antennal socket. (3) Antennal socket 0.31 mm high. (4) Clypeus 1.2 times higher than wide and apex less acute than in the female. Pubescence stronger than in the female. (5) Gena with half of the width of the eye at the level of the ocular sinus. Mesosoma. (1) Mesoscutum 1.1 times longer than wide. Metasoma. (1) Metasomal tergum I 1.7 times longer than broad. (2) Metasomal tergum II 0.8 times longer than wide. Posterior margin with slight emargination in the center. Genitalia (Fig. 6f–j). Paramere 2 mm long and 0.81 mm wide; parameral spine with one fifth of the paramere, straight and with long bristles; lobe with rounded apex and slightly curved downwards. Aedeagus 1.72 mm long; enlarged valve with a small emargination in the tip; apical portion 0.66 mm long and curved to the venter, ventral margin with denticles directed for the anterior region; denticulation with large and conical denticles in the basal and apical thirds and small denticles with widened bases in the median third, more sclerotized than the rest of the apical portion; small-sized bristles with alveolar base, closer in the lower half and sparse in the upper half; median expansion with one denticle and with acute apex; lateral apodeme not flattened dorsoventrally at the apex; basal apodeme arched to the venter. Digitus 3.4 times longer than wide; apical process little curved in the region of the upper half and with bristles of alveolar base small and sparse; rounded anteroventral lobe with a strip of black scale-like bristles crossing it obliquely at the base of the digitus; bristles absent in the lower margin and basal articulation. Cuspis approximately 0.44 mm long, with 16 black scale-like bristles on the lateral lobe, and long bristles with alveolar base and close throughout the area of the cuspis, except sparse in the central region and on the ventral margin.

Some A. obidensis specimens found in São Gabriel da Cachoeira (Amazonas) and Parque Nacional Serra da Mocidade (Roraima) are darker, like A. paraensis of coloration transitional between the yellow form and the black and yellow forms, which also occurs in these regions.

Unknown.

A. obidensis resembles A. paraensis, but it is distinguished by the lamella along the anterior margin of the pronotum which is very elevated (one third of the height of antennal socket) in A. obidensis, and low (one fifth of the height of antennal socket) in A. paraensis; pronotum with prominence absent in front of the fovea, but slight prominence in front of the fovea in A. paraensis; angulate gena with enlarged lower and upper region, but gena with only enlarged lower region in A. paraensis; parameral spine straight, but spine curved upwards in A. paraensis; aedeagus with lateral apodeme not flattened dorsoventrally at the apex, but apex of the lateral apodeme flattened in A. paraensis.

The label information of the male specimen described is Brazil, AM, Itacoatiara, Mil Madeireira. 16.xii.1999, Malaise, J. Vidal Leg.

♀, TYPE / Obidos / Polybia obidensis Ducke, ♀ typ. / MUSEUM PARIS, Brésil, Obidos, A. Ducke 1904 (MNHN, Paris), record MNHN, Paris EY25586 (Fig. 4a, b, c). Type specimen analyzed by images.

PARALECTOTYPE / Surinam, ex.coll. Fruhstorfer / spec. Typ. / Polybia paraensis luctuosa Schlz. ♀ an ♀, W. A. Schulz det. / Schulz Coll., 1908-157. / B.M. TYPE, HYM., 18.767b / (1 ♀, NHM). Polybia paraensis luctuosa is a junior synonym of Polybia obidensis Ducke, 1904, redescribed species here.

We examined 102 females and three males for A. obidensis; see supplementary material S1.

Brazil: Acre, Amapá, Amazonas, Mato Grosso, Pará, Roraima (new record); French Guiana; Guyana; Suriname (Fig. 10c).

Pará, Brazil.

Anterior wing of 13–15 mm; eyes with very small-sized and sparse bristles; angulate gena with enlarged lower region; pronotal lamella low on the anterior margin, one fifth of the height of antennal socket; axillary fossa with anterior margin directed to the anterior region; posterior submedian translucent mark not inserted in a depression.

