Enicospilus Stephens, 1835

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, European Journal of Taxonomy 873 (1), pp. 1-151 : 5-14

publication ID

https://doi.org/ 10.5852/ejt.2023.873.2133

publication LSID

lsid:zoobank.org:pub:099F6A49-6516-4F39-9E20-A077A5D60535

DOI

https://doi.org/10.5281/zenodo.8040466

persistent identifier

https://treatment.plazi.org/id/03E3AC44-FFFB-FF87-9767-F91B024C8C27

treatment provided by

Felipe

scientific name

Enicospilus Stephens, 1835
status

 

Genus Enicospilus Stephens, 1835 View in CoL View at ENA

Enicospilus Stephens, 1835: 126 View in CoL ; 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 View in CoL ; type species: Banchogastra niger Ashmead, 1900 , by original designation.

Pycnophion Ashmead, 1900: 87 View in CoL ; type species: Pycnophion molokaiensis Ashmead, 1900 View in CoL , 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 View in CoL , by monotypy.

Abanchogastra Perkins, 1902: 141 View in CoL ; type species: Abanchogastra debilis Perkins, 1902 View in CoL , 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 View in CoL ), by monotypy.

Ophiomorpha Szépligeti, 1905: 34 ; type species: Ophion curvinervis Cameron, 1886 (= Enicospilus cameronii Dalla Torre, 1901 View in CoL ), 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 View Fig ), 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 View in CoL

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 View Fig ); clypeus in profile strongly convex, subnasute ( Fig. 26b, d View Fig ) ................................................. E. exaggeratus Chiu, 1954 View in CoL

– 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 View Fig ) ................................................................................................ 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 View in CoL (in part)

– Fore wing 1m-cu&M evenly curved, without short ramellus ........................................................... 5

5. Mandible strongly twisted by ca 80° ( Fig. 24b View Fig ); fore wing 1 st subdiscal cell with sparse setae ( Fig. 24f View Fig ); fore wing length 12.5–15.0 mm ..................................................... E. erythrocerus (Cameron, 1905) View in CoL

– Mandible twisted by 20°–30° ( Fig. 35b View Fig ); fore wing 1 st subdiscal cell generally with denser setae at least on anterior 0.4 ( Fig. 35f View Fig ); fore wing length 19.0–23.0 mm .......... E. grandis (Cameron, 1905) View in CoL

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 View Fig ) ............. ...................................................................................................... E. eastopi Gauld & Mitchell, 1981 View in CoL

– 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 View Fig ); 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 View Fig ) .................................................................... E. bakerielli Gauld & Mitchell, 1981 View in CoL

– Interocellar area black ( Figs 17c View Fig , 78c View Fig ); 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 View Fig , 78f View Fig ) ..................................................................................................................... 9

9. Fore wing proximal sclerite larger, distance from proximal sclerite to vein 2r&RS less than its maximum diameter ( Fig. 17f View Fig ); 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 View Fig ); 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 View Fig , 19f View Fig , 37f View Fig , 39f View Fig , 47f View Fig , 61f View Fig , 76f View Fig ) or entirely absent ( Figs 31f View Fig , 64f View Fig ) ...................................................................................................................... 12

– Proximal sclerite quite widened, usually triangular (other shapes in E. dasychirae View in CoL , E. nigronotatus View in CoL and E. pseudoconspersae View in CoL ) 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 View Fig ) ................................................. E. biharensis Townes, Townes & Gupta, 1961 View in CoL

– 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 View Fig ) ....................................................... E. corculus ( Tosquinet, 1903) View in CoL

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 View Fig ); metasoma from T5 onwards blackish .......................................................................... E. fusiformis Chiu, 1954 View in CoL

– Fore wing vein 1m-cu&M sinuous and more or less uniformly thickened, ICI = 0.75–1.09 ( Fig. 64f View Fig ); metasomal tergites reddish brown, sometimes from T3 onwards with dorsal line and ventral parts blackish ...................................................................................................... E. pungens ( Smith, 1874) View in CoL

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 View Fig ) ...................................................................................... E. iapetus Gauld & Mitchell, 1981 View in CoL

