Redescription of the types of species of Anastatus Motschulsky , 1859 ( Hymenoptera : Chalcidoidea : Eupelmidae ) described by

Six species of Anastatus Motschulsky, 1859 (Hymenoptera: Eupelmidae) were described from China in Chinese by J.K. Sheng and coauthors in 1997 and 1998: A. dexingensis, A. fl avipes, A. fulloi, A. huangi, A. meilingensis and A. shichengensis. This represents almost half the species of Anastatus recorded from China, but no keys were given to differentiate the species and the original descriptions included only simple line drawings to illustrate the species. Because recognition of these species is critical prior to clarifying the Anastatus fauna of China and of the eastern Palaearctic and Oriental regions, we have redescribed the six species in detail in English based on original type material, illustrating the species through macrophotography of type material and providing a key to differentiate females of the species.


Introduction
Anastatus Motschulsky, 1859 (Hymenoptera: Chalcidoidea: Eupelmidae) is an economically important genus whose members are mostly egg parasitoids and who are natural enemies of many pest insects in

Material and methods
This study is based on a total of 512 pinned specimens, almost all Anastatus, found in six boxes of the Sheng collection at Jiangxi Agricultural University, Nanchang, China.Although some specimens had escaped serious damage, 149 (29.1%) were completely destroyed and more than half of the remainder were poorly preserved, presenting damage to various extents by mildew or partly eaten by insects.The state of preservation of many of the specimens complicates accurate redescription of some characters, particularly exact wing setal patterns.Further complicating the study of Sheng's collection was that many specimens lacked labels, including type labels indicating the designated holotype, allotype or paratypes, and there was not a consistent one-to-one match between each description record and each specimen, which made it difficult to check against the original literature.
The first step in studying Sheng's collection was an attempt to find any specimen with the same label data as recorded in the original descriptions.Sheng sometimes only labeled the first specimen in a column, the rest not bearing any labels.Because all the specimens lacked type labels, we considered the female labeled with "sp.nov." as the holotype of the respective species of that name.All specimens were examined with a stereoscope, and those females that possessed the same characteristics as the female considered to be the holotype, and that were consistent with Sheng's original descriptions, were considered as part of the type series for that species.Males were associated based on position with females in the collection, identical collection labels, and comparison with original descriptions.Redescriptions of the species are based on all specimens considered to form the type series.Observations were made using a fiber optic illuminator and a Nikon SMZ-U stereo microscope.Specimens were photographed with a Leica DC500 digital camera attached to a Leica Z16 APO macroscope and the serial images obtained combined with Auto Montage.Type specimens of A. splendens Nikol'skaya, 1952 (Canadian National Collection of Insects, Arachnids & Nematodes, Ottawa) and A. formosanus Crawford, 1913 (United States National Entomological Collection, Washington, D.C.) were also examined for comparison.
Measurements for structure follow Gibson (1989).A Nikon SMZ-U microscope fitted with a 14 mm ocular grid having 140 divisions was used for measurements.Fore wings of females and males were all taken at a magnification of 60; antennae, distance between ocelli and inner orbit measurements were taken at a magnification of 100.
CoLor.Head dark with metallic purple luster on frons and temple, gena and occiput metallic green; ocelli dark brown; maxillary and labial palpi dark brown to black (Figs 2-3).Antenna (Figs 3,5) with scape yellowish-brown and pedicel dark brown with metallic purple luster (flagella missing but presumably dark based on original description).Pronotum (Fig. 4) almost black, with purple and bluishgreen luster under some angles, except median groove dark brown; and posterolateral corner black anterior to spiracle.Mesoscutum (Fig. 4) with anterior convex part of medial lobe black with metallic purple to green luster, posterior concave part with metallic greenish-blue luster; lateral lobe dark with metallic green luster.Scutellar-axillar complex (Fig. 4) black with greenish-blue metallic luster.Tegula, acropleuron and prepectus dark brown (Fig. 6).Front leg (Fig. 7) with coxa and femur dark brown with metallic luster, tibia and basal two tarsomeres dark brown, apical three tarsomeres brown; middle and hind legs dark brown except mesotarsus and apex of metafemur brown.Metanotum and propodeum (Figs 1, 4) dark brown.Gaster (Fig. 1) dark brown to black with base whitish.
Legs.Profemur (Fig. 7) with ventral margin evenly arched basally, but with a sharp, tooth-like angulation in apical quarter.Middle leg with spur as long as basitarsus; tibia with apical pegs in a patch; basal four tarsomeres with black pegs on each side.Hind leg with basitarsus as long as following three tarsomeres combined.

