Anastatus (Anastatus) mantoidae, Motschulsky, 1859

Gibson, Gary A. P., 2020, Redescription of Anastatus mantoidae Motschulsky, the type species of Anastatus Motschulsky 1859, and Anastatus echidna (Motschulsky), the type species of Cacotropia Motschulsky 1863, with respect to taxonomy of Anastatus (Hymenoptera: Eupelmidae: Eupelminae), Zootaxa 4748 (3), pp. 485-513 : 488-497

A. (Anastatus) mantoidae Motschulsky

Figs 1–6

Anastatus mantoidae Motschulsky, 1859: 116–117 . Lectotype ♀ [ZMMU] designated by Bouček, 1988: 550.

Anastatus mantoidae ; Motschulsky, 1863: fig. 11.

Podagrion mantoidae ; Dalla Torre, 1898: 369; Schmiedeknecht, 1909: 117; Pruthi & Mani, 1940: 3, fig. 2 [misidentifications].

Anastatus mantoidae ; Gahan & Fagan, 1923: 12 [generic status]; Mani, 1938: 42 [catalogue]; Islam & Hayat, 1986: 57 [catalogue]; Mani, 1989: 691–692, fig. 160 [figure reproduced from Pruthi & Mani, 1940, fig. 2 and labelled as A. mantoidae ; misidentification]; Grissell, 1995: 242 [nomenclatural history].

A. (Anastatus) mantoidae ; Narendran, 2009: 85–86, 94, fig. 39 [key].

Type material examined. Four cards with a total of ten females and two males:

1. Card ( ZMMU) with three females ( Fig. 1A ): lectotype ♀ mounted by ventral surface in front of two other fe- male paralectotypes, the card with “ ♀ L-Type” printed in black ink beside the lectotype on one side and an unrelated hind leg glued to the card on the other side; lectotype uncontorted, entire, but apices of both fore wings missing; original labels—“ ♀ ” [small square label] / “[?]” [small circular label with pin through center obscuring writing] / “ Aús Man | - tis Eiern | Ceylon” [rectangular label]; subsequent labels—“front ♀ | LECTO- | TYPE” [purplebordered circular label] / “ ♀ LECTOTYPE | (front one) | Anastatus | mantoidae Motsch. | det. Z. Bouček, 1979 ” [rectangular label] / “Zool. Museum | Moscow Univ. | Russia ” [red rectangular label]. One paralectotype almost complete, the other missing its gaster and most appendages.

2. Card ( ZMMU) with six, variably well preserved females ( Fig. 1C ): two mostly complete mounted by ventral surface, one lacking gaster mounted by ventral surface, one without mesosoma with gaster mounted by ventral surface but head mounted with mouthparts faced dorsally, one without head mounted by dorsal surface, and one lacking gaster mounted laterally. Original labels similar to lectotype card.

3. Card ( ZMMU) with two males ( Fig. 3A ): mounted by ventral surface, the anterior-most male complete except antennae lacking beyond pedicels; posterior-most male lacking right antenna beyond scape and left fore and hind wing detached and glued to card on either side of anterior-most male. Original labels similar to lectotype label except with ♂ instead of ♀ symbol.

4. Card ( BMNH) with one female in pieces glued to rectangular card, labelled “ CEYLON: | Mt. Nurra Ellia, / aus Mantis Eiern / ex coll. Motschulsky | (coll. Nietner) / PARA- | LECTO- | TYPE ” [blue border circular label] / “parts of a paralec- | totype of ♀ | Anastatus mantoidae / NHMUK 010838800”, consisting of one complete flagel- lum beyond fl1 on card, head attached by right side to card with both antennae complete to fl1, mesosoma mounted dorsoventrally without wings, legs or propodeum, but right fore wing and one front, middle and hind leg glued individually to card.

