The Nepalese species of the genus Enicospilus Stephens, 1835 (Hymenoptera, Ichneumonidae, Ophioninae): a preliminary revision and identification key to species

A total of 10 species of Enicospilus (Hymenoptera, Ichneumonidae, Ophioninae) have previously been reported from Nepal. Six new species are described here (E. alleni Shimizu sp. nov., E. kakanicus Shimizu sp. nov., E. nepalensis Shimizu sp. nov., E. nikami Shimizu sp. nov., E. phulchokiensis Shimizu sp. nov., and E. tangi Shimizu sp. nov.), and 10 are newly recorded (E. ashbyi Ashmead, 1904, E. bifasciatus (Uchida, 1928), E. capensis (Thunberg, 1824), E. flavocephalus (Kirby, 1900), E. formosensis (Uchida, 1928), E. grammospilus (Enderlein, 1921), E. pudibundae (Uchida, 1928), E. purifenestratus (Enderlein, 1921), E. yonezawanus (Uchida, 1928), and E. zebrus Gauld & Mitchell, 1981) from Nepal. A preliminary identification key to the Nepalese species of Enicospilus is provided. The elevational pattern of Nepalese Enicospilus is briefly discussed. Enicospilus purifenestratus is also recorded for the first time from Brunei.

Enicospilus Stephens, 1835 is the largest genus of Ophioninae and predominantly tropical, with more than 700 species worldwide (e.g. Broad and Shaw 2016;Gadallah et al. 2017;Shimizu 2017;Johansson 2018). Enicospilus has been considered to be polyphyletic (Gauld 1985b), but there are no phylogenetic studies with comprehensive taxon sampling.
Nepal is a landlocked country between India and China's Tibet Autonomous Region (26°22'N-30°27'N, 80°4'E-88°12'E) in the central part of the Himalaya, about 800 km in latitudinal length and 140 km in longitudinal length (RAOnline 2019). Dramatic changes of altitude (from less than 100 m to more than 8,000 m) along the short longitudinal span in Nepal have created a very diverse climatic and topographic environment as well as a uniquely very rich species diversity of flora and fauna (Savada 1991;MFSC 2014;RAOnline 2019). Furthermore, Nepal is located between the Oriental and Palaearctic regions and is a melting pot of species originating from both regions (MFSC 2014). Therefore, Nepal is an interesting and important place to study biodiversity and biogeography. However, no researchers have studied Ophioninae of Nepal, although Gauld and Mitchell's (1981) great regional revision for Indo-Papuan Ophioninae included a few specimens and species from Nepal. Hence, only 10 species of Enicospilus have been recorded in Nepal (Gauld and Mitchell 1981), whilst 107 species have been reported from China and 73 from India , indicating a high potential species diversity of Enicospilus in Nepal.
This study aims to (1) review all previously recorded species in Nepal, (2) describe new Nepalese species, (3) newly record species from Nepal, (4) provide a preliminary identification key to the Nepalese species, and (5) briefly discuss the biogeography of the Nepalese fauna and species relationships with elevation.

Material and methods
A total of 707 specimens of Nepalese species of Enicospilus were examined, 148 of which are from Nepal and 559 from other countries (e.g. Brunei, China, India, Japan, Laos, and Taiwan). Specimens were observed using a stereoscopic microscope (SMZ1500, Nikon, Tôkyô, Japan). Photographs were taken using a single lens reflex camera (α7II, Sony, Tôkyô, Japan) with a micro-lens (LAOWA 25 mm F2.8 2.5-5× ULTRA MACRO, Anhui Changgeng Optics Technology Co., Ltd, Hefei, China and A FE 50mm F2.8 Macro SEL50M28, Sony, Tôkyô, Japan) and 2× teleconverter lens (SEL20TC, Sony, Tôkyô, Japan), captured in RAW format, developed using Adobe Lightroom Creative Cloud, and stacked using Zerene Stacker. All figures were edited in Adobe Illustrator 2019 and Photoshop Creative Cloud.
Morphological terms follow those of Broad et al. (2018). Legs and wings are described separately from the mesosoma. The lower face is defined as the area between the ventral margin of the clypeus and of the antennal sockets. Terms for surface sculpture follow Eady (1968) and Gauld and Mitchell (1981). Terms for wing veins and cells follow Broad et al. (2018) and those for characters of the discosubmarginal cell follow Gauld and Mitchell (1981) (Fig. 1). 'Sclerites' refer to the sclerites of the fore wing fenestra, which are differentiated as the proximal, central and distal sclerites, all or none of which might be absent in any one species. The indices follow Shimizu and Lima (2018) and Shimizu et al. (2019) and are listed below.
Indices for head GOI (geno-orbital index) = maximum breadth of eye in profile / maximum breadth of gena in same line Indices for fore wing AI (alar index) = length of 1m-cu&M between 2m-cu and bulla / length of 2rs-m CI (cubital index) = length of CU between 1m-cu&M and 2cu-a / length of 2cu-a DI (discoidal index) = maximum vertical distance between CU (between 2cu-a and 2m-cu) and 1m-cu&M / length of CU between 2cu-a and 2m-cu ICI (intercubital index) = length of 2rs-m / length of M between 2m-cu and 2rs-m SDI (second discoidal index) = length of CU between 2cu-a and 2m-cu / length of CU between M&RS and 1m-cu&M SI (sinuousness index) = maximum length between 1m-cu&M and a straight line connecting the intersection of M, 2m-cu, and 1m-cu&M and the intersection of 1m-cu&M and CU / distance between the intersection of M, 2m-cu, and 1m-cu&M and the intersection of 1m-cu&M and CU SRI (second recurrent index) = length of 2m-cu / length of CU between 2cu-a and 2m-cu Indices for hind wing NI (nervellar index) = length of CU between M and cu-a / length of cu-a RI (radial index) = length of RS between RA and rs-m / length of rs-m Indices for metasoma DMI (dorsal metasomal index) = length of dorsum of tergite 2 / length of dorsum of tergite 3 PI (petiolar index) = distance between base of tergite 1 and anterior margin of spiracle / distance between posterior margin of spiracle and apex of tergite 1 THI (thyridium index) = distance between anterior margin of tergite 2 and anterior margin of thyridium / maximum diameter of thyridium Wing characters are especially important for identifying ophionine species, but wings are almost always folded, wrinkled, and/or crooked. For accurate measurements of wing characters, the left wings have been removed from the body, placed between microscope slides in 99.9% ethanol, and photographed. Then, wings have been enclosed in paraffin paper, and the whole thing pinned under the respective specimen. Measurements were taken from photos using the software, Img-Measure ver. 1.14.
