The specific name is dedicated to the collector of the holotype, Mike Allen, who collected many specimens of Nepalese Hymenoptera that are now in NHMUK.

1♀: Nepal.

Type series: holotype ♀, Chautasa (6,000 ft), Nepal, 24.IX.1983, M.G. Allen leg. (NHMUK) (Fig. 3).

Nepal.

Female (Holotype) (Fig. 3). Body length ca 29.5 mm.

Head with GOI = 2.9 (Fig. 3C). Lower face 0.7× as wide as high, very finely coriaceous with fine punctures and setae (Fig. 3B). Clypeus 1.9× as wide as high, shiny and very finely coriaceous with fine punctures and setae, almost flat in profile, and its lower margin acute (Fig. 3B, C). Malar space 0.2× as long as basal mandibular width (Fig. 3B, C). Mandible weakly twisted by ca 25°, moderately long, evenly narrowed, its outer surface flat and smooth without a diagonal groove or a diagonal line of punctures (Fig. 3B, C). Upper mandibular tooth 1.4× as long as lower one (Fig. 3B). Frons, vertex and gena moderately shiny with fine setae (Fig. 3B–D). Posterior ocellus large and almost touching eye (Fig. 3B–D). Ventral end of occipital carina joining oral carina. Left antenna complete with 64 flagellomeres, and right antenna apically incomplete with 53 flagellomeres; first flagellomere 1.7× as long as second; 20^th flagellomere 1.6× as long as wide.

Mesosoma entirely very weakly shiny or not (Fig. 3E). Pronotum entirely striate. Mesoscutum 1.5× as long as its maximum width, very closely coriaceous with dense setae, very weakly shiny, evenly rounded in profile, and its anterior margin almost truncate in dorsal view and rounded in profile (Fig. 3E). Notauli absent (Fig. 3E). Scutellum moderately convex, very closely coriaceous with setae, with lateral longitudinal carinae reaching posterior end (Fig. 3E). Epicnemium densely punctate with setae. Epicnemial carina weak, almost straight and inclined to anterior, its dorsal end not reaching anterior margin of mesopleuron (Fig. 3E). Mesopleuron entirely closely longitudinally strigose (Fig. 3E). Submetapleural carina very strongly broadened anteriorly and forming a lobe. Metapleuron densely punctate to reticulate with setae, moderately swollen (Fig. 3E). Propodeum very strongly and abruptly declivous in profile; anterior transverse carina complete; pleural carina almost absent; anterior area longitudinally striate; spiracular area almost smooth with setae and strongly shiny; posterior area moderately subconcentrically strigose with a pair of strong posterior carinae laterally; propodeal spiracle elliptical, its outer margin not joining pleural carina by a ridge (Fig. 3E).

Wings. Fore wing length ca 19.5 mm with AI = 1.0, CI = 0.4, DI = 0.3, ICI = 0.6, SDI = 1.6, SI = 0.1, SRI = 0.2; vein 1m-cu&M almost evenly curved; vein 2r&RS very slightly sinuous and RS evenly curved; fenestra and sclerites of discosubmarginal cell as in Figure 3F; proximal sclerite triangular, confluent with distal sclerite, strongly pigmented; central sclerite entirely weakly sclerotised, very weakly pigmented partially, positioned in anterodistal part of fenestra; distal sclerite present proximally and absent distally; proximal corner of marginal cell uniformly setose; posterodistal corner of second discal cell ca 110°; posterodistal corner of subbasal cell ca 95°; vein 1cu-a antefurcal to M&RS by 0.3× 1cu-a length (Fig. 3F). Hind wing with NI = 2.8, RI = 2.2; vein RS straight; vein RA with 10 uniform hamuli.

Legs. Outer surface of fore tibia with scattered short spines. Hind leg with coxa in profile 1.8× as long as deep; basitarsus 2.0× as long as second tarsomere; fourth tarsomere 0.6× as long as third tarsomere and 2.9× as long as wide; tarsal claw simply pectinate.

Metasoma with DMI = 1.4, PI = 2.7, THI = 3.5; dorsal margin of tergite 1 slightly sinuous in profile; thyridium elliptical (Fig. 3A).

Colour (Fig. 3). Entirely testaceous except for yellow eye orbit and vertex, apex of mandible black. Wings hyaline; fore wing sclerites and pterostigma testaceous; veins testaceous to brown.

Variation. Unknown, only known from the holotype.

Male. Unknown

The affinities of E. alleni sp. nov. are unclear, but it may be related to the E. flavicaput group. However, E. alleni sp. nov. is a very distinctive species, readily distinguished by many characters, such as the elongate lower face (Fig. 3B), sculpture of the mesosoma (Fig. 3E), shape of propodeum (Fig. 3E), and two vestigial central sclerites of the fore wing fenestra (Fig. 3F).

Figure 3. 

Enicospilus alleni Shimizu sp. nov., ♀, holotype. A. Habitus; B. Head, frontal view; C. Head, lateral view; D. Head, dorsal view; E. Mesosoma, lateral view; F. Central part of fore wing.

11♀♀4♂♂: Nepal (1♂), India (11♀♀1♂), Taiwan (2♂♂).

Type series: lectotype of Henicospilus tainanensis Uchida, 1928, ♂, Tainan, Taiwan, S. Takano leg. (SEHU); holotype of Enicospilus concavus Chiu, 1954, ♂, Taihoku, Taiwan, 24.I.1932, J. Sonan leg. (TARI).

Non-type series: 1♂, Kathmandu (1,350 m), Nepal, VII.1983, M.G. Allen leg. (LT) (NHMUK) (Fig. 4); 9♀♀1♂, Patancheru, Andhra Pradesh, India, VII (7♀♀1♂) and VIII (2♀♀).1980, Bhatnagar leg. (LT) (NHMUK); 1♀, Jeypore, Orissa, India, IX.1958, P.S. Nathan leg. (EMUS); 1♀, Nilgira Hills, India, V. 1953, P.S. Nathan leg. (CNC).

Australasian and Oriental regions (Yu et al. 2016). Newly recorded from Nepal.

Head (Fig. 4B–D): GOI = 2.1–2.4; lower face 0.7–0.8× as wide as high; clypeus flat to slightly convex in profile, its lower margin subacute; mandible rather weakly twisted by 25–30°, moderately long, evenly tapered, its outer surface without a diagonal structure; upper mandibular tooth 1.2–1.5× as long as lower one; posterior ocellus almost touching eye; antenna with 45–56 flagellomeres and 20^th flagellomere 1.6–1.9× as long as wide.

Mesosoma (Fig. 4E): mesopleuron longitudinally punctostriate to striate; scutellum with lateral longitudinal carinae reaching posterior end and convergent posteriorly; metapleuron punctostriate; propodeum declivous, its posterior area moderately reticulate, outer margin of propodeal spiracle not joining pleural carina by a ridge.

Wings (Fig. 4F): fore wing with AI = 0.7–1.2, CI = 0.2–0.3, ICI = 0.5–0.7, SDI = 1.2–1.3; fore wing vein 1m-cu&M evenly curved, 2r&RS almost straight; fenestra and sclerites of discosubmarginal cell of fore wing as in Figure 4F; fenestra of fore wing not very long and its anterodistal corner distinctly separated from proximal end of vein RS; proximal sclerite triangular, confluent with distal one, strongly pigmented; central sclerite usually partially strongly pigmented and sclerotised, strongly pigmented part linear and parallel to vein 2r&RS, positioned in anterodistal part of fenestra; distal sclerite present proximally and vestigial distally; proximal corner of marginal cell of fore wing uniformly setose; vein 1cu-a antefurcal to subinterstitial to M&RS by less than 0.1× 1cu-a length.

Colour (Fig. 4): body including interocellar area entirely testaceous; wings hyaline.

Enicospilus ashbyi is similar to E. pallidus (Taschenberg, 1875) and separated from it by a few characters of the central sclerite (pigmented part of central sclerite narrower in E. ashbyi and wider in E. pallidus). However, the sclerite characters (e.g. the shape and degree of sclerotisation of the central sclerite) exhibit a wide range of variation within both species, suggesting that there are cryptic species and that integrative taxonomy is needed to define species limits in this complex.

Figure 4. 

Enicospilus ashbyi Ashmead, 1904, ♂. A. Habitus; B. Head, frontal view; C. Head, lateral view; D. Head, dorsal view; E. Mesosoma, lateral view; F. Central part of fore wing.

7♀♀3♂♂: Nepal (2♀♀), Taiwan (5♀♀3♂♂).

Type series: holotype of Henicospilus bifasciatus Uchida, 1928, ♀, Baibara, Taiwan, Uchida leg. (SEHU).

Non-type series: 2♀♀, Godaveri (1,550–1,700 m), Nepal, 1.VI.1984, M.G. Allen leg. (LT) (NHMUK) (Fig. 5); 1♂, Bukai, Taiwan, 13.VI.1934, L. Gressitt leg. (NHMUK); 1♂, Horisha, Taiwan, V. 1927, Sonan leg. (TARI); 1♀, Musha, Taiwan, IV.1938, Sonan leg. (TARI); 1♂, Shinten, Taiwan, IV.1921, Sonan leg. (TARI); 2♀♀, Taihoku, Taiwan, I. 1924, Sonan leg. (TARI); 1♀, Taipei, Taiwan, V.1950, Chiu leg. (TARI).

Oriental region (Yu et al. 2016). Newly recorded from Nepal.

Head (Fig. 5B–D): GOI = 3.1–3.4; lower face 0.6–0.7× as wide as high; clypeus moderately convex in profile, its lower margin acute; mandible rather strongly twisted by 55–65°, moderately long, evenly tapered, its outer surface without a diagonal structure; upper mandibular tooth 1.2–1.3× as long as lower one; posterior ocellus close to eye; antenna with 54–56 flagellomeres and 20^th flagellomere 3.1–3.3× as long as wide.

Mesosoma (Fig. 5E): mesopleuron rather coarsely longitudinally punctostriate to striate; scutellum with lateral longitudinal carinae reaching posterior end and convergent posteriorly; metapleuron rather coarsely striate; propodeum evenly weakly rounded, its posterior area moderately reticulate, outer margin of propodeal spiracle joining pleural carina by a ridge.

Wings (Fig. 5F): fore wing with AI = 0.7–1.0, CI = 0.4–0.5, ICI = 0.2, SDI = 1.1–1.2; fore wing vein 1m-cu&M evenly curved, 2r&RS almost straight, centrally broadened; fenestra and sclerites of discosubmarginal cell of fore wing as in Figure 5F; fenestra of fore wing not very long and its anterodistal corner distinctly separated from proximal end of vein RS; proximal sclerite triangular, confluent with distal one, strongly pigmented; central sclerite rather weakly pigmented and sclerotised, drop-shaped and its major axis parallel to vein 2r&RS, positioned in mediodistal part of fenestra; distal sclerite present proximally and vestigial distally; proximal corner of marginal cell of fore wing very sparsely setose, almost glabrous; vein 1cu-a subinterstitial to antefurcal to M&RS by less than 0.2× 1cu-a length.

Colour (Fig. 5): body entirely pale yellow with black marks on mesosoma, interocellar area, and posterior segments of metasoma; wings hyaline with two strongly infumate patches in the central part of the discosubmarginal cell (from anterior end of M&RS to base of 1m-cu&M) and the central part of the marginal cell (from anterocentral margin to base of RS).

Enicospilus bifasciatus is a very distinctive species and no closely related species are currently known. Hence, it is easily distinguished from all Enicospilus by many characters listed in the above diagnosis as well as identification key, such as two strongly infumate patches in the central part of the discosubmarginal cell (from anterior end of M&RS to base of 1m-cu&M) and the central part of the marginal cell (from anterocentral margin to base of RS) of fore wing, characteristic shape of sclerites of discosubmarginal cell of fore wing (cf. Fig. 5F), very sparsely setose and almost glabrous proximal corner of marginal cell of fore wing, and small value of ICI (= 0.2).

Figure 5. 

Enicospilus bifasciatus (Uchida, 1928), ♀. A. Habitus; B. Head, frontal view; C. Head, lateral view; D. Head, dorsal view; E. Mesosoma, lateral view; F. Central part of fore wing.

11♀♀1♂: Nepal (10♀♀1♂), India (1♀).

Type series: holotype of Henicospilus horsfieldi var glabratus Morley, 1913 (= Enicospilus biharensis Townes, Townes & Gupta, 1961), ♀, Bihar, Chapra, India (NHMUK, Type 3b.1266).

Non-type series: 1♀, Dotslghst (900 m), Nepal, 7.VII.1983, M.G. Allen leg. (Figs 2I, 6); 2♀♀, Kathmandu (4,300′), Nepal, VIII.1981, M.G. Allen leg.; 1♀, Kathmandu (1,300 m), Nepal, X.1982, M.G. Allen leg. (LT); 1♀, Kathmandu (1,350 m), Nepal, VII.1983, M.G. Allen leg. (LT); 1♀, Kathmandu (1,400 m), Nepal, IX.1983, M.G. Allen leg. (LT); 1♂, Kathmandu (1,500 m), Nepal, III.1983, M.G. Allen leg. (LT); 1♀, Sec. Vegetation (1,390 m), B. Embassy, Kathmandu, Nepal, VI.1983, M.G. Allen leg. (LT); 2♀♀, Dolalghat (900 m), Nepal, 7.VII.1983, M.G. Allen leg.; 1♀, Chautara (6,000′), Nepal, 24.IX.1983, M.G. Allen leg. (all NHMUK).

Eastern Palaearctic and Oriental regions (Yu et al. 2016). Gauld and Mitchell (1981) recorded this species from Nepal.

Head (Fig. 6B–D): GOI = 2.5–3.2; lower face 0.6–0.8× as wide as high; clypeus almost flat in profile, its lower margin acute; mandible moderately twisted by 30–40°, moderately long, evenly strongly tapered, its outer surface without a diagonal structure; upper mandibular tooth 1.2–1.3× as long as lower one; posterior ocellus almost touching eye; antenna with 53–64 flagellomeres and 20^th flagellomere 1.7–2.1× as long as wide.

Mesosoma (Fig. 6E): mesopleuron densely punctate to closely longitudinally punctostriate; scutellum with lateral longitudinal carinae reaching posterior end and convergent posteriorly; metapleuron densely punctate; propodeum declivous, its posterior area moderately reticulate, outer margin of propodeal spiracle not joining pleural carina by a ridge.

Wings (Fig. 6F): fore wing with AI = 0.6–0.7, CI = 0.3–0.5, ICI = 0.7–0.8, SDI = 1.2–1.4; fore wing vein 1m-cu&M evenly curved, 2r&RS almost straight; fenestra and sclerites of discosubmarginal cell of fore wing as in Figure 6F; fenestra of fore wing not long and its anterodistal corner distinctly separated from proximal end of vein RS; proximal sclerite linear, separated from or vestigially confluent with distal one, weakly pigmented; central sclerite absent; distal sclerite present proximally, vestigial to moderately strong distally; proximal corner of marginal cell of fore wing uniformly setose; vein 1cu-a antefurcal to M&RS by 0.2–0.3× 1cu-a length.

Colour (Fig. 6): body including interocellar area entirely testaceous; wings hyaline to slightly infuscate.

Enicospilus biharensis is similar to E. maruyamanus, E. nikami sp. nov., E. pudibundae, and E. transversus, but can be distinguished from E. maruyamanus, E. nikami sp. nov., and E. transversus by the evenly curved fore wing vein 1m-cu&M (Fig. 6F) (more or less sinuous in E. maruyamanus, E. nikami sp. nov., and E. transversus, e.g. as in Figure 19F), and from E. nikami sp. nov. and E. pudibundae by the proximally complete pectination of the hind tarsal claw (Fig. 2I) (proximally incomplete in E. nikami sp. nov. and E. pudibundae).

