Enicospilus Stephens 1835
Creators: name: Pham, Nhi Thi
name: Pham, Phu Van
name: Matsumoto, Rikio
name: Shimizu, So
name: Broad, Gavin R.
Publisher: name: Zenodo
Year: 2023
Subjects: Biodiversity
Taxonomy
Animalia
Arthropoda
Insecta
Hymenoptera
Ichneumonidae
Enicospilus

description: Genus Enicospilus Stephens, 1835
Enicospilus Stephens, 1835: 126; type species: Ophion combustus Gravenhorst, 1829, by subsequent designation (Horstmann 2005: 1264).
Henicospilus Agassiz, 1846: 138; unjustified emendation.
Allocamptus Förster, 1869: 150; type species: Ophion undulatus Gravenhorst, 1829, by subsequent designation (Thomson 1888: 1189).
Dispilus Kriechbaumer, 1894: 309; type species: Ophion (Dispilus) natalensis Kriechbaumer, 1894, by monotypy.
Pleuroneurophion Ashmead, 1900: 86; type species: Pleuroneurophion hawaiiensis Ashmead, 1900, by original designation.
Banchogastra Ashmead, 1900: 87; type species: Banchogastra niger Ashmead, 1900, by original designation.
Pycnophion Ashmead, 1900: 87; type species: Pycnophion molokaiensis Ashmead, 1900, by original designation.
Cymatoneura Kriechbaumer, 1901a: 22; type species: Ophion undulatus Gravenhorst, 1829, by subsequent designation (Viereck 1914: 8).
Pterospilus Kriechbaumer, 1901b: 156; type species: Ophion (Enicospilus) dubius Tosquinet, 1896, by subsequent designation (Viereck 1914: 126); junior homonym of Pterospilus Rondani, 1856.
Trispilus Kriechbaumer, 1901b: 156; type species: Ophion (Enicospilus) trimaculatus Tosquinet, 1896, by monotypy.
Abanchogastra Perkins, 1902: 141; type species: Abanchogastra debilis Perkins, 1902, by monotypy.
Metophion Szépligeti, 1905: 28; type species: Metophion bicolor Szépligeti, 1905, by subsequent designation (Viereck 1914: 94).
Ceratospilus Szépligeti, 1905: 28; type species: Ceratospilus biroi Szépligeti, 1905, by monotypy.
Atoponeura Szépligeti, 1905: 34; type species: Atoponeura concolor Szépligeti, 1905 (= Enicospilus atoponeurus Cushman, 1947), by monotypy.
Ophiomorpha Szépligeti, 1905: 34; type species: Ophion curvinervis Cameron, 1886 (= Enicospilus cameronii Dalla Torre, 1901), by subsequent designation (Hooker 1912): 134; junior homonym of Ophiomorpha Nilsson, 1836.
Cryptocamptus Brèthes, 1909: 230; unnecessary replacement name for Allocamptus Förster, 1869.
Amesospilus Enderlein, 1914: 222; type species: Ophion unicallosus Vollenhoven, 1878, by original designation.
Eremotyloides Perkins, 1915: 530; type species, Eremotylus orbitalis Ashmead, 1901, by monotypy.
Schizospilus Seyrig, 1935: 79; type species: Schizospilus divisus Seyrig, 1935, by original designation.
Diagnosis
Species of Enicospilus can be distinguished from other Ophioninae by the following characters: fore wing discosubmarginal cell usually with extensive glabrous area (referred to as a ʻfenestraʼ), often with one or more sclerites, and sometimes with a ʻquadraʼ in the middle of the fenestra (Fig. 1), of more or less different texture from the outer margins; mandibles apically more or less tapered and slightly to strongly twisted; inner surface of fore tibial spur without membranous flange. Species of Enicospilu s can be confused with the genus Dicamptus Szépligeti, 1905 and some species of Leptophion Cameron, 1901 because they share the characteristic fore wing fenestra and sclerites, but they can easily be distinguished by the weakly to strongly tapered and twisted mandible (mandible very weakly tapered and never strongly twisted in Dicamptus and Leptophion).
Key to the Vietnamese species of Enicospilus
This key is modified after Gauld & Mitchell (1981) and Shimizu et al. (2020). Supporting characters are given in square brackets.