The degree of sexual dimorphism of Ophioninae is almost always very small, and most species can morphologically readily be distinguished without needing to dissect male genitalia (Gauld 1984). Therefore, the male genitalia is not dissected and new species are described based on holotype of both sexes in the present study, like previous ophionine studies (e.g. Gauld 1988;Broad and Shaw 2016;Shimizu 2017). Head: mandible bidentate apically and weakly to strongly tapered and twisted (e.g. Fig. 2A-D); ocelli moderately to very large, and posterior ocellus often close to or touching eye (e.g. Figs 3B-D, 5B-D, 7B-D); occipital carina complete; antennae longer than fore wing length (e.g. Figs 5A, 12A, 16A), usually with more than 50 flagellomeres.
Wings (e.g. Figs 1, 6F, 7F, 19F, 28F, 31B, D, F): pterostigma of fore wing fairly slender; vein 1m-cu&M of fore wing usually without ramulus; vein 2r&RS of fore wing usually more or less broadened proximally and/ or centrally, straight, sinuous, or bowed, not proximally abruptly angled; discosubmarginal cell of fore wing with fenestra, and often also with one or more sclerites; vein RS of hind wing usually straight and rarely weakly curved; vein RA of hind wing usually with 4-12 uniform hamuli.
Legs: inner mesal surface of fore tibial spur without a membranous flange; outer distal margin of mid and hind trochantelli usually simple and without a decurved tooth; hind tarsal claw moderately to strongly curved with pectinae, usually all pecten are more or less uniform shape and length and a distal one is not significantly longer than true apex of claw (e.g. Fig. 2I, J).
Metasoma (e.g. Figs 3A, 9A): very slender; tergite 1 with spiracle clearly far behind the middle; thyridium moderately to strongly developed, and oval to ellipsoidal; ovipositor straight and usually short, its length less than posterior depth of metasoma.
Differential diagnosis. Adult wasps of Enicospilus are moderately to very large insects and distinguished from other genera of Ophioninae by the following combination of character states: inner mesal surface of the fore tibial spur lacking a membranous flange; mandibles more or less narrowed apically and moderately to strongly twisted (e.g. Fig. 2A-D); fore wing discosubmarginal cell with a fenestra (e.g. Fig. 31B, D, F), extensive glabrous area, and often one or more sclerotised and pigmented sclerites and/or quadra (e.g. Figs 3F, 15F, 27F); posterior transverse carina of mesosternum complete.
The fore wing fenestra and sclerites are usually reliable characters for recognising Enicospilus species. However, similar sclerites of the fore wing fenestra are also known in the genus Dicamptus Szépligeti, 1905 and rarely in the genus Leptophion Cameron, 1901. Enicospilus species are distinguished from both Dicamptus and Leptophion by the mandibles (i.e. mandible always weakly to strongly tapered and twisted in Enicospilus, but very weakly tapered and not twisted in Dicamptus and Leptophion).
Biology. Species belonging to Enicospilus are koinobiont endoparasitoids of Lepidoptera, such as Noctuidae (e.g. Gauld and Mitchell 1981;Gauld 1985bGauld , 1988Broad and Shaw 2016;Broad et al. 2018). Adult female wasps usually lay eggs within late instar larvae of Lepidoptera, with some exceptions. Broad et al. (2018) summarised the biology of Ophioninae including Enicospilus. Both sexes of adults are very frequently attracted to the light and considered to be nocturnal or crepuscular (e.g. Shimizu and Maeto 2016;Shimizu 2017).

Identification key to Enicospilus species of Nepal
This is a preliminary key to the Nepalese species of Enicospilus because there are potentially many more unrecorded or undescribed species in Nepal and its adjacent areas.  5D). Fore wing with fenestra moderately long, its anterodistal corner separated from proximal end of the vein RS by more than 1.0× of 2rs-m; ICI = 0.2, SDI = 1.1-1.2; posterodistal corner of the second discal cell ca 65°; central sclerite drop-shaped, its major axis parallel to 2r&RS; vein 2rs-m bowed (Fig. 5F).