– Marginal cell of fore wing uniformly setose, without quadra, ICI = 0.65 or less .............................. ............................................................................................................ E. pudibundae ( Uchida, 1928) View in CoL

18. Marginal cell of fore wing proximally with a glabrous area ( Figs 37f View Fig , 47f View Fig ) .................................. 19

– Marginal cell of fore wing proximally at most sparsely but uniformly setose, without a glabrous area ( Fig. 61f View Fig ) ......................................................................................................................................... 20

19. Fore wing vein 2r&RS relatively straight, relatively uniformly thickened, AI = 0.9 ( Fig. 37f View Fig ); mesopleuron puncto-striate ventrally ( Fig. 37e View Fig ); fore wing length 18.7 mm ..................................... ...................................................................................................... E. hedilis Gauld & Mitchell, 1981 View in CoL

– Fore wing vein 2r&RS sinuous, abruptly narrowed distally, AI = 1.36 ( Fig. 47f View Fig ); mesopleuron striate ventrally ( Fig. 47e View Fig ); fore wing length 12.8 mm ........................................... E. maritus Roman, 1913 View in CoL

20. Fore wing vein 1m-cu&M and 2r&RS strongly sinuous ( Fig. 61f View Fig ) ................................................... ...................................................................................................... E. plicatus ( Brullé, 1846) View in CoL (in part)

– Fore wing vein 1m-cu&M evenly arcuate, 2r&RS weakly sinuous ( Fig. 76f View Fig ) ................................... ................................................................................................................... E. transversus Chiu, 1954 View in CoL

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 View in CoL ) ( Figs 42f View Fig , 56f View Fig , 63f View Fig ) .......................................................................................................................... 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 View Fig ); body dark reddish brown with black markings ( Fig. 56a View Fig ) ............................................. E. nigronotatus Cameron, 1903 View in CoL

– Distal sclerite absent, SRI = 0.3; body reddish brown without black markings ................................. ..................................................................................... E. pseudoconspersae ( Sonan, 1927) View in CoL (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 View Fig ; SDI = 0.90–1.05 ..................................................... E. javanus ( Szépligeti, 1910) View in CoL

– 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 View Fig , 63f View Fig ; SDI at least 1.30 ................ 25

25. Proximal sclerite of fore wing fenestra usually entirely weakly pigmented and half-moon-shaped ( Fig. 63f View Fig ) ..................................................................... E. pseudoconspersae ( Sonan, 1927) View in CoL (in part) – Proximal sclerite of fore wing fenestra partly to entirely strongly pigmented and drop-shaped ( Fig. 20f View Fig ) ............................................................................................. E. dasychirae Cameron, 1905 View in CoL

26. Hind tarsal claws with distal pecten projecting beyond apical tooth .................................................. .................................................................................................... E. mythrus Gauld & Mitchell, 1981 View in CoL

– Hind tarsal claws simple, distal pecten not projecting ................................................................... 27

27. Interocellar area uniformly black ( Figs 21c View Fig , 57c View Fig ) .......................................................................... 28

– Interocellar area yellowish-brown to orange ( Figs 27c View Fig , 45c View Fig , 83c View Fig ) ................................................. 29

28. Fore wing with SDI = 1.07 or less, ICI = 0.35–0.36, vein 2r&RS centrally incrassate ( Fig. 57f View Fig ); metapleuron reticulate, at least on posterior 0.5 ( Fig. 57e View Fig ); mesoscutum and mesopleuron with black markings ( Fig. 57e View Fig ) ............................................................................ E. nigropectus Cameron, 1905 View in CoL

– Fore wing with SDI = 1.28–1.37, ICI = 0.53–0.59, vein 2r&RS more or less evenly thickened ( Fig. 21f View Fig ); metapleuron striate ( Fig. 21e View Fig ); mesoscutum and mesopleuron without black markings .. ............................................................................................................... E. dolosus ( Tosquinet, 1896) View in CoL

29. Outer mandibular surface with a diagonal groove extending from dorsal proximal corner to between base of teeth ( Figs 27b View Fig , 45b View Fig ) ........................................................................................................... 30