1. Fore wing fenestra without sclerites, at very most with an indistinct yellowish thickening or quadra in membrane ..................................................................................................................................... 2
– Fore wing fenestra with more or less distinct sclerites, sometimes also with quadra ...................... 6
2. Fore wing fenestra virtually occluded by large and elongate quadra (Fig. 26f); clypeus in profile strongly convex, subnasute (Fig. 26b, d) ................................................. E. exaggeratus Chiu, 1954
– Fore wing fenestra without quadra; clypeus weakly to moderately convex ..................................... 3
3. Fore wing 1m-cu&M centrally widened; 2r&RS rather stout, distally fairly abruptly tapered (Fig. 9f) ................................................................................................ E. atoponeus Cushman, 1947
– Fore wing 1m-cu&M not at all widened; 2r&RS less stout, more evenly tapered distally .............. 4
4. Fore wing 1m-cu&M angled medially, with short ramellus ........... E. plicatus Brullé, 1846 (in part)
– Fore wing 1m-cu&M evenly curved, without short ramellus ........................................................... 5
5. Mandible strongly twisted by ca 80° (Fig. 24b); fore wing 1 st subdiscal cell with sparse setae (Fig. 24f); fore wing length 12.5–15.0 mm ..................................................... E. erythrocerus (Cameron, 1905)
– Mandible twisted by 20°–30° (Fig. 35b); fore wing 1 st subdiscal cell generally with denser setae at least on anterior 0.4 (Fig. 35f); fore wing length 19.0–23.0 mm .......... E. grandis (Cameron, 1905)
6. Fore wing with SDI <0.94 ................................................................................................................ 7
– Fore wing with SDI ≥ 0.95 .............................................................................................................. 10
7. Hind tarsal claws simple, distal pecten not projecting; fore wing without any darkened patches, marginal cell sparsely setose proximally, proximal and distal sclerites confluent (Fig. 22f) ............. ...................................................................................................... E. eastopi Gauld & Mitchell, 1981
– Hind tarsal claws with distal pecten projecting beyond apical tooth; fore wing marginal cell proximally darkened, evenly setose, proximal and distal sclerites separated ..................................................... 8
8. Interocellar area yellow to weakly darkened (Fig. 11c); mandible outer surface with a weak diagonal setose groove or with a tuft of long setae; fore wing fenestra darkened between proximal and distal sclerites (Fig. 11f) .................................................................... E. bakerielli Gauld & Mitchell, 1981
– Interocellar area black (Figs 17c, 78c); mandible outer surface without a diagonal setose groove or tuft of long setae; fore wing fenestra without any darkened patches between proximal and distal sclerites (Figs 17f, 78f) ..................................................................................................................... 9
9. Fore wing proximal sclerite larger, distance from proximal sclerite to vein 2r&RS less than its maximum diameter (Fig. 17f); DMI = 1.2 .............................................. E. circuliscleritalis sp. nov.
– Fore wing proximal sclerite smaller, distance from proximal sclerite to vein 2r&RS about 1.9 × its maximum diameter (Fig. 78f); DMI = 1.3–1.4 ........................................................... E. trui sp. nov.
10. Fore wing fenestra with central sclerite entirely absent, at most with a quadra discernible ...........11
– Fore wing fenestra with central sclerite present, usually strongly pigmented ................................ 38
11. Proximal sclerite narrow, linear, weakly pigmented (Figs 14f, 19f, 37f, 39f, 47f, 61f, 76f) or entirely absent (Figs 31f, 64f) ...................................................................................................................... 12
– Proximal sclerite quite widened, usually triangular (other shapes in E. dasychirae, E. nigronotatus and E. pseudoconspersae) and usually strongly pigmented ........................................................... 21
12. Fore wing with CI less than 0.5 ...................................................................................................... 13
– Fore wing with CI at least 0.5 ......................................................................................................... 14
13. Fore vein 2r&RS virtually straight, vein 1m-cu&M evenly arcuate, AI less than 0.9; fore wing length 10–16 mm (Fig. 14f) ................................................. E. biharensis Townes, Townes & Gupta, 1961
– Fore wing vein 2r&RS moderately sinuous, vein 1m-cu&M strongly sinuous, AI more than 0.9; fore wing length 18–20 mm (Fig. 19f) ....................................................... E. corculus (Tosquinet, 1903)
14. Proximal alar sclerite entirely absent .............................................................................................. 15