Head with GOI = 2.9 (Fig. 3C). Lower face 0.7× as wide as high, very finely coriaceous with fine punctures and setae (Fig. 3B). Clypeus 1.9× as wide as high, shiny and very finely coriaceous with fine punctures and setae, almost flat in profile, and its lower margin acute (Fig.  3B, C). Malar space 0.2× as long as basal mandibular width (Fig. 3B, C). Mandible weakly twisted by ca 25°, moderately long, evenly narrowed, its outer surface flat and smooth without a diagonal groove or a diagonal line of punctures (Fig. 3B, C). Upper mandibular tooth 1.4× as long as lower one (Fig. 3B). Frons, vertex and gena moderately shiny with fine setae (Fig. 3B-D). Posterior ocellus large and almost touching eye (  Mesosoma entirely very weakly shiny or not (Fig. 3E). Pronotum entirely striate. Mesoscutum 1.5× as long as its maximum width, very closely coriaceous with dense setae, very weakly shiny, evenly rounded in profile, and its anterior margin almost truncate in dorsal view and rounded in profile (Fig. 3E). Notauli absent (Fig. 3E). Scutellum moderately convex, very closely coriaceous with setae, with lateral longitudinal carinae reaching posterior end (Fig. 3E). Epicnemium densely punctate with setae. Epicnemial carina weak, almost straight and inclined to anterior, its dorsal end not reaching anterior margin of mesopleuron (Fig. 3E). Mesopleuron entirely closely longitudinally strigose (Fig. 3E). Submetapleural carina very strongly broadened anteriorly and forming a lobe. Metapleuron densely punctate to reticulate with setae, moderately swollen (Fig. 3E). Propodeum very strong-ly and abruptly declivous in profile; anterior transverse carina complete; pleural carina almost absent; anterior area longitudinally striate; spiracular area almost smooth with setae and strongly shiny; posterior area moderately subconcentrically strigose with a pair of strong posterior carinae laterally; propodeal spiracle elliptical, its outer margin not joining pleural carina by a ridge (Fig. 3E).
Legs. Outer surface of fore tibia with scattered short spines. Hind leg with coxa in profile 1.8× as long as deep; basitarsus 2.0× as long as second tarsomere; fourth tarsomere 0.6× as long as third tarsomere and 2.9× as long as wide; tarsal claw simply pectinate.
Distribution. Australasian and Oriental regions . Newly recorded from Nepal.
Colour (Fig. 4): body including interocellar area entirely testaceous; wings hyaline. Differential diagnosis. Enicospilus ashbyi is similar to E. pallidus (Taschenberg, 1875) and separated from it by a few characters of the central sclerite (pigmented part of central sclerite narrower in E. ashbyi and wider in E. pallidus). However, the sclerite characters (e.g. the shape and degree of sclerotisation of the central sclerite) exhibit a wide range of variation within both species, suggesting that there are cryptic species and that integrative taxonomy is needed to define species limits in this complex.   evenly tapered, its outer surface without a diagonal structure; upper mandibular tooth 1.2-1.3× as long as lower one; posterior ocellus close to eye; antenna with 54-56 flagellomeres and 20 th flagellomere 3.1-3.3× as long as wide.
Colour (Fig. 5): body entirely pale yellow with black marks on mesosoma, interocellar area, and posterior segments of metasoma; wings hyaline with two strongly infumate patches in the central part of the discosubmarginal cell (from anterior end of M&RS to base of 1m-cu&M) and the central part of the marginal cell (from anterocentral margin to base of RS).
Differential diagnosis. Enicospilus bifasciatus is a very distinctive species and no closely related species are currently known. Hence, it is easily distinguished from all Enicospilus by many characters listed in the above diagnosis as well as identification key, such as two strongly infumate patches in the central part of the discosubmarginal cell (from anterior end of M&RS to base of 1m-cu&M) and the central part of the marginal cell (from anterocentral margin to base of RS) of fore wing, characteristic shape of sclerites of discosubmarginal cell of fore wing (cf. Fig. 5F), very sparsely setose and almost glabrous proximal corner of marginal cell of fore wing, and small value of ICI (= 0.2).  Distribution. Eastern Palaearctic and Oriental regions ). Gauld and Mitchell (1981) recorded this species from Nepal.
Differential diagnosis. Enicospilus capensis is most similar to E. insularis and distinguished from it by the not clearly delineated central sclerite ( Fig. 7F) (well delineated in E. insularis), but diagnostic characters for these species are not strongly supported and need more study. Enicospilus capensis also more or less resembles E. ramidulus, but distinguished from it by the densely punctate and submatt to matt meso-and metapleurae (Fig. 7E) (meso-and metapleurae moderately punctate and never submatt to matt in E. ramidulus). Distribution. Australasian and Oriental regions ). Gauld and Mitchell (1981) recorded this species from Nepal.  (Fig. 8B-D): GOI = 2.9-3.1; lower face 0.6-0.7× as wide as high; clypeus weakly convex in profile, its lower margin subacute; mandible moderately twisted by 30-40°, moderately long, evenly tapered, its outer surface without a diagonal structure; upper mandibular tooth 1.3-1.6× as long as lower one; posterior ocellus Mesosoma (Fig. 8E): mesopleuron rather coarsely longitudinally striate; scutellum with lateral longitudinal carinae reaching anterior 0.8-1.0 and convergent posteriorly; metapleuron rather coarsely striate to strigose; propodeum evenly rounded to slightly declivous, its posterior area coarsely reticulate, outer margin of propodeal spiracle joining pleural carina by a strong ridge.
Colour (Fig. 8): body including interocellar area entirely testaceous; wings hyaline to weakly infuscate. Differential diagnosis. Enicospilus flavicaput is most similar to E. kanshirensis but can be distinguished from it by the slender central sclerite (Fig. 8F) (central sclerite stouter in E. kanshirensis, as in Figure 14F), and larger body size (i.e. fore wing length more than 17.0 mm in E. flavicaput but less than 15.0 mm in E. kanshirensis).
