Revision of the Oriental and East Palaearctic genus Coryptilus Gibson (Hymenoptera, Chalcidoidea, Eupelmidae) Author Fusu, Lucian Corresponing author Author Kawano, Taisuke Author Park, Duk-Young Insect Biosystematics Laboratory, Department of Agricultural Biotechnology, Seoul National University, Seoul 08826, Republic of Korea. Email: dypark 91 @ snu. ac. kr text Zootaxa 2019 2019-08-19 4657 2 317 332 journal article 22542 10.11646/zootaxa.4657.2.5 8f8ea846-51e9-4b8f-9c6c-cb4467e09a37 1175-5326 3772307 D5822F40-4141-405F-8521-79ABE3153141 Coryptilus longicervix Kawano & Fusu sp. nov. Figs 32–55. Coryptilus sp.; Gibson 1995: 189 . Etymology . From the Latin longus (long) and cervix (neck), in reference to the elongate pronotum that differentiates it from C. indicus and C. circalatus . Type material . Holotype ♀ : JAPAN , HONSHU, Yamaguchi-ken , Yashisro-jima , Higashi-agenosho , Adaka ( OEṄĦAEẤTẆẤdz ) [Yashisro-jima, Higashi-agenosho, Adaka in Japanese], 10 Nov. 2014 , Yoshimitsu Higashiura / HOLOTYPUS Coryptilus longicervix Kawano et Fusu, T. Kawano det., 2017 ( ELKU ) . Condition of holotype : left front leg missing, glued by right side on triangular point, with metasoma glued on another triangular point, and left antenna, right fore leg and left fore wing mounted on slide; uncontorted. Metasoma broken after taking the picture in figure 32. Paratypes : 22♀ . JAPAN : Honshu , Aichi-ken , Nisshin-shi , Akaike , 30.ix.2006 , K. Yamagishi leg. (sw) [sweeping] ( 1♀ ELMU ) ; Yamaguchi-ken , Yashiro-jima , Yura , 14.x.2014 , Yoshimitsu Higashiura ( 1♀ ELKU ) ; Kyushu , Oita-ken Tsukumi , 30.viii.1955 , T . Tachikawa (sw) [sweeping] ( 1♀ ELKU ) ; Hyuga [former name for Miyazakiken], Aoidake , 9.x.1950 , Y. Hirashima ( 1♀ ELKU ) ; Miyazaki , Kitago , Inohae , 31°43’N 131°23’E , 21.ix. 1996 , 200m, L. Masner , ss J-55 ( 1♀ CNC , no. 2017-32 ) . SOUTH KOREA : GB [ Gyeongsangbuk-do ], Gyeongsan-si , Daehak-ro 280, Youngman University, N 35°49’11.6” E 128°45’53.6”, L. Fusu ( 1♀ AICF , no. 2017-25 ). Gyeong- sangbuk-do, Gyeongsan-si, Daehak-ro 280, Yeungnam Univ., 35°49′30″N 128°45′39″E , 20.ix.–3.x.2015 , J.W. Lee ( 1♀ YNU ); ditto , 35°49′12.9″N 128°51.9″E, 29.vii.–7.ix.2015 , J.W. Lee ( 2♀ YNU , one AICF with permis- sion from YNU ) ; Gyeongsangnam-do , Jinju-si , Ibanseong-myeon , Daecheon-ri , Gyeongnam For. Env. Res. Inst. , 35°9′39.7″N 128°17′41.3″E , 30.viii.–16.ix.2012 , J.H. Hwang ( 1♀ YNU ) ; Ulsan-si , Nam-gu , Ok-dong Grand Park , 35°31′50.5″N 129°17′2.3″E , 18.viii.–29.viii.2013 , N.S. Bang ( 1♀ YNU ) . TAIWAN : N. Taiwan , Taipei , 1♀ 26– 29.ix.1960 , 1♀ 24.x.1960 , 1♀ 5–10.xi.1960 , 2♀ 22–28.xi.1960 , 1♀ 15–30.ix.1963 , 1♀ 1–15.x.1963 , K. S. Lin ( TARI ) ; Hsintien , Taipei , 15–30.ix.1963 , K. S. Lin ( 1♀ TARI ) ; Wufeng , 400m , Hsinchu Hsien , 14–16.vii.1982 , K. C. Chou & C. C. Pan ( 1♀ TARI ) ; E. Taiwan , Fuli , Hwalien-Tungho , Taiting Hsien , 9–11.xi.1982 , K. C. Chou & S. P. Huang ( 1♀ TARI ) ; S. Taiwan , Kenting , Pingtung Hsien , 5–9.viii.1982 , K. C. Chou & C. N. Lin ( 1♀ TARI ) ; Kenting , Pingtung Hsien , 5–9.xii.1982 , S. C. Lin & S. P. Huang ( 1♀ TARI ) . FIGS 16–2 ­ 3 . Coryptilus indicus female paratype 2017-19 (photo LF ): 16 , lateral habitus; 17 , head two-thirds; 18–2­0 , head in lateral, dorsal, and frontal view; 2­1 , antenna; 2­2­ , fore wing; 2­3 , acropleuron. Scale bars represent 1 mm in 16 and 0.2 mm in 17–23 . FIGS 2­4 – 31 . Coryptilus indicus female paratype 2017-17 (photo LF ): 2­4 , lateral habitus; 2­5 , dorsal habitus; 2­6 , mesosoma, dorsal; 2­7 , metasoma and wings; 2­8 , mid leg; 2­9 , hind leg; 30 , fore wing; 31 , mesoscutellum and propodeum. Scale bars represent 1 mm in 24 and 25, and 0.2 mm in 26–31 . Description. Female. Length without ovipositor 2.8–3.3 mm ( holotype 2.9 mm ). Colour. Head (Figs 36–38) dark, frontovertex and upper facing part of parascrobal region with faint bronze and coppery reflections; face and gena green to bluish-green, metallic, and scrobal depression brighter green; lower face and gena additionally with bronze to violet reflections. Mandible dark-yellow basally, dark-brown apically. Maxillary and labial palpi dark brown. Scape yellowish, dark-brown basally and along ventral and dorsal margins (Fig. 40). Pedicel dark brown, narrowly pale-yellow apically. Flagellum