– Proximal alar sclerite discernible, weakly to moderately pigmented ............................................. 16
15. Fore wing vein 1m-cu&M centrally angulate and broadened, ICI = 0.62–0.74 (Fig. 31f); metasoma from T5 onwards blackish .......................................................................... E. fusiformis Chiu, 1954
– Fore wing vein 1m-cu&M sinuous and more or less uniformly thickened, ICI = 0.75–1.09 (Fig. 64f); metasomal tergites reddish brown, sometimes from T3 onwards with dorsal line and ventral parts blackish ...................................................................................................... E. pungens (Smith, 1874)
16. Hind tarsal claws lacking pecten proximally .................................................................................. 17
– Hind tarsal claws with pecten extending proximally ...................................................................... 18
17. Marginal cell of fore wing proximally with a glabrous area, quadra moderately large, ICI = 0.71–0.83 (Fig. 39f) ...................................................................................... E. iapetus Gauld & Mitchell, 1981
– Marginal cell of fore wing uniformly setose, without quadra, ICI = 0.65 or less .............................. ............................................................................................................ E. pudibundae (Uchida, 1928)
18. Marginal cell of fore wing proximally with a glabrous area (Figs 37f, 47f) .................................. 19
– Marginal cell of fore wing proximally at most sparsely but uniformly setose, without a glabrous area (Fig. 61f) ......................................................................................................................................... 20
19. Fore wing vein 2r&RS relatively straight, relatively uniformly thickened, AI = 0.9 (Fig. 37f); mesopleuron puncto-striate ventrally (Fig. 37e); fore wing length 18.7 mm ..................................... ...................................................................................................... E. hedilis Gauld & Mitchell, 1981
– Fore wing vein 2r&RS sinuous, abruptly narrowed distally, AI = 1.36 (Fig. 47f); mesopleuron striate ventrally (Fig. 47e); fore wing length 12.8 mm ........................................... E. maritus Roman, 1913
20. Fore wing vein 1m-cu&M and 2r&RS strongly sinuous (Fig. 61f) ................................................... ...................................................................................................... E. plicatus (Brullé, 1846) (in part)
– Fore wing vein 1m-cu&M evenly arcuate, 2r&RS weakly sinuous (Fig. 76f) ................................... ................................................................................................................... E. transversus Chiu, 1954
21. Proximal margin of proximal sclerite of fore wing fenestra distinctly separated from proximal margin of fenestra by more than width of proximal sclerite (except separated 0.75–1.0 × in E. dasychirae) (Figs 42f, 56f, 63f) .......................................................................................................................... 22
– Proximal margin of proximal sclerite of fore wing fenestra joining or close to proximal margin of fenestra, if separated then by less than half width of proximal sclerite .......................................... 26
22. Fore wing with both ICI and CI greater than 0.65; very large insects, fore wing length 20 mm or more ................................................................................................................................................ 23
– Fore wing with either or both ICI and CI less than 0.65; variously sized insects .......................... 24
23. Distal sclerite present, not confluent with proximal sclerite, SRI = 0.2 (Fig. 56f); body dark reddish brown with black markings (Fig. 56a) ............................................. E. nigronotatus Cameron, 1903
– Distal sclerite absent, SRI = 0.3; body reddish brown without black markings ................................. ..................................................................................... E. pseudoconspersae (Sonan, 1927) (in part)
24. Proximal and distal sclerites of fore wing fenestra strongly confluent and distal sclerite strongly sclerotised; confluent proximal and distal sclerites of fore wing fenestra usually shaped like a letter ‘Pʼ, as in Fig. 42f; SDI = 0.90–1.05 ..................................................... E. javanus (Szépligeti, 1910)
– Proximal sclerite of fore wing fenestra isolated and distal sclerite absent or vestigial; proximal sclerite of fore wing fenestra half-moon or drop-shaped, as in Figs 20f, 63f; SDI at least 1.30 ................ 25
25. Proximal sclerite of fore wing fenestra usually entirely weakly pigmented and half-moon-shaped (Fig. 63f) ..................................................................... E. pseudoconspersae (Sonan, 1927) (in part) – Proximal sclerite of fore wing fenestra partly to entirely strongly pigmented and drop-shaped (Fig. 20f) ............................................................................................. E. dasychirae Cameron, 1905