Wings (Fig. 9F): fore wing with AI = 0.4-1.5, CI = 0.6-0.8, ICI = 0.4-0.6, SDI = 1.1-1.2; fore wing vein 1m-cu&M centrally strongly angulated and broadened, 2r&RS almost straight; fenestra and sclerites of discosubmarginal cell of fore wing as in Figure 9F; fenestra of fore wing not very long and its anterodistal corner distinctly separated from proximal end of vein RS; proximal sclerite almost oval, isolated and not touching margin of fenestra, strongly pigmented; central sclerite strongly pigmented and sclerotised, linear and parallel to distal margin of the fenestra, positioned in mediodistal part of the fenestra; distal sclerite absent; proximal corner of marginal cell of fore wing almost uniformly setose; vein 1cu-a subinterstitial to antefurcal to M&RS by less than 0.2× 1cu-a length.
Colour ( Fig. 9): body including interocellar area entirely pale yellow with pale brown posterior segments of metasoma; wings hyaline. Differential diagnosis. Enicospilus flavocephalus is a very distinctive species, but its body size, colour pattern, and profile are very similar to E. xanthocephalus. Enicospilus flavocephalus is easily distinguished from E. xanthocephalus by many characters, such as the pale yellow interocellar area (Fig. 9B, D) (black in E. xanthocephalus) and centrally abruptly angled and broadened fore wing vein 1m-cu&M ( Fig. 9F)  synonymised by Gauld and Mitchell (1981: 424).
Wings (Fig. 10F): fore wing with AI = 0.2-0.6, CI = 0.2-0.9, ICI = 0.5-0.6, SDI = 1.1-1.3; fore wing vein 1m-cu&M weakly sinuous, 2r&RS almost straight; fenestra and sclerites of discosubmarginal cell of fore wing as in Figure 10F; fenestra of fore wing not very long and its anterodistal corner distinctly separated from proximal end of vein RS; proximal sclerite triangular, confluent with distal one, strongly pigmented; central sclerite moderately to strongly pigmented and sclerotised, linear and parallel to distal margin of fenestra, positioned in distal part of fenestra; distal sclerite present proximally and vestigial to absent distally; proximal corner of marginal cell of fore wing uniformly setose; vein 1cu-a subinterstitial to antefurcal to M&RS by less than 0.2× 1cu-a length.
Differential diagnosis. Enicospilus formosensis is a distinctive species and can easily be distinguished by many characters, such as the wide face (Fig. 10B), shape of the central sclerite (Fig. 10F), more or less subquadrate scutellum (Fig. 2E), as listed in the diagnosis. Distribution. Oriental region ). Newly recorded from Nepal.
Wings (Fig. 11F): fore wing with AI = 0.8-1.4, CI = 0.5-0.6, ICI = 0.4-0.5, SDI = 1.4-1.5; fore wing vein 1m-cu&M almost evenly curved, 2r&RS weakly bowed centrally; fenestra and sclerites of discosubmarginal cell of fore wing as in Figure 11F; fenestra of fore wing not very long and its anterodistal corner distinctly separated from proximal end of vein RS; proximal sclerite not triangular, confluent with distal one, weakly to strongly pigmented; central sclerite weakly to strongly pigmented and sclerotised, linear and parallel to vein 2r&RS, positioned in central part of fenestra; distal sclerite weak; proximal corner of marginal cell of fore wing uniformly setose; vein 1cu-a subinterstitial to antefurcal to M&RS by less than 0.1× 1cu-a length.
Differential diagnosis. Enicospilus grammospilus is a very distinctive species on account of its characteristic shape of fore wing vein 2r&RS and sclerites as in Figure 11F. No similar species are recognised and is very easily distinguished from all other Enicospilus species by the characters summarised in the above diagnosis, such as concentrically striate posterior area of propodeum and characteristic shape of sclerites of discosubmarginal cell of fore wing (cf. Fig. 11F).
Wings (Fig. 12F): fore wing with AI = 1.1-1.8, CI = 0.2-0.5, ICI = 0.4-0.6, SDI = 1.0-1.1; fore wing vein 1m-cu&M evenly curved, 2r&RS almost straight; fenestra and sclerites of discosubmarginal cell of fore wing as in Figure 12F; fenestra of fore wing not very long and its anterodistal corner distinctly separated from proximal end of vein RS; proximal sclerite not triangular, confluent with distal one, at least anteriorly very strongly pigmented; central sclerite absent; distal sclerite more or less strong and rather thick; proximal corner of marginal cell of fore wing more or less uniformly setose; vein 1cu-a subinterstitial to antefurcal to M&RS by less than 0.1× 1cu-a length.
Colour (Fig. 12): body including interocellar area entirely testaceous; wings hyaline to slightly infumate. Differential diagnosis. Enicospilus javanus is distinctive and one of the most easily distinguishable species on account of the proximally extended fore wing fenestra and the shape of the sclerites (cf. Fig. 12F).  (Figs 2F, 13).
Legs. Outer surface of fore tibia with scattered spines. Hind leg with coxa in profile 1.9× as long as deep; basitarsus 2.0× as long as second tarsomere; fourth tarsomere 0.6× as long as third tarsomere and 4.2× as long as wide; tarsal claw simply pectinate.
Variation. Unknown. Female. Unknown. Differential diagnosis. Enicospilus kakanicus sp. nov. is similar to and can be confused with E. longitarsis Tang, 1990, E. tangi sp. nov., and E. yonezawanus (Uchida, 1928). These species all belong to the E. ramidulus complex and share the following characters: outer surface of mandible with a diagonal setose deep groove between its dorsoproximal corner and base of mandibular apical teeth (e.g. Fig. 2B, D), fore wing fenestra without central sclerite (e.g. Figs 13F, 25F, 27F), and proximal sclerite triangular (e.g. Figs 13F, 25F, 27F). Enicospilus kakanicus sp. nov. is distinguished from the above species by the rather short lateral longitudinal carinae of the scutellum, i.e. reaching the anterior 0.6 of the scutellum in E. kakanicus sp. nov., as in Figure  2F, but almost always reaching the posterior end of the scutellum in E. longitarsis Tang, 1990, E. tangi sp. nov., and E. yonezawanus, as in, e.g., Figure 2H, and also by the characters used in the above key, such as width of lower face, mandibular shape and length, and surface sculptures of metapleuron. (Uchida, 1928) Enderlein, 1921; synonymised by Gauld and Mitchell (1981: 459).