– Outer mandibular surface without a diagonal groove ( Figs 40b View Fig , 75b View Fig ) ........................................... 32

30. Fore wing length 17.5 mm, proximal sclerite evenly tapered to join distal sclerite ( Fig. 45f View Fig ); antenna with 73 flagellomeres; metapleuron finely punctate ( Fig. 45e View Fig ) .................. E. longitarsis Tang, 1990 View in CoL

– 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 View Fig , 83f View Fig ); antenna with 54–61 flagellomeres; metapleuron puncto-striate ( Fig. 27e View Fig ) .................................................................................................................. 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 View Fig ) .................................................................................... E. fittoni Nikam, 1980 View in CoL

– 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 View Fig ) ........................................... E. yonezawanus ( Uchida, 1928) View in CoL

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 View Fig ); mesoscutum with three black stripes ............................ E. teleus Gauld & Mitchell, 1981 View in CoL

– Smaller body size, fore wing length not exceed 21 mm with fenestra much longer, at least 4.5 × proximal sclerite ( Figs 40f View Fig , 46f View Fig , 68f View Fig ); 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 View Fig ) ..................................................................................... E. insinuator ( Smith, 1860) View in CoL

– Mesoscutum with posterior 0.5 black; tegula with posterior half black; metapleuron coarsely strigose ( Figs 46e View Fig , 68e View Fig ) ................................................................................................................................ 35

35. Mandible twisted by about 80° ( Fig. 68b View Fig ); metasomal tergites exceptionally long, T 4 in profile 1.4× as long as high; metasoma from T3 onwards entirely black ( Fig. 68a View Fig ) .............................................. ........................................................................................................ E. rogus Gauld & Mitchell, 1981 View in CoL

– Mandible twisted by about 20° ( Fig. 46b View Fig ); metasomal tergites not exceptionally long, T 4 in profile less than 1.2× as long as high; T4 with pale yellow patch ( Fig. 46a View Fig ) ................. E. maai Chiu, 1954 View in CoL

36. Metapleuron puncto-striate ( Fig. 71e View Fig ); fore wing length 10.5–12.5 mm, ICI = 0.41–0.62 ................ .............................................................................................................. E. shinkanus ( Uchida, 1928) View in CoL

– Metapleuron striate to strigose ( Fig. 23e View Fig ); 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 View Fig ) ......................................................... E. purifenestratus ( Enderlein, 1921) View in CoL

– Lower face 0.75–0.82 × as wide as long; fore wing with distal sclerite indistinct, AI = 0.59–0.76 ( Fig. 23f View Fig ) ................................................................................................ E. enicospilus Nikam, 1972 View in CoL

38. Interocellar area black or dark brown, contrasting with vertex ( Figs 32c View Fig , 67c View Fig ) ............................. 39

– Interocellar area yellowish-brown to orange .................................................................................. 45

39. Clypeus nasute, strongly convex ( Fig. 67b, d View Fig ) ............ E. riukiuensis ( Matsumura & Uchida, 1926) View in CoL

– Clypeus not nasute, flat to moderately convex ............................................................................... 40

40. Lower face 0.8 × as wide as long ( Fig. 32b View Fig ); clypeus strongly convex; mandible wide and long, medially 0.6× as broad as base, two teeth equal in length ( Fig. 32b View Fig ); 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 View Fig ) .......................... ................................................................................................................... E. gasteralis Nikam, 1980 View in CoL

– 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 View Fig ); mesosoma reddish with black markings ( Fig. 54e View Fig ) ............................................................................................................ E. nigristigma Cushman, 1937 View in CoL

– 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 View Fig , 49f View Fig ) ................................................................................................................................... 43

– Fore wing with darkened patches; marginal cell proximally sparsely setose to glabrous; AI = 0.60–0.82 ( Figs 13f View Fig , 18f View Fig ) ................................................................................................................................. 44

43. Face and mesosoma entirely reddish brown ( Fig. 6b, e View Fig ); mesopleuron puncto-striate to finely striate; metapleuron coriaceous to finely striate ( Fig. 6e View Fig ); fore wing with central sclerite of similar size and shape to proximal sclerite, positioned in distal half of fenestra ( Fig. 6f View Fig ) ........................................... ................................................................................................................... E. aequiscleritalis sp. nov.