26. Hind tarsal claws with distal pecten projecting beyond apical tooth .................................................. .................................................................................................... E. mythrus Gauld & Mitchell, 1981
– Hind tarsal claws simple, distal pecten not projecting ................................................................... 27
27. Interocellar area uniformly black (Figs 21c, 57c) .......................................................................... 28
– Interocellar area yellowish-brown to orange (Figs 27c, 45c, 83c) ................................................. 29
28. Fore wing with SDI = 1.07 or less, ICI = 0.35–0.36, vein 2r&RS centrally incrassate (Fig. 57f); metapleuron reticulate, at least on posterior 0.5 (Fig. 57e); mesoscutum and mesopleuron with black markings (Fig. 57e) ............................................................................ E. nigropectus Cameron, 1905
– Fore wing with SDI = 1.28–1.37, ICI = 0.53–0.59, vein 2r&RS more or less evenly thickened (Fig. 21f); metapleuron striate (Fig. 21e); mesoscutum and mesopleuron without black markings .. ............................................................................................................... E. dolosus (Tosquinet, 1896)
29. Outer mandibular surface with a diagonal groove extending from dorsal proximal corner to between base of teeth (Figs 27b, 45b) ........................................................................................................... 30
– Outer mandibular surface without a diagonal groove (Figs 40b, 75b) ........................................... 32
30. Fore wing length 17.5 mm, proximal sclerite evenly tapered to join distal sclerite (Fig. 45f); antenna with 73 flagellomeres; metapleuron finely punctate (Fig. 45e) .................. E. longitarsis Tang, 1990
– Fore wing length 10.2–14.5 mm, proximal sclerite not confluent with distal sclerite or with point of confluence narrow and indistinct (Figs 27f, 83f); antenna with 54–61 flagellomeres; metapleuron puncto-striate (Fig. 27e) .................................................................................................................. 31
31. Fore wing vein M&RS interstitial to 1cu-a, fenestra about 2.0 × area of proximal sclerite, distal sclerite stout, strong (Fig. 27f) .................................................................................... E. fittoni Nikam, 1980
– Fore wing vein M&RS postfurcal to 1cu-a, fenestra about 3.0× area of proximal sclerite, distal sclerite vestigial or weak and narrow (Fig. 83f) ........................................... E. yonezawanus (Uchida, 1928)
32. Margin of propodeal spiracle joined to lateral carina by a raised flange ........................................ 33
– Margin of propodeal spiracle not joined to lateral carina by a raised flange .................................. 36
33. Large body size, fore wing length 21.2–22.3 mm with fenestra short, about 3.0× proximal sclerite (Fig. 75f); mesoscutum with three black stripes ............................ E. teleus Gauld & Mitchell, 1981
– Smaller body size, fore wing length not exceed 21 mm with fenestra much longer, at least 4.5 × proximal sclerite (Figs 40f, 46f, 68f); mesoscutum with posterior 0.5 black or with a median black stripe ............................................................................................................................................... 34
34. Mesoscutum with a median black stripe; tegula reddish brown; metapleuron from punctate to puncto-striate (Fig. 40e) ..................................................................................... E. insinuator (Smith, 1860)
– Mesoscutum with posterior 0.5 black; tegula with posterior half black; metapleuron coarsely strigose (Figs 46e, 68e) ................................................................................................................................ 35
35. Mandible twisted by about 80° (Fig. 68b); metasomal tergites exceptionally long, T 4 in profile 1.4× as long as high; metasoma from T3 onwards entirely black (Fig. 68a) .............................................. ........................................................................................................ E. rogus Gauld & Mitchell, 1981
– Mandible twisted by about 20° (Fig. 46b); metasomal tergites not exceptionally long, T 4 in profile less than 1.2× as long as high; T4 with pale yellow patch (Fig. 46a) ................. E. maai Chiu, 1954
36. Metapleuron puncto-striate (Fig. 71e); fore wing length 10.5–12.5 mm, ICI = 0.41–0.62 ................ .............................................................................................................. E. shinkanus (Uchida, 1928)
– Metapleuron striate to strigose (Fig. 23e); fore wing length 13.8–18.6 mm, ICI = 0.68–0.85 ...... 37
37. Lower face 0.71–0.74 × as wide as long; fore wing with distal sclerite strong, joining proximal sclerite, AI = 0.41–0.58 (Fig. 65f) ......................................................... E. purifenestratus (Enderlein, 1921)
– Lower face 0.75–0.82 × as wide as long; fore wing with distal sclerite indistinct, AI = 0.59–0.76 (Fig. 23f) ................................................................................................ E. enicospilus Nikam, 1972