Colour (Fig. 14): body including interocellar area entirely testaceous; wings hyaline. Differential diagnosis. Enicospilus kanshirensis is most similar to E. flavicaput but can be distinguished from it by the stouter central sclerite (Fig. 14F) (central sclerite slender in E. flavicaput as in Figure 8F), and smaller body size (i.e. fore wing length less than 15.0 mm in E. kanshirensis but more than 17.0 mm in E. flavicaput).
Colour (Fig. 16): body including interocellar area entirely testaceous; wings hyaline. Differential diagnosis. Some species of Oriental Enicospilus (e.g. E. fusiformis and E. unicolor) have a centrally broadened distal sclerite and lack proximal and central sclerites, as in Figure 16F. Among them, E. lineolatus is most similar to E. unicolor, but distinguished by the narrower distal sclerite than that of E. unicolor and testaceous fore wing pterostigma and sclerite (brown in E. unicolor).
Wings (Fig. 17F): fore wing with AI = 0.4-1.1, CI = 0.3-0.5, ICI = 0.4-0.5, SDI = 1.1-1.4; fore wing vein 1m-cu&M more or less evenly curved, 2r&RS almost straight; fenestra and sclerites of discosubmarginal cell of fore wing as in Figure 17F; fenestra of fore wing not very long and its anterodistal corner distinctly separated from proximal end of vein RS; proximal sclerite triangular, strongly confluent with distal one, strongly pigmented; central sclerite moderately to strongly pigmented and sclerotised, usually well-delineated oval, positioned in antero-to medio-distal part of fenestra; distal sclerite more or less evenly strong from proximal to distal; proximal corner of marginal cell of fore wing uniformly setose; vein 1cu-a subinterstitial to antefurcal to M&RS by less than 0.3× 1cu-a length.
Colour (Fig. 17): body including interocellar area entirely testaceous with black posterior segments of metasoma; wings hyaline. Differential diagnosis. Enicospilus melanocarpus is very similar to E. sauteri, but distinguished by the uniformly setose marginal cell of the fore wing (Fig. 17F) (marginal cell of fore wing proximally glabrous in E. sauteri) and the oval central sclerite (Fig. 17F) (central sclerite linear in E. sauteri). Many species were synony-mised with E. melanocarpus under Gauld's conservative species criteria, but their wide distribution and considerable range of morphological variation indicate this name includes many species. Therefore, further researches are needed to reveal the true species diversity under the name 'melanocarpus'. Etymology. The specific name is derived from the type locality.
Legs. Outer surface of fore tibia without dense and long spines. Hind leg with coxa in profile 1.7× as long as deep; basitarsus 2.0× as long as second tarsomere; fourth tarsomere 0.6× as long as third tarsomere and 3.5× as long as wide; tarsal claw simply pectinate.
Male. Unknown. Differential diagnosis. Enicospilus nepalensis sp. nov. is probably closely related to or belongs to the E. ramidulus complex. Among the complex, E. nepalensis sp. nov. is most closely related to E. tricorniatus Rao & Nikam, 1970 based on the rather small ocelli relative to other Enicospilus (posterior ocellus separated from eye by more than 0.3× its own maximum diameter) (e.g. Fig. 18B-D), highly shiny body (e.g. Fig. 18A-E), shape of body (e.g. Fig. 18A), shape and position of the fore wing veins and sclerites (e.g. Fig. 18F), distribution, etc. However, E. nepalensis sp. nov. is readily distinguishable from E. tricorniatus by the following characters: lower face more or less elongate and 0.8× as wide as high (Fig. 18B) (lower face subquadrate to transverse and 1.0-1.1× as wide as high in E. tricorniatus), the central sclerite weakly sclerotised and pigmented (Fig. 18F) (moderately to strongly sclerotised and pigmented in E. tricorniatus), moderate-sized, fore wing length 10.0-11.0 mm (small, fore wing length less than 8.5 mm in E. tricorniatus), posterior ocellus separated from eye by 0.3× its own maximum diameter (Fig. 18B-D) (posterior ocellus separated from eye by almost its own maximum diameter in E. tricorniatus).

Figs 2J, 19
Etymology. The specific name is dedicated to Dr P.K. Nikam who studied Ophioninae as well as other groups of Hymenoptera mainly of India.
Legs. Outer surface of fore tibia with very few spines. Hind leg with coxa in profile 1.7× as long as deep; basitarsus 2.2× as long as second tarsomere; fourth tarsomere 0.6× as long as third tarsomere and 2.6× as long as wide; tarsal claw simply pectinate except lacking pecten proximally.
