Research Article |
Corresponding author: Franciélle Dias de Oliveira ( fdoliveira1@gmail.com ) Corresponding author: Angélica Maria Penteado-Dias ( angelica@power.ufscar.br ) Academic editor: Kees van Achterberg
© 2020 Franciélle Dias de Oliveira, Angélica Maria Penteado-Dias.
This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
Citation:
de Oliveira FD, Penteado-Dias AM (2020) An update of the genera Idiasta Foerster and Rhacalysia Cameron (Hymenoptera, Braconidae, Alysiinae) and the descriptions of new species from the Neotropical Region. ZooKeys 976: 109-130. https://doi.org/10.3897/zookeys.976.56751
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Taxonomic combinations have been made involving the two genera Idiasta Foerster and Rhacalysia Cameron. Four new species are described from Brazil: Idiasta rupina sp. nov., Rhacalysia ampla sp. nov., Rhacalysia jatai sp. nov., and Rhacalysia monteiroi sp. nov. Dichotomous identification keys to the Neotropical species of Idiasta and Rhacalysia are provided. Phaenocarpa delicata Papp, 1969 is included in Rhacalysia and is a new combination.
Alysiini, Brazil, parasitic wasp, parasitoid, taxonomy
Idiasta Foerster, 1863 and Rhacalysia Cameron, 1910 are both genera of the tribe Alysiini (Braconidae, Alysiinae). Members of Idiasta possess the largest set of plesiomorphic characteristics within the Phaenocarpa complex (
The genus Rhacalysia includes four described species: Rhacalysia congoensis Fischer, 1993, R. delicata (Papp, 1969) comb. nov., R. profundinigra Fischer, 1999, and R. rufobalteata Cameron, 1910. The biology of Rhacalysia is unknown. Rhacalysia congoensis is known from Republic of the Congo (
The genus Rhacalysia was treated as a synonym of Idiasta by
Rhacalysia delicata was originally included in Phaenocarpa (Papp, 1969), later attributed to Idiasta (
Two specimens of Idiasta and eight of Rhacalysia were studied. Idiasta rupina sp. nov. was collected in a grassland environment characterized as rupestrian grassland (campo rupestre) (
Digital Scanning Electronic Microscope (SEM) photographs of uncoated specimens were taken with a FEI Quanta 250 SEM in a low vacuum mode. Color digital photographs were taken with a Leica M165C stereomicroscope, using a Leica DFC295 HD camera, LEICA APPLICATION SUITE software version 3.7. ADOBE PHOTOSHOP CS5 Extended version 12.1. was used for minor corrections of images and for the preparation of plates.
Idiasta (Alysia) maritima Haliday, 1838
Mandibles with three teeth, ventral and diagonal ridges well developed. First flagellar segment equal or shorter than second. Fore wing pterostigma broad, discrete, wedge-shaped; 2-SR vein longer than 3-SR. Hind wing with m-cu present, often well developed, M+CU generally equal to or longer than 1-M.
Muscidae (Diptera).
Cosmopolitan.
Holotype
pinned, female, (
Idiasta rupina is distinct from other Neotropical species in having the eyes glabrous, notauli incomplete, metanotum with high median flange, m-cu of fore wing interstitial, and fore wing cu-a postfurcal.
Female (holotype) (Fig.
Head. 1.6× as wide as long; 1.8× as wide as face, 1.6× as wide as mesosoma, 3.3× as wide as apex of first metasomal tergite; slightly wider at eyes than temples in dorsal view. Eye glabrous, 1.2× as high as wide, 2.6× as wide as temple in lateral view (Fig.
Mesosoma. 1.4× as long as high, 1.9× as long as wide, 1.7× as high as head. Pronotum in dorsal view with small but distinct pronope, crenulate in posterior margin; in lateral view, crenulate in upper middle area. Mesoscutum 1.1× as wide as long, scattered setae present along notauli. Notauli deep, crenulate anteriorly, absent posteriorly. Mesoscutal pit deep, slightly elongate, occupying a little less than 1/5 extent of mesoscutum. Scutellar sulcus 2.5× as wider as long, with well-developed mid ridge and some weak ridges at posterior margin of lateral areas. Scutellar disc smooth, setiferous; parascutellar area weakly rugulose posteriorly, with setae near scutellar sulcus. Metanotum setiferous anteriorly, in dorsal view rugose medially, depressed lateral fields crenulate; mid ridge complete and two well-developed median lateral ridges; metanotum in lateral view with high median flange. Mesopleuron with scattered setae in ventral area, antero-basal margin crenulate towards anterior subalar area; posterior margin crenulate. Precoxal sulcus deep, long, widely crenulate, separated from posterior margin (Fig.