Figure 6. 

Enicospilus biharensis Townes, Townes & Gupta, 1961, ♀. A. Habitus; B. Head, frontal view; C. Head, lateral view; D. Head, dorsal view; E. Mesosoma, lateral view; F. Central part of fore wing.

66♀♀43♂♂ and 3 unsexed: Nepal (1♀), India (57♀♀41♂♂), Japan (1♂), Kenya (2♀♀1♂ and 1 unsexed), Madagascar (1♀ and 1 unsexed), Malaysia (1♀), Saudi Arabia (1 unsexed), South Africa (1♀), Uganda (2♀♀), Zimbabwe (1♀).

Type series: holotype of Henicospilus yanagiharai Sonan, 1940, ♂, Kitadaitô-jima, Okinawa Pref., Ryûkyûs, Japan, 18.III.1939, M. Yanagihara leg. (TARI); holotype of Enicospilus fossatus Chiu, 1954, ♀, Jahore, Malaysia, 1.X.1916, J. Sonan leg. (TARI); holotype of Henicospilus euxoae Wilkinson, 1928, ♀, Salisbury, Zimbabwe, 31.XII.1927, J.I. Roberts leg. (from Euxoa) (NHMUK, Type 3b.1289).

Non-type series: 1♀, Chitwan (200 m), Terai, Nepal, 12–13.III.1983. M.G. Allen leg. (Fig. 7); 1♀5♂♂, Coimbatore, India, III–IV.1935, P.S. Nathan leg.; 55♀♀36♂♂, Andhra Pradesh, Patancheru, India, I (26♀♀19♂♂), II (1♂), VII (1♀), IX (1♀), X (2♀♀), XI (7♀♀4♂♂), XII (18♀♀12♂♂).1980, Bhatnagar leg. (LT) (all NHMUK); 1♀, Agra, India, IX.1955, V.K. Gupta leg. (EMUS); 1 unsexed, Asir, Suda, Saudi Arabia, 5.VII.1962, G. Popoy leg.; 1 unsexed, Bekily, Madagascar, VIII.1933, A. Seyrig leg.; 1♀, Madagascar, XII.1920, A. Seyrig leg.; 2♀♀, Ruwenzori Range, Ibanda, Uganda, 20–21.VIII (1♀), 4–12.IX (1♀).1952, D.S. Fletcher leg.; 1♂, Nairobi, Kenya, 10–12.XII.1952, C.G.M. de Worms leg.; 1♀, Kenya; 1♀ and 1 unsexed, Kabete, Kenya, III.1929, H.E. Box leg. (all NHMUK); 1♀, Grahamstown, South Africa, 10–12.III.1971, F. Gess leg. (CNC).

Afrotropical, Australasian, Oceanic, and Oriental regions (Yu et al. 2016). Newly recorded from Nepal.

Head (Fig. 7B–D): GOI = 1.5–2.0; lower face 0.8–1.0× as wide as high; clypeus rather strongly convex in profile, its lower margin impressed; mandible weakly twisted by 10–20°, long, proximally tapered and distally parallel sided, its outer surface with a diagonal setose groove between its dorsoproximal corner and base of mandibular apical teeth; upper mandibular tooth 2.5–3.0× as long as lower one; posterior ocellus separated from eye by 0.1–0.2× its own maximum diameter; antenna with 44–66 flagellomeres and 20^th flagellomere 1.6–2.0× as long as wide.

Mesosoma (Fig. 7E): mesopleuron densely punctate, submatt to matt; scutellum with lateral longitudinal carinae reaching posterior end and convergent posteriorly; metapleuron densely punctate, submatt to matt; propodeum declivous, its posterior area moderately reticulate, outer margin of propodeal spiracle not joining pleural carina by a ridge.

Wings (Fig. 7F): fore wing with AI = 0.4–0.8, CI = 0.3–0.6, ICI = 0.4–0.6, SDI = 1.3–1.5; fore wing vein 1m-cu&M slightly sinuous, 2r&RS almost straight; fenestra and sclerites of discosubmarginal cell of fore wing as in Figure 7F; fenestra of fore wing not long and its anterodistal corner distinctly separated from proximal end of vein RS; proximal sclerite triangular, not confluent with distal one, strongly pigmented; central sclerite rather weakly to strongly pigmented and sclerotised, and ill-delineated oval to semicircular, positioned in antero- to mediodistal part of the fenestra; distal sclerite absent proximally and strong distally; proximal corner of marginal cell of fore wing approximately uniformly setose; vein 1cu-a subinterstitial to antefurcal to M&RS by less than 0.3× 1cu-a length.

Colour (Fig. 7): body including interocellar area entirely yellow- to red-brown; wings hyaline.

Enicospilus capensis is most similar to E. insularis and distinguished from it by the not clearly delineated central sclerite (Fig. 7F) (well delineated in E. insularis), but diagnostic characters for these species are not strongly supported and need more study. Enicospilus capensis also more or less resembles E. ramidulus, but distinguished from it by the densely punctate and submatt to matt meso- and metapleurae (Fig. 7E) (meso- and metapleurae moderately punctate and never submatt to matt in E. ramidulus).

Figure 7. 

Enicospilus capensis (Thunberg, 1822), ♀. A. Habitus; B. Head, frontal view; C. Head, lateral view; D. Head, dorsal view; E. Mesosoma, lateral view; F. Central part of fore wing.

5♀♀ and 1 unsexed: Brunei (3♀♀), Indonesia (1♀), Myanmar (1♀), Sri Lanka (1 unsexed); no Nepalese specimens were examined.

Type series: holotype of Enicospilus xanthocephalus Cameron, 1907 (= Henicospilus flavicaput Morley, 1912), ♀, Haundraw Valley, Tenasserim, Myanmar, VIII.1894, C.T. Bingham leg. (NHMUK, Type 3b.1233).

Non-type series: 1♀, U. Temburong (1,500 m), Bukit Retak, Brunei, IV.1981, I.D. Gauld leg. (Fig. 8); 1♀, Montane forest (1,618 m), Bukit Retak, Brunei, V.1979, I.D. Gauld leg.; 1♀, Pagon Ridge, Pagon, Brunei, II.1982, G. Allen leg.; 1♀, Perliawatte (1,200–1,500 m), Mt Gede, West Java, Indonesia, I.1938; 1 unsexed, near Mahiyangana, Badulla Dist., Sri Lanka, 24.V.1974, Gans & Prasanna leg. (all NHMUK).

Australasian and Oriental regions (Yu et al. 2016). Gauld and Mitchell (1981) recorded this species from Nepal.

Head (Fig. 8B–D): GOI = 2.9–3.1; lower face 0.6–0.7× as wide as high; clypeus weakly convex in profile, its lower margin subacute; mandible moderately twisted by 30–40°, moderately long, evenly tapered, its outer surface without a diagonal structure; upper mandibular tooth 1.3–1.6× as long as lower one; posterior ocellus close to eye; antenna with 71–76 flagellomeres and 20^th flagellomere 2.3–2.5× as long as wide.

Mesosoma (Fig. 8E): mesopleuron rather coarsely longitudinally striate; scutellum with lateral longitudinal carinae reaching anterior 0.8–1.0 and convergent posteriorly; metapleuron rather coarsely striate to strigose; propodeum evenly rounded to slightly declivous, its posterior area coarsely reticulate, outer margin of propodeal spiracle joining pleural carina by a strong ridge.

Wings (Fig. 8F): fore wing with AI = 0.3–0.4, CI = 0.2–0.4, ICI = 0.6–0.7, SDI = 1.2–1.4; fore wing vein 1m-cu&M weakly sinuous, 2r&RS almost straight; fenestra and sclerites of discosubmarginal cell of fore wing as in Figure 8F; fenestra of fore wing not long and its anterodistal corner distinctly separated from proximal end of vein RS; proximal sclerite triangular, confluent with distal one, strongly pigmented; central sclerite strongly pigmented and sclerotised, linear and parallel to vein 2r&RS, positioned in anterodistal part of fenestra; distal sclerite present proximally and vestigial to absent distally; proximal corner of marginal cell of fore wing uniformly setose; vein 1cu-a antefurcal to M&RS by 0.1–0.2× 1cu-a length.

Colour (Fig. 8): body including interocellar area entirely testaceous; wings hyaline to weakly infuscate.

Enicospilus flavicaput is most similar to E. kanshirensis but can be distinguished from it by the slender central sclerite (Fig. 8F) (central sclerite stouter in E. kanshirensis, as in Figure 14F), and larger body size (i.e. fore wing length more than 17.0 mm in E. flavicaput but less than 15.0 mm in E. kanshirensis).

Figure 8. 

Enicospilus flavicaput (Morley, 1912), ♀. A. Habitus; B. Head, frontal view; C. Head, lateral view; D. Head, dorsal view; E. Mesosoma, lateral view; F. Central part of fore wing.

13♀♀11♂♂ and 1 unsexed: Nepal (2♀♀1♂), Australia (5♀♀2♂♂ and 1 unsexed), Brunei (2♀♀1♂), Japan (1♂), Singapore (1♀), Taiwan (3♀♀6♂♂).

Type series: lectotype of Ophion flavocephalus Kirby, 1900, ♂, Flying Fish Cove, Christmas Island, Australia, C.W. Andrews leg. (NHMUK, Type 3b.1273); holotype of Henicospilus albicaput Morley, 1912, ♂, Mackay, Queensland, Australia (NHMUK, Type 3b.1254); holotype of Henicospilus similis Matsumura & Uchida, 1926, ♂, Okinawa, Ryûkyûs, Japan, S. Sakaguchi leg. (SEHU).

Non-type series: 1♀, Kathmandu (1,350 m), Nepal, VII.1983, M.G. Allen leg. (LT) (Fig. 9); 1♀, Kathmandu (1,300 m), Nepal, XI.1982, M.G. Allen leg. (LT); 1♂, Pokhara, Nepal, VIII.1982, M.G. Allen leg. (LT); 5♀♀, Christmas Island, Australia, 1939; 1 unsexed, Christmas Island, Australia, 1898, C.W. Andrews leg.; 1♂, Ulu Temburong (300 m), Base camp hut, Brunei, 16.II–9.III.1982, M.C. Day leg.; 1♀, Pagon, Pagon Ridge, Brunei, II.1982, M.G. Allen leg.; 1♀, Bukit Retak (1,618 m), Montane forest, Brunei, IX.1979, I.D. Gauld leg.; 1♀, Singapore, 1908, H.N. Ridley leg. (all NHMUK); 1♀, Wanfeng Hill, Taichung, Taiwan, VII.1984, K.S. Lin & K.C. Chou leg. (MsT); 1♂, Kukuan (730 m), Taichung, Taiwan, 14–17.X.1980, K.S. Lin & C.H. Wang leg,; 1♀1♂, Pingtung, Taiwan, IV.1961, K.S. Lin leg. (LT); 1♀1♂, Silo, Yunlin, Taiwan, V.1961, K.S. Lin leg. (LT); 1♂, Lishan, Taichung, Taiwan, 14.IX.1978; 2♂♂, Chung-ying, Taiwan, III.1961, S.C. Chiu leg. (all TARI).

Australasian, Oceanic, and Oriental regions (Yu et al. 2016). Newly recorded from Nepal.

Head (Fig. 9B–D): GOI = 2.5–2.9; lower face 0.5–0.7× as wide as high; clypeus very slightly convex in profile, its lower margin subacute to blunt; mandible moderately twisted by 25–35°, moderately long, more or less evenly tapered, its outer surface without a diagonal structure; upper mandibular tooth 1.1–1.2× as long as lower one; posterior ocellus almost touching eye; antenna with 45–51 flagellomeres and 20^th flagellomere 1.8–2.3× as long as wide.

Mesosoma (Fig. 9E): mesopleuron punctate to longitudinally punctostriate; scutellum with lateral longitudinal carinae reaching posterior end and convergent posteriorly; metapleuron moderately strigose to striate; propodeum evenly rounded, its posterior area rather finely reticulate, outer margin of propodeal spiracle joining pleural carina by a ridge.

Wings (Fig. 9F): fore wing with AI = 0.4–1.5, CI = 0.6–0.8, ICI = 0.4–0.6, SDI = 1.1–1.2; fore wing vein 1m-cu&M centrally strongly angulated and broadened, 2r&RS almost straight; fenestra and sclerites of discosubmarginal cell of fore wing as in Figure 9F; fenestra of fore wing not very long and its anterodistal corner distinctly separated from proximal end of vein RS; proximal sclerite almost oval, isolated and not touching margin of fenestra, strongly pigmented; central sclerite strongly pigmented and sclerotised, linear and parallel to distal margin of the fenestra, positioned in mediodistal part of the fenestra; distal sclerite absent; proximal corner of marginal cell of fore wing almost uniformly setose; vein 1cu-a subinterstitial to antefurcal to M&RS by less than 0.2× 1cu-a length.

Colour (Fig. 9): body including interocellar area entirely pale yellow with pale brown posterior segments of metasoma; wings hyaline.

Enicospilus flavocephalus is a very distinctive species, but its body size, colour pattern, and profile are very similar to E. xanthocephalus. Enicospilus flavocephalus is easily distinguished from E. xanthocephalus by many characters, such as the pale yellow interocellar area (Fig. 9B, D) (black in E. xanthocephalus) and centrally abruptly angled and broadened fore wing vein 1m-cu&M (Fig. 9F) (evenly curved in E. xanthocephalus).

Figure 9. 

Enicospilus flavocephalus (Kirby, 1900), ♀. A. Habitus; B. Head, frontal view; C. Head, lateral view; D. Head, dorsal view; E. Mesosoma, lateral view; F. Central part of fore wing.

2♀♀2♂♂ and 1 unsexed: Nepal (1♂), Brunei (1♂), India (1 unsexed), Japan (1♀), Taiwan (1♀).

Type series: holotype of Henicospilus formosensis Uchida, 1928, ♀, Baibara, Taiwan, 15.VI.1926, Y. Saito & Kikuchi leg. (SEHU); holotype of Enicospilus saepis Chiu, 1954, ♀, Nara, Honshû, Japan, 17.VIII.1918, J. Sonan leg. (TARI).

Non-type series: 1♂, mixed forest (1,550 m), Godaveri, Nepal, 6.V.1984, M.G. Allen leg. (LT) (Figs 2E, 10); 1♂, Ulu Temburong (1,000 m), Brunei, II.1980, M.G. Allen leg.; 1 unsexed, NW Himalaya, Dalhousie, India, 8.VII.1965, Tikar leg. (all NHMUK).

Eastern Palaearctic and Oriental regions (Yu et al. 2016). Newly recorded from Nepal.

Head (Fig. 10B–D): GOI = 2.2–2.4; lower face 0.8–0.9× as wide as high; clypeus moderately convex in profile, its lower margin subacute to blunt; mandible weakly twisted by 10–20°, moderately long, evenly tapered, its outer surface without a diagonal structure; upper mandibular tooth 1.2–1.4× as long as lower one; posterior ocellus close to eye; antenna with 66–69 flagellomeres and 20^th flagellomere 2.1–2.2× as long as wide.