Other material examined. INDONESIA: Java, c.c. Widodaren, Tanggoel, VIII.1936, H.J. de Fluiter, Anastatus sp. Ch. Ferrière det. (4♀, 1 ♂ BMNH; male CNC Photo 2019-65). Sumatra, East Sumatra, Asahan, 50–60 m, X/1079 [?], F. Schneider, 1934-36, from Gambir [ Rubiaceae : Uncaria ]-pests (7♀, 1♂ BMNH; male CNC Photo 2019-67), spec. 3B (1♀ BMNH), must be close to mantoidae Motsch. ! (1♀ BMNH). THAILAND (all QSBG, on indefinite loan to CNC): Chaiyaphum, Tat Tone NP, dry dipterocarp forest, near swamp at Sab samboon forest unit, 16°01.059’N 101°58.603’E, 674m, 5–12.XI.2006, Tawit Jaruphan, Malaise trap, T1137 (1♀; CNC Photo 2019-66). Loei, Phu Kradueng, Forest Protection Unit Loei.5 (Phakbung), 16°50.540’N 101°41.663’E, 406m, 11–12.II.2007, Wuthicahi Kwanjam, pan trap, T1492 (1♀). Nakhon Nayok, Khao Yai NP, Lam Takong viewpoint, 14°25.762’N 101°23.527’E, 732m, 12–19.X.2006, Wirat Sook-kho, Malaise Trap, T744 (1♀, exoskeleton only, DNA voucher LFMM00039_0101, extracted by L. Fusu and J.-Y. Rasplus). Phetchabun, Khao Kho NP, mixed deciduous forest, 16°32.5612’N 101°02.479’E, 537m, 26.XII.2006 – 2.I.2007, Somchai Chachumnan & Sainksington, Malaise trap, T1187(1♀).

Description of type specimens. FEMALE ( Figs 1 , 2 ). Length about 3.8 mm. Head with frontovertex ( Fig. 1F ) and parascrobal region and scrobal depression above level of dorsal angle of interantennal prominence ( Fig. 1D ) dark with slight reddish-coppery luster, but scrobes, lower part of parascrobal region, and most of interantennal prominence ( Fig. 1D ) green or with slight blue luster under some angles of light, and gena, temple and occiput variably distinctly green under different angles of light ( Figs 1 D–F). Antenna pale, yellow or pedicel and flagellum brownish-yellow but similar in colour to scape ( Figs 1E , 2A ). Labiomaxillary complex with palpi yellow (observed from one specimen). Pronotum ( Fig. 2C ) mostly brown dorsally but lateral panel partly green to bluish. Mesonotum ( Fig. 2C ) with mesoscutum anteriorly or at least convex part of mesoscutal medial lobe to near posterior angle brown, but otherwise green to bluish depending on angle of light; scutellar-axillar complex similarly dark brown as most of medial mesoscutal lobe, without distinct metallic luster. Tegula ( Figs 2C, D ) and prepectus ( Fig. 2D ) brown. Mesopleurosternum ( Fig. 2D ) with setose region of mesopectus mostly brown but acropleuron green to bluish over about anterior half and brownish over about posterior half. Metanotum ( Fig. 2E ) brown. Propodeum ( Fig. 2E ), at least under some angles of light, extensively green to blue, most conspicuously on callus. Legs ( Fig. 1B ) with femora and tibiae mostly brown but trochanters, trochantelli, knees, tibiae apically and tarsi much paler, more yellowish. Fore wing ( Fig. 2B ) discal region with distinct, posteriorly widened, brownish-infuscate region behind stigmal and postmarginal veins extending posteriorly to at least level of medial fold (cf. Fig. 5A : mdf), but basally uniformly hyaline or at most very slightly and inconspicuously infuscate near parastigma, and with all setae dark, hair-like; vein complex yellow. Gaster ( Figs 1A, B ) dark brown except Gt1 apically and Gt2 whitish-hyaline so in dorsal view with subbasal white band (cf. Fig. 4B ), and with St1–St3 similarly whitish-hyaline (cf. Fig. 4A ); syntergal flange ( Fig. 2F : stf) yellow to yellowish-hyaline; ovipositor sheaths yellow ( Fig. 2F ).

