Anchieta fumosellus (Westwood, 1867)

Figs 10, 11

Mantispa (Trichoscelia) fumosella Westwood, 1867: 504. Holotype: male, Amazonia (OUMNH), specimen examined.

Anisoptera fumosella (Westwood, 1867). Gerstaecker (1888).

Trichoscelia fumosella Westwood, 1867. Enderlein (1910).

Anchieta fumosella (Westwood, 1867). Penny (1982 a).

Anchieta nobilis Navás, 1909: 484, male, female. Lectotype, sex not indicated, Brazil (MNHN). Synonymized with Anchieta fumosella by Penny (1982 a): 4019, High resolution images examined.

Trichoscelia nobilis (Navás, 1909). Enderlein (1910).

Material examined.

Holotype of Mantispa (Trichoscelia) fumosella.

[Brazil] • ♂; “ fumosella Westwood, male ”, “ Platymantispa fumosella (Westwood, 1867) ”, “ Holotype male, Trichoscelia fumosella Westwood, 1867, R. G. Beard 1968, now belongs in genus Platymantispa Rehn ”, “ Type Westwood, Trans. Entomol. Soc. 1865, p. 504, Coll. Hope Oxon ”, “ Holotype male, Genitalia of Trichoscelia fumosella Westwood, 1867, prep. R. G. Beard, 1968 ”, “ over? fumosella Westw. ”, “ Type Neur: No. 13, Mantispa Trichoscelia fumosella Westw. HOPE Dept. Oxford ”; Terminalia cleared and stored in a microvial; OUMNH.

Lectotype of Anchieta nobilis.

Brazil • ♂; Goiás, Jataí; Sep. – Nov. 1897; Anchieta nobilis Navás, Longin Navas det. 1907; Lectotype ♂ designated by R. G. Beard, 1968; MNHN.

Paralectotype of Anchieta nobilis.

Brazil • ♀; Goiás, Jataí; Anchieta nobilis Navás, Longin Navas det. 1907; Lectoallotype ♂ designated by R. G. Beard, 1968; MNHN.

Other material.