Mesosoma (Fig. 10E): mesopleuron moderately punctate; scutellum with lateral longitudinal carinae reaching anterior 0.8 or more and weakly convergent posteriorly so that subquadrate (Fig. 2E); metapleuron punctate with isolated striae; propodeum declivous in profile, its posterior area coarsely irregularly wrinkled, sometimes with posterior transverse carina laterally, outer margin of propodeal spiracle joining pleural carina by a ridge or not.

Wings (Fig. 10F): fore wing with AI = 0.2–0.6, CI = 0.2–0.9, ICI = 0.5–0.6, SDI = 1.1–1.3; fore wing vein 1m-cu&M weakly sinuous, 2r&RS almost straight; fenestra and sclerites of discosubmarginal cell of fore wing as in Figure 10F; fenestra of fore wing not very long and its anterodistal corner distinctly separated from proximal end of vein RS; proximal sclerite triangular, confluent with distal one, strongly pigmented; central sclerite moderately to strongly pigmented and sclerotised, linear and parallel to distal margin of fenestra, positioned in distal part of fenestra; distal sclerite present proximally and vestigial to absent distally; proximal corner of marginal cell of fore wing uniformly setose; vein 1cu-a subinterstitial to antefurcal to M&RS by less than 0.2× 1cu-a length.

Colour (Fig. 10): body including interocellar area testaceous; wings weakly infumate.

Enicospilus formosensis is a distinctive species and can easily be distinguished by many characters, such as the wide face (Fig. 10B), shape of the central sclerite (Fig. 10F), more or less subquadrate scutellum (Fig. 2E), as listed in the diagnosis.

Figure 10. 

Enicospilus formosensis (Uchida, 1928), ♂. A. Habitus; B. Head, frontal view; C. Head, lateral view; D. Head, dorsal view; E. Mesosoma, lateral view; F. Central part of fore wing.

14♀♀3♂♂: Nepal (1♀), Indonesia (1♂), Brunei (13♀♀2♂♂).

Type series: holotype of Dicamptus grammospilus Enderlein, 1921, ♂, Soekaranda, Sumatra, Indonesia, Dohrn leg. (IZPAN) [photos examined].

Non-type series: 1♀, Pokhara (950 m), Nepal, VII–VIII.1983, M.G. Allen leg. (LT) (Fig. 11); 2♀♀1♂, Montane Forest (1,618 m), Bukit Retak, Brunei, IX.1979, I.D. Gauld leg.; 8♀♀1♂, Bukit Retak (1,500 m), U. Temburgon, Brunei, IV.1981, I.D. Gauld leg.; 2♀♀, Pagon (1,700 m), U. Temburong, Brunei, IV.1981, I.D. Gauld leg.; 1♀, Pagon Ridge, Pagon, Brunei, II.1982, M.G. Allen leg. (all NHMUK).

Oriental region (Yu et al. 2016). Newly recorded from Nepal.

Head (Fig. 11B–D): GOI = 2.5–2.7; lower face 0.7–0.8× as wide as high; clypeus almost flat in profile, its lower margin acute; mandible moderately twisted by 20–30°, moderately long, proximally tapered and distally almost subparallel sided, its outer surface without a diagonal structure; upper mandibular tooth 1.4–1.5× as long as lower one; posterior ocellus (almost) touching eye; antenna with 58–62 flagellomeres and 20^th flagellomere 1.7–1.9× as long as wide.

Mesosoma (Fig. 11E): mesopleuron punctate dorsally and rather closely longitudinally punctostriate to striate ventrally; scutellum with lateral longitudinal carinae reaching anterior 0.8 or more and convergent posteriorly; metapleuron rather closely striate; propodeum declivous in profile, its posterior area concentrically striate, outer margin of propodeal spiracle not joining pleural carina by a ridge.

Wings (Fig. 11F): fore wing with AI = 0.8–1.4, CI = 0.5–0.6, ICI = 0.4–0.5, SDI = 1.4–1.5; fore wing vein 1m-cu&M almost evenly curved, 2r&RS weakly bowed centrally; fenestra and sclerites of discosubmarginal cell of fore wing as in Figure 11F; fenestra of fore wing not very long and its anterodistal corner distinctly separated from proximal end of vein RS; proximal sclerite not triangular, confluent with distal one, weakly to strongly pigmented; central sclerite weakly to strongly pigmented and sclerotised, linear and parallel to vein 2r&RS, positioned in central part of fenestra; distal sclerite weak; proximal corner of marginal cell of fore wing uniformly setose; vein 1cu-a subinterstitial to antefurcal to M&RS by less than 0.1× 1cu-a length.

Colour (Fig. 11): body including interocellar area entirely testaceous; wings hyaline.

Enicospilus grammospilus is a very distinctive species on account of its characteristic shape of fore wing vein 2r&RS and sclerites as in Figure 11F. No similar species are recognised and is very easily distinguished from all other Enicospilus species by the characters summarised in the above diagnosis, such as concentrically striate posterior area of propodeum and characteristic shape of sclerites of discosubmarginal cell of fore wing (cf. Fig. 11F).

Figure 11. 

Enicospilus grammospilus (Enderlein, 1921), ♀. A. Habitus; B. Head, frontal view; C. Head, lateral view; D. Head, dorsal view; E. Mesosoma, lateral view; F. Central part of fore wing.

44♀♀4♂♂: Nepal (5♀♀2♂♂), Brunei (30♀♀2♂♂) India (2♀♀), Papua New Guinea (4♀♀), Singapore (1♀), Sri Lanka (2♀♀).

Non-type series: 2♀♀1♂, Kakani (2,000 m), Nepal, VIII.1982, M.G. Allen leg. (LT); 3♀♀, Kathmandu (1,350 m), Nepal, VII.1983, M.G. Allen leg. (LT) (Fig. 12); 1♂, Pokhara, Nepal, VIII.1982, M.G. Allen leg. (LT); 2♀♀1♂, Gn. Pagon (1,700 m), U. Temburong, Brunei, IV.1981, I.D. Gauld leg.; 24♀♀, Bukit Retak (1,500 m), U. Temburong, Brunei, IV.1981, I.D. Gauld leg.; 1♂, Montane forest (1,618 m), Bukit Retak, Brunei, IX.1979, I.D. Gauld leg.; 2♀♀, Pagon Ridge, Pagon, Brunei, II.1982, I.D. Gauld leg.; 2♀♀, 1′ forest (500 m), U. Temburong, Brunei, IV.1981, I.D. Gauld leg.; 1♀, Thekkadi, Periyar Dam, Travancore, India, 6–10.V.1937; 1♀, Andhra Pradesh, Patancheru, India, XII.1980, Bhatnagar leg. (LT); 1♀, Wau (1,200 m), Morobe Dist., Papua New Guinea, X.1979, I.D. Gauld leg.; 3♀♀, Mt Lawes (400 m), Port Moresby, Papua New Guinea, 5.III–12.V.1963, W.W. Brandt leg.; 1♀, Singapore, 1901, H.N. Kidley leg.; 1♀, Peradeniya, Sri Lanka, 25.VII.1919, N.K. Jardine leg.; 1♀, Matale (1,500 m), Sri Lanka, 10.V.1919, N.K. Jardine leg. (all NHMUK).

Australasian, Eastern Palaearctic, and Oriental regions (Yu et al. 2016). Gauld and Mitchell (1981) recorded this species from Nepal.

Head (Fig. 12B–D): GOI = 2.7–3.2; lower face 0.7–0.8× as wide as high; clypeus moderately convex in profile, its lower margin blunt; mandible moderately twisted by 15–30°, moderately long, evenly tapered, its outer surface without a diagonal structure; upper mandibular tooth 1.2–1.5× as long as lower one; posterior ocellus almost touching eye; antenna with 55–62 flagellomeres and 20^th flagellomere 2.0–2.5× as long as wide.

Mesosoma (Fig. 12E): mesopleuron dorsally punctate to longitudinally striate and ventrally longitudinally striate; scutellum with lateral longitudinal carinae reaching posterior end and convergent posteriorly; metapleuron striate; propodeum declivous, its posterior area moderately reticulate, outer margin of propodeal spiracle joining pleural carina by a ridge.

Wings (Fig. 12F): fore wing with AI = 1.1–1.8, CI = 0.2–0.5, ICI = 0.4–0.6, SDI = 1.0–1.1; fore wing vein 1m-cu&M evenly curved, 2r&RS almost straight; fenestra and sclerites of discosubmarginal cell of fore wing as in Figure 12F; fenestra of fore wing not very long and its anterodistal corner distinctly separated from proximal end of vein RS; proximal sclerite not triangular, confluent with distal one, at least anteriorly very strongly pigmented; central sclerite absent; distal sclerite more or less strong and rather thick; proximal corner of marginal cell of fore wing more or less uniformly setose; vein 1cu-a subinterstitial to antefurcal to M&RS by less than 0.1× 1cu-a length.

Colour (Fig. 12): body including interocellar area entirely testaceous; wings hyaline to slightly infumate.

Enicospilus javanus is distinctive and one of the most easily distinguishable species on account of the proximally extended fore wing fenestra and the shape of the sclerites (cf. Fig. 12F).

Figure 12. 

Enicospilus javanus (Szépligeti, 1910), ♀. A. Habitus; B. Head, frontal view; C. Head, lateral view; D. Head, dorsal view; E. Mesosoma, lateral view; F. Central part of fore wing.

The specific name is derived from the type locality, Kakani, Nepal.

1♂: Nepal.

Type series: holotype ♂, Kakani (2,000 m), Nepal, VIII.1982, M.G. Allen leg. (LT) (NHMUK) (Figs 2F, 13).

Nepal.

Male (Holotype) (Fig. 13). Body length ca 24.0 mm.

Head with GOI = 2.8 (Fig. 13C). Lower face 0.7× as wide as high, moderately punctate with setae, strongly shiny (Fig. 13B). Clypeus 1.5× as wide as high, moderately punctate with setae, moderately convex in profile, and its lower margin impressed (Fig. 13B, C). Malar space 0.4× as long as basal mandibular width (Fig. 13B, C). Mandible moderately twisted by ca 30°, moderately long, more or less evenly tapered, its outer surface with a diagonal setose groove between its dorsoproximal corner to base of mandibular apical teeth (Fig. 13B, C). Upper mandibular tooth 2.0× as long as lower one (Fig. 13B). Frons, vertex and gena strongly shiny with fine setae (Fig. 13B–D). Posterior ocellus close to eye (Fig. 13B–D). Ventral end of occipital carina joining oral carina. Antenna with 73 flagellomeres; first flagellomere 1.7× as long as second; 20^th flagellomere 2.8× as long as wide.

Mesosoma entirely moderately to strongly shiny with setae (Fig. 13E). Pronotum punctate with wrinkles centrally. Mesoscutum 1.4× as long as its maximum width, finely punctate with setae, and evenly rounded in lateral profile (Fig. 13E). Notauli absent (Fig. 13E). Scutellum moderately convex, finely punctate with setae, with lateral longitudinal carinae reaching anterior 0.6 (Fig. 2F). Epicnemium densely punctate. Epicnemial carina strongly present, evenly and moderately curved to anterior, its dorsal end reaching anterior margin of mesopleuron (Fig. 13E). Mesopleuron entirely punctate with longitudinal wrinkles or striae (Fig. 13E). Submetapleural carina almost parallel sided (Fig. 13E). Metapleuron entirely finely punctate with setae (Fig. 13E). Propodeum weakly evenly rounded in profile; anterior transverse carina complete; anterior area longitudinally striate; spiracular area finely punctate with setae; posterior area moderately longitudinally strigose centrally and irregularly wrinkled to reticulate laterally; propodeal spiracle elliptical, its outer margin not joining pleural carina by a ridge (Fig. 13E).

Wings. Fore wing length ca 17.0 mm with AI = 0.4, CI = 0.4, DI = 0.3, ICI = 0.5, SDI = 1.4, SI = 0.1, SRI = 0.3; vein 1m-cu&M almost evenly curved; vein 2r&RS straight and RS evenly curved; fenestra and sclerites of discosubmarginal cell as in Figure 13F; proximal sclerite triangular, confluent with distal sclerite, strongly pigmented; central sclerite absent; distal sclerite more or less entirely present from base to apex, weakly to moderately pigmented; proximal corner of marginal cell evenly setose; posterodistal corner of second discal cell ca 95°; posterodistal corner of subbasal cell ca 95°; vein 1cu-a antefurcal to M&RS by 0.2× 1cu-a length (Fig. 13F). Hind wing with NI = 1.8, RI = 1.8; vein RS straight; vein RA with six uniform hamuli.

Legs. Outer surface of fore tibia with scattered spines. Hind leg with coxa in profile 1.9× as long as deep; basitarsus 2.0× as long as second tarsomere; fourth tarsomere 0.6× as long as third tarsomere and 4.2× as long as wide; tarsal claw simply pectinate.

Metasoma with PI = 3.8, DMI = 1.2, THI = 3.6; dorsal margin of tergite 1 not sinuous; thyridium elongate (Fig. 13A).

Colour (Fig. 13). Entirely testaceous except for apex of mandible black. Wings hyaline; sclerites of fenestra and veins testaceous.

Variation. Unknown.

Female. Unknown.

Enicospilus kakanicus sp. nov. is similar to and can be confused with E. longitarsis Tang, 1990, E. tangi sp. nov., and E. yonezawanus (Uchida, 1928). These species all belong to the E. ramidulus complex and share the following characters: outer surface of mandible with a diagonal setose deep groove between its dorsoproximal corner and base of mandibular apical teeth (e.g. Fig. 2B, D), fore wing fenestra without central sclerite (e.g. Figs 13F, 25F, 27F), and proximal sclerite triangular (e.g. Figs 13F, 25F, 27F). Enicospilus kakanicus sp. nov. is distinguished from the above species by the rather short lateral longitudinal carinae of the scutellum, i.e. reaching the anterior 0.6 of the scutellum in E. kakanicus sp. nov., as in Figure 2F, but almost always reaching the posterior end of the scutellum in E. longitarsis Tang, 1990, E. tangi sp. nov., and E. yonezawanus, as in, e.g., Figure 2H, and also by the characters used in the above key, such as width of lower face, mandibular shape and length, and surface sculptures of metapleuron.

Figure 13. 

Enicospilus kakanicus Shimizu sp. nov., ♂. A. Habitus; B. Head, frontal view; C. Head, lateral view; D. Head, dorsal view; E. Mesosoma, lateral view; F. Central part of fore wing.

3♀♀3♂♂: Nepal (1♂), India (1♀), Indonesia (1♀1♂), Taiwan (1♀1♂).

Type series: holotype of Henicospilus kanshirensis Uchida, 1928, ♂, Kanshirei [= Gauziling], Tainan, Taiwan, 15.IV.1908, S. Matsumura leg. (SEHU).

Non-type series: 1♂, Dharan Sal & 2^y forest (330m), Terai, Nepal, 14–15.XI.1983, M.G. Allen leg. (Fig. 14); 1♀, Anamalai Hills (3,500′), Cinchona, India, V.1957, P.S. Nathan leg.; 1♂, Tjigaeha, Mt Djampang, West Java, Indonesia, I.1938, K.M. Walsh leg.; 1♀, Tengah, Mt Tjioeng, Djampang Mts, West Java, Indonesia, I.1938, K.M. Walsh leg.; 1♀, Sunmoon Lake, Taiwan, 22–29.IX.1970, Shui-Chen Chiu leg. (all NHMUK).

Australasian and Oriental regions (Yu et al. 2016). Gauld and Mitchell (1981) recorded this species from Nepal.