Variation. Unknown Male. Unknown Differential diagnosis. Enicospilus nikami sp. nov. is similar to E. biharensis, E. maruyamanus, E. pudibundae, and E. transversus and these species are rather difficult to separate from each other. However, E. nikami sp. nov. can be distinguished from E. biharensis, E. maruyamanus and E. transversus by the proximally incomplete pectinae of hind tarsal claw (Fig. 2J) (hind tarsal claw completely pectinate from its base to apex in E. biharensis, E. maruyamanus and E. transversus, as in e.g. Figure 2I), from E. biharensis and E. pudibundae by the sinuous fore wing vein 1m-cu&M (Fig. 19F) (1m-cu&M evenly curved in E. biharensis and E. pudibundae as in Figures 6F, 23F), from E. maruyamanus by the entirely moderately punctate mesopleuron (Fig. 19E) (mesopleuron entirely longitudinally punctostriate in E. maruyamanus) and the angle of posterodistal corner of second discal cell (i.e. ca 90° in E. nikami sp. nov. as in Figure 19F, but ca 115° in E. maruyamanus), and from E. transversus by the entirely moderately punctate mesopleuron (Fig. 19E)  Etymology. The specific name is derived from the type locality.
Legs. Outer surface of fore tibia with sparse spines. Hind leg with coxa in profile 2.0× as long as deep; basitarsus 2.0× as long as second tarsomere; fourth tarsomere 0.6× as long as third tarsomere and 4.6× as long as wide; tarsal claw simply pectinate.
Mesosoma (Fig. 21E): mesopleuron entirely punctate; scutellum with lateral longitudinal carinae reaching posterior end and convergent posteriorly; metapleuron punctate; propodeum evenly weakly rounded, its posterior area moderately reticulate, outer margin of propodeal spiracle not joining pleural carina by a ridge.  Figure 21F; fenestra of fore wing not very long and its anterodistal corner distinctly separated from proximal end of vein RS; proximal sclerite triangular, separated from distal one, strongly pigmented; central sclerite partially strongly pigmented and sclerotised, ill-delineated oval, positioned in almost medio-distal part of fenestra; distal sclerite strong distally; proximal corner of marginal cell of fore wing uniformly setose; vein 1cu-a subinterstitial to antefurcal to M&RS by less than 0.2 1cu-a length.
Colour (Fig. 21): body including interocellar area entirely red-brown; wings hyaline. Differential diagnosis. As mentioned under E. laqueatus, four Oriental species of Enicospilus (E. laqueatus, E. pseudantennatus, E. vestigator, and E. tripartitus) have similar fenestra, sclerites, and fore wing veins (e.g. Figs 15F, 21F, 26F). Among them, E. pseudantennatus is distinguished from E. laqueatus by the flat outer surface of the mandible (outer surface of mandible with a diagonal deep setose groove between dorsoproximal corner and base of mandibular apical teeth in E. laqueatus), from E. tripartitus by the not densely setose and proximally more or less flat outer mandibular surface (outer surface of mandible with very dense setae and sharp and rather deep proximal concavity in E. tripartitus, as in Figure 2C), and from E. vestigator by the weakly twisted mandible (10-20°) (mandible strongly twisted by 60-80° in E. vestigator).
Colour (Fig. 22): body including interocellar area entirely testaceous; wings hyaline. Differential diagnosis. Enicospilus pseudoconspersae is one of the most distinctive and easily distinguishable species among the Oriental species of Enicospilus on account of the characteristic isolated and weakly pigmented semicircular proximal sclerite (Fig. 22F). There are no known morphologically similar species.
Diagnosis. Head (Fig. 23B-D): GOI = 2.6-2.8; lower face 0.7× as wide as high; clypeus almost flat in profile, its lower margin acute to subacute; mandible weakly twisted by 10-20°, moderately long, evenly tapered, its outer surface without a diagonal structure; upper mandibular tooth 1.2-1.5× as long as lower one; posterior ocellus (almost) touching eye; antenna with Wings (Fig. 23F): fore wing with AI = 0.5-1.0, CI = 0.5-0.7, ICI = 0.5-0.7, SDI = 1.4-1.5; fore wing vein 1m-cu&M evenly curved, 2r&RS almost straight; fenestra and sclerites of discosubmarginal cell of fore wing as in Figure 23F; fenestra of fore wing not very long and its anterodistal corner distinctly separated from proximal end of vein RS; proximal sclerite more or less linear, very weakly confluent with distal one or not, very weakly to strongly pigmented; central sclerite absent; distal sclerite more or less weak to absent; proximal corner of marginal cell of fore wing sparsely to uniformly setose; vein 1cu-a subinterstitial to antefurcal to M&RS by less than 0.2× 1cu-a length.
Differential diagnosis. Enicospilus pudibundae resembles E. biharensis, E. maruyamanus, E. nikami sp. nov., and E. transversus, but can be distinguished from E. biharensis, E. maruyamanus, and E. transversus by the proximally incomplete pectination of the hind tarsal claw (pectination of hind tarsal claw complete from base to apex of the claw in E. biharensis, E. maruyamanus, and E. transversus, as in e.g. Figure 2I) and also from E. maruyamanus, E. nikami sp. nov., and E. transversus by the evenly curved fore wing vein 1m-cu&M ( Fig. 23F) (1m-cu&M more or less sinuous in E. maruyamanus, E. nikami sp. nov. and E. transversus, as in e.g. Figure 19F). The Nepalese and some other Oriental specimens exhibit a rather wider proximal sclerite and sparser setosity in the proximal corner of the fore wing fenestra than the holotype and Eastern Palaearctic specimens, suggesting that the Oriental specimens are potentially cryptic species. However, at present, I have not enough evidence to describe them as a new species and tentatively follow Gauld and Mitchell's (1981) species criteria. Distribution. Australasian, Eastern Palaearctic, and Oriental regions . Newly recorded from Nepal and Brunei.
