Afrotropical, Australasian, Holarctic, Neotropical, Oceanic, and Oriental regions (Yu et al. 2016).

According to the available evidence, species of Enicospilus are koinobiont endoparasitoids of usually late instar Lepidoptera larvae, but sometimes ovipositing in early instars (see summary in Broad et al. 2018). Many lepidopteran families, such as Noctuidae, Notodontidae and Saturniidae, are recorded as hosts (e.g., Gauld and Mitchell 1978, 1981; Gauld 1988; Broad and Shaw 2016). They are frequently collected at light and usually considered to be nocturnal or crepuscular (e.g., Gauld and Mitchell 1981; Gauld 1988; Shimizu and Maeto 2016; Shimizu 2017). Some Enicospilus are parasitoids of economically important Lepidoptera pests (e.g., Nagatomi et al. 1972; Kusigemati 1976). For instance, Enicospilus signativentris (Tosquinet, 1903) is a parasitoid of Poaceae pests, such as the noctuid moths Anadevidia peponis Fabricius, 1775, Autographa nigrisigna Walker, 1857 and Trichoplusia intermixta (Warren, 1913) (Kusigemati 1976, 1981; Kusigemati and Tanaka 1992); and E. sakaguchii (Matsumura & Uchida, 1926) is also known as a parasitoid of the rice pests Sesamia turpis (Butler, 1879) and S. inferens (Walker, 1856) (Nagatomi et al. 1972). Hence, Ophioninae could potentially be useful for agriculture as biocontrol agents (e.g., Gauld and Mitchell 1978, 1981; Gauld 1988).

Enicospilus species are moderately to very large insects, fore wing length usually 10.0–30.0 mm, with ophionoid facies. Easily distinguishable from other Ophioninae by the following characters: fore wing discosubmarginal cell with extensive glabrous area (fenestra), often with one or more sclerites (e.g., Fig. 4); mandibles narrow, slightly to strongly twisted (e.g., Fig. 2); inner surface of fore tibial spur lacking membranous flange. Enicospilus species can be confused with the genus Dicamptus but easily distinguished by the weakly to strongly tapered and twisted mandible (mandible very weakly tapered and never strongly twisted in Dicamptus). A key to the Japanese genera of Ophioninae has also been provided by Shimizu and Watanabe (2017).

Middle- to large-sized wasps (fore wing length usually 10.0–30.0 mm).

Head. Clypeus flat to strongly convex in profile, ventral margin acute, blunt, or impressed. Mandible weakly to strongly tapered and twisted, usually moderately long, outer surface with or without diagonal setose groove or line of punctures, and bidentate apically. Frons, vertex and gena shiny and smooth. Ocelli usually very large and posterior ocellus often close to or touching eye. Occipital carina usually complete, ventrally reaching oral carina or not. Antennae usually longer than fore wing, with usually more than 50 flagellomeres.

Mesosoma entirely weakly to moderately shiny with setae. Pronotum finely punctate or diagonally wrinkled and not specialised. Mesoscutum shiny and punctate to smooth with setae, evenly rounded in profile, and notauli usually absent. Scutellum moderately convex and usually with lateral longitudinal carinae. Epicnemium usually densely punctate with setae. Epicnemial carina present, straight to curved, inclined to curved to anterior margin of mesopleuron. Posterior transverse carina of mesosternum usually complete. Propodeum evenly rounded or declivous in profile; anterior transverse carina usually complete; anterior area longitudinally striate; spiracular area usually smooth; posterior area reticulate, wrinkled, striate, or rugose; and posterior transverse carina usually absent.

Wings. Fore wing pterostigma fairly slender; vein 1m-cu&M evenly curved, angulate or sinuate, usually without a ramulus; vein 2r&RS usually more or less widened and sinuate; discosubmarginal cell usually with bare fenestra, often with one or more sclerotised sclerites. Hind wing vein RS usually straight and rarely weakly curved; vein RA usually with 4–12 uniform hamuli.

Legs. Inner mesal surface of fore tibial spur without membranous flange. Outer distal margin of mid and hind trochantelli usually simple without decurved tooth. Hind tarsal claw moderately to strongly curved and usually simply pectinate.

Metasoma very slender. Spiracle of T1 far behind middle. Thyridium well developed. Ovipositor straight and almost always not longer than posterior depth of metasoma.

Colour. General body colour usually entirely testaceous, with posterior metasomal segments sometimes darker, but body sometimes entirely dark brown to black or pale. Wings usually entirely hyaline or weakly infuscate, but wings with strong infumate area in a few species; fenestra always hyaline; sclerites weakly to strongly pigmented amber.

We summarise the especially important diagnostic characters to identify Enicospilus species below.

Head (Fig. 2). The head provides many good characters to define species, as many previous authors have indicated (e.g., Gauld and Mitchell 1978, 1981; Gauld 1988; Schwarzfeld and Sperling 2014; Johansson and Cederberg 2019). Among them, the width of lower face as well as of clypeus, colour of interocellar area (or stemmaticum), shape of clypeus, and mandibular characters are especially useful and easy to use.