Fore wing. Approximately as long as body. Pterostigma 3.4× as long as wide, 2.25× as wide as vein r length; r 0.3× as long as 2-SR, arising distad midpoint of pterostigma; submarginal cell 2.6× as long as high; 2-SR 2.5× as long as r-m, 1.4× as long as 3-SR; 3-SR 2.65× as long as r, 1.8× as long as r-m; SR1 5.15× as long as 3-SR; 2-CU1 1.1× as long as m-cu, this interstitial; cu-a postfurcal by distance less than its length; subdiscal cell closed, nearly parallel-sided; CU1a arising slightly above middle of subdiscal cell (Fig.
Hind wing. With three hamuli, 5.7× as long as wide; vein 1-M 1.15× as long as M+CU, 2.0× as long as 1r-m; m-cu interstitial, spectral (Fig.
Legs. Hind femur 6.35× as long as wide. Hind tibia 11.2× as long as its maximum subapical width, 0.9× as long as hind tarsus. First segment of hind tarsus 1.7× as long as second segment.
Metasoma. First metasomal tergite 1.6× as long as apical width; apex 1.5× as wide as base; strongly strigose surface, dorsal carinae convergent and uniting in basal third, continuing as a distinct median carina to apex, dorsope deep. Ovipositor 1.3× as long as hind tibia, 1.45× as long as mesosoma; straight and strongly directed upwards (Fig.
Color. Dark brown to black. Gena and mandibles brown, mandibles lighter in apical third; scape, pedicel, and basal half of first flagellar segment yellow brown; flagellar segments 2–13 brown, 14–19 white, 20–32 dark brown. Propleuron, ventral mesopleuron, and tegulae brown. Second and third metasomal tergites brownish yellow. Legs yellow, gradually darkening towards apex; coxa and trochanter pale yellow; hind tibia and hind tarsus brown. Wings hyaline, venation and pterostigma brown.
Male. Similar to female but body length 2.4 mm; hind wing 2.0 mm; head 3.0× as wide as apex of first metasomal tergite; antenna with 36 flagellar segments; first flagellar segment 3.9× as long as wide; second flagellar segment 7.95× as long as wide, 1.8 × length of first segment; third flagellar segment 7.0× as long as wide, 1.4× length of first segment; maxillary palp 2.4× as long as head height. Mesoscutum slightly longer than wide; scutellar sulcus 2.25× as wide as long. Fore wing with 2-SR vein 1.9× as long as r-m, 1.6× as long as 3-SR; 3-SR 1.2× as long as r-m. Hind wing 1-M 0.8× as long as M+CU, 2.6× as long as 1r-m. Hind femur 5.15× as long as wide; first segment of hind tarsus 1.5× as long as second segment. Antenna brown except yellowish basal half of first flagellar segment; wing venation and pterostigma light brown.
The species name refers to the ecosystem from which the studied material was collected.
Brazil, State of Minas Gerais, São Roque de Minas, rupestrian grassland.
Idiasta rupina and I. dixi are related Neotropical species and share the notauli absent posteriorly, fore wing cu-a postfurcal, and hind wing m-cu not tubular. The color pattern of the body is also similar. However, Idiasta rupina can be differentiated by the glabrous eye (with sparse setae in I. dixi), high median flange of the metanotum (absent in I. dixi), fore wing m-cu interstitial (slightly antefurcal in I. dixi). Additionally, Idiasta rupina differ in the following quantitative ratios: eye 2.5 × as wide as temple (3.1× in I. dixi); maxillary palp ca. twice as long as head height (1.4 × in I. dixi); sulcus scutellar 2.5× as wide as long (1.4× in I. dixi); fore wing vein 3-SR 2.6 × as long as r (3.5× in I. dixi); SR1 5.1× as long as 3-SR (4.5× in I. dixi); ovipositor 1.4× as long as mesosoma (ca. 1.0× in I. dixi).