Notes
Of the original type material, only one labeled female and one unlabeled female were found.The labeled female, here considered to be the holotype, has three labels: "Dexing, Jiangxi, host: egg of Dendrolimus kikuchii", "1984, YU Yunxiang", and "Anastatus dexingensis sp.nov.Det.Sheng Jinkun".The holotype lacks both antennae beyond the pedicels, the left protibia, the left protarsus, both left wings and the right hind wing; the right fore wing is only partially preserved.The unlabeled female, here considered to be a paratype, lacks both antennae beyond the anelli, the hind wings, the front legs (lost accidently when the first author examined the specimen), the right protibia and protarsus; both fore wings are only partially preserved.

Remarks
The most important characters of females of this species are as follows: 1) profemur ventrally with a sharp tooth at apical quarter (Fig. 7) (not at apical third as stated in original description); 2) scrobal depression with scrobes shallow, lateral margin very weak dorsally (Fig. 5); 3) mesoscutum with less setae and with a different pattern (Fig. 4) than for A. shichengensis (Fig. 45), and with anterior convex part of medial lobe V-shaped; 4) posterior concave part of mesoscutum very short, only one-quarter the length of the scutellum (Fig. 4).The paratype originally had its left front leg, and the tooth of the profemur was clearly evident, but the leg was lost accidently when the first author examined the specimen.In the key of Kalina (1981), A. dexingensis keys out to A. splendens, but in A. splendens the posterior concave part of mesoscutum is as long as the scutellum (Nikol'skaya 1952), and the profemur is not toothed.Sheng & Wang, 1997 Figs 8-18 Anastatus flavipes Sheng & Wang in Sheng et al., 1997: 60-61, figs 10-13.
Legs.Profemur (Fig. 8) with ventral margin evenly arched, not toothed.Middle leg (Fig. 9) with spur as long as basitarsus; tibia with apical pegs in a patch; basal four tarsomeres with black pegs on each margin.Hind leg with basitarsus as long as following two tarsomeres combined.

Notes
Of the original type material, 18 specimens were found that resemble the original description and illustrations provided for A. flavipes.One labeled female in good condition, here considered to be the holotype, has two labels: "Shangrao, Jiangxi Prov., host: egg of Dendrolimus kikuchii (Drury)", and "15-25 June 1984, YU Jingting".One labeled male in good condition, here considered to be the allotype, has two labels the same as the holotype.Other specimens considered as paratypes: 6 ♀♀, 4 ♂♂, Dexing (part of Shangrao City) or Shangrao, Jiangxi Prov., 15-25 June 1984, YU Jingting; 1 ♂, Yushan, Jiangxi Prov., 17 Nov. 1984, YU Jingting; 1 ♀, Liantang, Jiangxi Prov., early August 1955; 4 ♀♀, Xintai, Shandong Prov., July -August 1987, FAN Di.All the specimens from Heilongjiang Province were destroyed.The label dates of the 12 specimens from Dexing and Shangrao labeled by Sheng are in conflict with the published data.We suspect that 15 June is abbreviated for 15-25 June.