Head ( Figs 1 D–F) with scrobal depression bare, but otherwise greenish parts with white setae, the setae hairlike on lateral surface and very slender-lanceolate on frontal surface, and with dark hair-like setae on dark parts; head in dorsal view ( Fig. 1F ) comparatively thick, subrectangular; HW: HH: HL: IOD = 8.4: 7.3: 5.5: 2.7; OOL: POL: LOL: MPOD = 0.5: 1.9: 1.3: 1.4; eye bare, EH: EW = 5.3: 3.8; scrobal depression ( Fig. 1D ) with lateral margin abrupt or raised as low ridge, and curved toward anterior ocellus but not margined dorsally, with obscurely delimited dorsal margin separated from anterior ocellus by distance about equal to two-thirds minimum diameter of ocellus; frontovertex ( Figs 1D, F ) with vertex mesh-like reticulate, ocellar triangle more or less imbricate, and frons lateral and anterior of anterior ocellus mesh-like coriaceous-pustulate or somewhat imbricate; scrobal depression ( Fig. 1D ) above level of interantennal prominence mesh-like pustulate to imbricate but scrobes mesh-like reticulate; interantennal prominence similarly reticulate to reticulate-rugulose as lower parascrobal region; clypeus comparatively more finely coriaceous, and gena lateral to clypeus irregularly roughened. Antenna ( Fig. 2A ) with scape ( Fig. 1E ) tubular, slightly curved, about 4.5× as long as greatest width; pedicel about 2× as long as apical width and greatest (ventral) length of fl1; fl1 subquadrate with dorsal length sometimes slightly less, and ventral length sometimes slightly greater, than apical width, and subsequent funiculars not unusually modified, increasingly widened apically such that fl2–fl4 longer than wide, fl5 subquadrate, and fl6–fl8 wider than long; clava about equal in length to com- bined length of fl6–fl8 plus apical two-thirds of fl5. Mandibles normal for genus, with tiny ventroapical tooth and broad dorsoapical margin.

Mesonotum ( Fig. 2C ) with mesoscutum uniformly covered with hair-like, brownish setae except mesoscutal lateral lobes bare dorsomedially; convex part of anteromedial lobe triangular, with slightly sinuate, posteriorly convergent sides extending slightly beyond half length of mesoscutum, and mostly transversely alutaceous-imbricate (cf. Fig. 4H ) to somewhat punctulate or strigose-imbricate anteriorly, but depressed posteromedial region between lateral lobes entirely, distinctly mesh-like reticulate; mesoscutal lateral lobe longitudinally strigose-reticulate posteriorly or with obscure longitudinal carina posteromedially near transscutal articulation, but dorsolongitudinally anterior of longitudinal sculpture more finely mesh-like coriaceous to transversely strigose relative to more coarsely mesh-like sculptured lateral and mesally inclined surfaces of lateral lobe; axilla mesh-like reticulate anteriorly to anteromedially, but obliquely strigose to strigose-reticulate posteriorly; scutellum low-convex with more or less concentric U-like striae so as to appear longitudinally strigose to strigose-reticulate on either side of median. Acropleuron ( Fig. 2D ) finely sculptured, mostly finely, longitudinally alutaceous-striate but becoming very shallowly mesh-like reticulate posteriorly. Fore wing ( Fig. 2B ) length about 3× width; mv almost 6× length of stv and about 0.8× length of costal cell, and pmv about 3× length of stv (approximate because apex of stv not clearly visible in any specimen); costal cell bare dorsally, but with complete row of setae ventrally, the setae in more than one row basally and apically; basal cell (cf. Fig. 5C : bac) and cubital area (cf. Fig. 5C : cua) uniformly setose but vanal region (cf. Fig. 5C : vna) bare; discal region uniformly setose. Profemur with ventral margin evenly curved apically; mesotibia with 6 apical pegs in patch rather than straight row (visible only on one leg of one specimen). Metanotum ( Fig. 2E ) normal for genus, with comparatively small and narrow, vertical, dorsally rounded dorsellum behind posterior surface of scutellum ventrally. Propodeum ( Fig. 2E ) normal for genus, with anterior margin V-like incised medially for reception of dorsellum and foramen carinately margined medially and broadly incurved almost to apex of V-like incision to differentiate more or less bowtie-like medial region (deeply concave, triangular region on either side of V-like medial region) and inclined surface of plical region mesal to spiracle relative to convex surface of callus lateral of spiracle; plical region apparently very finely mesh-like coriaceous (region not clearly visible or clean in any specimen); callus setose laterally, with white hair-like setae.