Rhacalysia rufobalteata Cameron, 1910
Enlarged paraclypeal fovea extending to eye. First flagellar segment shorter than second. Fore wing venation complete; 2-RS vein longer than 3-SR, m-cu antefurcal to interstitial. Metasomal terga unsculptured beyond the first tergite, not forming a carapace.
Unknown.
Afrotropical, Asian, and Neotropical Regions.
Holotype
pinned, female (
Rhacalysia ampla can be differentiated from other species of genus by the notauli incomplete, precoxal sulcus sculptured only in anterior fourth, fore wing with m-cu interstitial, CU1a arising below middle of subdiscal cell, and hind wing with three hamuli.
Female (Fig.
Head. 1.5–1.85× as wide as long; 1.7× as wide as face, 1.5–1.6× as wide as mesosoma, ca. 2.2× as wide as apex of first metasomal tergite; slightly wider at eyes than temples in dorsal view. Eye glabrous, 1.0–1.1× as high as wide, 2.9–3.0× as wide as temples in lateral view (Fig.
Mandible 3-dentate (Figs
Mesosoma. 1.3–1.4× as long as high, 1.9–1.95× as long as wide, 2.0–2.4× as high as head. Pronotum in dorsal view with distinct pronope, crenulate laterally; smooth in lateral view. Notauli deep, narrow, crenulate anteriorly, absent posteriorly (Fig.
Fore wing. 0.9–1.1× as long as body. Pterostigma 3.6–3.75× as long as wide, 1.9–2.2× as wide as vein r length; r 0.2–0.3× as long as 2-SR, arising distad midpoint of pterostigma; submarginal cell 2.5–2.7× as long as high; 2-SR 2.5–2.7× as long as r-m, 1.1–1.3× as long as 3-SR; 3-SR 3.1–3.5× as long as r, 2.1–2.3× as long as r-m; SR1 3.5–4.0× as long as 3-SR; 2-CU1 1.3–1.4× as long as m-cu, this interstitial; cu-a postfurcal by distance ca. equal to its length; subdiscal cell closed, expanded distally, CU1a arising below middle of subdiscal cell (Fig.
Hind wing. With three hamuli, 1.3–1.4× as long as wide; vein 1-M 1.2× as long as M+CU, 1.3–1.6× as long as 1r-m; m-cu antefurcal, strongly nebulous for most of its length, tubular basally near its insertion, nearly reaching wing margin (Fig.
Legs. Hind femur 5.7–6.1× as long as wide. Hind tibia 11.4–12.1× as long as its maximum subapical width, 1.0–1.1× as long as hind tarsus. First segment of hind tarsus 1.5–1.7× as long as second segment.
Metasoma. First metasomal tergite ca. as long as apical width; apex 2.0–2.1× as wide as base; smooth surface, dorsal carinae strongly convergent, uniting in basal third, continuing as distinct median carina but not reaching to apex; dorsope deep (Fig.
Color. Mostly dark brown. Mandibles light brown to yellow, darker at base. Clypeus, scape, pedicel, scutellum, and metanotum brown to light brown. Flagellar segment 17–20 whitish (holotype). Mesonotum brown to reddish brown. Propodeum and metapleuron yellowish to dark orange. First metasomal tergite orange to yellowish orange, base of terga 2 sometimes orange, other tergites brown. Tegula, ovipositor, and most of legs yellow. Trochanter and tronchantellus pale yellow, telotarsus darkened; hind leg with distal tibia and tarsus brown. Wings hyaline; venation and pterostigma light brown to brown.
Male. Unknown.
The species name refers to the form of the paraclypeal fovea.
Brazil, State of Rio de Janeiro, Teresópolis, dense ombrophilous forest.
Rhacalysia ampla is similar morphologically to R. delicata, with which it shares many characteristics. Members Rhacalysia ampla can be differing by the precoxal sulcus weakly sculptured only in the anterior fourth of mesopleuron (Fig.
Holotype
pinned, female, (
Rhacalysia jatai can be recognizable by the notauli and precoxal sulcus entirely smooth, fore wing with m-cu interstitial, CU1a arising at middle of subdiscal cell, hind wing with four hamuli; ovipositor 2.2× as long as mesosoma, body yellow (without brown parts).
Female (holotype) (Fig.