Brazil – Bahia • 2 ♂; Encruzilhada; 15°32'25"S, 40°50'12"W; 800 m; 10–12 Dec. 2007; J. A. Rafael, P. C. Grossi, D. R. Parizotto leg.; light trap; Anchieta fumosella (Westwood, 1867), det. R. J. P. Machado; INPA. • 1 ♂; Encruzilhada; 960 m; Nov. 1972; Seabra and Alvarenga leg.; DZUP. – Minas Gerais • 1 ♀; Feb. 1932; R. Bandens, J. Blaser leg.; MCZ. • 1 ♂; same data as for preceding, Passos; Mar. 1961; C. Elias leg.; det. L. A. Stange; FSCA. – Paraná • 1 ♀; São José dos Pinhais; 25°36'18"S, 49°11'37"W; 1–30 Nov. 2019; A. C. Domahovski leg.; malaise; DZUP. • 1 ♀; same data as for preceding; 17–31 Dec. 2016; A. C. Domahovski leg.; sweep; DZUP. • 1 ♂; Campo Largo, Estrada do Cerne, Km 45; 22 Nov. 1979; Expedition Dep. Zoo UFPR leg.; DZUP. • 1 ♀; Ponta Grossa, Pq. Est. Vila Velha; 25°14'S, 49°59'W; 23 Nov. 2001; G. A. R. Melo; DZUP. – Rio de Janeiro • 1 ♂; Nova Friburgo, Sans Souci; 1050 m; Nov. 2004; P. Grossi leg.; light; coleçao E. & P. Grossi; Anchieta fumosella (Westwood, 1867), det. R. J. P. Machado; INPA. • 1 ♂; Itatiaia, P. N. Itatiaia, casa pesquisador; 22°27'21"S, 44°36'30"W; 05 Dec. 2015; A. P. M. Santos and D. M. Takiya leg.; light trap; DZUP. – Santa Catarina • 1 ♀; Blumenau; F. Müller leg.; McLachlan Coll. B. M.; NHMUK, 1938-674, BMNH (E) - 1241392. • 1 ♀; same data as for preceding; Blumenau, Virgil; 1987; v. d. Weele leg.; det. L. A. Stange; FSCA. • 1 ♂; Nova Teutonia; 27°11'S, 58°23'W; 300–500 m; Jan. 1971; F. Plaumann leg.; INPA. – São Paulo • 1 ♀; Jardim Botanico; 18 Dec. 1987; R. L. Jeane leg.; MCZ. • 1 ♂ 1 ♀; Same data as for preceding; Luiz Antonio, Estação Ecológica Jataí; 21°36'47"S, 47°45'04"W; 01 Oct. 2008; R. Lara leg.; light trap; DZUP. • 2 ♀; same data as for preceding; 16 Sep. 2009, Lara and team leg; Malaise; DZUP. • 2 ♂; same data as for preceding; 13 Feb. 2008; R. Lara and team leg.; light trap; DZUP. • 2 ♂; same data as for preceding; 12 Mar. 2008; R. Lara and team leg.; DZUP. • 1 ♂; same data as for preceding; 17 Sep. 2008; R. Lara and team leg.; DZUP. • 3 ♂ 1 ♀; same data as for preceding; 01 Oct. 2008; R. Lara and team leg.; DZUP. • 2 ♂; same data as for preceding; 29 Oct. 2008; R. Lara and team leg.; DZUP. • 3 ♂; same data as for preceding; 27 Sep. 2008; R. Lara and team leg.; DZUP. • 2 ♂ 3 ♀; same data as for preceding; 28 Jan. 2009; R. Lara and team leg.; DZUP. • 5 ♂; same data as for preceding; 28 Feb. 2009; R. Lara and team leg.; DZUP. • 1 ♂ 1 ♀; same data as for preceding; 18 Mar. 2009; R. Lara and team leg.; DZUP. • 2 ♂; same data as for preceding; 01 Apr. 2009; R. Lara and team leg.; DZUP. • 1 ♀; same data as for preceding; 09 Apr. 2009; R. Lara and team leg.; DZUP. • 1 ♂; same data as for preceding; 15 Apr. 2009; R. Lara and team leg.; DZUP. • 1 ♂; same data as for preceding; 29 Apr. 2009; R. Lara and team leg.; DZUP. • 1 ♂; same data as for preceding; 27 May. 2009; R. Lara and team leg.; DZUP. • 1 ♂; same data as for preceding; 30 Sep. 2009; R. Lara and team leg.; DZUP. • 1 ♂; same data as for preceding; 15 Oct. 2009; R. Lara and team leg.; DZUP. • 2 ♂; same data as for preceding; 29 Oct. 2009; R. Lara and team leg.; DZUP. • 1 ♀; same data as for preceding; 08 Dec. 2009; R. Lara and team leg.; DZUP. • 1 ♂; São Luiz Paraitinga, PESM nucleo St Virginia; 22°19'18"S, 45°05'43"W; 21 Nov. 2011; N. W. Perioto and team leg.; Malaise (7); DZUP.

Diagnosis.

The body coloration pattern of this species may be yellow with dark brown stripes or nearly completely dark brown. The forewing membrane is pale amber, with darker area on distal ½ of costal field, subcostal space, radial space, area between R + M and CuP, and area adjacent to RP. The hind wing is short and narrowly oval. On the male genitalia, the tergite IX has a posterolateral tuft of long setae, which surpasses the posterior margin of ectoproct. The sternite IX is blunt, sclerotized, with lateral margins concave. The gonocoxite IX is thin, short, and sinuous, with posterior apex blade-shaped, with sharp tip, sometimes slightly lanceolate, or with two or three tiny, preapical processes. On the female genitalia, the gonapophyses VIII form a posteromedially projected, tubular process with blunt to pointed apex.

Description.

Measurements. Male (n = 5). Forewing length: 11.0– 12.8 mm; Hind wing length: 6.6–8.57 mm. Female (n = 3): Forewing length: 7.7–12.15 mm; Hind wing length: 4.6–7.63 mm.