Gaster ( Figs 1 A–C) normal for genus, about as long as mesosoma; posterior margin of Gt1 much more deeply incised than only slightly emarginate Gt2–Gt4, Gt5 with posterior margin broadly incurved, and Gt6 with posterior margin broadly rounded ( Fig. 2F ); Gt1 and Gt2 apparently smooth and shiny but at least subsequent tergites finely mesh-like coriaceous; Gt1 bare dorsally but subsequent tergites with at least two transverse rows of setae; syntergal flange ( Fig. 2F : stf) with broadly rounded posterior margin, transverse but at most only about twice as wide as its medial length; ovipositor sheaths protruding only slightly beyond apex of syntergal flange.

MALE ( Fig. 3 ). Length about 3 mm. Head ( Figs 3C, E ) with similar colour pattern as female, dark dorsally with very slight reddish luster on vertex and frons to about level of dorsal margin of scrobal depression and medially within scrobal depression below anterior ocellus, but otherwise apparently mostly green (face not clearly visible in frontal view). Antenna ( Fig. 3D ) uniformly yellow. Pronotum concealed by head. Mesonotum ( Figs 3F, G ) mostly green but mesal angles of axillae and scutellum broadly reddish-violaceous dorsolongitudinally and with some coppery luster laterally ( Fig. 3F ). Prepectus and acropleuron ( Fig. 3G : pre, ac) mostly greenish, mesepimeron ( Fig. 3G : ep2) blue to purple and at least lateral and apparently ventral surface of mesepisternum greenish except for diagonal brown band within femoral depression that diverges near midlength to form a Y-like yellowish-brown band. Tegula ( Fig. 3G ) yellowish to yellowish-brown. Fore wing ( Figs 3B, H ) hyaline with yellowish vein complex. Legs beyond coxae entirely yellowish except outer surface of metafemur ( Fig. 3B : fm3) extensively dark ventrolongitudinally. Propodeum ( Fig. 3I ) blue to purple. Gaster dark brown.

Head ( Figs 3C, E ) with similar setal distribution and colour pattern as for female; HW: HL: IOD = 8.0: 5.0: 3.5 (head height not measurable because of position of head); OOL: POL: LOL: MPOD = 0.8: 2.7: 1.4: 1.5; eye sparsely microsetose, EH: EW = 4.6: 3.6 (malar space not measurable because of position of head); scrobal depression ( Figs 3C, E ) bell-shaped, with lateral margins abruptly margined only over about ventral half but extended obscurely to near anterior ocellus; sculpture similar to female with vertex ( Fig. 3E ) transversely reticulate-rugulose, ocellar triangle more imbricate, frons on either side of anterior ocellus mesh-like coriaceous, scrobal depression above interantennal prominence ( Figs 3C, E ) mesh-like reticulate to reticulate-rugulose and scrobes transversely rugulosestrigose (sculpture of interantennal prominence, parascrobal region and lower face not clearly visible). Antenna ( Fig. 3D ) with all flagellomeres subequally wide, together forming robust-filiform flagellum; setal and mps patterns typical for genus, i.e. each flagellomere microsetose and with numerous, dense mps in multiple rows per flagellomere; accurate measurements of scape, pedicel and flagellomeres not possible because of mounting position but fl2 the longest flagellomere and not conspicuously curved, and subsequent flagellomeres decreased in length such that at least apical funicular quadrate and clava somewhat shorter than combined length of apical three funiculars. Mandibles not visible.