Head. 1.7× as wide as long; 1.7× as wide as face, 1.6× as wide as mesosoma; 1.9 × as wide as apex of first metasomal tergite; slightly wider at eyes than temples in dorsal view. Eye glabrous, ca. as high as wide, 2.5× as wide as temple in lateral view (Fig.
Mesosoma. 1.3× as long as high, 2.0× as long as wide, 2.2× as high as head. Pronotum smooth (Fig.
Fore wing. 1.05× as long as body. Pterostigma 4.0× as long as wide, 2.2× as wide as vein r length; r 0.2× as long as 2-SR, arising distad midpoint of pterostigma; submarginal cell 2.25× as long as high; 2-SR 2.4× as long as r-m, 1.3× as long as 3-SR; 3-SR 3.8× as long as r, 1.8× as long as r-m; SR1 3.8× as long as 3-SR; 2-CU1 1.2 × as long as m-cu; m-cu interstitial; cu-a postfurcal by distance slightly shorter than its length (Fig.
Hind wing. With four hamuli, 4.4× as long as wide; vein 1-M 0.9× as long as M+CU, 1.6× as long as 1r-m; m-cu antefurcal, nebulous but heavily pigmented.
Legs. Hind femur 5.2× as long as wide. Hind tibia 11.2× as long as its maximum subapical width, 1.1× as long as hind tarsus. First segment of hind tarsus 1.95× as long as second segment.
Metasoma. First metasomal tergite 0.9× as long as apical width; apex 2.0× as wide as base; smooth surface, dorsal carinae converging anteriorly but not extending as median carina (Fig.
Color. Yellow. Ocellar field and epicranial suture dark brown (Fig.
Male. Unknown.
The name of species refers to locality of collection of material for study.
Brazil, State of São Paulo, Luiz Antônio, seasonal forest.
Rhacalysia jatai is morphologically similar to R. monteiroi and both species shares the hind wing with four hamuli, as well as several other features. Rhacalysia jatai can be differentiated by the follow quantitative ratios: third flagellar segment 4.9× as long as wide (4.5–4.6× in R. monteiroi), 1.4× length of first segment (1.0–1.1 × in R. monteiroi); eye 2.5× as wide as temple (1.5–2.0× in R. monteiroi); vein 3-SR of fore wing 3.8× as long as r (3.0–3.1× in R. monteiroi); hind femur 5.2 × as long as wide (6.2–6.7× in R. monteiroi); hind tibia 11.1× as long as its maximum apical width (12.2–12.7× in I. monteiroi); ovipositor 2.0× as long as hind tibia (1.3–1.5 × in R. monteiroi), 2.2× as long as mesosoma (1.4–1.7× in R. monteiroi). In addition, in Rhacalysia jatai the body is entirely yellowish (Figs
Holotype
pinned, female, (
Diagnosis. Rhacalysia monteiroi can be recognized by the notauli incomplete, fore wing with m-cu interstitial, CU1a arising at middle or slightly below middle of subdiscal cell, hind wing with four hamuli, hind femur 6.2–6.7× as long as wide, coloration of body mixed between yellowish and brown parts.
Female (Fig.
Head. 1.7–1.9× as wide as long; 1.7–1.9× as wide as face, 1.4–1.6× as wide as mesosoma; 2.1× as wide as apex of first metasomal tergite; slightly wider at eyes than temples in dorsal view. Eye glabrous, 1.1–1.2× as high as wide, 1.5–2.0× as wide as temples in lateral view (Figs
Mesosoma. 1.3–1.4× as long as high, 1.9× as long as wide, 2.4× as high as head. Pronotum with pronope relatively large, slightly crenulate laterally and with some setae in dorsal view; smooth to slightly crenulate in lateral view. Mesoscutum 1.1× as wide as long, scattered setae, smooth to weakly crenulate in postero-lateral margins. Notauli deep, smooth to weakly crenulate anteriorly, absent posteriorly (Figs
Fore wing. 1.0–1.2× as long as body. Pterostigma 3.6–4.1× as long as wide, 2.1–2.3× as wide as vein r length; r 0.2–0.25× as long as 2-SR, arising distad midpoint of pterostigma; submarginal cell 2.5–3.5× as long as high; 2-SR 2.4–2.7× as long as r-m, 1.3–1.6× as long as 3-SR; 3-SR 3.0–3.1× as long as r, 1.7–2.0× as long as r-m; SR1 3.8–4.0× as long as 3-SR; 2-CU1 1.0–1.35× as long as m-cu, this interstitial; cu-a postfurcal by distance ca. equal to its length; subdiscal cell closed, slightly expanded distally, CU1a arising at middle to slightly below middle of subdiscal cell (Fig.