Remarks
Females of A. flavipes resemble those of A. fulloi, but the body is yellowish-brown (Fig. 9) rather than metallic green (Fig. 24); scrobal depression with lateral margin not distinct dorsally, but about 1.5-2.0× diameter of ocellus from anterior ocellus (Fig. 12) rather than 0.8-1.10 × diameter of ocellus from anterior ocellus (Fig. 20); and fore wing with basal infuscate setose region about 2.2 × wider than hyaline cross-band (Fig. 10) rather than 3 × wider than hyaline cross-band behind MV (Fig. 23).Males differ from those of A. fulloi by the antennal clava being as long as the apical three funiculars, much shorter than the apical four funiculars (Figs 15,18).In the key of Kalina (1981), females of A. flavipes key out to A. formosanus, but in A. formosanus the lateral margins of the scrobal depression reach the anterior ocellus, the frontovertex is as wide as an eye, and the anterior convex part of the medial lobe is 0.78 × the total length of the mesoscutum.Sheng & Wang, 1997 Figs CoLor.Head (Figs 19-21) dark with coppery-green metallic luster on, gena, temple and occiput, and frons with metallic purple luster; ocelli light brown, maxillary and labial palpi dark brown to black.Antenna  with scape brown, pedicel brownish-black with metallic bluish-green luster, flagellum brownish-black.Pronotum brown with purple luster under some angles, and posterolateral corner black anterior to spiracle.Mesoscutum (Fig. 24) with anterior convex part of medial lobe metallic golden reddish-green, posterior concave part dark brown with coppery green metallic luster; lateral lobe brown with metallic purple luster.Scutellar-axillar complex (Fig. 24) metallic green to purple.Tegula and prepectus brown, acropleuron dark brown with metallic purple luster.Legs mostly dark brown, tarsi, tibiae apically, and knees brown.Metanotum and propodeum dark brown.Gaster (Fig. 23) black with base yellowish-brown.
Body.Pronotum in dorsal view acute-triangular, smooth to very finely coriaceous, concave posteromedially and bare at anterior margin of collar, one long seta and four short setae posteriorly anterior to each spiracle; with distinct median groove.Mesoscutum (Fig. 24) with anterior convex part of medial lobe shield-like with sides subparallel basally and abruptly convergent apically, 0.75 × total length of mesoscutum, longer than broad, coarsely punctate-reticulate and bare, but posterior concave part strongly concave, smooth to reticulate, with white long setae in a radial pattern; lateral lobe inclined and imbricate.Scutellar-axillar complex (Fig. 24) convex, coarsely punctate-reticulate, scutellum 1.1-1.2× as long as broad.Acropleuron with anterior one-third finely coriaceous and with translucent short hair-like setae, posterior two-thirds finely strigose.Macropterous; fore wing (Fig. 23) 1.7 mm in length, about 3 × as long as wide, with apex rounded and extending beyond gaster; basal plate with a row of five dark setae; basal cell sparsely setose with thinner setae than setae of disc; costal cell ventrally with a row of setae along length, and dorsally bare; submarginal vein with a row of seven white setae; disc with brownish infuscation basally and apically of hyaline cross-band behind MV except about apical one-fifth slightly more hyaline along wing margin, with basally infuscate region 3 × wider than hyaline cross-band; disc densely setose with comparatively thick brown setae except for white setae of hyaline cross-band; hyaline cross-band curved, not extending to junction of MV and STV; SMV: MV: PMV: STV= 32-35: 27-29: 12-15: 5-6.
Legs.Profemur with ventral margin evenly arched, not toothed.Middle leg with spur as long as basitarsus, tibia with pegs in a patch; basal four tarsomeres with black pegs on each side.Hind leg with basitarsus as long as following three tarsomeres combined.
gaster.Shorter than mesosoma, ovipositor sheath exserted for distance slightly shorter than length of syntergum.

Notes
Of the original type material, 16 specimens were found that resemble the original description and illustrations provided for A. fulloi.One female labeled with "Anastatus fulloi", here considered to be the holotype, has another two labels: "Meiling, Jiangxi Prov.host: Erthesina fullo (Thunberg)", and "5 July 1980, Sheng Jinkun".One male in good condition, here considered to be the allotype, has two labels: "Mt.Wuyi, Jiangxi Prov." and "10 July 1980, SHENG Jinkun".Other specimens considered as paratypes: 1 ♀, Mt. Wuyi, Jiangxi Prov., 10 July 1980, SHENG Jinkun;3 ♀♀, Meiling, Jiangxi Prov., 10 July 1980, SHENG Jinkun;5 ♀♀, 1 ♂, Meiling, Jiangxi Prov., 5 July 1980, SHENG Jinkun;2 ♀♀, Meiling, Jiangxi Prov., 19 July 1980, SHENG Jinkun;1 ♀, Xinjian, Jiangxi Prov., 30 September 1991, ZHAO Fengxia.Associated with these specimens are five leaf-remnants with attached egg shells collected on 5 July 1980.All the specimens were in one box in mixed series; we did not find any specimen from Mt. Wuyi collected on 16 July 1980.The specimens from Mt. Wuyi have handwritten labels, but the specimens from Meiling have a printed locality label and a written date label.It seems Sheng incorrectly labeled the Meiling specimens because he could not be in two such widely distant places (Mt.Wuyi is more than 300 Km from Meiling) in one day (10 July 1980).We suspect that he collected the bug egg masses on 5 July 1980, and the eupelmids then emerged successively.