38. Interocellar area black or dark brown, contrasting with vertex (Figs 32c, 67c) ............................. 39
– Interocellar area yellowish-brown to orange .................................................................................. 45
39. Clypeus nasute, strongly convex (Fig. 67b, d) ............ E. riukiuensis (Matsumura & Uchida, 1926)
– Clypeus not nasute, flat to moderately convex ............................................................................... 40
40. Lower face 0.8 × as wide as long (Fig. 32b); clypeus strongly convex; mandible wide and long, medially 0.6× as broad as base, two teeth equal in length (Fig. 32b); fore wing with SDI = 1.0, CI = 0.72; metasomal tergites long and narrow, T5 1.5–1.6 × as long as high (Fig. 32a) .......................... ................................................................................................................... E. gasteralis Nikam, 1980
– Lower face narrower, about 0.6–0.7 × as wide as long; clypeus convex; mandible narrow, medially less than 0.6 × as wide as base, upper tooth longer than lower; fore wing with SDI = 1.0–1.27, CI less than 0.55; metasomal tergites shorter and wider, T5 less than 1.2 × as long as high ............... 41
41. Fore wing length 17.6–23.0 mm, central sclerite weak, C-shaped, formed from the distal periphery of an extensive quadra, ICI = 0.8–1.0 (Fig. 54f); mesosoma reddish with black markings (Fig. 54e) ............................................................................................................ E. nigristigma Cushman, 1937
– Fore wing length 7.5–23.5 mm, central sclerite weak to strong, varied in shaped, fenestra without quadra, ICI less than 0.5; mesosoma yellow with or without black markings black markings ...... 42
42. Fore wing entirely hyaline; marginal cell proximally evenly and densely setose; AI = 1.55–2.32 (Figs 6f, 49f) ................................................................................................................................... 43
– Fore wing with darkened patches; marginal cell proximally sparsely setose to glabrous; AI = 0.60–0.82 (Figs 13f, 18f) ................................................................................................................................. 44
43. Face and mesosoma entirely reddish brown (Fig. 6b, e); mesopleuron puncto-striate to finely striate; metapleuron coriaceous to finely striate (Fig. 6e); fore wing with central sclerite of similar size and shape to proximal sclerite, positioned in distal half of fenestra (Fig. 6f) ........................................... ................................................................................................................... E. aequiscleritalis sp. nov.
– Face and mesosoma with black markings (Fig. 49b, e); mesopleuron and metapleuron coarsely striate (Fig. 49e); fore wing with central sclerite much smaller than proximal sclerite and positioned in center of fenestra (Fig. 49f) ............................................................................ E. melanothoracicus sp. nov.
44. Antenna with 55–57 flagellomeres; fore wing darkened in marginal cell, marginal cell proximally glabrous, central sclerite linear, curved, subparallel to distal sclerite, SDI = 1.16–1.22 (Fig. 18f) ... .......................................................................................................... E. concentralis Cushman, 1937
– Antenna with 49–53 flagellomeres; fore wing darkened in marginal and discosubmarginal cells, marginal cell proximally sparsely setose, central sclerite transverse with proximal part broadened and distal part narrow (Fig. 13f) .......................................................... E. bifasciatus (Uchida, 1928)
45. Outer mandibular surface with a diagonal groove extending from upper proximal corner to between base of teeth (Figs 51b, 69b) ........................................................................................................... 46 – Outer mandibular surface without a diagonal groove (Figs 52b, 73b) ........................................... 54
46. Central sclerite linear, more or less parallel to distal sclerite (Fig. 51f); metapleuron strongly rugose (Fig. 51e) .................................................................................... E. nathani Gauld & Mitchell, 1981
– Central sclerite varied in shaped, but never parallel to distal sclerite; metapleuron punctate to puncto-striate ............................................................................................................................................... 47
47. Proximal and distal sclerites confluent (Figs 10f, 48f, 69f, 74f) .................................................... 48
– Proximal sclerite not confluent with distal sclerite (Figs 4f, 12f, 44f) ........................................... 52
48. Lower face 0.82 × as wide as long (Fig. 38b); lateral longitudinal carina of scutellum present on anterior 0.6; central sclerite shaped almost as a footprint with distal end broadened (Fig. 38f) ........ ................................................................................................................................... E. hiepi sp. nov.