Hind wing. With four hamuli, 4.1–4.9× as long as wide; vein 1-M 1.0–1.2× as long as M+CU, 1.4–1.7× as long as 1r-m; m-cu antefurcal, heavily nebulous, tubular basally or not, almost reaching wing margin (Fig.
Legs. Hind femur 6.2–6.7× as long as wide. Hind tibia 12.2–12.7× as long as its maximum subapical width, 1.1–1.2× as long as hind tarsus. First segment of hind tarsus 1.7–1.8× as long as second segment.
Metasoma. First metasomal tergite 1.0–1.1× as long as apical width; apex 1.8–1.9 × as wide as base, strigose to slightly strigose; dorsal carinae converging in basal third, extending posteriorly as median carina incomplete or median carina absent (Fig.
Color. Body parts vary between dark brown to yellow (Figs
Male. Unknown.
The species is named in honor of Ricardo Ferreira Monteiro, the collector of the studied material.
Brazil, State of Rio de Janeiro, Teresópolis, dense ombrophilous forest.
Rhacalysia monteiroi shares many morphological characteristics with R. jatai; both species can be differentiated by the coloration pattern of body, relative length of the ovipositor, and relative length/wide of the posterior femur (see above).
The three specimens of R. monteiroi studied here vary considerably in some characteristics. In short, the sculpturing of face (striate or rugulose above the clypeus), notauli (smooth or weakly crenulate), precoxal sulcus (smooth or crenulate), and first metasomal tergite (with or without median carina); the coloration of vertex, pronotum, metanotum, propodeum, and metapleuron yellowish or brown (Figs
Members of the genera Idiasta and Rhacalysia can be morphologically differentiated as follows: paraclypeal fovea not extending to eye in Idiasta (Fig.
1 | Fore wing patterned with several dark spots or bands, M+CU1 of fore wing very weak, not, or only weakly, pigmented for much of its length; notauli complete to mesoscutal pit; metanotum with high flange. Body length 3.0–5.0 mm. Mexico, Holarctic | I. maritima (Haliday) (♀♂) |
– | Fore wing either hyaline or dusky but never patterned with spots or bands; M+CU1 of fore wing well-developed, usually strongly pigmented throughout; variable development of notauli; metanotum with or without high flange | 2 |
2 | Notauli complete and rugose; fore wing with CU1a arising well below middle of first subdiscal cell, cu-a interstitial or postfurcal; metanotum with high flange. Body length 2.5–4.0 mm. Mexico | I. euryzona Wharton (♀♂) |
– | Notauli incomplete, not reaching mesoscutal pit (as Fig. |
3 |
3 | Eye with scattered setae, maxillary palp 1.4× as long as head height; scutellar sulcus 1.4× as wide as long; metanotum without high flange; 3-SR of fore wing 3.4× as long as r, m-cu slightly antefurcal (2-SR+M present); ovipositor ca. as long as mesosoma. Colombia | I. dixi Dix (♀) |
– | Eye glabrous; maxillary palp 2.0–2.4× as long as head height; scutellar sulcus 2.2–2.5× as wide as long; metanotum with high flange; 3-SR of fore wing 2.6× as long as r, m-cu interstitial (2-SR+M absent); ovipositor 1.45× as long as mesosoma. Body length 2.4–2.7 mm. Brazil (Figs |
I. rupina sp. nov. (♀♂) |
1 | Fore wing with vein CU1a interstitial. Body length 2.4 mm. Republic of the Congo. | R. congoensis Fischer (♀♂) |
– | Fore wing with variable CU1a insertion but not interstitial | 2 |
2 | Notauli complete and heavily sculptured; face with two wide and sculptured longitudinal sulcus lateral to mid ridge. Body length 4.3 mm. India | R. rufobalteata Cameron (♀♂) |
– | Notauli smooth or incomplete (Figs |
3 |
3 | Clypeus slightly wider than high; vein m-cu of fore wing antefurcal; scutellar sulcus with 7 longitudinal ridges; first metasomal tergite longitudinally striate, median carinae present; body entirely black. Body length 5.0 mm. India | R. profundinigra Fischer (♀♂) |