Remarks
Females most closely resemble A. flavipes and key out to A. formosanus using Kalina (1981), but can be differentiated by the features discussed under the latter species.Males have the clava as long as the apical four funiculars or longer (Fig. 25) whereas males of A. flavipes have the clava as long as apical three funiculars but much shorter than the apical four funiculars (Figs 15,17).Sheng & Yu, 1998 Figs 28-33 Anastatus huangi Sheng & Yu, 1998: 6-7, fig. 2.
Body.Pronotum in dorsal view acute-triangular, very finely coriaceous, concave postero-medially, four short setae at anterior margin of collar and only one seta posteriorly anterior to each spiracle; median groove shallow.Mesoscutum (Fig. 31) with anterior convex part of medial lobe V-shaped, about 0.64 × total length of mesoscutum, coarsely punctate-reticulate and slightly convex, but posteriorly concave part smooth and shiny, with white long setae directed laterally; lateral lobe inclined and finely reticulate.Scutellar-axillar complex (Fig. 32) convex, longitudinally coarsely reticulate; scutellum 1.63 × longer than wide.Acropleuron (Fig. 28) with fine longitudinally aligned coriaceous sculpture, anterior portion almost bare.Brachypterous; fore wing (Fig. 32) about 0.6 mm in length, 4 × as long as wide, with apex rounded and extending to middle of gaster; basal plate with one dark brown seta; basal cell sparsely setose with much thinner setae than setae of disc; costal cell ventrally with few setae, and dorsally bare; disc with dark setae and brownish infuscation basally and apically of posteriorly tapered hyaline cross-band with much thinner white setae behind marginal vein; SMV extending half-length of wing, MV slightly shorter than submarginal vein, STV and PMV absent.
Legs.Profemur with ventral margin evenly arched, without tooth.Middle leg (Fig. 33) with spur as long as basitarsus; tibia with apical pegs in a patch, basal four tarsomeres with black pegs on each margin.Hind leg (Fig. 33) with basitarsus as long as following four tarsomeres combined.

Notes
Only one unlabeled female, here interpreted as the holotype, was found that resembles the original description and illustrations provided for A. huangi.This specimen is entire except for its left funicle and clava.

Remarks
The most important characters of females of this species are as follows: 1) head in dorsal view with width 1.43 × length; 2) clava slightly longer than the length of apical three funiculars (Fig. 30); 3) fore wing only extending to middle of gaster (Fig. 28); 4) mesoscutum with anterior convex part of medial lobe triangular, 0.64 × total length of mesoscutum (Fig. 31).Females resemble those of A. meilingensis, but are differentiated by the characters discussed under the latter species.In the key of Kalina (1981), A. huangi keys out to A. brevipennis Ashmead, 1904 andA. gastropachae Ashmead, 1904.However, females of A. brevipennis have the head in dorsal view 1.84 × as wide as long and the clava is shorter than the total length of the apical three funiculars (Kalina 1981), whereas in A. gastropachae the anterior convex part of the medial lobe of mesoscutum is 0.56 × the total length of the mesoscutum and the head in dorsal view is 1.96 × as wide as long (Kalina 1981).Sheng & Yu, 1998 Figs 34-40 Anastatus meilingensis Sheng & Yu, 1998: 5-6, fig. 1.
Legs (Fig. 36).Profemur with ventral margin evenly arched, without tooth.Middle leg with spur slightly shorter than basitarsus; mesotibia with apical pegs in a patch; basal four tarsomeres with black pegs on each side.Hind leg with basitarsus as long as following three tarsomeres combined.

Notes
Of the original type material, only four unlabeled females were found that resemble the original description and illustration provided for A. meilingensis.One complete female is here interpreted as holotype of the species.A second female lacks the funicle and clava of both antennae, and left front leg and right metatarsi.The third female lacks the right protarsi, and the fourth female lacks its left antenna, right funicle and clava, and right mesotarsus.