Head (Fig. 14B–D): GOI = 2.8–3.1; lower face 0.7× as wide as high; clypeus moderately convex in profile, its lower margin subacute to blunt; mandible moderately twisted by 20–30°, moderately long, evenly tapered, its outer surface without a diagonal structure; upper mandibular tooth 1.4–1.6× as long as lower one; posterior ocellus close to eye; antenna with 63–66 flagellomeres and 20^th flagellomere 2.1–2.3× as long as wide.

Mesosoma (Fig. 14E): mesopleuron entirely closely to rather coarsely longitudinally striate; scutellum with lateral longitudinal carinae reaching anterior 0.8 or more and convergent posteriorly; metapleuron rather coarsely striate to strigose; propodeum weakly declivous, its posterior area coarsely reticulate to concentrically striate, outer margin of propodeal spiracle joining pleural carina by a ridge.

Wings (Fig. 14F): fore wing with AI = 0.4–0.5, CI = 0.2–0.3, ICI = 0.5–0.7, SDI = 1.1–1.3; fore wing vein 1m-cu&M moderately sinuate, 2r&RS almost straight; fenestra and sclerites of discosubmarginal cell of fore wing as in Figure 14F; fenestra of fore wing not very long and its anterodistal corner distinctly separated from proximal end of vein RS; proximal sclerite almost triangular, confluent with distal one, strongly pigmented; central sclerite strongly pigmented and sclerotised, linear and parallel to vein 2r&RS, positioned in anterodistal part of fenestra; distal sclerite present proximally and vestigial to absent distally; proximal corner of marginal cell of fore wing uniformly setose; vein 1cu-a antefurcal to M&RS by 0.1–0.2× 1cu-a length.

Colour (Fig. 14): body including interocellar area entirely testaceous; wings hyaline.

Enicospilus kanshirensis is most similar to E. flavicaput but can be distinguished from it by the stouter central sclerite (Fig. 14F) (central sclerite slender in E. flavicaput as in Figure 8F), and smaller body size (i.e. fore wing length less than 15.0 mm in E. kanshirensis but more than 17.0 mm in E. flavicaput).

Figure 14. 

Enicospilus kanshirensis (Uchida, 1928), ♂. A. Habitus; B. Head, frontal view; C. Head, lateral view; D. Head, dorsal view; E. Mesosoma, lateral view; F. Central part of fore wing.

29♀♀7♂♂ and 2 unsexed: Nepal (3♀♀4♂♂), India (2♀♀1♂), Taiwan (23♀♀2♂♂ and 2 unsexed), Zambia (1♀).

Type series: holotype of Enicospilus leetoni Chiu, 1954, ♀, Taihoku, Taiwan, 1.IX.1925, J. Sonan leg. (TARI).

Non-type series: 1♂, Gokarna (1,450 m), Nepal, VI.1983, M.G. Allen leg. (Fig. 15); 1♀, Kathmandu (1,350 m), Nepal, VII.1983, M.G. Allen leg. (LT); 1♂, Kathmandu, Nepal, M.G. Allen leg.; 2♀♀, Kakani (2,070 m), Nepal, VII.1983, M.G. Allen leg. (LT); 1♂, Kakani (2,000 m), Nepal, VIII.1982, M.G. Allen leg. (LT); 1♂, Phulchoki (2,500 m), Nepal, IX.1982, M.G. Allen leg. (LT); 2♀♀, Delhi, India, 14.XI.1967 (1♀), 5.III.1968 (1♀); 1♂, U.P. Garjia, India, 22.IV.1967, Gupta leg. (all NHMUK); 1♀1♂ and 1unsexed, Taitung, Taiwan, 31.V–6.VI (1♀), 7–13.VI (1♂), 1–14.XI (1 unsexed).1971 (MsT); 21♀♀1♂ and 1 unsexed, Kuanhsi, Taiwan, 16.VIII (1♂), 19.VIII (1 unsexed), 29.VIII (2♀♀), IX (13♀♀), 10.X (3♀♀), 24–30.XII (1♀).1968, 11–17.III.1969 (1♀), 30.VIII.1970 (1♀) (MsT) (all TARI); 1♀, 15 km E Lusaka, Zambia, 22–31.I.1980, R.A. Beaver leg. (NHMUK).

Afrotropical and Oriental regions (Yu et al. 2016). Gauld and Mitchell (1981) recorded this species from Nepal.

Head (Fig. 15B–D): GOI = 2.9–3.1; lower face 0.7–0.8× as wide as high; clypeus moderately convex in profile, its lower margin acute; mandible weakly twisted by 10–25°, moderately long, evenly tapered, its outer surface with a diagonal setose groove between its dorsoproximal corner and base of mandibular apical teeth; upper mandibular tooth 1.3–1.4× as long as lower one; posterior ocellus almost touching eye; antenna with 56–62 flagellomeres and 20^th flagellomere 2.0–3.0× as long as wide.

Mesosoma (Fig. 15E): mesopleuron punctate to longitudinally punctostriate; scutellum with lateral longitudinal carinae reaching posterior end and convergent posteriorly; metapleuron punctostriate; propodeum weakly declivous in profile, its posterior area moderately reticulate, outer margin of propodeal spiracle joining pleural carina by a ridge.

Wings (Fig. 15F): fore wing with AI = 0.4–0.6, CI = 0.4, ICI = 0.4–0.6, SDI = 1.2–1.4; fore wing vein 1m-cu&M almost evenly curved or very slightly sinuous, 2r&RS almost straight; fenestra and sclerites of discosubmarginal cell of fore wing as in Figure 15F; fenestra of fore wing not very long and its anterodistal corner distinctly separated from proximal end of vein RS; proximal sclerite triangular, separated from distal one, strongly pigmented; central sclerite strongly pigmented, sclerotised, well-delineated D-shaped to semi-circular, positioned in almost mediodistal part of fenestra; distal sclerite absent proximally and more or less strong distally; proximal corner of marginal cell of fore wing uniformly setose; vein 1cu-a antefurcal to M&RS by 0.1–0.3× 1cu-a length.

Colour (Fig. 15): body including interocellar area entirely testaceous; wings hyaline.

Enicospilus laqueatus, E. pseudoantennatus, E. vestigator, and E. tripartitus share similar fenestra, sclerites, and fore wing venation (e.g. Figs 15F, 21F, 26F). However, E. laqueatus can be readily separated from E. pseudoantennatus, E. vestigator, and E. tripartitus by a diagonal setose deep groove of the outer surface of the mandible between its dorsoproximal corner and base of mandibular apical teeth (outer mandibular surface without a distinct diagonal setose deep groove in E. pseudoantennatus, E. vestigator, and E. tripartitus, as in e.g. Figure 2C).

Figure 15. 

Enicospilus laqueatus (Enderlein, 1921), ♂. A. Habitus; B. Head, frontal view; C. Head, lateral view; D. Head, dorsal view; E. Mesosoma, lateral view; F. Central part of fore wing.

88♀♀15♂♂ and 3 unsexed: Nepal (10♀♀4♂♂), Australia (1♀), Brunei (2♀♀), India (34♀♀7♂♂ and 1 unsexed), Japan (17♀♀), Papua New Guinea (2♀♀), Sri Lanka (1♀), Taiwan (21♀♀4♂♂ and 2 unsexed).

Type series: holotype of Enicospilus uniformis Chiu, 1954, ♀, Taihoku, Taiwan, 14.IV.1921, S. Aoki leg. (TARI); holotype of Enicospilus flatus Chiu, 1954, ♀, Taihoku, Taiwan, 28.V.1931, J. Sonan leg. (TARI); lectotype of Enicospilus unicornis Rao & Nikam, 1969, ♂, Himayatbagh, Aurangabad, Maharashtra, India, VIII.1968, Nikam leg. (NHMUK, Type 3b.2858).

Non-type series: 1♂, Kathmandu (4,300′), Nepal, VIII.1981, M.G. Allen leg. (Fig. 16A–E); 1♀, Kathmandu (4,300′), Nepal, VIII.1982, M.G. Allen leg.; 1♀2♂♂, Phulchoki (2,000 m), Nepal, VIII.1982, M.G. Allen leg. (LT) (Fig. 16F by ♂); 2♀♀, Godavarl (6,000′), Kathmandu, Nepal, 1–2 (1♀), 3 (1♀).VIII.1967 (MsT); 1♀, Godavarl (5,000′), Kathmandu, Nepal, 10.VIII.1967 (MsT); 1♀, near Simra (180 m), Adhabhar, Nepal, 23–28.VIII.1967 (MsT); 1♀, Kakani (2,070 m), Nepal, VII.1983, M.G. Allen leg. (LT); 2♀♀, Kakani (2,000 m), Nepal, VIII.1982, M.G. Allen leg. (LT); 1♀1♂, Kathmandu (1,350 m), Nepal, VII.1983, M.G. Allen leg. (LT); 1♀, Victoria, Toolangi, Australia, I–II.1983, Farrugia & Gauld leg.; 2♀♀, U. Temburong (1,500 m), Bukit Retak, Brunei, IV.1881, I.D. Gauld leg.; 34♀♀6♂♂ and 1 unsexed, Andhra Pradesh, Patancheru, India, VI (1♀), VII (1♀), VIII (4♀♀), IX (27♀♀5♂♂ and 1 unsexed), X (1♀).1980 (1♂), Bhatnagar leg. (LT) (all NHMUK); 17♀♀, Hitsujigaoka (43°00'N, 141°24'E), Sapporo, Hokkaidô, Japan, 16–23.VIII (1♀), 30.VIII–6.IX (1♀).2007, 28.VII–4.VIII (1♀), 4–11 (6♀♀), 11–18 (4♀♀).VIII, 1–8.IX (4♀♀).2008, K. Konishi leg. (MsT) (EUM); 1♀, Kokoda (365 m), Papua New Guinea, VI.1933, L.E. Cheesman leg.; 1♀, Wau (1,200 m), Morobe District, Papua New Guinea, 24–26.II.1963, J. Sedlacek leg. (MsT); 1♀, Peak View Motel (550 m), Kandy, Sri Lanka, 15–24.I.1970, Davis & Rowe leg. (all NHMUK); 3♀♀3♂♂ and 2 unsexed, Karen, Taiwan, 6–14.V (1♀1♂ and 1 unsexed), 26.VIII–4.XI (1♀2♂♂).1972, 16–22.IV.1973 (1♀ and 1 unsexed) (MsT); 16♀♀1♂, Wanfeng Hill, Taichung, Taiwan, I (2♀♀1♂), II (1♀), IV (10♀♀), V (3♀♀).1984, K.S. Lin & K.C. Chou leg. (MsT) (all TARI).

Australasian, Eastern Palaearctic, Oceanic, and Oriental regions (Yu et al. 2016). Gauld and Mitchell (1981) recorded this species from Nepal.

Head (Fig. 16B–D): GOI = 2.2–2.7; lower face 0.7–0.8× as wide as high; clypeus almost flat in profile, its lower margin acute to subacute; mandible rather weakly twisted by 10–20°, moderately long, proximally tapered and distally more or less parallel sided, its outer surface without a diagonal structure; upper mandibular tooth 1.3–1.5× as long as lower one; posterior ocellus almost touching eye; antenna with 51–61 flagellomeres and 20^th flagellomere 1.9–2.2× as long as wide.

Mesosoma (Fig. 16E): mesopleuron punctate; scutellum with lateral longitudinal carinae reaching at least anterior 0.8 and convergent posteriorly; metapleuron punctate to punctostrigose; propodeum weakly declivous, its posterior area moderately reticulate, outer margin of propodeal spiracle not joining pleural carina by a ridge.

Wings (Fig. 16F): fore wing with AI = 0.5–0.9, CI = 0.5–0.9, ICI = 0.7–1.0, SDI = 1.3–1.5; fore wing vein 1m-cu&M moderately sinuous, 2r&RS almost straight; fenestra and sclerites of discosubmarginal cell of fore wing as in Figure 16F; fenestra of fore wing not long and its anterodistal corner distinctly separated from proximal end of vein RS; proximal and central sclerites absent; distal sclerite strong and more or less centrally broadened; proximal corner of marginal cell of fore wing uniformly setose; vein 1cu-a interstitial to antefurcal to M&RS by less than 0.3× 1cu-a length.

Colour (Fig. 16): body including interocellar area entirely testaceous; wings hyaline.

Some species of Oriental Enicospilus (e.g. E. fusiformis and E. unicolor) have a centrally broadened distal sclerite and lack proximal and central sclerites, as in Figure 16F. Among them, E. lineolatus is most similar to E. unicolor, but distinguished by the narrower distal sclerite than that of E. unicolor and testaceous fore wing pterostigma and sclerite (brown in E. unicolor).

Figure 16. 

Enicospilus lineolatus (Roman, 1913), ♂. A. Habitus; B. Head, frontal view; C. Head, lateral view; D. Head, dorsal view; E. Mesosoma, lateral view; F. Central part of fore wing.

105♀♀21♂♂ and 6 unsexed: Nepal (5♀♀2♂♂), Australia (1♀), China (1♀), Maldives (1♂), India (26♀♀), Indonesia (4♀♀2♂♂ and 1 unsexed), Japan (2♀♀), Malaysia (1♀), Papua New Guinea (7♀♀1♂), Philippines (7♀♀), Singapore (1 unsexed), Sri Lanka (8♀♀), Taiwan (43♀♀15♂♂ and 4 unsexed).

Type series: holotype of Enicospilus reticulatus Cameron, 1902, ♂, Hulule, Maldive Islands, 20.VI.1900 (NHMUK, Type 3b.1268); holotype of Eniscospilus (sic) melanocarpus Cameron, 1905, ♀, Sri Lanka (NHMUK, Type 3b.1234); lectotype of Henicospilus turneri Morley, 1912, ♀, Mackay, Queensland, Australia, 1899, Turner leg. (NHMUK, Type 3b.1261); holotype of Henicospilus atricornis var. zeylanicus Morley, 1913, ♀, Kandy, Sri Lanka, 11.VII.1910, Green leg. (NHMUK, Type 3b.2098); holotype of Enicospilus quintuplex Chiu, 1954, ♀, Shaowu, Fukien, China, 8.X.1945, S.H. Chao leg. (TARI).