– Lower face less than 0.76× as wide as long; lateral longitudinal carina of scutellum present on at least anterior 0.8; central sclerite oval, elongate to linear or crescentic ......................................... 49
49. Mandible with a conspicuous brush of long stout setae on diagonal groove; fore wing with central sclerite crescentic, fenestra with distinct quadra (Fig. 74f); metapleuron shiny with fine punctures (Fig. 74e) ..................................................................................................... E. strigilatus Tang, 1990
– Mandible with fine pubescence in diagonal groove; fore wing with central sclerite oval, elongate to linear, fenestra without distinct quadra; metapleuron less shiny to mat, densely punctate to diagonally striate ............................................................................................................................................... 50
50. Fore wing with marginal cell glabrous proximally (Fig. 69f) ............... E. sauteri (Enderlein, 1921)
– Fore wing with marginal cell uniformly setose, without any glabrous area (Figs 10f, 48f) ........... 51
51. Central sclerite oval (Fig. 48f); terminal metasomal tergites black (Fig. 48a) ................................... ........................................................................................................ E. melanocarpus Cameron, 1905
– Central sclerite elongately oval (Fig. 10f); metasomal tergites reddish brown, without black markings (Fig. 10a) ..................................................................................................... E. bacillaris Wang, 1997
52. Central sclerite moderately large, weakly sclerotized (Fig. 4f); upper mandibular tooth about 2.0× as long as lower tooth (Fig. 4b); metapleuron with diagonal striae (Fig. 4e) ..................................... ...................................................................................................... E. aciculatus (Taschenberg, 1875)
– Central sclerite small to large, strongly pigmented; upper mandibular tooth about 1.3–1.6 × as long as lower tooth; metapleuron punctate with isolated striae to puncto-striae or coriaceous ............. 53
53. Central sclerite fairly large, D-shaped, separated from proximal sclerite by less than 2.0× its own maximum diameter (Fig. 44f) ............................................................ E. laqueatus (Enderlein, 1921)
– Central sclerite moderately small, circular to oval, separated from proximal sclerite by more than 3.0 × its own maximum diameter (Fig. 12f) .......................................... E. bharatensis Nikam, 1980
54. Proximal sclerite entirely absent (Figs 16f, 60f) ............................................................................ 55
– Proximal sclerite present, usually strongly pigmented ................................................................... 56
55. Fore wing with vein 2r&RS strongly sinuous, abruptly narrow and straight at distal 0.15; fenestra with two medium-sized sclerites in distal part ............................... E. pinguivena (Enderlein, 1921)
– Fore wing with vein 2r&Rs weakly sinuous, distal 0.4 narrow and straight; fenestra with one small sclerite in distal part ............................................................................. E. centraliscleritiger sp. nov.
56. Fore wing fenestra long and anterodistal corner (sub) interstitial to postfurcal to RS (Figs 52f, 58f, 73f, 84f) .......................................................................................................................................... 57
– Fore wing fenestra short to moderately long and anterodistal corner antefurcal to RS more than 0.4× length of 2rs-m (Figs 5f, 29f, 34f, 59f) .......................................................................................... 60
57. Margin of propodeal spiracle not joined to lateral carina by a raised flange; fore wing ICI about 0.41–0.43; [body yellow orange with mesoscutum, T3–4 ventrally and T5–8 entirely darkish brown to black] ............................................................................................. E. nigribasalis (Uchida, 1928)
– Margin of propodeal spiracle joined to lateral carina by a raised flange; fore wing ICI at least 0.47 ......................................................................................................................................................... 58
58. Fore wing with darkened areas (Fig. 84f); [mesosoma and metasomal tergites with yellow and black pattern] .......................................................................................... E. zebrus Gauld & Mitchell, 1981
– Fore wing hyaline, without darkened areas .................................................................................... 59
59. Metapleuron puncto-striate; fore wing with central sclerite small, weak, circular to oval (Fig. 73f); metasoma from T5 onwards black .................................................... E. stenophleps Cushman, 1937
– Metapleuron strigose (Fig. 58e); fore wing with central sclerite C-shaped, formed from the distal periphery of an extensive quadra (Fig. 58f); T3–4 dorsally and T5 onwards entirely black .............. ......................................................................................................... E. pallidistigma Cushman, 1937