dark brown, with sensorial region of clava paler, yellowish-white apically. Mesosoma (Figs 32, 33, 45, 46) mostly dark brown to black with weak bronze to violet luster under some angles of light except anterolateral surface of pronotum with green luster; mesonotum with blu- ish-green to bronze reflection, more evident on posterior depressed region (Figs 33, 50); tegula brown; propodeal callus with blue luster anteromedially and violet luster latero-posteriorly (Fig. 35). Fore wing veins pale yellow to light brown. Front leg (Fig. 45) dark brown, except trochanter, trochantellus and tarsus basally paler. Mid leg (Fig. 43) almost uniformly dark brown with apico-dorsal dark yellow macula on femur; mesotibial pegs and spur brown; mesotarsus with basitarsus and 2 nd tarsomere brown, sometimes both lighter dorsally, 3 rd and 4 th tarsomeres brownish yellow, and apical tarsomere brown except yellowish basally. Hind leg (Fig. 44) dark brown with part of coxa, trochanter, trochantellus, and first four tarsomeres paler, yellowish brown. Metasoma (Figs 41, 51) dark brown, with faint purple luster under some angles of light, except sometimes Gt1 paler in anterior half and with a basal pair of brown maculae (Fig. 41); syntergal flange translucent. Ovipositor sheaths dark brown, narrowly brownish-yellow apically. Head. In dorsal view (Fig. 37) 1.3–1.4× as wide as long and 1.6–1.7× as wide as mesoscutum; temples 0.16× eye length; interocular distance 0.3× head width; distance between prominent apex of parascrobal region and anterior FIGS 32­–40. Coryptilus longicervix female (photo TK for holotype and TARI specimen, LF for others): 32­ , lateral habitus ( holotype ); 33 , thorax, dorsal ( holotype ); 34 , lateral habitus in a light bleached specimen (TARI); 35 , propodeum ( holotype ); 36–38 , head in lateral, dorsal, and frontal view (LF 2017-32 ); 39 , fore wing (LF 2017-25 ); 40 , antenna (LF 2017-32 ). Scale bars represent 1 mm in 32 and 34, and 0.2 mm in 33, 35–40. margin of eye as long as MPOD. Head in lateral view (Fig. 36) subtriangular, 1.1× as long as high, with parascrobal region prominent and abruptly bent at about midheight. Head in frontal view (Fig. 38) 0.6–0.7× as high as broad, with about dorsal margin of torulus in line with lower orbit. Ocelli in a right-angled to slightly acute triangle, with POL 6× OOL and OOL 0.6–0.8× MPOD. Eye 1.3–1.4× as long as broad, with short, comparatively dense pilosity. Malar space 0.5× eye length; malar sulci converging downwards; mouth opening 1.1–1.3× as broad as malar space. Scrobal depression deep (Figs 37, 38), about as wide as high, carinately margined laterally and dorsally, reticulate. Vertex transversely reticulate, with more isodiametric cells behind ocellar triangle; occiput along upper margin with a transverse belt of white, semierect, curved setae and a tuft of black, upturned setae on upper corner behind eye; frontovertex and dorsally facing part of parascrobal region densely setose with long, erect, lanceolate, white setae. Upper face, parascrobal region and gena reticulate, with short, prostrate setae. Lower face and interantennal region imbricate-alutaceous to granular, setose similar to gena. Antenna (Fig. 40) with scape 2.8–3.4× as long as broad and 0.8–0.9× eye length, compressed laterally, with prostrate white setae. Pedicel plus flagellum 1.3× head width; pedicel in lateral view 2.0–2.5× as long as broad, 1.9–2.3× as long as fl1. Flagellum basally narrower than pedicel; fl1 short, 1.1× as long as broad, fl2 elongate, 1.8–1.9× as long as broad; both fl3 and fl4 the longest, fl3 1.1–1.2× as long as fl2, 1.8–1.9× as long as broad; fl4–fl8 1.5–1.7, 0.9–1.0, 0.6–0.8, 0.6–0.7, and 0.5–0.6× as long as broad respectively; clava 2.0–2.2× as long as broad. FIGS 41–44. Coryptilus longicervix female (photo TK for holotype , LF for others): 41 , metasoma, dorsal ( holotype ); 42­ , acropleuron (LF 2017-25 ); 43 , mid leg (LF 2017-32 ); 44 , hind leg (LF 2017-32 ). Scale bars represent 0.2 mm . Mesosoma. Elongate, in dorsal view (Fig. 33) 1.8× as long as broad. Pronotum in dorsal view divided by deep median groove, with collar comparatively elongate, 0.4× as long as broad, imbricate to reticulate, densely covered with long, black, bristle-like setae on pronotal neck and with sparse white setae on