Non-type series: 1♂, Godaveri (1,550–1,700 m), Nepal, VI.1983, M.G. Allen leg. (LT) (Fig. 17); 1♂, Pokhara (950 m), Nepal, VII–VIII.1983, M.G. Allen leg. (LT); 1♀, montane and oak forest (2,760 m), Phulchoki, Nepal, VIII.1983, M.G. Allen leg. (LT); 1♀, Phulchoki (2,500 m), Nepal, IX.1982, M.G. Allen leg. (LT); 1♀, Phulchoki (2,000 m), Nepal, VIII.1982, M.G. Allen leg. (LT); 1♀, Kathmandu (1,350 m), Nepal, VII.1983, M.G. Allen leg. (LT); 1♀, Kakani (2,000 m), Nepal, VIII.1982, M.G. Allen leg. (LT); 18♀♀, Pradesh, Patancheru, India, I (1♀), II (1♀), VIII (2♀♀), IX (5♀♀), X (9♀♀).1980, Bhatnagar leg. (LT); 8♀♀, Mysore, Mudigere, India, X–XI.1979, J.S. Noyes leg.; 3♀♀, Radjamandula, Java, Indonesia, XI.1937, K.M. Walsh leg.; 1♂ and 1 unsexed, Mt Djampang, Tjigaeha, Java, Indonesia, I.1938, K.M. Walsh leg.; 1♀, Mt Melang, Djampang Wetan, Java, Indonesia, VIII.1937, K.M. Walsh leg.; 1♂, Gunung Gede, Lebak Sioe, Java, Indonesia, IX.1937, K.M. Walsh leg. (all NHMUK); 1♀, Buzena, Nago City, Kunigami County, Okinawa-hontô, Okinawa Pref., Japan, 15.IV.1991, M. Hayashi leg.; 1♀, Uebaru, Nakijin Vil., Kunigami County, Okinawa-hontô, Okinawa Pref., Japan, 23.IV.1991, M. Hayashi leg. (LT) (all NIAES); 1♀, Clearing, Perak, Malaysia, 1.VI.1941, F. Gerald leg.; 5♀♀1♂, Bulolo forestry Reserve, Papua New Guinea, IX.1979, I.D. Gauld leg.; 2♀♀, Wau (1,000 m), Morobe, Papua New Guinea, X.1979, I.D. Gauld leg.; 7♀♀, Bay Bay, Leyte, Visca forest, Philippines, VIII (1♀), 30.VIII–4.IX (4♀♀), 5–13.IX (2♀♀).1980, L. Tuangganr leg.; 1 unsexed, Singapore, 2.XII.1967, C.G. Roche leg.; 6♀♀, Peak View Motel, Kandy, Kan. Dist. Sri Lanka, 7–14 (3♀♀), 14–24 (3♀♀).I.1970, Davis & Rowe leg. (all NHMUK); 5♀♀6♂♂ and 4 unsexed, Karen, Taiwan, 25.VI–1.VII (2♂♂), 17–23.VIII (3 unsexed), 26.VIII–4.XI (1♀), 12–19 (2♀♀), 14–19 (1♂).XI.1972, 16–22.IV.1973 (2♀♀3♂♂ and 1 unsexed) (MsT); 12♀♀2♂♂, Sunmoon Lake, Taichung, Taiwan, 2.X (1♀), 8.XI (1♀).1968, 9 (1♀), 31 (1♂).IV, 11 (1♂), 14 (1♀).VII, 1 (3♀♀), 2–8 (2♀♀), 7–13 (1♀), 9–15 (1♀).IX, 4–10.XI (1♀).1969 (MsT); 26♀♀7♂♂, Wufeng, Taichung, Taiwan, 25–28.VI (2♀♀), 1–3 (1♀1♂), 7–11 (1♀), 16–20 (1♂).VII, 19–26.X (1♀), 29.X–5.XI. (2♀♀), 5–10 (3♀♀), 10–15 (1♀2♂♂), 17–22 (1♀1♂).XI, 27.XI–3.XII.1979 (2♀♀1♂), 7–14 (1♀1♂), 15–21 (1♀), 20–26 (2♀♀).XII.1979, 1–4 (3♀♀), 4–11 (1♀), 25–31 (2♀♀).I.1980, 9–20.II (2♀♀).1980, K.C. Chou leg. (all TARI).

Australasian, Eastern Palaearctic, Oceanic, and Oriental regions (Yu et al. 2016). Gauld and Mitchell (1981) recorded this species from Nepal.

Head (Fig. 17B–D): GOI = 2.5–3.1; lower face 0.7–0.8× as wide as high; clypeus slightly to strongly convex in profile, its lower margin acute; mandible weakly twisted by 10–20°, moderately long, evenly tapered, its outer surface without a diagonal structure; upper mandibular tooth 1.2–1.5× as long as lower one; posterior ocellus almost touching eye; antenna with 53–65 flagellomeres and 20^th flagellomere 1.8–2.4× as long as wide.

Mesosoma (Fig. 17E): mesopleuron punctate to longitudinally punctostriate; scutellum with lateral longitudinal carinae reaching posterior end and convergent posteriorly; metapleuron punctate to punctostriate; propodeum almost evenly rounded, its posterior area moderately reticulate, outer margin of propodeal spiracle not joining pleural carina by a ridge.

Wings (Fig. 17F): fore wing with AI = 0.4–1.1, CI = 0.3–0.5, ICI = 0.4–0.5, SDI = 1.1–1.4; fore wing vein 1m-cu&M more or less evenly curved, 2r&RS almost straight; fenestra and sclerites of discosubmarginal cell of fore wing as in Figure 17F; fenestra of fore wing not very long and its anterodistal corner distinctly separated from proximal end of vein RS; proximal sclerite triangular, strongly confluent with distal one, strongly pigmented; central sclerite moderately to strongly pigmented and sclerotised, usually well-delineated oval, positioned in antero- to medio-distal part of fenestra; distal sclerite more or less evenly strong from proximal to distal; proximal corner of marginal cell of fore wing uniformly setose; vein 1cu-a subinterstitial to antefurcal to M&RS by less than 0.3× 1cu-a length.

Colour (Fig. 17): body including interocellar area entirely testaceous with black posterior segments of metasoma; wings hyaline.

Enicospilus melanocarpus is very similar to E. sauteri, but distinguished by the uniformly setose marginal cell of the fore wing (Fig. 17F) (marginal cell of fore wing proximally glabrous in E. sauteri) and the oval central sclerite (Fig. 17F) (central sclerite linear in E. sauteri). Many species were synonymised with E. melanocarpus under Gauld’s conservative species criteria, but their wide distribution and considerable range of morphological variation indicate this name includes many species. Therefore, further researches are needed to reveal the true species diversity under the name ‘melanocarpus’.

Figure 17. 

Enicospilus melanocarpus Cameron, 1905, ♂. A. Habitus; B. Head, frontal view; C. Head, lateral view; D. Head, dorsal view; E. Mesosoma, lateral view; F. Central part of fore wing.

The specific name is derived from the type locality.

2♀♀: Nepal.

Type series: holotype ♀, Pokhara (950 m), Nepal, VII–VIII.1983, M.G. Allen leg. (LT) (NHMUK) (Figs 2A, 18); paratype ♀, same label and repository as holotype.

Nepal.

Female (Holotype) (Fig. 18). Body length ca 16.5 mm.

Head with GOI = 2.5 (Fig. 18C). Lower face 0.8× as wide as high, finely punctate with setae, strongly shiny (Fig. 18B). Clypeus 1.6× as wide as high, finely punctate with setae, moderately convex in profile, its lower margin impressed (Fig. 18B, C). Malar space 0.4× as long as basal mandibular width (Fig. 18B, C). Mandible weakly twisted by ca 15°, moderately long, its proximal half evenly narrowed and distal half subparallel sided, its outer surface entirely almost flat with long and rather stout setae (Figs 2A, 18B, C). Upper mandibular tooth 1.7× as long as lower one, very slender and cylindrical (Figs 2A, 18B). Frons, vertex and gena strongly shiny with fine setae (Fig. 18B–D). Posterior ocellus rather small and separated from eye by 0.3× its own maximum diameter (Fig. 18B–D). Ventral end of occipital carina joining oral carina. Antenna with 49 flagellomeres; first flagellomere 1.6× as long as second; 20^th flagellomere 2.3× as long as wide.

Mesosoma entirely strongly shiny with setae (Fig. 18E). Pronotum punctostriate dorsally and finely coriaceous ventrally (Fig. 18E). Mesoscutum 1.5× as long as its maximum width, almost smooth with very fine punctures with setae, and evenly rounded in profile (Fig. 18E). Notauli absent (Fig. 18E). Scutellum moderately convex, almost smooth with very fine and sparse punctures with setae, with lateral longitudinal carinae reaching posterior end (Fig. 18E). Epicnemium from densely strigose dorsally to densely punctate ventrally with setae. Epicnemial carina present, evenly curved to anterior, its dorsal end not reaching anterior margin of mesopleuron (Fig. 18E). Mesopleuron finely punctate dorsally and longitudinally punctostriate to strigose ventrally (Fig. 18E). Submetapleural carina almost parallel sided centrally and weakly broadened anteriorly (Fig. 18E). Metapleuron moderately punctate with setae (Fig. 18E). Propodeum evenly rounded in profile; anterior transverse carina complete centrally, its lateral end almost joining pleural carina; anterior area longitudinally striate; spiracular area almost smooth with very fine and sparse punctures and setae; posterior area rather finely subconcentrically striate; propodeal spiracle elliptical, its outer margin not joining pleural carina by a ridge (Fig. 18E).

Wings. Fore wing length ca 11.0 mm with AI = 0.4, CI = 0.3, DI = 0.4, ICI = 0.4, SDI = 1.2, SI = 0.2, SRI = 0.3; vein 1m-cu&M almost evenly curved; vein 2r&RS slightly sinuous and RS evenly curved; fenestra and sclerites of discosubmarginal cell as in Figure 18F; proximal sclerite triangular, not confluent with distal sclerite, very strongly pigmented; central sclerite small and its major diameter subequal to thickness of vein 2r&RS, suboval, weakly sclerotised and pigmented, positioned in posterodistal part of fenestra; distal sclerite moderately pigmented; proximal corner of marginal cell evenly setose; posterodistal corner of second discal cell ca 95°; posterodistal corner of subbasal cell ca 95°; vein 1cu-a slightly antefurcal to M&RS by 0.1× 1cu-a length (Fig. 18F). Hind wing with NI = 1.2, RI = 1.7; vein RS straight; vein RA with 6 uniform hamuli.

Legs. Outer surface of fore tibia without dense and long spines. Hind leg with coxa in profile 1.7× as long as deep; basitarsus 2.0× as long as second tarsomere; fourth tarsomere 0.6× as long as third tarsomere and 3.5× as long as wide; tarsal claw simply pectinate.

Metasoma with PI = 2.8, DMI = 1.3, THI = 2.5; dorsal margin of tergite 1 more or less sinuous; thyridium elongate (Fig. 18A).

Colour (Fig. 18). Entirely testaceous except for apex of mandible, posterior part of T5, and T6–8 black. Wings hyaline; proximal sclerite brown; central and distal sclerites amber; veins brown.

Variations (n = 2): body length 15.5–16.5 mm; head with GOI = 2.4–2.5; clypeus 1.6–1.7× as wide as high; malar space 0.3–0.4× as long as basal mandibular width; mandible twisted by 15–25°; upper mandibular tooth 1.6–2.1× as long as lower one; antenna with first flagellomere 1.6–1.7× as long as second; pronotum punctostriate dorsally and finely coriaceous ventrally or entirely almost smooth to weakly coriaceous with very sparse and fine punctures; metapleuron sparsely to moderately punctate; fore wing length 10.0–11.0 mm; hind coxa in profile 1.6–1.7× as long as deep; fourth tarsomere 3.5–3.7× as long as wide; metasoma with PI = 2.7–2.8; THI = 2.5–2.8; mandible proximally testaceous and apically black or entirely dark brown to black.

Male. Unknown.

Enicospilus nepalensis sp. nov. is probably closely related to or belongs to the E. ramidulus complex. Among the complex, E. nepalensis sp. nov. is most closely related to E. tricorniatus Rao & Nikam, 1970 based on the rather small ocelli relative to other Enicospilus (posterior ocellus separated from eye by more than 0.3× its own maximum diameter) (e.g. Fig. 18B–D), highly shiny body (e.g. Fig. 18A–E), shape of body (e.g. Fig. 18A), shape and position of the fore wing veins and sclerites (e.g. Fig. 18F), distribution, etc. However, E. nepalensis sp. nov. is readily distinguishable from E. tricorniatus by the following characters: lower face more or less elongate and 0.8× as wide as high (Fig. 18B) (lower face subquadrate to transverse and 1.0–1.1× as wide as high in E. tricorniatus), the central sclerite weakly sclerotised and pigmented (Fig. 18F) (moderately to strongly sclerotised and pigmented in E. tricorniatus), moderate-sized, fore wing length 10.0–11.0 mm (small, fore wing length less than 8.5 mm in E. tricorniatus), posterior ocellus separated from eye by 0.3× its own maximum diameter (Fig. 18B–D) (posterior ocellus separated from eye by almost its own maximum diameter in E. tricorniatus).

Figure 18. 

Enicospilus nepalensis Shimizu sp. nov., ♀, holotype. A. Habitus; B. Head, frontal view; C. Head, lateral view; D. Head, dorsal view; E. Mesosoma, lateral view; F. Central part of fore wing.

The specific name is dedicated to Dr P.K. Nikam who studied Ophioninae as well as other groups of Hymenoptera mainly of India.

1♀: Nepal.

Type series: holotype ♀, Kathmandu (1,300 m), Nepal, XI.1982, M.G. Allen leg. (LT) (NHMUK) (Figs 2J, 19).

Nepal.

Female (Holotype) (Fig. 19). Body length ca 23.0 mm.

Head with GOI = 2.9 (Fig. 19C). Lower face 0.6× as wide as high, shiny, rather finely punctate with setae (Fig. 19B). Clypeus 1.6× as wide as high, sparsely and finely punctate with setae, almost flat in profile, and its lower margin acute (Fig. 19B, C). Malar space 0.2× as long as basal mandibular width (Fig. 19B, C). Mandible weakly twisted by ca 25°, long, proximally strongly narrowed, centrally to apically subparallel sided, its outer surface flat and smooth without a diagonal groove and line of punctures (Fig. 19B, C). Upper mandibular tooth 1.3× as long as lower one (Fig. 19B). Frons, vertex and gena moderately shiny with fine setae (Fig. 19B–D). Posterior ocellus almost touching eye (Fig. 19B–D). Ventral end of occipital carina joining oral carina. Antenna with 59 flagellomeres; first flagellomere 1.9× as long as second; 20^th flagellomere 1.5× as long as wide.

Mesosoma entirely moderately shiny with setae (Fig. 19E). Pronotum finely coriaceous with punctures to closely strigose (Fig. 19E). Mesoscutum 1.5× as long as its maximum width, finely punctate with setae, strongly shiny, and evenly rounded in profile (Fig. 19E). Notauli absent (Fig. 19E). Scutellum moderately convex, with lateral longitudinal carinae almost reaching posterior end, and moderately punctate with setae (Fig. 19E). Epicnemium densely punctate with setae. Epicnemial carina present, evenly weakly curved to anterior, its dorsal end not reaching anterior margin of mesopleuron (Fig. 19E). Mesopleuron entirely moderately punctate, and ventral margin longitudinally finely strigose (Fig. 19E). Submetapleural carina broadened anteriorly (Fig. 19E). Metapleuron diagonally closely strigose (Fig. 19E). Propodeum declivous in profile; anterior transverse carina complete centrally, its lateral end not joining pleural carina; pleural carina absent posteriorly; anterior area longitudinally striate medially and smooth laterally; spiracular area finely coriaceous; posterior area concentrically striate; propodeal spiracle elliptical, its outer margin not joining pleural carina by a ridge (Fig. 19E).

Wings. Fore wing length ca 15.0 mm with AI = 0.5, CI = 0.6, DI = 0.3, ICI = 0.8, SDI = 1.5, SI = 0.1, SRI = 0.3; vein 1m-cu&M moderately sinuous; vein 2r&RS very slightly bowed but almost straight, and RS evenly curved; fenestra and sclerites of discosubmarginal cell as in Figure 19F; proximal sclerite linear, weakly pigmented and virtually unsclerotised so that vestigial, separated from distal sclerite; central sclerite absent; distal sclerite almost absent but anterodistal part slightly pigmented; proximal corner of marginal cell uniformly setose; posterodistal corner of second discal cell ca 90°; posterodistal corner of subbasal cell ca 65°; vein 1cu-a antefurcal to M&RS by 0.2× 1cu-a length (Fig. 19F). Hind wing with NI = 2.6, RI = 1.7; vein RS straight; vein RA with 11 uniform hamuli.