Size. (1) Head 2.01 mm long, 3.35 mm high, and 3.71 mm wide; (2) mesosoma 6 mm long, anterior wing 14.78 mm long, and posterior wing 10.37 mm long; (3) metasoma 10.8 mm long. Head. (1) Lateral ocelli with 0.27 mm and median ocellus with 0.29 mm of diameter, inserted in a declivity of the vertex and the lateral ocelli separated from the eyes for two times its diameter. (2) Compound eyes with very small-sized and sparse bristles. (3) Frons with interantennal space with 1.7 times the height of antennal socket. Anterior tentorial fovea closer to the antennal socket than to the eye. Central region of the frons with very long bristles. (4) Antennal socket 0.37 mm high. (5) Clypeus as high as wide, contact with eyes for a distance of approximately the height of antennal socket and lateral lobe not touching the eye. Clypeus with long bristles, but with very long bristles on the apical margin. (6) Gena wider than half of the width of the eye at the level of the ocular sinus. Mesosoma. (1) Anterior lamella of pronotum with height of one fifth of the height of antennal socket. Pronotal fovea with ellipsoid shape, shallow, and with slight anterior prominence. (2) Mesoscutum convex and 1.2 times longer than wide. (3) Tegula 1.6 times longer than wide. (4) Axillary fossa with anterior margin directing to the anterior region. (5) Propodeum with translucent posterior submedian mark, anterior to the propodeal valve, not inserted in a depression. Propodeal valve complete and expanded, median region with half of the height of antennal socket. (6) Anterior wing with prestigma as long as wide. (7) Posterior wing with 13 hamuli. Metasoma. (1) Metasomal tergum I 1.5 times longer than broad. Tergum with angulation in the posterior third, in lateral view. (2) Metasomal tergum II 0.7 times longer than broad and with a row of very long bristles on the posterior margin. Color. Yellowish-brown in general. Yellow: lateral of the vertex, lateral of the frons, interantennal elevation, lateral and the lower margin of the clypeus, mandibles, gena, anterior half of the lower quarter of the gena, malar space, band contouring the posterior margin of the pronotum, tegula, spot anterior to the scrobal furrow of the mesepisternum, longitudinal submedian band and thin lateral band in the mesoscutum, axilla, metanotum, submedian band in the propodeum, margin anterior to the propodeal valve, upper region of the metapleural basalar area, apex of the femora, tibiae and tarsi, bands on the posterior margins of the metasomal terga I–V, in the anterior margin of the sternum II, and in the posterior margins of the sterna II–IV. Brown: a spot that extends from the vertex to the middle of the frons, transversal band in the pronotum, scrobal furrow, region posterior to the scrobal furrow, metapleural basalar area, lateral bands of the propodeum, and metasomal terga I and II. Dark brown: ocellar area, mesoscutum, posterior half of the scutellum, posterior margin of the metanotum, bands in furrow and in anterior marginal of the propodeum, and metasomal terga III and IV. Reddish-brown: lower margin of the clypeus and mandibular teeth. Reddish yellow antenna. Wings with yellowish-brown in cells and in venation, except reddish-brown in the veins C, Sc+R, M+Cu, M, at the beginning of the Cu and pterostigma.

Size. (1) Head 1.40 mm long, 3.08 mm high, and 3.47 mm wide; (2) mesosoma 5.28 long, anterior wing 14.2 mm long, and posterior wing 9.9 mm long; (3) metasoma 10.5 mm long. Head. (1) Lateral ocelli with 0.24 mm and median ocellus with 0.28 mm of diameter, and lateral ocelli separated from the eyes for 1.7 times its diameter. (2) Frons with interantennal space with 1.5 times the height of antennal socket. Anterior tentorial fovea closer to the eye than to the antennal socket. (3) Antennal socket with 0.34 mm high. (4) Clypeus 1.2 times higher than wide and apex less acute than in the female. Pubescence stronger than in the female. (5) Gena with half of the width of the eye at the level of the ocular sinus. Mesosoma. (1) Mesoscutum 1.1 times longer than wide. (2) Propodeum without translucent posterior submedian mark. Propodeal valve with a median region with two third of the height of antennal socket. (3) Anterior wing with prestigma as long as wide. (4) Posterior wing with 14 hamuli. Metasoma. (1) Metasomal tergum I 1.7 times longer than broad. (2) Metasomal tergum II 0.8 times longer than wide. Genitalia (Fig. 6k–o). Paramere 2.3 mm long and 0.9 mm wide; parameral spine with one fifth of the paramere, curved upwards and with long bristles; lobe with rounded apex and not curved. Aedeagus 1.68 mm long; enlarged valve with a small emargination in the tip; apical portion 0.72 mm long and curved to the venter, ventral margin with denticles directed for the anterior region; denticulation with large and conical denticles in the basal and apical thirds and denticles reduced in the middle third, more sclerotized than the rest of the apical portion; small-sized bristles with alveolar base, closer in the lower half and sparse in the upper half; median expansion without denticles and with acute apex; lateral apodeme flattened dorsoventrally at the apex; sinuous basal apodeme. Digitus 2.7 times longer than wide; apical process little curved in the region of the upper half and with bristles of alveolar base small and sparse; rounded anteroventral lobe with a strip of black scale-like bristles crossing it obliquely at the base of the digitus; bristles absent in the lower margin and in the basal articulation. Cuspis approximately 0.46 mm long, with 26 black scale-like bristles on the lateral lobe, and long bristles with alveolar base and close throughout the area of the cuspis, except sparse in the central region.