Colour (Fig. 24F): body including interocellar area entirely testaceous; wings hyaline. Differential diagnosis. Enicospilus purifenestratus is very similar to E. urocerus Gauld & Mitchell, 1981, but distinguished from it by the unswollen segments 3 and 4 of the maxillary palp (segments 3 and 4 of the maxillary palp swollen in E. urocerus) and thinner distal sclerite (Fig. 24F)  Etymology. The specific name is dedicated to Dr Yuqing Tang who described E. longitarsis, which is morphologically the most similar species to the one that is hereby described, and has contributed to the taxonomy of Ophioninae in Asia, represented by the monograph of Chinese Enicospilus (Tang 1990 Head with GOI = 2.5 (Fig. 25C). Lower face 0.9× as wide as high, moderately punctate with setae and shiny (Fig. 25B). Clypeus 1.7× as wide as high, moderately punctate with setae, moderately convex in profile, lower margin impressed (Fig. 25B, C). Malar space 0.4× as long as basal mandibular width (Fig. 25B, C). Mandible weakly twisted by ca 25°, very long, proximally strongly narrowed, centrally to apically subparallel sided, its outer surface with a diagonal setose deep groove between dorsoproximal corner to base of mandibular apical teeth (Figs 2B, 25B, C). Upper mandibular tooth 2.1× as long as lower one, stouter than lower one (Figs 2B, 25B). Frons, vertex and gena moderately shiny with fine setae ( Fig. 25B-D). Posterior ocellus close to eye, separated from eye by less than 0.1× its own maximum diameter (Fig. 25B-D). Ventral end of occipital carina joining oral carina. Antenna incomplete apically, right antenna with 64 flagellomeres and left antenna with 65 flagellomeres; first flagellomere 1.9× as long as second; 20 th flagellomere 2.2× as long as wide.
Legs. Ventral 0.7 of outer surface of fore tibia with rather dense spines. Hind leg with coxa in profile 1.8× as long as deep; basitarsus 2.0× as long as second tarsomere; fourth tarsomere 0.7× as long as third tarsomere and 5.0× as long as wide; tarsal claw simply pectinate.
Diagnosis. Head (Figs 2C, 26B-D): GOI = 2.2-2.9; lower face 0.7-0.8× as wide as high; clypeus moderately to strongly convex in profile, its lower margin more or less blunt; mandible rather weakly twisted by 10-20°, moderately long, proximally tapered and distally parallel sided, its outer surface flat but with conspicuous dense setae and a proximal deep concavity; upper mandibular gent posteriorly; metapleuron densely punctate as mesopleuron; propodeum weakly declivous, its posterior area moderately reticulate, outer margin of propodeal spiracle not joining pleural carina by a ridge.
Wings (Fig. 26F): fore wing with AI = 0.3-0.6, CI = 0.3-0.4, ICI = 0.5-0.7, SDI = 1.2-1.6; fore wing vein 1m-cu&M almost evenly curved to slightly sinuous, 2r&RS almost straight; fenestra and sclerites of discosubmarginal cell of fore wing as in Figure 26F; fenestra of fore wing not very long and its anterodistal corner distinctly separated from proximal end of vein RS; proximal sclerite triangular, separated from distal one, strongly pigmented; central sclerite strongly pigmented and sclerotised, well-delineated oval and its major axis parallel to distal margin of fenestra, positioned in mediodistal part of fenestra; distal sclerite absent to weak; proximal corner of marginal cell of fore wing uniformly setose; vein 1cu-a subinterstitial to antefurcal to M&RS by less than 0.2× 1cu-a length.
Colour (Fig. 26): body including interocellar area entirely reddish brown; wings hyaline. Differential diagnosis. Four Oriental Enicospilus species, E. laqueatus, E. pseudantennatus, E. vestigator, and E. tripartitus, have similar fenestra, sclerites, and fore wing veins (e.g. Figs 15F, 21F, 26F), as mentioned under E. laqueatus and E. pseudantennatus. Among them, E. tripartitus is readily distinguishable from other species by the outer mandibular surface: outer surface of mandible with very dense setae and sharp and rather deep proximal concavity in E. tripartitus (Figs 2C, 26B, C), but more or less flat proximally with scattered setae in E. laqueatus (Fig. 15B, C) E. pseudantennatus (Fig. 21B, C) and E. vestigator. (Uchida, 1928 clypeal structure, shape of fore wing sclerites, and surface sculpture of mesopleuron, but can be confused with E. kakanicus sp. nov., E. longitarsis, and E. tangi sp. nov. However, E. yonezawanus is distinguishable from E. kakanicus sp. nov. by the complete lateral longitudinal carinae of the scutellum (lateral longitudinal carinae of the scutellum posteriorly absent in E. kakanicus sp. nov., as in Figure 2F), from E. longitarsis and E. tangi sp. nov. by the scattered spines on the outer fore tibial surface (spines rather dense in E. longitarsis and E. tangi sp. nov.) and the separated proximal and distal sclerites (Fig. 27F) (proximal and distal sclerites confluent in E. longitarsis and E. tangi sp. nov. as in e.g. Figure 25F). Distribution. Oriental region ). Newly recorded from Nepal.
Mesosoma (Fig. 28E): mesopleuron punctate to rather coarsely longitudinally striate; scutellum with lateral longitudinal carinae reaching posterior end and convergent posteriorly; metapleuron rather coarsely striate; propodeum evenly weakly rounded, its posterior area more or less coarsely irregularly wrinkled with strong posterior transverse carina laterally, outer margin of propodeal spiracle joining pleural carina by a ridge.