collar (Figs 33, 36). Mesoscutum concave with indistinct median and lateral lobes, and with median lobe 1.1–1.2× as long as broad, lateral lobes with incurved carinate line; mostly reticulate except lateral lobes imbricate-alutaceous changing to reticulate towards posterior depressed area where cells smaller than those of lateral lobes; with brown bristle-like setae on median and lateral lobes but with sparse, white, hair-like setae on posterior depressed area (Fig. 33). Mesoscutellar-axillar complex 1.1–1.2× as long as broad, 0.6× as long as mesoscutum; mesoscutellum reticulate; axilla reticulate; covered with black bristle-like setae pointing forward (Fig. 33). Metanotum with dorsellum as a narrow vertical flange. Propodeum (Fig. 35) anteromedially with V-like carinate emargination and foramen Λ-like carinate, the anteromedian and foraminal carinae connected mesally; with translucent region between foraminal carinae and foramen; with elongate comma-like plical furrow. Callar region slightly convex with finely coriaceous anteromedian region and smooth latero-posterior region. Prepectus (Fig. 49) divided by vertical furrow into coriaceous anterior region and reticulate posterior region. Tegula elongate, alutaceous, with adpressed black, bristle-like setae. Acropleuron (Figs 42, 49) with antero- and posterodorsal regions coarsely reticulate, central and ventral regions shiny, finely alutaceous; anteriorly with a line of long, white, lanceolate setae along acropleural sulcus. Mesepisternum strongly reticulate anteriorly, smooth otherwise, with sparse, long, white, hair-like setae. Profemur with 3 long lanceolate setae in a row ventroapically. Mesofemur posteroventrally with a line of 4 or 5 erect, slightly lanceolate setae, of which only distal-most seta longer and about as long as tibia height (Fig. 43); mesotibia with 4 apical pegs in 1 row; basitarsus to tarsomere 4 ventrally with 16, 6 or 7 ( 6 in holotype ), 3, and 2 pegs in row on each side, respectively; those on basitarsus in serrate row (Fig. 52). Metafemur basally with a group of long, appressed, hair-like setae, but otherwise inconspicuously setose (Fig. 44). FIGS 45–50. Coryptilus longicervix female (photo DYP): 45 , lateral habitus; 46 , dorsal habitus; 47–48 , head in lateral and dorsal view; 49 , prepectus and acropleuron; 50 , mesoscutum to metanotum, dorsal. Scale bars represent 1 mm in 45 and 46 and 0.2 mm in 47–50. Fore wing reduced in size (Figs 39, 53), extended to about base of Gt5, 4.3–4.6× as long as broad and narrowovoidal with subparallel margins; strongly infuscate and with dense, bristle-like dark brown setae except for white setae on broad hyaline region behind apical half of submarginal vein and on posteriorly tapered oblique streak behind distal third of marginal vein. Basal cell setose except variably extensively bare at the angle between basal fold and submarginal vein (Fig. 39). Costal cell 9.1–11.6× as long as broad, 1.3–1.5× length of marginal vein. Marginal vein 4.0–4.2× as long as stigmal vein (measured including uncus). Stigmal vein straight to curved; stigma with obscure, short, thick uncus. Postmarginal vein 1.4–1.6× as long as stigmal vein, and 0.3–0.4× as long as marginal vein. Metasoma broadest at level of cerci or slightly before, length excluding ovipositor sheaths 1.2× that of mesosoma and 2.0–2.5× as long as broad, with white setae except asetose dorsally on Gt1–Gt4; tergites comparatively shiny, Gt1 and Gt2 coriaceous to finely reticulate; following tergites with similar but stronger sculpture, imbricatecoriaceous. Gt1 narrowly emarginate to incised, posterior margin of Gt2–Gt5 broadly emarginate, Gt6 outcurved. Ovipositor sheaths short, extended slightly beyond apex of metasoma, about 0.5× as long as metatibia. Male. Unknown. Variation. The white streak on the fore wing can be narrower than shown in figures 39 and 53. The specimens from Taiwan (in TARI) have a much lighter mesosoma. However, this is probably an artefact due to light-bleaching because the oldest specimens (collected in 1960) are the lightest (Fig. 34), those from 1963 are slightly darker, and those from 1983 are almost as dark as those from Japan and South Korea . Similar light-bleaching was documented also in other small-bodied Hymenoptera ( Popovici et al . 