– | Clypeus 1.6–2.0× as wide as long; vein m-cu of fore wing interstitial (Fig. |
4 |
4 | Hind wing with three hamuli; metanotum without high flange; body color mostly brown. | 5 |
– | Hind wing with four hamuli; metanotum with high flange; body yellowish or at least with distinctly yellow-orange parts. | 6 |
5 | Eye 2.0–2.2× as wide as temple; mesosoma 1.7× as high as head; hind femur ca. 5.0× as long as wide; fore wing with pterostigma 3.0× as wide as vein r length, cu-a slightly postfurcal, CU1a arising at middle of subdiscal cell; ovipositor ca. twice as long as hind tibia. Body length 3.5–5.4 mm. Argentina, Brazil, Colombia, Mexico, Panama, Peru, and Venezuela | R. delicata (Papp) (♀♂) |
– | Eye ca. 3.0× as wide as temple; mesosoma 2.0–2.4× as high as head; hind femur 5.7–6.1× as long as wide; fore wing with pterostigma 1.9–2.4× as wide as vein r length, cu-a postfurcal by distance ca. equal to its length, CU1a arising below of subdiscal cell; ovipositor 1.0–1.2× as long as hind tibia. Body length 3.1–4.2 mm. Brazil (Figs |
R. ampla sp. nov. (♀) |
6 | Vein 3-SR of fore wing 3.8× as long as r; hind femur 5.2× as long as wide; hind tibia 11.2× as long as its maximum subapical width; ovipositor 2.0× as long as hind tibia, 2.2× as long as mesosoma; body color yellowish. Body length 3.8 mm. Brazil (Figs |
R. jatai sp. nov. (♀) |
– | Vein 3-SR of fore wing 3.0–3.1× as long as r; hind femur 6.2–6.7× as long as wide; hind tibia 12.2–12.7× as long as its maximum subapical width; ovipositor 1.4–1.5× as long as hind tibia, 1.5–1.7× as long as mesosoma; mesosoma with distinctly brown parts. Body length 3.4–4.1 mm. Brazil (Figs |
R. monteiroi sp. nov. (♀) |
The wing venation pattern of Idiasta is widely maintained in Rhacalysia. The morphological support for the retention of the generic status of Rhacalysia has been the enlarged paraclypeal fovea (
The insertion antefurcal of vein m-cu of fore wing (and therefore 2-SR+M present) was considered diagnostic characteristic of Idiasta by
In addition to the enlarged paraclypeal fovea, all known species of Rhacalysia shares the follow characteristics: fore wing with pterostigma distinct and wide, vein r arising from its distal middle, vein r shorter than pterostigma width, cu1 postfurcal (Figs
Studies covering more Rhacalysia specimens in the future, should provide a clearer and more conclusive morphological delineation in relation to the genus Idiasta, especially if together the analysis of molecular data. Nevertheless, as well as
Considering our records, the distribution of Idiasta is slightly altered: the genus is no longer recorded from Argentina, Peru, and Venezuela. In turn, considering delicata species as Rhacalysia, the genus Rhacalysia is now known from many countries of the Neotropical Region.
We are thankful to FAPESP (Fundação de Amparo à Pesquisa do Estado de São Paulo), CNPq (Conselho Nacional de Desenvolvimento Científico e Tecnológico), CAPES (Coordenadoria de Aperfeiçoamento de Pessoal de Nível Superior), INCT–HYMPAR (Instituto Nacional de Ciência e Tecnologia dos Hymenoptera Parasitoides) for financial support; COTEC (Comissão Técnico-Científica) for permits the collection at the Estação Ecológica do Jataí; to Parque Nacional da Serra dos Órgãos and Parque Nacional da Serra da Canastra for permits the collection of material; to Nelson W. Periotto, Ricardo F. Monteiro and Airton S. Soares for collecting the material; to Carlos V. S. Gomes and the Laboratório de Ecologia de Insetos of Universidade Federal do Rio de Janeiro for the specimens loan collected in the Parque Nacional da Serra dos Órgãos; to Luciana B. R. Fernandes for taking and editing the electron micrographs. The manuscript was improved thanks to the helpful comments of the reviewer and subject editor.