Wings (Fig. 28F): fore wing with AI = 0.5, CI = 0.3-0.4, ICI = 0.4-0.5, SDI = 1.4-1.5; fore wing vein 1m-cu&M very slightly sinuous, 2r&RS almost straight; fenestra and sclerites of discosubmarginal cell of fore wing as in Figure 28F; fenestra of fore wing very long and its anterodistal corner very close to proximal end of vein RS; proximal sclerite triangular, confluent with distal one, strongly pigmented; central sclerite moderately pigmented and sclerotised, ill-delineated semicircular to oval, its major axis parallel to distal margin of fenestra, positioned in very distal and slightly anterior part of fenestra; distal sclerite entirely moderately pigmented; proximal corner of marginal cell of fore wing uniformly setose; vein 1cu-a antefurcal to M&RS by 0.1× 1cu-a length.
Colour (Fig. 28): body entirely black with pale yellow patterns, interocellar area not infuscate; wings hyaline but fore wing with three strongly infumate areas around anterocentral part of discosubmarginal cell, proximal part of second discal cell, and central part of marginal cell. Differential diagnosis. Gauld and Mitchell (1981) suggested that E. zebrus is related to the E. signativentris species-group and very close to E. biumbratus (Morley, 1912) on body and wing colour pattern as well as other characters, but E. zebrus is distinguished from E. biumbratus by many characters, such as the longer fore wing fenestra (Fig. 28F), smaller and semicircular to oval central sclerite (Fig. 28F), etc.

Species inquirendae and pending taxonomic acts
Some morphospecies and species-groups listed below are tentatively treated as species inquirendae pending taxonomic acts. Two morphospecies (Enicospilus sp. 1 (Fig. 29) and Enicospilus sp. 2 (Fig. 30)) are likely to be undescribed species, but the only available specimens are in poor condition, so they are not be described here. Also, the E. erythrocerus species-group is currently taxonomically challenging. Type specimens must be re-examined and integrative taxonomic methods should be included to delimit and redefine species. Three morphospecies are included in Nepalese specimens of the E. erythrocerus group (Fig. 31), and at least one of these is potentially an undescribed species.
Comments. The mandibular structure of this species indicates it is associated with the E. ramidulus complex. Enicospilus sp. 1 does not key out to any species in Gauld and Mitchell's (1981) and Tang's (1990) keys and is possibly an undescribed species. It may potentially be found to be closely related to E. choui Tang, 1990 or E. sinicus Tang, 1990. However, only one broken specimen is known, so I tentatively treat this species as a species inquirenda. Material examined. 1♂: Nepal. 1♂,Terai (200 m), Chitwan, Nepal, 12-13.III.1983, M.G. Allen leg. (NHMUK) (Fig. 30).
Comments. The material examined is not in bad condition except for incomplete antennae. However, antennal characters are often useful and important for distinguishing Enicospilus species, as previous studies suggested (e.g. Broad and Shaw 2016). Therefore, antennae should be complete to describe a new species.
The affinities of this species are not clear, but, as with Enicospilus sp. 1, it also does not key out to any species in Gauld and Mitchell's (1981) or Tang's (1990) keys, indicating that it is potentially an undescribed species.   Comments. The E. erythrocerus species-group is moderately large and one of the most taxonomically confusing groups within Enicospilus. It consists of rather large wasps with the fore wing fenestra lacking any trace of sclerites, SDI more than 1.2, lateral longitudinal carinae of the scutellum almost always reaching the posterior end, moderately sized fore wing fenestra, etc. In this study, I examined 27 Nepalese specimens of this species-group and recognised at least three morphospecies (Fig. 31). However, further research is needed to identify or describe them. Therefore, I tentatively treat all specimens of the E. erythrocerus species-group as species inquirenda.

Discussion
Many species of Nepalese Enicospilus were recognised from middle elevations, and the median value of elevation for 83% of Nepalese Enicospilus fauna is between 950-2,070 m (Fig. 32). These species are generally widely distributed in the mountainous areas of the (sub)tropical Oriental region and, in some species, such as E. lineolatus and E. yonezawanus, also in the Eastern Palaearctic region. On the other hand, three species (i.e. E. capensis, E. kanshirensis, and E. pudibundae) have been collected only at lower elevations, from 200-330 m (Fig. 32). These species are also widely distributed in the Oriental region, as with the middle-elevation species, and in particular E. capensis is a very widespread Old World species known from the Afrotropical, Australasian, Oceanic, and Oriental regions (Gauld and Mitchell 1981). However, E. zebrus has been collected only at higher elevations (Fig. 32), above 2,500 m, suggesting that this species is restricted to the northern high-elevational margin of the continental Oriental region and endemic to the southern slope and eastern highlands of the Himalayas. This is a preliminary study of the Nepalese fauna of Enicospilus, as well as of Ophioninae; the sample size is small and the sampling bias of the materials used in the present study is not known, but trends of elevational distribution patterns are indicated. These elevational and distribution patterns of Nepalese Enicospilus species are fairly consistent with those proposed by Gauld and Mitchell (1981) and Gauld (1985a).
The Nepalese fauna of Enicospilus has trebled through this study, even though it is a preliminary work. Based on species represented by more than two specimens, no endemicity of the Nepalese fauna is recognised, with most species common to other Oriental countries. Moreover, several common Oriental Enicospilus species, such as E. abdominalis (Szépligeti, 1906), E. aciculatus (Taschenberg, 1875), E. concentralis Cushman, 1937, E. dasychirae Cameron, 1905, E. dolosus (Tosquinet, 1896, E. exaggeratus Chiu, 1954, E. nigropectus Cameron, 1905 riukiuensis (Matsumura & Uchida, 1926), E. shinkanus (Uchida, 1928) and E. signativentris (Tosquinet, 1903), have not been found in Nepal yet, but they may be present in the country. Considering the Enicospilus fauna of adjacent areas of Nepal and that of the Old World, at least 60 species are potentially found in Nepal. Therefore, additional studies and greater sampling efforts are needed to reveal the true Enicospilus diversity in Nepal.