60. Fore wing vein 1m-cu&M centrally angled, CI = 0.73–0.84 (Fig. 29f) ............................................. ........................................................................................................... E. flavocephalus (Kirby, 1900)
– Fore wing vein 1m-cu&M sinuous or evenly arcuate, CI at most 0.65 .......................................... 61
61. Fore wing vein 2r&RS with a weak, central, anterior angulation (Fig. 34f); [sclerites weak to moderately strong, central sclerite linear, situated in the middle of fenestra and subparallel to 2r&RS] ............. .................................................................................................... E. grammospilus (Enderlein, 1921)
– Fore wing vein 2r&RS without a central, anterior angulation, sometimes with posterior bulb ..... 62
62. Fore wing with proximal sclerite moderately weak, blade-like in shape (Fig. 59f) ........................... ....................................................................................................................... E. pantanae Tang, 1990
– Fore wing with proximal sclerite strongly pigmented, triangular .................................................. 63
63. Mandible very strongly twisted by ca 85°, two teeth equal in length (Fig. 5b); median lobe and posterior part of mesoscutum black ............................................................. E. acutus Shimizu, 2020
– Mandible torsion various, not very strongly twisted, upper tooth more or less longer than lower tooth; colour of mesoscutum various ........................................................................................................ 64
64. Propodeum with distinct posterior transverse carina arising from pleural carina and centrally incomplete (Fig. 72e) .................................................................. E. signativentris (Tosquinet, 1903)
– Propodeum without even a lateral vestige of posterior transverse carina ....................................... 65
65. Fore wing basal and discosubmarginal cells with short and sparse setae, separated by more than their own length, marginal cell sparsely setose to glabrous proximally, vein 2r&RS with posterior bulb medially, vein M&RS interstitial to 1cu-a (Figs 15f, 66f) .............................................................. 66
– Fore wing basal and discosubmarginal cells with longer and denser setae, separated by less than their own length, marginal cell uniformly setose; vein 2r&RS without posterior bulb, M&RS interstitial to or postfurcal to 1cu-a .................................................................................................................. 67
66. Mesoscutum entirely reddish brown; central sclerite circular to oval (Fig. 66f) ................................ ....................................................................................................... E. rhetus Gauld & Mitchell, 1981
– Mesoscutum reddish brown with three black stripes; central sclerite elongate oval (Fig. 15f) ......... ........................................................................................................................ E. bulbipennis sp. nov.
67. Fore wing with both ICI and CI smaller than 0.5 (Fig. 53f) ....................... E. nigristernalis sp. nov.
– Fore wing with ICI greater than 0.5, CI various ............................................................................. 68
68. Fore wing CI = 0.45 or more .......................................................................................................... 69
– Fore wing CI less than 0.40 ............................................................................................................ 71
69. Fore wing with central sclerite C-shaped and parallel to distal sclerite, discosubmarginal cell with conspicuous long line of setae (Fig. 30f); [lower face 0.82–0.87× as broad as long] ........................ ............................................................................................................ E. formosensis (Uchida, 1928)
– Fore wing with central sclerite subtriangular to circular, discosubmarginal cell without conspicuous long line of setae (Figs 33f, 70f) ..................................................................................................... 70
70. Lower face 0.82 × as wide as long (Fig. 33b); fore wing with proximal sclerite about 0.5× as high as maximum width, central sclerite oval (Fig. 33f); S6–8 of male without long stout erect setae on posterior margins ............................................................................................. E. gialaiensis sp. nov.
– Lower face 0.67–0.75 × as wide as long (Fig. 70b); fore wing with proximal sclerite about 0.7 × as high as maximum width, central sclerite subtriangular to circular (Fig. 70f); S6–8 of male with long stout erect setae on posterior margins ............................................................ E. selmatos Chiu, 1954
71. Fore wing partly darkened, central sclerite oval, with maximum diameter smaller than distance from central sclerite to vein 2r&RS (Fig. 79f) .................................................................. E. tuani sp. nov.