Coloration (Fig. 10). Head. Mostly black, vertexal region with small pale brown mark above compound eye; with lateral rows of dark brown setae. Postgena with the inner ½ dark brown, outer ½ yellowish with dark brown suffusions. Antennal scape pale brown ventrally, dorsally dark brown, pedicel dark brown; flagellum brown to dark brown, except for ~ 8–10 yellow apical flagellomeres, some specimens with flagellum completely dark; toruli surrounded by pale brown. Clypeus with dark amber posterolateral corners; labrum with pale brown margins; mandible dark amber; maxillary palpus brown; labium black except ligula, pale brown, labial palpus brown, palpimacula pale. Thorax. Pronotum black. Episternum brown, postfurcasternum dark brown. Meso- and metanotum blackish brown, with pale brown setae; pteropleura mostly blackish brown, with orange areas on anepisterna, with brown setae. Foreleg. Coxa mostly dark reddish brown with yellowish area at apex; trochanter dark brown with pale brown ventral band. Femur mostly brown with paler areas on outer surface and small pale brown area dorsally at level of anteroventral primary process; tibia mostly dark reddish brown, with pale brown basal area, clavate setae pale; basitarsus pale amber with amber apex, second to fourth tarsomere pale brown. Mid- and hind legs. Mid-leg with brown coxa and trochanter, femur, and tibia pale brown at base and apex, the rest of the surface brown; tarsomeres completely pale brown. Hind leg with coxa, trochanter and femur brown, with interspersed pale and dark brown setae; tibia mostly pale brown with darker median area; tarsus pale brown. Wings. Forewing membrane pale amber, with darker area on distal ½ of costal field, radial space, subcostal space, area between R + M and CuP, and area adjacent to RP; with small dark amber spot posteriorly on anal region; pterostigma pale brown with pale posterior and distal margins; venation mostly brown with R + M, proximal ½ of CuA and A 3 darker. Hind wing pale amber, base of subcostal space and anterior ½ of ra-rp 1 slightly darker; pterostigma amber or pale brown, venation brown. Abdomen. Dark brown, with pale brown setae.

Chromatic variation (Fig. 10). Head. Mainly yellow, vertexal region with dark brown posterior transverse band and arc-shaped marking on frontal sutures; frons dark brown. Antennal scape bicolor, yellow on proximal ½, dark brown on distal ½; flagellum yellow to pale brown, setae pale to dark brown. Labrum pale brown, clypeus yellow; mandible yellow with pale amber apex; maxillary palpus amber; labium bicolor, yellow on mentum and brown on ligula, labial palpus amber. Thorax. Pronotum brown with lateral yellow stripes; episternum pale brown, with yellow margins; postfurcasternum pale brown medially, yellow on margins. Mesonotum dark brown with yellow lateral stripes; metanotum dark brown with medium yellow areas; pteropleura yellow with longitudinal brown bands, setae pale yellow. Foreleg. Coxa mostly yellow, with brown areas on basal ½; trochanter bicolor, yellow ventrally, brown dorsally. Femur mostly yellow, anterior surface with irregularly shaped brown marking extending from center to apex; posterior surface with a brown J-shaped marking on medial region; tibia yellow on basal ½, brown to dark brown on distal ½; basitarsus pale amber, remaining tarsomeres yellow. Mid and hind legs. Mid-leg mostly yellow with dark brown spots on trochanter and tibia. Hind leg with coxa bicolor, trochanter brown, femur mostly brown with yellow base and apex; tibia mainly yellow with brown posteromedian region. Abdomen. Brown, with yellow anterior and posterior margins of sclerites.