Legs. Outer surface of fore tibia with very few spines. Hind leg with coxa in profile 1.7× as long as deep; basitarsus 2.2× as long as second tarsomere; fourth tarsomere 0.6× as long as third tarsomere and 2.6× as long as wide; tarsal claw simply pectinate except lacking pecten proximally.

Metasoma with PI = 3.2, DMI = 1.3, THI = 2.5; dorsal margin of tergite 1 not sinuous; thyridium elongate (Fig. 19A).

Colour (Fig. 19). Entirely testaceous except for apex of mandible black. Wings hyaline; sclerites of fore wing fenestra very slightly pigmented, testaceous; veins black to testaceous.

Variation. Unknown

Male. Unknown

Enicospilus nikami sp. nov. is similar to E. biharensis, E. maruyamanus, E. pudibundae, and E. transversus and these species are rather difficult to separate from each other. However, E. nikami sp. nov. can be distinguished from E. biharensis, E. maruyamanus and E. transversus by the proximally incomplete pectinae of hind tarsal claw (Fig. 2J) (hind tarsal claw completely pectinate from its base to apex in E. biharensis, E. maruyamanus and E. transversus, as in e.g. Figure 2I), from E. biharensis and E. pudibundae by the sinuous fore wing vein 1m-cu&M (Fig. 19F) (1m-cu&M evenly curved in E. biharensis and E. pudibundae as in Figures 6F, 23F), from E. maruyamanus by the entirely moderately punctate mesopleuron (Fig. 19E) (mesopleuron entirely longitudinally punctostriate in E. maruyamanus) and the angle of posterodistal corner of second discal cell (i.e. ca 90° in E. nikami sp. nov. as in Figure 19F, but ca 115° in E. maruyamanus), and from E. transversus by the entirely moderately punctate mesopleuron (Fig. 19E) (mesopleuron entirely longitudinally striate in E. transversus).

Figure 19. 

Enicospilus nikami Shimizu sp. nov., ♀, holotype. A. Habitus; B. Head, frontal view; C. Head, lateral view; D. Head, dorsal view; E. Mesosoma, lateral view; F. Central part of fore wing.

The specific name is derived from the type locality.

1♀: Nepal.

Type series: holotype ♀, Phulchoki, M.G. Allen leg. (NHMUK) (Figs 2G, 20).

Nepal.

Female (Holotype) (Fig. 20). Body length ca 21.5 mm.

Head with GOI = 2.9 (Fig. 20C). Lower face 0.7× as wide as high, rather finely punctate with setae, strongly shiny (Fig. 20B). Clypeus 1.3× as wide as high, finely punctate with setae, moderately convex in profile, and its lower margin impressed (Fig. 20B, C). Malar space 0.4× as long as basal mandibular width (Fig. 20B, C). Mandible weakly twisted by ca 20°, moderately long, evenly narrowed, its outer surface with a diagonal setose deep groove between its dorsoproximal corner to base of mandibular apical teeth (Fig. 20B, C). Upper mandibular tooth 1.6× as long as lower one, slender and cylindrical (Fig. 20B). Frons, vertex and gena strongly shiny with fine setae (Fig. 20B–D). Posterior ocellus almost touching eye (Fig. 20B–D). Ventral end of occipital carina joining oral carina. Antenna with 64 flagellomeres; first flagellomere 1.7× as long as second; 20^th flagellomere 2.2× as long as wide.

Mesosoma entirely strongly shiny with setae (Fig. 20E). Pronotum finely punctate dorsally and strigose to rugose ventrally (Fig. 20E). Mesoscutum 1.5× as long as its maximum width, almost smooth with very fine punctures with setae, and evenly rounded in profile (Fig. 20E). Notauli absent (Fig. 20E). Scutellum moderately convex, anterior 0.4 transversely striate, anterior 0.4–0.5 punctate, and posterior 0.5 longitudinally strigose, with lateral longitudinal carinae reaching posterior end (Figs 2G, 20E). Epicnemium densely punctate with setae. Epicnemial carina present, evenly curved to anterior, its dorsal end close to anterior margin of mesopleuron (Fig. 20E). Mesopleuron entirely finely punctate, longitudinally strigose ventrally (Fig. 20E). Submetapleural carina weakly evenly broadened anteriorly (Fig. 20E). Metapleuron entirely finely punctate with setae (Fig. 20E). Propodeum almost evenly rounded in profile; anterior transverse carina complete; anterior area longitudinally striate; spiracular area almost smooth with very fine and sparse punctures with setae; posterior area rather finely irregularly rugose; propodeal spiracle elliptical, its outer margin not joining pleural carina by a ridge.

Wings. Fore wing length ca 13.5 mm with AI = 0.4, CI = 0.4, DI = 0.4, ICI = 0.5, SDI = 1.2, SI = 0.1, SRI = 0.3; vein 1m-cu&M weakly sinuous; vein 2r&RS almost straight and RS evenly curved; fenestra and sclerites of discosubmarginal cell as in Figure 20F; proximal sclerite triangular, confluent with distal sclerite, moderately pigmented; central sclerite rather small and its minor diameter smaller than thickness of vein 2r&RS, elliptical, moderately sclerotised and pigmented, positioned in mediodistal part of fenestra; distal sclerite moderately pigmented; proximal corner of marginal cell evenly setose; posterodistal corner of second discal cell ca 105°; posterodistal corner of subbasal cell ca 85°; vein 1cu-a subinterstitial to M&RS (Fig. 20F). Hind wing with NI = 1.3, RI = 1.7; vein RS straight; vein RA with 7 uniform hamuli.

Legs. Outer surface of fore tibia with sparse spines. Hind leg with coxa in profile 2.0× as long as deep; basitarsus 2.0× as long as second tarsomere; fourth tarsomere 0.6× as long as third tarsomere and 4.6× as long as wide; tarsal claw simply pectinate.

Metasoma with PI = 2.9, DMI = 1.4, THI = 2.9; dorsal margin of tergite 1 weakly sinuous; thyridium elongate (Fig. 20A).

Colour (Fig. 20). Entirely testaceous except for apex of mandible black. Wings hyaline; sclerites amber; veins brown.

Variation. Unknown

Male. Unknown

Mandibular structure and mesosoma sculpture of E. phulchokiensis sp. nov. indicate that it belongs to the E. ramidulus complex. Enicospilus phulchokiensis sp. nov. runs to couplet 230 (including E. melanocarpus and E. xavius) of Gauld and Mitchell’s (1981: 143) key, and to couplet 61 (including E. melanocarpus and E. sauteri) of Tang’s (1990: 34, 181) key. However, E. phulchokiensis sp. nov. is distinguishable from E. melanocarpus, E. sauteri and E. xavius by the strongly sculptured scutellum (Fig. 2G), sinuous fore wing vein 1m-cu&M (Fig. 20F), and entirely testaceous metasoma without black posterior segments (Fig. 20A). Moreover, E. phulchokiensis sp. nov. is possibly related to E. puncticulatus Tang, 1990 and its related species-group, but distinguished from E. puncticulatus by the confluent proximal and distal sclerites (Fig. 20F) (separated in E. puncticulatus).

Figure 20. 

Enicospilus phulchokiensis Shimizu sp. nov., ♀, holotype. A. Habitus; B. Head, frontal view; C. Head, lateral view; D. Head, dorsal view; E. Mesosoma, lateral view; F. Central part of fore wing.

6♀♀6♂♂: Australia (5♀♀6♂♂), Indonesia (1♀). No Nepalese specimens were examined.

Type series: paratypes of Enicospilus pseudantennatus Gauld, 1977, 1♀, Paramatta, NSW, Australia, 16.I.1921 (EMUS); 5♂♂,Tambourine Mts, SE Queensland, Australia, 1–9.V.1935, R.E. Turner leg.; 1♂, Cabramatta, NSW, Australia, 6.IV.1963, M. Nikitin leg.; 1♀, Merrylands, NSW, Australia, 25.XI.1964, M. Nikitin leg. (all NHMUK).

Non-type series: 1♀, D.P.I Research Stn, Gatton, SE Queensland, Australia, 13–21.IV.1981 (MsT) (Fig. 21); 1♀, Mt Tanbourine, Queensland, Australia, 12–18.X.1978, I.D. Galloway leg.; 1♀, Canberra, ACT, Australia, IX.1981, I.D. Gauld leg. (all NHMUK); 1♀, Ambon, Indonesia, 29.IX.1960, A.M.R. Wegner leg. (EMUS).

Australasian, Oceanic and Oriental regions (Yu et al. 2016). Gauld and Mitchell (1981) recorded this species from Nepal.

Head (Fig. 21B–D): GOI = 2.2–2.8; lower face 0.7–0.8× as wide as high; clypeus moderately convex in profile, its lower margin impressed; mandible rather weakly twisted by 10–20°, moderately long, evenly tapered, its outer surface without a diagonal structure; upper mandibular tooth 1.3–1.6× as long as lower one; posterior ocellus close to eye; antenna with 56–63 flagellomeres and 20^th flagellomere 2.2–2.4× as long as wide.

Mesosoma (Fig. 21E): mesopleuron entirely punctate; scutellum with lateral longitudinal carinae reaching posterior end and convergent posteriorly; metapleuron punctate; propodeum evenly weakly rounded, its posterior area moderately reticulate, outer margin of propodeal spiracle not joining pleural carina by a ridge.

Wings (Fig. 21F): fore wing with AI = 0.3–0.6, CI = 0.3–0.4, ICI = 0.5–0.7, SDI = 1.2–1.4; fore wing vein 1m-cu&M moderately sinuous, 2r&RS almost straight; fenestra and sclerites of discosubmarginal cell of fore wing as in Figure 21F; fenestra of fore wing not very long and its anterodistal corner distinctly separated from proximal end of vein RS; proximal sclerite triangular, separated from distal one, strongly pigmented; central sclerite partially strongly pigmented and sclerotised, ill-delineated oval, positioned in almost medio-distal part of fenestra; distal sclerite strong distally; proximal corner of marginal cell of fore wing uniformly setose; vein 1cu-a subinterstitial to antefurcal to M&RS by less than 0.2 1cu-a length.

Colour (Fig. 21): body including interocellar area entirely red-brown; wings hyaline.

As mentioned under E. laqueatus, four Oriental species of Enicospilus (E. laqueatus, E. pseudantennatus, E. vestigator, and E. tripartitus) have similar fenestra, sclerites, and fore wing veins (e.g. Figs 15F, 21F, 26F). Among them, E. pseudantennatus is distinguished from E. laqueatus by the flat outer surface of the mandible (outer surface of mandible with a diagonal deep setose groove between dorsoproximal corner and base of mandibular apical teeth in E. laqueatus), from E. tripartitus by the not densely setose and proximally more or less flat outer mandibular surface (outer surface of mandible with very dense setae and sharp and rather deep proximal concavity in E. tripartitus, as in Figure 2C), and from E. vestigator by the weakly twisted mandible (10–20°) (mandible strongly twisted by 60–80° in E. vestigator).

Figure 21. 

Enicospilus pseudantennatus Gauld, 1977, ♀. A. Habitus; B. Head, frontal view; C. Head, lateral view; D. Head, dorsal view; E. Mesosoma, lateral view; F. Central part of fore wing.

7♀♀7♂♂: Nepal (5♀♀4♂♂), China (1♀1♂), Japan (1♂), Taiwan (1♀1♂).

Type series: holotype of Henicospilus pseudoconspersae Sonan, 1927, ♂, Taihoku, Taiwan, 25.IV.1927, J. Sonan leg. (TARI); holotype of Henicospilus mushanus Uchida, 1928, ♀, Musha, Taiwan, 24.VII.1925, Matsumura (SEHU); holotype of Enicospilus tenuinubeculus Chiu, 1954, ♀, Fukien, Shaown, China, 23–29.V.1944, H.F. Chao leg. (TARI).

Non-type series: 1♀, Kakani (2,070 m), Nepal, VII.1983, M.G. Allen leg. (LT) (Fig. 22); 1♀, Kathmandu (1,300 m), Nepal, XI.1982, M.G. Allen leg. (LT); 1♀, Godaveri (1,550–1,700 m), Nepal, V.1983, M.G. Allen leg. (LT); 1♂, Godaveri (5,000′), Nepal, 5.VIII.1967; 1♀, Phulchoki (2,000 m), Nepal, VIII.1982, M.G. Allen leg. (LT); 1♀, Kathmandu (4300′), Nepal, VIII.1982; 1♂, Sal & 2y forest (330 m), Dharan, Terai, Nepal, 14–15.XI.1983, M.G. Allen leg.; 1♂, Godavari, Kathmandu, Nepal, 5.VIII.1967; 1♂, l mi. S of Ulleri (5,500–7,000′), Nepal, 16.V.1954, J. Ouinlan leg.; 1♂, China (all NHMUK); 1♂, Hiji agricultural road (85 m, 26°43'16.8"N, 128°10'43.4"E), Hiji, Kunigami Village, Kunigami County, Okinawa-hontô, Okinawa Pref., Japan, 3–4.VII.2016, S. Shimizu et al. leg. (LT) (NIAES).

Eastern Palaearctic and Oriental regions (Yu et al. 2016). Gauld and Mitchell (1981) recorded this species from Nepal.

Head (Fig. 22B–D): GOI = 2.8–3.1; lower face 0.6–0.7× as wide as high; clypeus almost flat in profile, its lower margin acute to subacute; mandible rather weakly twisted by 15–25°, moderately long, proximally tapered and distally approximately parallel sided, its outer surface without a diagonal structure; upper mandibular tooth 1.2–1.4× as long as lower one; posterior ocellus almost touching eye; antenna with 52–65 flagellomeres and 20^th flagellomere 1.7–2.3× as long as wide.

Mesosoma (Fig. 22E): mesopleuron punctate to longitudinally punctostriate; scutellum with lateral longitudinal carinae reaching posterior end and convergent posteriorly; metapleuron punctostriate; propodeum evenly rounded, its posterior area moderately reticulate, outer margin of propodeal spiracle not joining pleural carina by a ridge.

Wings (Fig. 22F): fore wing with AI = 0.7–0.9, CI = 0.6–0.7, ICI = 0.4–0.6, SDI = 1.3–1.4; fore wing vein 1m-cu&M moderately sinuous, 2r&RS almost straight; fenestra and sclerites of discosubmarginal cell of fore wing as in Figure 22F; fenestra of fore wing not very long and its anterodistal corner distinctly separated from proximal end of vein RS; proximal sclerite semicircular, isolated and not touching margin of fenestra, almost always (very) weakly pigmented; central sclerite absent; distal sclerite absent or vestigial; proximal corner of marginal cell of fore wing uniformly setose; vein 1cu-a antefurcal to M&RS by 0.2–0.3× 1cu-a length.

Colour (Fig. 22): body including interocellar area entirely testaceous; wings hyaline.

Enicospilus pseudoconspersae is one of the most distinctive and easily distinguishable species among the Oriental species of Enicospilus on account of the characteristic isolated and weakly pigmented semicircular proximal sclerite (Fig. 22F). There are no known morphologically similar species.

Figure 22. 

Enicospilus pseudoconspersae (Sonan, 1927), ♀. A. Habitus; B. Head, frontal view; C. Head, lateral view; D. Head, dorsal view; E. Mesosoma, lateral view; F. Central part of fore wing.

18♀♀2♂♂: Nepal (2♀♀1♂), Brunei (3♀♀), India (1♀), Japan (12♀♀1♂).

Type series: lectotype of Henicospilus pudibundae Uchida, 1928, ♂, Sapporo, Hokkaidô, Japan, 4.VI.1925, Tamanuki leg. (emerged from Dasychira pudibunda L.) (SEHU).