– Face and mesosoma with black markings ( Fig. 49b, e View Fig ); mesopleuron and metapleuron coarsely striate ( Fig. 49e View Fig ); fore wing with central sclerite much smaller than proximal sclerite and positioned in center of fenestra ( Fig. 49f View Fig ) ............................................................................ 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 View Fig ) ... .......................................................................................................... E. concentralis Cushman, 1937 View in CoL

– 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 View Fig ) .......................................................... E. bifasciatus ( Uchida, 1928) View in CoL

45. Outer mandibular surface with a diagonal groove extending from upper proximal corner to between base of teeth ( Figs 51b View Fig , 69b View Fig ) ........................................................................................................... 46 – Outer mandibular surface without a diagonal groove ( Figs 52b View Fig , 73b View Fig ) ........................................... 54

46. Central sclerite linear, more or less parallel to distal sclerite ( Fig. 51f View Fig ); metapleuron strongly rugose ( Fig. 51e View Fig ) .................................................................................... E. nathani Gauld & Mitchell, 1981 View in CoL

– 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 View Fig , 48f View Fig , 69f View Fig , 74f View Fig ) .................................................... 48

– Proximal sclerite not confluent with distal sclerite ( Figs 4f View Fig , 12f View Fig , 44f View Fig ) ........................................... 52

48. Lower face 0.82 × as wide as long ( Fig. 38b View Fig ); lateral longitudinal carina of scutellum present on anterior 0.6; central sclerite shaped almost as a footprint with distal end broadened ( Fig. 38f View Fig ) ........ ................................................................................................................................... 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 View Fig ); metapleuron shiny with fine punctures ( Fig. 74e View Fig ) ..................................................................................................... E. strigilatus Tang, 1990 View in CoL

– 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 View Fig ) ............... E. sauteri ( Enderlein, 1921) View in CoL

– Fore wing with marginal cell uniformly setose, without any glabrous area ( Figs 10f View Fig , 48f View Fig ) ........... 51

51. Central sclerite oval ( Fig. 48f View Fig ); terminal metasomal tergites black ( Fig. 48a View Fig ) ................................... ........................................................................................................ E. melanocarpus Cameron, 1905 View in CoL

– Central sclerite elongately oval ( Fig. 10f View Fig ); metasomal tergites reddish brown, without black markings ( Fig. 10a View Fig ) ..................................................................................................... E. bacillaris Wang, 1997 View in CoL

52. Central sclerite moderately large, weakly sclerotized ( Fig. 4f View Fig ); upper mandibular tooth about 2.0× as long as lower tooth ( Fig. 4b View Fig ); metapleuron with diagonal striae ( Fig. 4e View Fig ) ..................................... ...................................................................................................... E. aciculatus ( Taschenberg, 1875) View in CoL

– 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 View Fig ) ............................................................ E. laqueatus ( Enderlein, 1921) View in CoL

– Central sclerite moderately small, circular to oval, separated from proximal sclerite by more than 3.0 × its own maximum diameter ( Fig. 12f View Fig ) .......................................... E. bharatensis Nikam, 1980 View in CoL

54. Proximal sclerite entirely absent ( Figs 16f View Fig , 60f View Fig ) ............................................................................ 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) View in CoL

– 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 View Fig , 58f View Fig , 73f View Fig , 84f View Fig ) .......................................................................................................................................... 57

– Fore wing fenestra short to moderately long and anterodistal corner antefurcal to RS more than 0.4× length of 2rs-m ( Figs 5f View Fig , 29f View Fig , 34f View Fig , 59f View Fig ) .......................................................................................... 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) View in CoL

– 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 View Fig ); [mesosoma and metasomal tergites with yellow and black pattern] .......................................................................................... E. zebrus Gauld & Mitchell, 1981 View in CoL

– Fore wing hyaline, without darkened areas .................................................................................... 59