Morphology (Fig. 10). Head. Diamond-shaped in frontal view, smooth, vertexal region domed above compound eyes; coronal suture discrete; paraocular area concave. Antenna moniliform, flagellum with 43–61 flagellomeres, discoidal in shape, those of distal ½ wider, except at apex narrower; all flagellomeres with medial ring of short setae. Compound eye hemispherical, as wide as ½ of interocular distance at toruli level. Thorax. Pronotum nearly as long as wide, with groove contiguous to lateral and distal margins; in lateral view, posterior margin slightly raised, the rest of the surface straight; entire surface with scattered, thick setae, arising flush the pronotal surface. Postfurcasternum quadrangular. Mesonotum slightly wider than long, with a few scattered fine setae, metanotum ~ 3 × as wide as long, mostly glabrous covered with microtrichia. Pteropleura covered with abundant fine and long setae. Foreleg. Coxa as long as femur, cylindrical, slightly expanded at preapical region, with abundant, fine and long setae; trochanter subtrapezoidal, densely setose, dorsally with a tuft of long and thick setae, anterior surface with blunt process. Femur robust, with abundant fine, long setae; closing surface with posteroventral row of integumentary specializations fully developed, composed of tubercle-shaped processes with conical Stitz organs, proximally with a more developed sub-basal, spine-shaped process; adjacent row of thickened setae with globular base reduced to distal ¾ or ½; anteroventral row of processes reduced to proximal region and apex, composed of tubercle-shaped processes, with conical setae; basal primary process present, spine-shaped, curved; adjacent row of thickened setae with globular base present on distal 4 / 5. Tibia almost as long as femur, curved, glabrous, closing surface with a row of prostrate setae; apical region of anterior surface with a patch of clavate setae. Basitarsus with long lanceolate process; basal ½ with clavate setae on anterior surface, ventrally with a row of prostrate setae. Mid- and hind legs. Mid- and hind leg covered with abundant fine setae; tibia unmodified, thin. Wings. Forewing oval, venation setose, trichosors present along wing margin except at base; costal space slightly widened medially, humeral vein sometimes forked, with 13–18 subcostal veinlets; pterostigma rectangular; subcostal space with single medially located crossvein; Sc vein abruptly bent posteriad at proximal margin of pterostigma to merge the RA; rarp 2 straight with 3–6 veins arising from it, three from rarp 1; M fused basally with R; RP base located near separation of M and R, M fork near such separation; 1 r-m located between RP base and M fork, forming a small trapezoidal cell; 6–8 gradate crossveins present; Cu deeply forked, CuA apically branched; CuP proximally angled, approaching A 1, forked slightly beyond 1 m-cu; A 1 simple, A 2 forked. Hind wing notably smaller and narrower than forewing; costal space narrow and reduced, with seven or eight subcostal veinlets; C and Sc fused at 1 / 3 of wing length; Sc vein abruptly curved posteriad at proximal margin of pterostigma, to merge the RA; subcostal space without crossveins; pterostigma elongated, narrow; radial space with single sinuous crossvein; two or three veins arising from rarp 1, 1–3 from rarp 2. M vein forked at level of R fork; Cu deeply forked, CuA gently bent, distally forked, first branch forked; CuP distally gently bent anteriorly, near posterior wing margin, with two or three branches; CuP sometimes fused to posterior margin, and then diverges again forming an arch; A 1 arched, A 2 short, simple. Abdomen. Medially widened, tergites without keeled processes.

Male genitalia (Fig. 11 A – E). Tergite IX notably narrower medially than laterally, lateral region with posteromedial tuft of long, thick setae, which surpass posterior margin of ectoproct. Sternites VIII and IX fused, fusion line moderately distinct; sternite IX posteroventrally projected, blunt, sclerotized, lateral margins concave. Gonocoxites IX thin, sinuous, as long as gonapophyses X, posterior apex curved laterally, flattened, blade-shaped, with sharp tip, sometimes slightly lanceolate, or with two or three tiny preapical processes. Ectoproct ovoid, with subparallel anterior and posterior margins; ventrally on inner surface with a more sclerotized region covered with 25–29 thick, conical setae on posterior ½ and thin, short setae on anterior ½. Gonocoxites X ventrally canaliculated, with anterior apex expanded and dorsally bent, posterior region with lateral and dorsal processes; base of gonostylus X thickened, triangular, concave in lateral view, the rest of the structure whip-shaped, short, with apical portion recurved. Gonapophyses X straight, narrow, with spatulate and dorsally recurved tips; gonapophyses joined by membrane, forming a V-shaped structure. Gonocoxites XI narrow, U-shaped, medial lobe expanded, composed of two concave plates medially joined by narrow bridge; lateral arms slightly arched, anterior apex curved.