Non-type series: 2♀♀, Kakani, Nepal, 1–30.V.1984, M.G. Allen leg. (Fig. 23); 1♂, Sal & 2^y forest (330 m), Dharan, Terai, Nepal, 14–15.XI.1983, M.G. Allen leg.; 2♀♀, U. Temburong (1,700 m), Gn. Pagon, Brunei, IV.1981, I.D. Gauld leg.; 1♀, U. Temburong (1,500 m), Bukit Retak, Brunei, IV.1981, I.D. Gauld leg. (all NHMUK); 1♀, Anamalai Hills (3,500′), Cinchona, India, IV.1956, P.S. Nathan leg. (CNC); 1♀, Takadomari, Fukagawa City, Hokkaidô, Japan, 5–19.VIII.2007, H. Hara leg. (MsT) (NSMT); 1♀, Yoshigahira, Niigata Pref., Japan, 25.VI.1954, K. Baba leg. (MNHA); 1♀, Kurokawa, Niigata Pref., Japan, 22.VI.1954, K. Baba leg. (MNHA); 1♀, Oyamada, Machida City, Tôkyô, Japan, IX.2008, S. Ohsato leg. (NHMUK); 1♀, Dokan-Shinmichi, Imperial Palace, Chiyoda Ward, Tôkyô, Japan, 27.VII–3.VIII.2010 (MsT) (NSMT); 1♀, Mt Futatabi-san, Kôbe City, Hyôgo Pref., Japan, 28.VIII.1990, N. Sugiura leg. (MNHA); 6♀♀, Mori, Tôjyô Town, Shôbara City, Hiroshima Pref., Japan, 21.VII.2015 (1♀), 6 (2♀♀), 8 (1♀). IX, 3.X (1♀).2016, 17.IX.2017 (1♀), N. Takashiba leg. (LT) (HMNH).

Eastern Palaearctic and Oriental regions (Yu et al. 2016). Newly recorded from Nepal.

Head (Fig. 23B–D): GOI = 2.6–2.8; lower face 0.7× as wide as high; clypeus almost flat in profile, its lower margin acute to subacute; mandible weakly twisted by 10–20°, moderately long, evenly tapered, its outer surface without a diagonal structure; upper mandibular tooth 1.2–1.5× as long as lower one; posterior ocellus (almost) touching eye; antenna with 54–59 flagellomeres and 20^th flagellomere 2.0–2.1× as long as wide.

Mesosoma (Fig. 23E): mesopleuron entirely punctate; scutellum with lateral longitudinal carinae reaching posterior end and convergent posteriorly; metapleuron punctate; propodeum weakly declivous, its posterior area irregularly wrinkled, outer margin of propodeal spiracle not joining pleural carina by a ridge.

Wings (Fig. 23F): fore wing with AI = 0.5–1.0, CI = 0.5–0.7, ICI = 0.5–0.7, SDI = 1.4–1.5; fore wing vein 1m-cu&M evenly curved, 2r&RS almost straight; fenestra and sclerites of discosubmarginal cell of fore wing as in Figure 23F; fenestra of fore wing not very long and its anterodistal corner distinctly separated from proximal end of vein RS; proximal sclerite more or less linear, very weakly confluent with distal one or not, very weakly to strongly pigmented; central sclerite absent; distal sclerite more or less weak to absent; proximal corner of marginal cell of fore wing sparsely to uniformly setose; vein 1cu-a subinterstitial to antefurcal to M&RS by less than 0.2× 1cu-a length.

Colour (Fig. 23): body including interocellar area entirely testaceous, sometimes posterior segments of metasoma weakly infuscate; wings hyaline to very slightly infuscate.

Enicospilus pudibundae resembles E. biharensis, E. maruyamanus, E. nikami sp. nov., and E. transversus, but can be distinguished from E. biharensis, E. maruyamanus, and E. transversus by the proximally incomplete pectination of the hind tarsal claw (pectination of hind tarsal claw complete from base to apex of the claw in E. biharensis, E. maruyamanus, and E. transversus, as in e.g. Figure 2I) and also from E. maruyamanus, E. nikami sp. nov., and E. transversus by the evenly curved fore wing vein 1m-cu&M (Fig. 23F) (1m-cu&M more or less sinuous in E. maruyamanus, E. nikami sp. nov. and E. transversus, as in e.g. Figure 19F). The Nepalese and some other Oriental specimens exhibit a rather wider proximal sclerite and sparser setosity in the proximal corner of the fore wing fenestra than the holotype and Eastern Palaearctic specimens, suggesting that the Oriental specimens are potentially cryptic species. However, at present, I have not enough evidence to describe them as a new species and tentatively follow Gauld and Mitchell’s (1981) species criteria.

Figure 23. 

Enicospilus pudibundae (Uchida, 1928), ♀. A. Habitus; B. Head, frontal view; C. Head, lateral view; D. Head, dorsal view; E. Mesosoma, lateral view; F. Central part of fore wing.

4♀♀4♂♂: Nepal (1♀4♂♂), Brunei (2♀♀), Singapore (1♀).

Non-type series: 1♀, Kathmandu (1,350 m), Nepal, VII.1983, M.G. Allen leg. (LT); 4♂♂, Phulchoki (2,000 m), Nepal, VIII.1982, M.G. Allen leg. (LT) (Fig. 24); 2♀♀, Seria, Brunei, XII.1979, Allen leg.; 1♀, Singapore, 1905, H.N. Ridley leg. (all NHMUK).

Australasian, Eastern Palaearctic, and Oriental regions (Yu et al. 2016). Newly recorded from Nepal and Brunei.

Head (Fig. 24B–D): GOI = 2.7–3.0; lower face 0.6–0.7× as wide as high; clypeus slightly convex in profile, its lower margin subacute to blunt; mandible weakly twisted by 10–20°, moderately long, proximally tapered and distally more or less parallel sided, its outer surface without a diagonal structure; upper mandibular tooth 1.3–1.5× as long as lower one; posterior ocellus almost touching eye; antenna with 56–59 flagellomeres and 20^th flagellomere 1.6–1.9× as long as wide.

Mesosoma (Fig. 24E): mesopleuron punctate to longitudinally punctostriate; scutellum with lateral longitudinal carinae reaching anterior 0.8 or more and convergent posteriorly; metapleuron punctostriate to striate; propodeum weakly declivous, its posterior area irregularly to subconcentrically wrinkled, outer margin of propodeal spiracle not joining pleural carina by a ridge.

Wings (Fig. 24F): fore wing with AI = 0.5–0.6, CI = 0.2–0.4, ICI = 0.6–0.8, SDI = 1.3–1.4; fore wing vein 1m-cu&M moderately sinuous, 2r&RS almost straight; fenestra and sclerites of discosubmarginal cell of fore wing as in Figure 24F; fenestra of fore wing not very long and its anterodistal corner distinctly separated from proximal end of vein RS; proximal sclerite triangular, confluent with distal one, strongly pigmented; central sclerite absent; distal sclerite more or less entirely pigmented; proximal corner of marginal cell of fore wing uniformly setose; vein 1cu-a antefurcal to M&RS by 0.1–0.3× 1cu-a length.

Colour (Fig. 24F): body including interocellar area entirely testaceous; wings hyaline.

Enicospilus purifenestratus is very similar to E. urocerus Gauld & Mitchell, 1981, but distinguished from it by the unswollen segments 3 and 4 of the maxillary palp (segments 3 and 4 of the maxillary palp swollen in E. urocerus) and thinner distal sclerite (Fig. 24F) (distal sclerite thicker in E. urocerus).

Figure 24. 

Enicospilus purifenestratus (Enderlein, 1921), ♂. A. Habitus; B. Head, frontal view; C. Head, lateral view; D. Head, dorsal view; E. Mesosoma, lateral view; F. Central part of fore wing.

The specific name is dedicated to Dr Yuqing Tang who described E. longitarsis, which is morphologically the most similar species to the one that is hereby described, and has contributed to the taxonomy of Ophioninae in Asia, represented by the monograph of Chinese Enicospilus (Tang 1990).

1♂: Nepal.

Type series: holotype ♂, Kakani (2,070 m), Nepal, 1–23.VIII.1983, M.G. Allen leg. (NHMUK) (Figs 2B, H, 25).

Nepal.

Male (Holotype) (Figs 2B, H, 25). Body length ca 24.5 mm.

Head with GOI = 2.5 (Fig. 25C). Lower face 0.9× as wide as high, moderately punctate with setae and shiny (Fig. 25B). Clypeus 1.7× as wide as high, moderately punctate with setae, moderately convex in profile, lower margin impressed (Fig. 25B, C). Malar space 0.4× as long as basal mandibular width (Fig. 25B, C). Mandible weakly twisted by ca 25°, very long, proximally strongly narrowed, centrally to apically subparallel sided, its outer surface with a diagonal setose deep groove between dorsoproximal corner to base of mandibular apical teeth (Figs 2B, 25B, C). Upper mandibular tooth 2.1× as long as lower one, stouter than lower one (Figs 2B, 25B). Frons, vertex and gena moderately shiny with fine setae (Fig. 25B–D). Posterior ocellus close to eye, separated from eye by less than 0.1× its own maximum diameter (Fig. 25B–D). Ventral end of occipital carina joining oral carina. Antenna incomplete apically, right antenna with 64 flagellomeres and left antenna with 65 flagellomeres; first flagellomere 1.9× as long as second; 20^th flagellomere 2.2× as long as wide.

Mesosoma entirely moderately shiny with setae (Fig. 25E). Pronotum punctate dorsally and punctostrigose centrally to ventrally (Fig. 25E). Mesoscutum 1.5× as long as its maximum width, densely and finely punctate with setae, rather weakly shiny, and evenly rounded in profile (Fig. 25E). Notauli absent (Fig. 25E). Scutellum moderately convex, moderately punctate with setae, with lateral longitudinal carinae almost reaching posterior end (Figs 2H, 25E). Epicnemium densely punctate with setae. Epicnemial carina present, evenly weakly curved to anterior, its dorsal end not reaching anterior margin of mesopleuron (Fig. 25E). Mesopleuron entirely weakly to moderately punctostriate to reticulate-strigose longitudinally (Fig. 25E). Submetapleural carina almost parallel sided centrally and weakly broadened anteriorly (Fig. 25E). Metapleuron densely punctostriate with setae (Fig. 25E). Propodeum almost evenly rounded in profile; anterior transverse carina complete centrally, its lateral end almost joining pleural carina; pleural carina vestigial; anterior area longitudinally striate; spiracular area strongly shiny and finely punctures with setae; posterior area rather moderately rugose; propodeal spiracle elliptical, its outer margin not joining pleural carina by a ridge (Fig. 25E).

Wings. Fore wing length ca 15.5 mm with AI = 0.4, CI = 0.4, DI = 0.3, ICI = 0.5, SDI = 1.3, SI = 0.1, SRI = 0.3; vein 1m-cu&M almost evenly curved; vein 2r&RS almost straight and RS evenly curved; fenestra and sclerites of discosubmarginal cell as in Figure 25F; proximal sclerite triangular, confluent with distal sclerite, moderately pigmented; central sclerite absent; distal sclerite weakly pigmented; proximal corner of marginal cell uniformly setose; posterodistal corner of second discal cell ca 95°; posterodistal corner of subbasal cell ca 90°; vein 1cu-a antefurcal to M&RS by 0.3× 1cu-a length (Fig. 25F). Hind wing with NI = 1.8, RI = 1.5; vein RS straight; vein RA with 6 uniform hamuli.

Legs. Ventral 0.7 of outer surface of fore tibia with rather dense spines. Hind leg with coxa in profile 1.8× as long as deep; basitarsus 2.0× as long as second tarsomere; fourth tarsomere 0.7× as long as third tarsomere and 5.0× as long as wide; tarsal claw simply pectinate.

Metasoma with PI = 2.8, DMI = 1.3, THI = 2.1; dorsal margin of tergite 1 slightly sinuous; thyridium elongate (Fig. 25A).

Colour (Fig. 25). Entirely testaceous except for apex of mandible black. Wings hyaline; proximal sclerite testaceous, distal sclerite very weakly pigmented; veins brown.

Variation. Unknown.

Female. Unknown.

Enicospilus tangi sp. nov. can be confused with E. kakanicus sp. nov., E. longitarsis, and E. yonezawanus, all of which belong to the E. ramidulus complex. Among these species, E. tangi sp. nov. is most closely related to E. longitarsis, and these species are distinguished from the other Oriental species of Enicospilus by the triangular proximal sclerite (e.g. Fig. 25F), the absence of the central sclerite (e.g. Fig. 25F), moderately large value of SDI (over 1.3) (e.g. Fig. 25F), a diagonal setose deep groove of the mandibular outer surface (e.g. Fig. 2B), moderately large fore wing fenestra (e.g. Fig. 25F), rather dense spines on the outer surface of the fore tibia, etc. Enicospilus tangi sp. nov. is distinguished from E. longitarsis by the following character states: scutellum narrowed posteriorly (Fig. 2H) (subquadrate in E. longitarsis); fore wing vein 1m-cu&M evenly curved (Fig. 25F) (slightly sinuous in E. longitarsis); lower face 0.9× as wide as high (Fig. 25B) (0.8 in E. longitarsis); GOI = 2.5 (Fig. 25C) (1.8 in E. longitarsis).

Figure 25. 

Enicospilus tangi Shimizu sp. nov., ♂, holotype. A. Habitus; B. Head, frontal view; C. Head, lateral view; D. Head, dorsal view; E. Mesosoma, lateral view; F. Central part of fore wing.

27♀♀10♂♂ and 2 unsexed: Nepal (24♀♀8♂♂ and 1unsexed), China (1♀), India (1♂), Japan (1 unsexed), Taiwan (2♀♀), unknown (1♂).

Type series: holotype of Enicospilus tripartitus Chiu, 1954, ♀, Taihoku, Taiwan, 27.VIII.1937, J. Sonan leg. (TARI); paratype of same species, 1♂, no data (NHMUK).

Non-type series: 24♀♀8♂♂, Kakani (2,000 m), Nepal, VIII.1982 (2♀♀), VI (4♀♀1♂), VII (4♀♀3♂♂), 1–23 (2♀♀2♂♂).VIII (3♀♀1♂), IX (4♀♀), X (2♀♀).1983, 1–30.V (1♀1♂), 1–14.VII (2♀♀).1984, M.G. Allen leg. (Figs 2C, 26); 1 unsexed, Sangu (ca 6,200′), Taplejung district, Nepal, 16–29.X.1961; 1♀, ShinKaiSi (1,340 m), Mt Omei, Szechuen, China; 1♂, Kangra Valley (1,370 m), India, X.1899, Dudgeon leg. (all NHMUK); 1 unsexed, Genka-yama, Okinawa-hontô, Okinawa Pref., Japan, 4.V.1964, T. Takara & T. Kakinohana leg. (MNHA); 1♀, Kuanhsi, Taiwan, 29.VIII.1968 (MsT) (TARI).

Eastern Palaearctic and Oriental regions (Yu et al. 2016). Gauld and Mitchell (1981) recorded this species from Nepal.

Head (Figs 2C, 26B–D): GOI = 2.2–2.9; lower face 0.7–0.8× as wide as high; clypeus moderately to strongly convex in profile, its lower margin more or less blunt; mandible rather weakly twisted by 10–20°, moderately long, proximally tapered and distally parallel sided, its outer surface flat but with conspicuous dense setae and a proximal deep concavity; upper mandibular tooth 1.2–1.6× as long as lower one; posterior ocellus close to eye; antenna with 55–66 flagellomeres and 20^th flagellomere 2.2–2.4× as long as wide.