Mesonotum ( Figs 3F, G ) with mesoscutum uniformly covered with hair-like, whitish to light brownish setae depending on angle of light; mesoscutal medial lobe comparatively deeply mesh-like reticulate-rugose, mesoscutal lateral lobe somewhat more finely sculptured than medial lobe but entirely roughened, punctate-reticulate to reticulate-imbricate; scutellar-axillar complex with axilla mostly mesh-like coriaceous to alutaceous, anteromesal angle only obscurely mesh-like reticulate, and scutellum mostly mesh-like coriaceous-reticulate to reticulate-imbricate posteriorly. Prepectus ( Fig. 3G : pre) shallowly mesh-like reticulate. Mesopleuron with acropleuron ( Fig. 3G : ac, cf. Fig. 6F : ac) mesh-like coriaceous to shallowly reticulate anteriorly but shiny and smooth posteriorly; mesepisternum ( Fig. 3G , cf. Fig. 6F : eps2) with mostly very fine, subeffaced sculpture except more distinctly reticulate an- teroventrally; mesepimeron ( Fig. 3G : epm2) smooth and shiny except narrowly along dorsal margin. Metapleuron ( Fig. 3G : pl3) smooth and shiny except narrowly dorsally. Fore wing setal pattern ( Fig. 3H ) apparently normal for genus, i.e. without large speculum behind parastigma (uncertain because of condition of specimens); costal cell bare dorsally, and ventrally with 1 row of setae along length; relative lengths of smv: mv: stv: pmv = 5.6: 3.8: 1.0: 2.3. Propodeum ( Fig. 3I ) with complete median carina and short, irregular striae extending anterolaterally from carina and long foramen, though smoother on either side of median carina anteriorly to spiracle.

Metasoma with petiole ( Fig. 3I ) transverse-lenticular with anterior margin slightly emarginate medially; gaster with Gt1 bare dorsally, remaining tergites with at least 1 transverse row of dark setae.

Non-type variation. FEMALE. Length 3.5–3.9 mm. Head colour pattern similar to type females except reddish-coppery luster sometimes more extensive over frontal surface, including parascrobal region and interantennal prominence ( Fig. 4C ); two of four females from Thailand have the flagellum and pedicel brown, distinctly darker than the scape ( Fig. 5D ), and all have brown palpi. Mesosomal colour pattern also similar to type females, but sometimes more extensively green, with mesoscutum more uniformly green ( Figs 4F, H ) with varying amounts of blue to purple within the depressed posteromedial part of the mesoscutal medial lobe, and mesopleurosternum often with setose region and acropleuron greenish over at least about anterior half ( Fig. 4G ) and acropleuron sometimes with some purple luster below tegula. Mesoscutum with posteromedial depressed part more or less uniformly, even if somewhat less densely, setose than convex part of the anteromedial lobe ( Fig. 4F ). The females from East Sumatra have entirely brownish discal setae similar to type females, whereas the Java females, depending on angle of light, can have a few whitish setae within the more hyaline region behind the marginal vein, and the females from Thailand have quite a distinct region of white setae behind the marginal vein ( Figs 5A, B ) similar to that described for A. motschulskyi ; all females have some white setae within the basal cell ( Fig. 5C : bac). In structure, the interorbital distance is about 0.27–0.33 head width, the POL is about 1.3–1.6 times the maximum diameter of a posterior ocellus and 1.6–2.0 times the minimum diameter of the ocellus, and the scrobal depression is separated from the anterior ocellus by a distance distinctly less than the longitudinal diameter of the ocellus ( Fig. 4E : dso). In sculpture, the convex part of the mesoscutal medial lobe is similarly transversely, though more finely alutaceous-imbricate ( Fig. 4H ) than type females ( Fig. 2C ), and the depressed posteromedial part of the medial lobe is much more finely sculptured, mostly mesh-like coriaceous to shallowly reticulate-coriaceous to partly smooth and shiny posteromedially ( Fig. 4F ).

MALE. Length 2.75–3.0 mm. The males from Java and Sumatra are also similar in colour pattern to type males, with the frontovertex dark with reddish-violaceous luster ( Figs 6C, D ) as for type males ( Figs 3C, E ), though the scrobal depression lacks the mediolongitudinal reddish line ( Fig. 6C ) and the frontal surface of the head is bright green or with very limited blue to purple lusters on the lower face under some angles of light ( Fig. 6C ); the labiomaxillary complex is yellow ( Fig. 6A ); the legs are almost completely yellow with the metafemur less obviously darkened than for type males, though still at least inconspicuously brownish with metallic luster ventroapically ( Fig. 6A ). In structure, mandibular dentition is not visible but the antenna is subequally wide with the flagellomeres forming a robust-filiform flagellum ( Fig. 5F ), and otherwise structurally similar to type males except the interantennal prominence is continued dorsally through most of the scrobal depression as a variably distinct median carina ( Fig. 6C : arrow); fore wing with the costal cell ( Fig. 5G ) similarly setose as type males, the basal region with the basal cell ( Fig. 5G : bac) setose but the cubital (cua) and vanal (vna) areas as well as the mediocubital fold (mcf) bare, and the discal region with the cubital fold ( Fig. 5G : cuf) bare for a short distance beyond the basal cell but otherwise setose, including behind the parastigma adjacent to the basal cell so a distinct speculum is lacking. In sculpture, the upper ( Fig. 6F : uep2) and lower mesepimeron ( Fig. 6F : lep2) and the metapleuron ( Fig. 6F : pl3) are shiny and at least extensively smooth, though with obscure sculpture dorsally on the metapleuron and somewhat more distinct sculpture over at most about the ventral half of the lower mesepimeron.