Female genitalia (Fig. 11 F – I). Sternite VII subrectangular, with the posterior margin broadly concave, densely setose towards posterior margin. Tergite VIII slightly narrower medially than laterally, enclosing the spiracle of the segment, lateral margin trapezoidal. Gonocoxites VIII as two medially fused, trapezoidal, concave plates; gonapophyses VIII as a convex plate, posteromedially projected, forming a tubular process with blunt to somewhat pointed apex; lateral part of gonapophyses VIII as a trapezoidal plate hidden by the tergite IX + ectoproct. Tergite IX + ectoproct elongated, lateral margin quadrangular. Gonocoxites IX elongated, sinuous, and narrow, as long as the last five abdominal segments together. Gonapophyses IX reduced to a pair of tiny sclerites hidden by membranes, situated behind the gonocoxites IX base. Bursa copulatrix narrow, short, membranous. Spermatheca complex and coiled; proximal section long and thin, forming four coils; medial section thicker than proximal section, short, forming a couple of convolutions; distal section wider than medial section, progressively expanded towards apex, sac-like, forming a convolution; fertilization canal duct, long, spiral-shaped, with four or five convolutions, covered by the proximal section of the spermatheca; fertilization canal elongated, thin, J-shaped, covered with microfilaments.

Distribution.

Brazil (Bahía, Distrito Federal, Goiás, Minas Gerais, Rio de Janeiro, Paraná, Santa Catarina, Rio Grande do Sul, São Paulo, Tocantins).

Remarks.

This species is so far known only exclusively from Brazil and has been previously recorded in the states of Bahia, Distrito Federal, Goiás, Minas Gerais, São Paulo, Rio de Janeiro, Paraná, Santa Catarina, Tocantins, and Rio Grande do Sul (Penny 1982 a; Penny and da Costa 1983; Carvalho and Corseuil 1991; Alvim et al. 2019; Schuster and Machado 2021). Herein, most of these records are confirmed based on the material examined. The holotype of Mantispa (Trichoscelia) fumosella lacks specific collecting data, but Westwood (1867) in the original description of the species indicated “ Amazonia ” as the collecting site. This information was questioned by Penny (1982 a) and herein such idea is corroborated, as this species is restricted to Central and Southwestern Brazil. Araújo et al. (2021) reported several specimens reared from artificial nests of bees in the Amazonian portion of the Brazilian state of Mato Grosso which might be A. fumosellus, however detailed examination of such material was not possible at present, so the presence of this species in that state will require corroboration (R. J. P. Machado, pers. observation).

Anchieta fumosellus expresses a Batesian mimicry pattern with wasps of the family Vespidae (Buys 2008; Rasmussen and Ardila-Camacho 2021). Penny (1982 a) realized that this species exhibits a considerable variation in the body color pattern. Some specimens can be completely dark brown, other can be yellow with dark brown stripes, while other can have intermediary colorations, tracing different species of vespid wasps as happens in the mantispid genus Climaciella Enderlein, 1910 (Rasmussen and Ardila-Camacho 2021). However, the male genitalia and wing coloration and venation between the yellow and dark forms are identical. This led to the synonym of A. nobilis (whose type has yellow body with dark stripes) with A. fumosella (in which the type has a dark brown body) (Penny 1982 a). Herein that hypothesis is confirmed, after a careful examination of the morphology of several specimens of both morphs. Based on the distribution and coloration pattern and variation of this species, the model for the dark morph is likely the wasp Agelaia vicina (de Saussure, 1854), whereas the model species for the yellow morph is probably Agelaia multipicta (Haliday, 1836) (R. Lopes, pers. comm.).

In the phylogeny of Symphrasinae, this species was recovered as an intermediary species between the braconid mimicking A. fasciatellus and the clade containing all the bee mimicking species. This evolutionary pattern matches with the phylogeny of Hymenoptera, in which the parasitoid wasps of the family Braconidae are part of a large clade that diverged first in the phylogeny of Hymenoptera, then, within Aculeata the Vespoidea had an early divergence, while the bees or Antophila evolved as a highly specialized group within Apoidea (Peters et al. 2017).

This species can be recognized by the pale amber area and the anterior region of the forewing, which resembles the folding of the forewings of vespid wasps, the short and narrow hind wing, and the male genitalia. The male genitalia of this species are similar to that of A. nebulosus as both have a tuft of long setae on the lateral region of the tergite IX, arched goncoxites XI, and narrow and short male gonocoxites IX lacking digitiform processes. By contrast, the female goncoxites + gonapophyses VIII are similar to those of A. fasciatellus, but the spermatheca is similar to that of the species included within the bee-mimicking clade.