– Fore wing hyaline (except somewhat darkened in case of E. abdominalis), central sclerite various, if oval or subtriangular then its maximum diameter larger than distance from central sclerite to vein 2r&RS ............................................................................................................................................. 72
72. Central sclerite linear or crescentic, formed from the sclerotized anterior, antero-distal or distal margin of large quadra; quadra occluding much of fenestra (Figs 28f, 36f, 43f, 55f, 80f) ........................ 73
– Central sclerite more or less oval or subtriangular; quadra, if present, smaller ............................. 80
73. Fore wing with AI = 0.29–0.53 ....................................................................................................... 74
– Fore wing with AI = 0.59–1.0 ......................................................................................................... 76
74. Metapleuron punctate or puncto-granulate, without any diagonal strigose ridges (Fig. 80e); fore wing with proximal sclerite high and narrow, about 2.2× as high as median width (Fig. 80f) ................... .......................................................................................................... E. urus Gauld & Mitchell, 1981
– Metapleuron with conspicuous diagonal strigose ridges (Figs 28e, 43e); fore wing with proximal sclerite moderately high and wider, about 1.5–1.7× as high as median width (Figs 28f, 43f) ...... 75
75. Central sclerite long and slender (Fig. 28f) .......................................... E. flavicaput (Morley, 1912)
– Central sclerite short and stout (Fig. 43f) ......................................... E. kanshirensis (Uchida, 1928)
76. Margin of propodeal spiracle connected to lateral carina by a raised flange, mesoscutum always with black markings ................................................................................................................................ 77
– Margin of propodeal spiracle not connected to lateral carina by a raised flange or rarely the two connected by weak creases; mesoscutum entirely reddish brown or with median black stripe ..... 78
77. Mandible twisted by ca 75° (Fig. 55b); mesoscutum with posterior half black ................................. ................................................................................................................ E. nigriventris Nikam, 1975
– Mandible twisted by 10–15° (Fig. 36b); mesoscutum with three black stripes .................................. ................................................................................................... E. hamatus Gauld & Mitchell, 1981
78. Body large, entirely reddish yellow; fore wing length 18.5–19.3 mm, central sclerite strongly sclerotized (Fig. 7f); hind wing vein RA with 9 hamuli; metapleuron with conspicuous diagonal strigose ridges (Fig. 7e) ................................................................. E. argus Gauld & Mitchell, 1981
– Body smaller, reddish yellow to reddish brown with at least fore wing costal vein black; fore wing length 12.7–17.2 mm, central sclerite weakly to strongly sclerotized; hind wing vein RA with 6–8 hamuli; metapleuron from puncto-striate to striate ........................................................................ 79
79. Central sclerite strongly pigmented as an ellipse parallel to vein 2r&RS, formed from antero-distal margin of quadra (Fig. 8f) .......................................................................... E. ashbyi Ashmead, 1904
– Central sclerite weakly to moderately strongly pigmented, C-shaped, formed from distal margin of quadra (Fig. 81f) ................................................................................... E. verticinus (Roman, 1913)
80. Mesosoma with black markings; T1–4 anteriorly pale yellow, posteriorly black; hind wing distally darkened ........................................................................................ E. abdominalis (Szépligeti, 1906)
– Mesosoma entirely reddish brown; T1–4 without alternative light and dark banding; hind wing entirely hyaline ............................................................................................................................................. 81
81. Fore wing with SRI = 0.33, posterodistal corner of second discal cell acute to right- angled (Figs 62f, 77f); propodeum finely and closely wrinkled; clypeus with apical margin impressed .................. 82
– Fore wing with SRI = 0.25 or less, posterodistal corner of second discal cell obtuse, at least 95° or more (Figs 41f, 82f); propodeum coarsely wrinkled to concentrically striate; clypeus with apical margin never impressed .................................................................................................................. 83
82. Mandible with a brush of closely spaced fine setae on outer surface (Fig. 77b); lower face uniformly orange-yellow, at most grading to yellow towards orbits (Fig. 77b); central sclerite oval (Fig. 77f) ...................................................................................................................... E. tripartitus Chiu, 1954
– Mandible with sparse scattered setae on outer surface; lower face reddish brown with discrete white-yellow orbital bands (Fig. 62b); central sclerite more or less circular (Fig. 62f) ............................... ........................................................................................................ E. pseudantennatus Gauld, 1977
83. Distal side of central sclerite conspicuously angulate (Fig. 41f) ........................................................ ......................................................................................................... E. ixion Gauld & Mitchell, 1981
– Distal side of central sclerite evenly rounded (Fig. 82f) ......................... E. vestigator (Smith, 1858)
description: Published as part of Pham, Nhi Thi, Pham, Phu Van, Matsumoto, Rikio, Shimizu, So & Broad, Gavin R., 2023, A review of the genus Enicospilus Stephens (Ichneumonidae: Ophioninae) from Vietnam, with descriptions of ten new species, pp. 1-151 in European Journal of Taxonomy 873 (1) on pages 5-14, DOI: 10.5852/ejt.2023.873.2133, http://zenodo.org/record/8038022