Distribution 1. * Indonesia (Java, Sumatra), Sri Lanka, * Thailand.

Host. Eggs of Mantidae (Mantodea) .

Remarks. Bouček (1988: 550) stated that the type material of A. mantoidae in ZMMU consisted of three cards with a total of nine females and two males. He did not mention anything about the BMNH female labelled as paralectotype, which although mounted on a more modern card certainly is part of the original type material. Because the specimen is in separate pieces glued to the card it is even possible that it might be a combination of body parts missing from various specimens on the other ZMMU cards. Bouček (1988) also stated that he had seen specimens of A. mantoidae that had been reared from mantid eggs in Sabah ( Malaysia) in northern Borneo. These specimens are undoubtedly the BMNH specimens described below as part of a new species that is very similar to A. mantoidae .

Females and males of A. mantoidae are differentiated most readily from those of A. motschulskyi by the features given in Tables 1 and 2, respectively (see ‘Remarks’ for A. motschulskyi for further discussion).

Figures

FIGURES 1A–F. Anastatus mantoidae, female type material. A, card with lectotype; B, lateral habitus (LT); C, card with paralectotypes. D–F, head (LT): D, frontal; E, lateral; F, dorsal. Abbreviation: LT = lectotype.

FIGURES 2A–F. Anastatus mantoidae, female type material. A, antenna (LT); B, fore wing (LT); C, mesonotum (PLT); D, mesosoma, lateral (LT); E, apex of scutellum to base of gaster, dorsal (PLT); F, apex of gaster, dorsal (PLT). Abbreviations: LT = lectotype, PLT = paralectotype, stf = syntergal flange, stl = syntergal flange length, stw = syntergal flange width.

FIGURES 3A–I. Anastatus mantoidae, paralectotype males. A, card with paralectotypes; B, lateral habitus; C, head, frontodorsal; D, antenna; E, head, dorsal; F, mesosoma, dorsal; G, mesosoma, lateral; H, fore wing; I, propodeum and petiole. Abbreviations: ac = acropleuron, ep2 = mesepimeron, fm3 = metafemur, pl3 = metapleuron, pre = prepectus, ti3 = metatibia.

FIGURES 4A–H. Anastatus mantoidae, female (2019-66). A, lateral habitus; B, dorsal habitus; C, head, frontal; D, head, dorsal; E, frontovertex; F, mesonotum; G, mesosoma, lateral; H, anterior half of mesoscutum. Abbreviation: dso = distance between scrobal depression and anterior ocellus.

FIGURES 5A–G. Anastatus mantoidae. A–D, female (2019-66): A, fore wing; B, central part of fore wing; C, basal part of fore wing; D, antenna. E–G, male (2019-65): E, fore wing; F, antenna; G, basal part of fore wing. Abbreviations: bac = basal cell, mcf = mediocubital fold, mdf = medial fold, cua = cubital area, cuf = cubital fold, vna = vanal area.

FIGURES 6A–F. Anastatus mantoidae, male. A, lateral habitus (2019-67); B, dorsal habitus (2019-65); C, head, frontal (2019- 65) [arrow points to mediolongitudinal ridge]; D, head, dorsal (2019-65); E, mesosoma, dorsal (2019-65); F, mesosoma, lateral (2019-67). Abbreviations: ac = acropleuron, eps2 = mesepisternum, lep2= lower mesepimeron, pl3 = metapleuron, pre = prepectus, uep2 = upper mesepimeron.