Angiopolybia pallens (Lepeletier, 1836)

Barroso, Paulo Cezar Salgado, Menezes, Rodolpho Santos Telles, de Oliveira, Marcio Luiz & Somavilla, Alexandre, 2022, A systematic review of the Neotropical social wasp genus Angiopolybia Araujo, 1946 (Hymenoptera: Vespidae): species delimitation, morphological diagnosis, and geographical distribution, Arthropod Systematics & amp; Phylogeny 80, pp. 75-97 : 75

Angiopolybia pallens (Lepeletier, 1836)

Figs 1a-f , 2a-c , 6a-e , 7a-g

Rhopalidia pallens Lepeletier, 1836: 539; Spinola, 1851: 63 (nest); de Saussure, 1854: 189 (synonym of Polybia pallipes (Olivier, 1792)); du Buysson, 1906: 342 (synonym of Apoica pallida var. pallens (Fabricius, 1804)); Ducke, 1910: 542 (specimen of the collection of Spinola = S. infernalis (de Saussure,1854)); Schulz, 1912: 60 (synonym: S. infernalis (de Saussure, 1854)).

Polistes rufina Erichson, 1848: 590; Spinola, 1851: 79 (synonym of Rhopalidia pallens Lepeletier, 1836).

Polybia infernalis de Saussure, 1854: 195, plate XXV: fig. 3 (in division My); Ducke, 1905a: 662 (synonym: P. ampullaria Möbius, 1856); Richards, 1943: 45 (invalid designation of the type species of Stelopolybia ); Richards, 1978: 232 (lectotype designation). Type locality: "Le Para", female ( MNHN, Paris) [examined by images].

Polybia ampullaria Möbius, 1856: 133, 155 (key identification of nests), 165 (plate VII), VII (figs. 1-8 - female, nest).

Eumenes flavopectus Provancher, 1888: 422.

Stelopolybia infernalis ; Ducke, 1910: 519 (key), 524 (synonym: Polybia ampullaria Möbius, 1856), 525 (fig. 12 - nest); Ducke, 1913: 331 (synonym: R. pallens Lepeletier, 1836).

Stelopolybia pallens ; Bequaert, 1944: 293 (key), 294 (synonym: Polybia infernalis Saussure, 1854, Polybia ampullaria Möbius, 1856, Polistes rufina Erichson, 1884, Eumenes flavopectus Provancher 1888).

Angiopolybia pallens ; Araujo, 1946: 169; Richards, 1978: 231 (key), 232 (description of male, and diagnose of female and male); Andena et al., 2007: 59 (phylogenetic discussion), 60 (table 2 - characters matrix), 61 (figs. 1A, 2A), 62 (figs. 3B, 5 - cladogram), 63 (key), 64 (locality of examined material); Carvalho et al., 2014, 2021 (evolutionary hypothesis of species distribution).

Stelopolybia (Angiopolybia) pallens ; Richards and Richards, 1951: 77 (list and notes about the nests).

Angiopolybia infernalis ; Overal, 1978: 9 (list of species).

Type locality.

Cayenne, French Guiana.

Diagnosis.

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.

Redescription of female (Fig. 1a, b, c).

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.

Redescription of male (Fig. 1d, e, f).

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.

Morphological variation (Fig. 7a-g).

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.

Nest (Fig. 2a-c).

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.

Comparative comments.

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.

Additional comments.

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.

Lectotype.

♀, 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.

Additional material examined.

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

Geographic distribution.

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 ).

Figures

Figure 1. Lectotype, female of Polybia infernalis de Saussure, 1856 [currently Angiopolybia pallens (Lepeletier, 1836)]: a. lateral view, b. dorsal view, c. head in frontal view. Male: d. head in frontal view, e. lateral view, f. dorsal view. Scale: 1 mm. Source: Agniele Touret-Alby MNHN for images a, b, and c.

Figure 2. Structure variation of the nest of Angiopolybia pallens: a. typical form in external view, b. and c. variation found in external and internal view, respectively.

Figure 6. Male genitalia of three Angiopolybia species. Angiopolybia pallens: a. paramere, b. aedeagus in left lateral view, c. aedeagus in ventral view, d. digitus, e. cuspis (image mirrored). Angiopolybia obidensis: f. paramere, g. aedeagus in left lateral view, h. aedeagus in ventral view, i. digitus, j. cuspis. Angiopolybia paraensis: k. paramere, l. aedeagus in left lateral view, m. aedeagus in ventral view, n. digitus, o. cuspis. Scale: 0.5 mm.

Figure 7. Morphological variation of Angiopolybia species. Angiopolybia pallens: a. Cayenne, French Guiana; b. Amazonas, Brazil; c. Bahia, Brazil; d. Sao Paulo, Brazil; e. Loreto, Peru; f. Napo, Ecuador; g. Chapare, Bolivia. Angiopolybia zischkai: h. Pipeline, Panama. Angiopolybia obidensis: i. Roraima, Brazil. Angiopolybia paraensis: j. Roraima (brown and yellow specimen), Brazil; k. Napo, Ecuador (brown specimen); l. Crique Alama, French Guiana (dark brown specimen). Scale: 1 mm.

Figure 10. Map of South and Central America showing the geographic distribution of Angiopolybia species: a. Angiopolybia pallens, b. Angiopolybia zischkai, c. Angiopolybia obidensis, and d. Angiopolybia paraensis.