59. Metapleuron puncto-striate; fore wing with central sclerite small, weak, circular to oval ( Fig. 73f View Fig ); metasoma from T5 onwards black .................................................... E. stenophleps Cushman, 1937 View in CoL

– Metapleuron strigose ( Fig. 58e View Fig ); fore wing with central sclerite C-shaped, formed from the distal periphery of an extensive quadra ( Fig. 58f View Fig ); T3–4 dorsally and T5 onwards entirely black .............. ......................................................................................................... E. pallidistigma Cushman, 1937 View in CoL

60. Fore wing vein 1m-cu&M centrally angled, CI = 0.73–0.84 ( Fig. 29f View Fig ) ............................................. ........................................................................................................... E. flavocephalus ( Kirby, 1900) View in CoL

– 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 View Fig ); [sclerites weak to moderately strong, central sclerite linear, situated in the middle of fenestra and subparallel to 2r&RS] ............. .................................................................................................... E. grammospilus ( Enderlein, 1921) View in CoL

– 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 View Fig ) ........................... ....................................................................................................................... E. pantanae Tang, 1990 View in CoL

– Fore wing with proximal sclerite strongly pigmented, triangular .................................................. 63

63. Mandible very strongly twisted by ca 85°, two teeth equal in length ( Fig. 5b View Fig ); 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 View Fig ) .................................................................. E. signativentris ( Tosquinet, 1903) View in CoL

– 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 View Fig , 66f View Fig ) .............................................................. 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 View Fig ) ................................ ....................................................................................................... E. rhetus Gauld & Mitchell, 1981 View in CoL

– Mesoscutum reddish brown with three black stripes; central sclerite elongate oval ( Fig. 15f View Fig ) ......... ........................................................................................................................ E. bulbipennis sp. nov.

67. Fore wing with both ICI and CI smaller than 0.5 ( Fig. 53f View Fig ) ....................... 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 View Fig ); [lower face 0.82–0.87× as broad as long] ........................ ............................................................................................................ E. formosensis ( Uchida, 1928) View in CoL

– Fore wing with central sclerite subtriangular to circular, discosubmarginal cell without conspicuous long line of setae ( Figs 33f View Fig , 70f View Fig ) ..................................................................................................... 70

70. Lower face 0.82 × as wide as long ( Fig. 33b View Fig ); fore wing with proximal sclerite about 0.5× as high as maximum width, central sclerite oval ( Fig. 33f View Fig ); 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 View Fig ); fore wing with proximal sclerite about 0.7 × as high as maximum width, central sclerite subtriangular to circular ( Fig. 70f View Fig ); S6–8 of male with long stout erect setae on posterior margins ............................................................ E. selmatos Chiu, 1954 View in CoL

71. Fore wing partly darkened, central sclerite oval, with maximum diameter smaller than distance from central sclerite to vein 2r&RS ( Fig. 79f View Fig ) .................................................................. E. tuani sp. nov.

– Fore wing hyaline (except somewhat darkened in case of E. abdominalis View in CoL ), 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 View Fig , 36f View Fig , 43f View Fig , 55f View Fig , 80f View Fig ) ........................ 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 View Fig ); fore wing with proximal sclerite high and narrow, about 2.2× as high as median width ( Fig. 80f View Fig ) ................... .......................................................................................................... E. urus Gauld & Mitchell, 1981 View in CoL

– Metapleuron with conspicuous diagonal strigose ridges ( Figs 28e View Fig , 43e View Fig ); fore wing with proximal sclerite moderately high and wider, about 1.5–1.7× as high as median width ( Figs 28f View Fig , 43f View Fig ) ...... 75

75. Central sclerite long and slender ( Fig. 28f View Fig ) .......................................... E. flavicaput ( Morley, 1912) View in CoL

– Central sclerite short and stout ( Fig. 43f View Fig ) ......................................... E. kanshirensis ( Uchida, 1928) View in CoL

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 View Fig ); mesoscutum with posterior half black ................................. ................................................................................................................ E. nigriventris Nikam, 1975 View in CoL

– Mandible twisted by 10–15° ( Fig. 36b View Fig ); mesoscutum with three black stripes .................................. ................................................................................................... E. hamatus Gauld & Mitchell, 1981 View in CoL