Remarks
Females of this species resemble those of A. huangi, but are differentiated by the following characters: fore wing much shorter, extending only slightly over middle of Mt2 compared to middle of gaster for A. huangi (cf. Figs 32,34); fore wing disc with apical one-third densely setose with yellowish-brown setae, without hyaline cross-band compared to fore wing disc of A. huangi with thinner brownish setae, and has a tapered hyaline cross-band behind marginal vein; anterior convex medial lobe rhomboidal compared to triangular for A. huangi with (cf.Figs 31, 38); hind leg with basitarsus as long as following three tarsomeres compared to following four tarsomeres in A. huangi (cf. Figs 33,36).In the key of Kalina (1981), A. meilingensis keys out to A. gastropachae, which is a brachypterous form of A. bifasciatus (Geoffroy, 1785) based on the rearings and synonym of Ishii (1938).The original description of A. gastropachae (Ashmead 1904) states that the fore wings are "fuscous, with a transverse band at base and [within] apical third hyaline", whereas A. meilingensis has the basal two-thirds hyaline and the apical one-third with yellowish-brown setae, which is similar to the fore wing color pattern of A. meilingensis females.Females of A. gastropachae determined in 2015 by L. Fusu in the Canadian National Collection of Insects, Arachnids & Nematodes (Ottawa) were examined by the senior author.Females have the medial lobe of mesoscutum triangular, the gaster longer than the mesosoma, and the scrobal depression distinctly delimited dorsally compared to A. gastropachae.Sheng & Wang, 1997 Figs CoLor.Head (Figs 42,44,46) dark with purple metallic luster on frons, and gena, lower face, temple and occiput with metallic green luster; ocelli, maxillary and labial palpi dark brown.Antenna (Figs 42,44,46) with scape yellowish-brown, pedicel dark brown to black with metallic purple luster, flagellum dark brown.Pronotum (Fig. 45) dark brown to black, with purple-green luster under some angles, median groove brown, and with posterolateral corner black anterior to spiracle.Mesoscutum (Fig. 45) with anterior convex part of medial lobe dark with golden-purple luster, posterior concave part smooth with metallic green luster, lateral lobe dark with purple luster.Scutellar-axillar (Fig. 45) metallic green.Tegula (Fig. 41) dark brown, prepectus with frontal surface black and lateral surface dark brown, and acropleuron (Fig. 41) black with metallic purple luster within basal anterior quarter.Legs (Fig. 41) dark brown to black except front and middle legs with tibiae and tarsi, and hind leg with tarsus light brown.Metanotum and propodeum black.Gaster (Fig. 41) black with base yellowish-brown.
Body.Pronotum (Fig. 45) in dorsal view acute-triangular, coriaceous, concave postero-medially, with five short setae along anterior margin on each side of collar and two long seta and six short setae posteriorly in front of each spiracle, median groove distinct.Mesoscutum (Fig. 45).Anterior convex part of medial lobe U-shaped, 0.66 × total length of mesoscutum, distinctly longer than broad, rugose-punctate and densely setose, and posterior portion strongly concave, smooth medially to imbricate laterally, with setae in radial pattern; lateral lobe inclined, imbricate.Scutellar-axillar complex (Fig. 45) convex, coarsely reticulate; scutellum 1.3 × as long as broad.Acropleuron (Fig. 41) imbricate to finely strigose, anterior one-quarter with short white hair-like setae.Macropterous, fore wing (Fig. 43) about 2.2-2.4 mm in length, 2.5 × as long as wide, with apex rounded and extending beyond gaster; basal plate with seven brown setae; basal cell sparsely setose with thinner setae than setae of disc; costal cell ventrally with a row of setae along length, and dorsally bare; submarginal vein with a row more than 12 white disc with brown infuscation basally and apically of hyaline cross-band behind MV except about apical one-fifth more hyaline along wing margin, with basally infuscate setose region 2 × wider than hyaline cross-band; disc densely setose with thick brown setae except for white setae of hyaline cross-band; hyaline cross-band curved, not extending to junction of MV and STV; SMV: MV: PMV: STV = 53-56: 33-36: 18-19: 9.
Legs.Profemur with ventral margin evenly arched basally, but expanded into blunt tooth within apical quarter.Middle leg (Fig. 41) with spur shorter than basitarsus; tibia with apical pegs in a patch; basal four tarsomeres with black pegs on either margin.Hind leg with basitarsus as long as following three tarsomeres combined.

Notes
Of the original type material, five females and two leaf-remnants with attached egg shells were found, all with labels by Sheng.One female labeled with "species name: Anastatus papillosae sp.nov.designated by Sheng Jinkun", here considered to be the holotype, has another two labels: "Shicheng, Jiangxi Prov., host: Tessaratoma papillosa" and "1979, Collector:".Another four females have the same collection label as the holotype.The holotype lacks both flagella.Of the paratypes, two are intact; one has lost both flagella, the middle legs, the left hind leg and right metatibia; one has only the head, left antenna, a broken right fore wing, as well as one middle and one hind leg remaining.
The published data for this species gives the date of collection as 1989, but the labels on the specimens have the printed year as "198_", and a written date "79", with "7" covering the number "8".Consequently, we suspect these specimens were collected in 1979, but that Sheng made a mistake with the published date.

Remarks
The most important differential characters of females are as follow: 1) profemur ventrally bearing a blunt tooth at apical quarter (Fig. 47); 2) anterior convex part of medial lobe U-shaped (Fig. 45); 3) posterior concave part of medial lobe with setae in two radial patterns (Fig. 45); 4) anellus wider than long, and 8 th flagellomere longer than wide; 5) SMV 1.3 × as long as MV.Females most closely resemble those of A. dexingensis, but A. dexingensis has a sharp profemural tooth (cf.Figs 7, 47) and the anterior convex part of medial lobe is 45).Using Kalina (1981), A. shichengensis keys out to A. splendens, but differs from the latter species in the same features given under remarks for A. dexingensis.