Genus Enicospilus Stephens, 1835

Enicospilus Stephens 1835: 126; type species, Ophion merdarius Gravenhorst sensu Stephens (= Ichneumon ramidulus Linneaus), by monotypy (Stephens 1845).

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); 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.

Distribution.

Worldwide except Antarctica (Yu et al. 2016).

Diagnosis.

Moderately to very large insects (fore wing length usually 9.0-30.0 mm).

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.

Mesosoma: pronotum unspecialised; notauli almost always absent; scutellum with lateral longitudinal carinae usually along more than 0.8 × its length (e.g. Fig. 2E-H); epicnemial carina present laterally (e.g. Figs 5E, 8E, 18E); posterior transverse carina of mesosternum complete; propodeum with anterior transverse carina usually more or less complete medially, anterior area long and longitudinally striate.

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.

Colour: body usually entirely testaceous, pale yellow to reddish brown (e.g. Figs 4A-E, 11A-E, 21A-E, 26A-E), sometimes posterior metasomal segments infuscate (e.g. Figs 9A, 17A, 18A); in some species body entirely brown to black, usually with testaceous to pale yellow patterns (e.g. Figs 5A-E, 28A-E); wings entirely hyaline or weakly infuscate (e.g. Figs 3F, 9F, 10F), rarely with strong infumate patches (e.g. Figs 5F, 28F); fenestra always hyaline (e.g. Figs 10F, 19F); sclerites hyaline to black (e.g. Figs 18F, 19F, 23F).

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 1985b, 1988; Broad 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).