Anterior wing of 13–15 mm; posterior wing of 13–19 hamuli; Angiopolybia paraensis occurs in three color variants, identified by Richards (1978) as A. paraensis morph paraensis (yellow specimens), A. paraensis morph ruficornis (brown and yellow specimens) and A. paraensis morph obscurior (brown specimens). Angiopolybia paraensis show small changes between yellow populations and between brown and yellow populations, such as a slightly darker color or some yellow marks, respectively, but they are well-defined. Despite the color variation, the morphological characteristics of female and male adults, and male genitalia used in the description are preserved in the three forms, so they should not be treated as subspecies or differentiated as morphs. Only a few A. paraensis populations, with dark brown color from French Guiana and Suriname, showed morphological variations such as the absence of very long bristles in the posterior margin of the metasomal tergum II and propodeum with a region anterior to the spiracle less projected in the metapleural basalar area. However, we believe that this evidence is still insufficient to justify a new species. Moreover, the analysis of species delimitation with molecular data (see below) showed that these variants are within the intraspecific limits of A. paraensis.

Described by Schulz (1903) as a spherical nest about 25 mm in diameter (possibly incorrect unit of measurement was used, with centimeter (cm) being the correct). Ducke (1910) complements the description of Schulz (1903), stating that the nest is composed of four overlapping combs, joined by a central pedicel, with a simple and very resistant envelope, and with transverse streaked with light and dark colors.

Angiopolybia paraensis resembles A. obidensis, but it is distinguished by the pronotum with low lamella in the anterior margin (one fifth of the height of antennal socket), being high lamella (one third of the height of antennal socket) in A. obidensis; pronotum with a slight prominence in front of the fovea, but prominence absent in A. obidensis; gena not enlarged in the upper region, but enlarged in the upper region in A. obidensis; parameral spine curved upwards, but straight parameral spine in A. obidensis; aedeagus with the apex of the lateral apodeme dorsoventrally flattened, but apex not flattened in A. obidensis.

Angiopolybia paraensis was described by Spinola (1851) and his type specimen (or specimens) was deposited in the Museo Regionale di Scienze Naturali di Torino (MRSN, Torino, Italy). Richards (1978) did not find any A. paraensis type specimen during his study about the social wasps of the Americas, and we did not receive any answer from the Museum about the type specimen. Additionally, on the online page Checklist of Epiponini wasps (http://iunh2.sci.ibaraki.ac.jp/wasp/Epiponini/epiponini.htm; consulted in 2021) produced by Dr. James M. Carpenter, the presence of the type specimen in the Museum’s collection is also uncertain. Based on this, the redescription of the species was carried out using a specimen identified by Ducke in 1909 and from the same locality of the type specimen (Brazil, Pará, 26.9.1901, Ducke / Polybia paraensis Spin. ♀, det. Ducke 1909 / Brazil., Mus.Goeldi., 1910-90. (1 ♀, NHM)). The information about the male specimen described is: BRA, Roraima, Amajarí, Serra do Tepequém, SESC Tepequém. 1–15.iii.2016 / Malaise grande, J.A. Rafael, F.F. Xavier Filho col.

Although some checklists treat A. paraensis as registered for Bahia state [for example, the Checklist of Epiponini wasps produced by Dr. James M. Carpenter (http://iunh2.sci.ibaraki.ac.jp/wasp/Epiponini/epiponini.htm; consulted in 2021) and Barbosa et al. (2016)], we did not confirm this information.

Without information.

We examined 79 females and six males for A. paraensis; see supplementary material S1.

Bolivia: Cochabamba, La Paz; Brazil: Acre, Amapá, Amazonas, Maranhão, Mato Grosso, Pará, Rondônia, Roraima; Colombia: Amazonas, Caquetá; Ecuador; French Guiana; Guyana; Peru: Cuzco, Huánuco, Junín, Loreto, Madre de Dios, Pasco, San Martin; Panama; Suriname; Venezuela: Amazonas, Bolívar (Fig. 10d).