Mesosoma (Fig. 26E): mesopleuron entirely more or less densely punctate and submatt; scutellum with lateral longitudinal carinae reaching posterior end and convergent posteriorly; metapleuron densely punctate as mesopleuron; propodeum weakly declivous, its posterior area moderately reticulate, outer margin of propodeal spiracle not joining pleural carina by a ridge.

Wings (Fig. 26F): fore wing with AI = 0.3–0.6, CI = 0.3–0.4, ICI = 0.5–0.7, SDI = 1.2–1.6; fore wing vein 1m-cu&M almost evenly curved to slightly sinuous, 2r&RS almost straight; fenestra and sclerites of discosubmarginal cell of fore wing as in Figure 26F; fenestra of fore wing not very long and its anterodistal corner distinctly separated from proximal end of vein RS; proximal sclerite triangular, separated from distal one, strongly pigmented; central sclerite strongly pigmented and sclerotised, well-delineated oval and its major axis parallel to distal margin of fenestra, positioned in mediodistal part of fenestra; distal sclerite absent to weak; proximal corner of marginal cell of fore wing uniformly setose; vein 1cu-a subinterstitial to antefurcal to M&RS by less than 0.2× 1cu-a length.

Colour (Fig. 26): body including interocellar area entirely reddish brown; wings hyaline.

Four Oriental Enicospilus species, E. laqueatus, E. pseudantennatus, E. vestigator, and E. tripartitus, have similar fenestra, sclerites, and fore wing veins (e.g. Figs 15F, 21F, 26F), as mentioned under E. laqueatus and E. pseudantennatus. Among them, E. tripartitus is readily distinguishable from other species by the outer mandibular surface: outer surface of mandible with very dense setae and sharp and rather deep proximal concavity in E. tripartitus (Figs 2C, 26B, C), but more or less flat proximally with scattered setae in E. laqueatus (Fig. 15B, C) E. pseudantennatus (Fig. 21B, C) and E. vestigator.

Figure 26. 

Enicospilus tripartitus Chiu, 1954, ♀. A. Habitus; B. Head, frontal view; C. Head, lateral view; D. Head, dorsal view; E. Mesosoma, lateral view; F. Central part of fore wing.

27♀♀8♂♂: Nepal (1♀), India (10♀♀), Indonesia (1♀), Japan (2♀♀7♂♂), Laos (8♀♀), Malaysia (4♀♀), Papua New Guinea (1♀), Taiwan (1♂).

Type series: lectotype of Henicospilus yonezawanus Uchida, 1928, ♀, Yonezawa, Yamagata Pref., Honshû, Japan, 23.VII.1919, S. Matsumura leg. (SEHU); holotype of E. microstriatellus Uchida, 1956, ♂, Sinmura, Amami-ôshima, Kagoshima Pref., Ryûkyûs, Japan, 7.IV.1954, T. Kumata leg. (SEHU).

Non-type series: 1♀, Godaveri (1,550–1,700 m), Nepal, IX.1983, M.G. Allen leg. (LT) (Figs 2D, 27); 10♀♀, Andhra Pradesh, Patanchneru, India, IX.1980, Rhatnagar leg. (LT); 1♀, Medan, L. Fulmek, Sumatra, Indonesia (all NHMUK); 1♂, Isa (32°8'29.3"N, 130°33'13.7"E), Kagoshima Pref., Kyûshû, Japan, 7.IX.2012, Y. Matsubara & K. Fukuda leg. (MsT) (NSMT); 5♂♂, Isa (32°6'41.8"N, 130°31'38.4"E), Kagoshima Pref., Kyûshû, Japan, 7.IX.2012, Y. Matsubara & K. Fukuda leg. (MsT) (CNC); 1♀, Hiji agricultural road (85 m, 26°43'16.8"N, 128°10'43.4"E), Hiji, Kunigami Vil., Kunigami County, Okinawa-hontô, Okinawa Pref., Ryûkyûs, Japan, 2–3.VII.2016, S. Shimizu et al. leg. (LT) (MNHA); 4♀♀, Phou Khoun (19.250697 N, 102.254204 E), Luang Phabang, Laos, 21–22.IV.2018, H. Yoshitomi leg. (EUM); 4♀♀, Sala Phou Khoun (19°25'7.57"N, 102°25'41.6"E), Luang Phabang, Laos, 21.IV.2018, K. Yasuda leg. (LT) (EUM); 1♀, Serdang, Malaysia, IX.1979, Khashiyah leg.; 1♀, Carambola Farm, Serdang, Selangor, Malaysia, XI.1979; 2♀♀, Serdang, Selangor, Malaysia, X (1♀) and XI (1♀).1979, I.D. Gauld leg.; 1♀, Morobe (1,000 m), Wau, Papua New Guinea, X.1979, I.D. Gauld leg. (all NHMUK); 1♂, Chihpen, Taitung, Taiwan, 17–18.II.1982, L.Y. Chou & K.C. Chou leg. (TARI).

Australasian, Eastern Palaearctic, and Oriental regions (Yu et al. 2016). Newly recorded from Nepal.

Head (Figs 2D, 27B–D): GOI = 2.9–3.2; lower face 0.7–0.8× as wide as high; clypeus moderately convex in profile, its lower margin impressed; mandible weakly twisted by 10–20°, moderately long, evenly tapered, its outer surface with a diagonal setose groove between its dorsoproximal corner and base of mandibular apical teeth; upper mandibular tooth 1.2–1.5× as long as lower one; posterior ocellus almost touching eye; antenna with 63–70 flagellomeres and 20^th flagellomere 2.0–2.2× as long as wide.

Mesosoma (Fig. 27E): mesopleuron punctate to rather closely longitudinally striate; scutellum with lateral longitudinal carinae reaching posterior end and convergent posteriorly; metapleuron punctate to striate; propodeum almost evenly rounded, its posterior area moderately reticulate, outer margin of propodeal spiracle joining pleural carina by a ridge.

Wings (Fig. 27F): fore wing with AI = 0.3–0.7, CI = 0.2–0.4, ICI = 0.4–0.6, SDI = 1.2–1.3; fore wing vein 1m-cu&M almost evenly curved to very slightly sinuous, 2r&RS almost straight; fenestra and sclerites of discosubmarginal cell of fore wing as in Figure 27F; fenestra of fore wing not very long and its anterodistal corner distinctly separated from proximal end of vein RS; proximal sclerite triangular, separated from distal one, strongly pigmented; central sclerite absent; distal sclerite absent proximally and weak distally; proximal corner of marginal cell of fore wing uniformly setose; vein 1cu-a antefurcal to M&RS by 0.1–0.3× 1cu-a length.

Colour (Fig. 27): body including interocellar area entirely testaceous; wings hyaline.

Enicospilus yonezawanus is one of the most common in the Eastern Palaearctic and Oriental regions and more or less distinctive species based on some characters, such as mandibular and clypeal structure, shape of fore wing sclerites, and surface sculpture of mesopleuron, but can be confused with E. kakanicus sp. nov., E. longitarsis, and E. tangi sp. nov. However, E. yonezawanus is distinguishable from E. kakanicus sp. nov. by the complete lateral longitudinal carinae of the scutellum (lateral longitudinal carinae of the scutellum posteriorly absent in E. kakanicus sp. nov., as in Figure 2F), from E. longitarsis and E. tangi sp. nov. by the scattered spines on the outer fore tibial surface (spines rather dense in E. longitarsis and E. tangi sp. nov.) and the separated proximal and distal sclerites (Fig. 27F) (proximal and distal sclerites confluent in E. longitarsis and E. tangi sp. nov. as in e.g. Figure 25F).

Figure 27. 

Enicospilus yonezawanus (Uchida, 1928), ♀. A. Habitus; B. Head, frontal view; C. Head, lateral view; D. Head, dorsal view; E. Mesosoma, lateral view; F. Central part of fore wing.

8♀♀3♂♂: Nepal (3♀♀2♂♂), China (2♀♀1♂), Myanmar (3♀♀).

Type series: holotype of Enicospilus zebrus Gauld & Mitchell, 1981, ♀, Mt Victoria (2,800 m), Myanmar, V.1938, G. Heinrich leg. (EMUS); paratypes of E. zebrus, 2♀♀, same data as holotype except for 2,400 m (NHMUK and EMUS).

Non-type series: 1♂, Choche Lekh (3,500 m), Chautara Dist., Nepal, 17.VI.1983, G. Robinson leg.; 1♂, Phulchoki peak (2,700 m), Nepal, X.1983, M.G. Allen leg. (LT); 1♀, Phulchoki (2,500 m), Nepal, IX.1982, M.G. Allen leg. (LT) (Fig. 28); 1♀, montane & oak forest (2,760 m), Phulchoki, Nepal, VIII.1983, M.G. Allen leg. (LT); 1♀, Nauling Lekh (9,000′), Gobre, Nepal, VI.1983, M.G. Allen leg. (LT); 2♀♀1♂, Yu Lung Mountain (3,200 m), Likiang, Yunnan, P.R. China, 15–21.VI.2009, A.C. Galsworthy leg. (LT) (all NHMUK).

Oriental region (Yu et al. 2016). Newly recorded from Nepal.

Head (Fig. 28B–D): GOI = 3.0–3.2; lower face 0.6–0.7× as wide as high; clypeus slightly convex in profile, its lower margin acute; mandible weakly twisted by 10–20°, moderately long, proximally tapered and distally more or less parallel sided, its outer surface without a diagonal structure; upper mandibular tooth 1.2–1.3× as long as lower one; posterior ocellus almost touching eye; antenna with 58–61 flagellomeres and 20^th flagellomere 2.6–2.7× as long as wide.

Mesosoma (Fig. 28E): mesopleuron punctate to rather coarsely longitudinally striate; scutellum with lateral longitudinal carinae reaching posterior end and convergent posteriorly; metapleuron rather coarsely striate; propodeum evenly weakly rounded, its posterior area more or less coarsely irregularly wrinkled with strong posterior transverse carina laterally, outer margin of propodeal spiracle joining pleural carina by a ridge.

Wings (Fig. 28F): fore wing with AI = 0.5, CI = 0.3–0.4, ICI = 0.4–0.5, SDI = 1.4–1.5; fore wing vein 1m-cu&M very slightly sinuous, 2r&RS almost straight; fenestra and sclerites of discosubmarginal cell of fore wing as in Figure 28F; fenestra of fore wing very long and its anterodistal corner very close to proximal end of vein RS; proximal sclerite triangular, confluent with distal one, strongly pigmented; central sclerite moderately pigmented and sclerotised, ill-delineated semicircular to oval, its major axis parallel to distal margin of fenestra, positioned in very distal and slightly anterior part of fenestra; distal sclerite entirely moderately pigmented; proximal corner of marginal cell of fore wing uniformly setose; vein 1cu-a antefurcal to M&RS by 0.1× 1cu-a length.

Colour (Fig. 28): body entirely black with pale yellow patterns, interocellar area not infuscate; wings hyaline but fore wing with three strongly infumate areas around anterocentral part of discosubmarginal cell, proximal part of second discal cell, and central part of marginal cell.

Gauld and Mitchell (1981) suggested that E. zebrus is related to the E. signativentris species-group and very close to E. biumbratus (Morley, 1912) on body and wing colour pattern as well as other characters, but E. zebrus is distinguished from E. biumbratus by many characters, such as the longer fore wing fenestra (Fig. 28F), smaller and semicircular to oval central sclerite (Fig. 28F), etc.

Figure 28. 

Enicospilus zebrus Gauld & Mitchell, 1981, ♀. A. Habitus; B. Head, frontal view; C. Head, lateral view; D. Head, dorsal view; E. Mesosoma, lateral view; F. Central part of fore wing.

1 unsexed: Nepal.

1 unsexed, Phulchoki (2,000 m), Nepal, VIII.1982, M.G. Allen leg. (LT) (NHMUK) (Fig. 29).

The mandibular structure of this species indicates it is associated with the E. ramidulus complex. Enicospilus sp. 1 does not key out to any species in Gauld and Mitchell’s (1981) and Tang’s (1990) keys and is possibly an undescribed species. It may potentially be found to be closely related to E. choui Tang, 1990 or E. sinicus Tang, 1990. However, only one broken specimen is known, so I tentatively treat this species as a species inquirenda.

Figure 29. 

Enicospilus sp. 1, unsexed, ♀. A. Habitus; B. Head, frontal view; C. Head, lateral view; D. Head, dorsal view; E. Mesosoma, lateral view; F. Central part of fore wing.

1♂: Nepal.

1♂, Terai (200 m), Chitwan, Nepal, 12–13.III.1983, M.G. Allen leg. (NHMUK) (Fig. 30).

The material examined is not in bad condition except for incomplete antennae. However, antennal characters are often useful and important for distinguishing Enicospilus species, as previous studies suggested (e.g. Broad and Shaw 2016). Therefore, antennae should be complete to describe a new species.

The affinities of this species are not clear, but, as with Enicospilus sp. 1, it also does not key out to any species in Gauld and Mitchell’s (1981) or Tang’s (1990) keys, indicating that it is potentially an undescribed species.

Figure 30. 

Enicospilus sp. 2, ♂. A. Habitus; B. Head, frontal view; C. Head, lateral view; D. Head, dorsal view; E. Mesosoma, lateral view; F. Central part of fore wing.

8♀♀19♂♂: Nepal.

1♀1♂, Godaveri (1,550–1,700 m), Nepal, VI (1♂), IX (1♀).1983, M.G. Allen leg. (LT); 1♂, Chautasa (6,000′), Nepal, 24.IX.1983, M.G. Allen leg. (Fig. 31A, B); 1♀, Kakani (2,070 m), Nepal, VII.1983, M.G. Allen leg. (LT); 1♀, secondary vegetation (1,500 m), Kathmandu, Nepal, 10.VI.1984, M.G. Allen leg. (LT); 1♂, Godaveri (1,550–1,700 m), Nepal, 5.VIII.1984, M.G. Allen leg. (LT) (Fig. 31C, D); 1♀, Kakani (2,070 m), Nepal, VII.1983, M.G. Allen leg. (LT); 3♀♀15♂♂, Kathmandu (1,300 m), Nepal, X (3♂♂), XI (3♀♀12♂♂).1982, M.G. Allen leg. (LT) (Fig. 31E, F, 1♀); 1♀, Kakani (2,070 m), Nepal, VII.1983, M.G. Allen leg. (LT); 1♂, Pokhara (950 m), Nepal, M.G. Allen leg. (LT) (all NHMUK).

The E. erythrocerus species-group is moderately large and one of the most taxonomically confusing groups within Enicospilus. It consists of rather large wasps with the fore wing fenestra lacking any trace of sclerites, SDI more than 1.2, lateral longitudinal carinae of the scutellum almost always reaching the posterior end, moderately sized fore wing fenestra, etc. In this study, I examined 27 Nepalese specimens of this species-group and recognised at least three morphospecies (Fig. 31). However, further research is needed to identify or describe them. Therefore, I tentatively treat all specimens of the E. erythrocerus species-group as species inquirenda.

Figure 31. 

Nepalese specimens of the Enicospilus erythrocerus species-group spp. A-B. ♂ from Chautasa: A. habitus, B. central part of fore wing; C-D. ♂ from Godaveri: C. habitus, D. central part of fore wing; E-F. ♀ from Kathmandu: E. habitus, F. central part of fore wing.