78. Body large, entirely reddish yellow; fore wing length 18.5–19.3 mm, central sclerite strongly sclerotized ( Fig. 7f View Fig ); hind wing vein RA with 9 hamuli; metapleuron with conspicuous diagonal strigose ridges ( Fig. 7e View Fig ) ................................................................. E. argus Gauld & Mitchell, 1981 View in CoL

– 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 View Fig ) .......................................................................... E. ashbyi Ashmead, 1904 View in CoL

– Central sclerite weakly to moderately strongly pigmented, C-shaped, formed from distal margin of quadra ( Fig. 81f View Fig ) ................................................................................... E. verticinus ( Roman, 1913) View in CoL

80. Mesosoma with black markings; T1–4 anteriorly pale yellow, posteriorly black; hind wing distally darkened ........................................................................................ E. abdominalis ( Szépligeti, 1906) View in CoL

– 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 View Fig , 77f View Fig ); 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 View Fig , 82f View Fig ); 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 View Fig ); lower face uniformly orange-yellow, at most grading to yellow towards orbits ( Fig. 77b View Fig ); central sclerite oval ( Fig. 77f View Fig ) ...................................................................................................................... E. tripartitus Chiu, 1954 View in CoL

– Mandible with sparse scattered setae on outer surface; lower face reddish brown with discrete white-yellow orbital bands ( Fig. 62b View Fig ); central sclerite more or less circular ( Fig. 62f View Fig ) ............................... ........................................................................................................ E. pseudantennatus Gauld, 1977 View in CoL

83. Distal side of central sclerite conspicuously angulate ( Fig. 41f View Fig ) ........................................................ ......................................................................................................... E. ixion Gauld & Mitchell, 1981 View in CoL

– Distal side of central sclerite evenly rounded ( Fig. 82f View Fig ) ......................... E. vestigator ( Smith, 1858) View in CoL

Kingdom

Animalia

Phylum

Arthropoda

Class

Insecta

Order

Hymenoptera

Family

Ichneumonidae

Loc

Enicospilus Stephens, 1835

Pham, Nhi Thi, Pham, Phu Van, Matsumoto, Rikio, Shimizu, So & Broad, Gavin R. 2023
2023
Loc

Schizospilus

Seyrig A. 1935: 79
1935
Loc

Eremotyloides

Perkins R. C. L. 1915: 530
1915
Loc

Amesospilus

Enderlein G. 1914: 222
1914
Loc

Cryptocamptus Brèthes, 1909: 230

Brethes J. 1909: 230
1909
Loc

Metophion Szépligeti, 1905: 28

Viereck H. L. 1914: 94
Szepligeti G. 1905: 28
1905
Loc

Ceratospilus Szépligeti, 1905: 28

Szepligeti G. 1905: 28
1905
Loc

Atoponeura Szépligeti, 1905: 34

Szepligeti G. 1905: 34
1905
Loc

Ophiomorpha Szépligeti, 1905: 34

Szepligeti G. 1905: 34
1905
Loc

Abanchogastra

Perkins R. C. L. 1902: 141
1902
Loc

Cymatoneura

Viereck H. L. 1914: 8
Kriechbaumer J. 1901: 22
1901
Loc

Pterospilus

Viereck H. L. 1914: 126
Kriechbaumer J. 1901: 156
1901
Loc

Trispilus

Kriechbaumer J. 1901: 156
1901
Loc

Dispilus

Ashmead W. H. 1900: 86
1900
Loc

Pleuroneurophion

Ashmead W. H. 1900: 86
1900
Loc

Banchogastra

Ashmead W. H. 1900: 87
1900
Loc

Pycnophion

Ashmead W. H. 1900: 87
1900
Loc

Allocamptus Förster, 1869: 150

Thomson C. G. 1888: 1189
Forster A. 1869: 150
1869
Loc

Henicospilus

Agassiz L. J. R. 1846: 138
1846
Loc

Enicospilus

Horstmann K. 2005: 1264
Stephens J. L. 1835: 126
1835
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