2018 , 2019 ). Some specimens have the mesoscutum more metallic violaceous to dark blue (Fig. 50) compared to the holotype (Fig. 33) but the intensity of the metallic colour depends also on the viewing angle and lighting. FIGS 51–55. Coryptilus longicervix female (photo DYP): 51 , metasoma, dorsal; 52­ , apex of mesotibia and mesotarsus; 53 , fore wing; 54 , antenna; 55 , propodeum. Scale bars represent 0.2 mm . Distribution. Japan (Honshu and Kyushu islands), South Korea , Taiwan . Remarks. Besides the more obvious differences mentioned in the first couplet of the key, females of C. longicervix differ from C . indicus and C. circalatus by having: mesoscutum with white setae on posterior depressed area; lateral lobes more imbricate-alutaceous than reticulate (Fig. 33); tegula with only dark adpressed setae; occiput with line of erect black setae above belt of white setae interrupted mesally (Fig. 37). Females of both C . indicus and C. circalatus have the mesoscutum entirely covered in dark, inconspicuous setae and reticulate lateral lobes (Figs 15, 26); tegula with scattered, erect, black, bristle-like setae and anteriorly with a few shorter, white, lanceolate setae; occiput with a complete line of black erect setae above belt of white setae (Figs 4, 19). Coryptilus longicervix further differs from C. indicus in having a dark brown mesosoma with comparatively stronger metallic luster; in C. indicus the mesosoma has the acropleuron and many dorsal parts dark orange or at least light brown and mostly non-metallic, except the pronotum dorsally and partly laterally is brown with dark green metallic luster and sometimes the mesoscutum and mesoscutellum are partly brown with faint blue-green and purple luster under some angles of light. FIG. 56. Distribution map of the genus Coryptilus : pink/light grey dots for C. indicus , black dots for C. circalatus , blue/medium grey dots for C. longicervix . Discussion . Coryptilus longicervix is present in what we recognize as both the Palaearctic and Oriental regions. Its Oriental distribution is established by its presence in lowland areas of Taiwan (Fig. 56). According to Kreft & Jetz (2013) Taiwan is included in a Palaearctic to Oriental transitional zone because it has both temperate ecosystems at higher altitudes and tropical forest on the lowlands ( Udvardy 1975 ). However, Holt et al . (2013) include Taiwan in the Oriental realm and not in their newly proposed Sino-Japanese realm which, according to Kreft & Jetz (2013) , is a transitional zone. Olson et al . (2001) include Taiwan in the Oriental realm (as Indo-Malayan). It is interesting that females of all three known species of Coryptilus have reduced wings, but wing reduction is not as extreme as in some species of Anastatus ( Gibson et al . 2012 , Peng et al . 2017 ) or Eupelmus Dalman (Fusu 2017) . In C. circalatus and C. indicus the fore wing, even if shortened, always reaches at least to Gt5 or Gt6 and is similar in shape to a normally developed wing ( cf . Figs 12 and 13). In C. circalatus most females have the fore wing at least slightly shortened but a few fully winged individuals are known, whereas in C. indicus all known females have shortened fore wings. In C. longicervix known females are brachypterous, with the fore wing reaching to the base of Gt5 so as to be similarly long as the reduced wings of females of the other two species, but differing conspicuously by being much narrower and less similar to a normally developed wing. It is also interesting that males of Coryptilus have not been collected so far. If similar to other Eupelminae , males of all three species should be macropterous. If they have a head shape that is at least slightly similar to females they should be easily recognizable. However, they could possess a non-modified head, and in this case they could look deceptively similar to Anastatus or Zaischnopsis males based on the phylogenetic hypotheses of Gibson (1995) , and hence would not be recognizable as Coryptilus without association through rearing or genetic markers. All the Anastatus males from the same collecting event with paratype no. 2017-25 of C. longicervix were DNA barcoded using the standard protocol for another project (L. Fusu, unpublished data) and they were all associated with females; hence none was a Coryptilus male.