Width of the lower face is usually stable within a species group and/or species, even if the species is widespread, and sometimes provides enough gaps between species, although a few species, such as E. capensis, exhibit considerable variation.

Although body colour can be very variable within species, the colour of the interocellar area is usually stable at the species level and a good diagnostic character.

The shape of the clypeus is also very useful. For instance, the nasute clypeus is one of the most critical diagnostic characters of E. riukiuensis and related species (Fig. 40D), and the flat and projecting clypeus of E. sakaguchii is distinctive (Fig. 41D). The shape of the ventral margin, i.e., acute, blunt, or impressed, is also a very useful diagnostic character.

Features of mandibles are some of the most important diagnostic characters of Enicospilus species. First, the outer mandibular surface sculpture, especially presence or absence of a diagonal setose groove or line of punctures between the dorsoproximal corner and base of the apical teeth, is important. For example, the outer mandible surface of E. ramidulus has a diagonal setose groove (Fig. 39B, D), but of E. pungens is smooth (Fig. 38B, D). Second, the torsion of the mandible is a useful character, although it is rather difficult to measure. For instance, the strongly twisted mandible of E. acutus sp. nov. is one of the most important diagnostic characters for this species (Fig. 11B, D). Finally, length and shape also provide good characters. For instance, the mandible of E. shikokuensis is very long, slender, strongly tapered proximally, and subparallel-sided distally (Fig. 44B, D), but of E. sakaguchii is very short, stout, and evenly tapered (Fig. 41B, D).

Some mandibular diagnostic characters at the species level, such as degree of torsion and length of teeth are possibly adaptive characters and have been considered to be related to modes of emergence from host insects; hence, these characters are usually easily modified and not phylogenetically restricted, so it is indeed useful for species level taxonomy.

Mesosoma (Fig. 3). Mesosomal characters are also very informative. Surface microsculuptures of meso- and metapleuron are rather stable within species and also show large gaps between species and/or species groups. For example, the mesopleuron is coarsely longitudinally striate and metapleuron coarsely rugose in E. nigristigma (Fig. 31E), but the meso- and metapleuron are evenly moderately punctate and strongly shiny in E. unctus sp. nov. (Fig. 50E). Propodeum characters are useful as well. The posterior transverse carina of the propodeum of E. signativentris is unique in the Japanese species (Fig. 46E), and globally few Enicospilus species possess this carina. In other cases, the posterior area can be entirely densely punctate to finely reticulate in E. limnophilus sp. nov. (Fig. 25E), but coarsely concentrically striate in E. insinuator (Fig. 19E), providing an easy means to separate them. In this way, the propodeum is useful for definition of the species or species group when combined with other characters. Characters of the scutellum are also sometimes good for species recognition. For example, the quadrate scutellum of E. formosensis contributes to its identification, with the scutellum more or less trapezoidal or triangular in most Enicospilus species. Additionally, the length of the lateral longitudinal carinae of the scutellum is almost always stable within species and useful, with very rare exceptions, such as in E. limnophilus sp. nov., that exhibits a very wide range of variation (length of the carinae varies from 0.1–1.0× scutellum length).

Wings (Fig. 4). Wing characters have probably been regarded as the most important diagnostic characters in nocturnal Ophioninae by many previous authors (e.g., Gauld 1977; Gauld and Mitchell 1981; Shimizu 2020). These are much easier to use than other characters and can easily be measured.

First, the number, shape, and position of sclerites of the fore wing fenestra is usually very useful in Enicospilus. The number of sclerites varies in Enicospilus species from zero to four, but is nearly always stable within a species. The shape and position of sclerites are also very diverse, and some previous research has suggested that some species exhibit a wide range of intraspecific variation (e.g., Gauld and Mitchell 1981). However, it is stable in many cases and, if a species shows a wide range of intraspecific variation, it is likely that cryptic species are involved.

Second, the shape and setosity of the fore wing fenestra is sometimes a very useful character. For example, among Japanese species, the fenestra of E. nigribasalis and E. stenophleps is long and its anterodistal corner interstitial to fore wing vein RS (Figs 30F, 47F), but in other Japanese species the fenestra is shorter and its anterodistal corner clearly antefurcal to RS (e.g., Figs 25F, 36F).

Finally, the length and shape of wing veins also offer very good diagnostic characters. For example, the shape of fore wing veins 1m-cu&M and 2r&RS, the position of fore wing vein 1cu-a, and values of indices (e.g., AI, CI, ICI) are useful to distinguish species.

Legs. Legs do not seem to provide many useful characters, but some characters, such as the density of spines on the outer surface of the fore tibia, and pectination of the hind tarsal claw, are useful in species definition. For example, E. maruyamanus and E. pudibundae are very difficult to distinguish from each other, but the hind tarsal claw of the former is entirely uniformly pectinate, and the latter is not pectinate proximally.

Metasoma (Fig. 5). The shape of the first metasomal segment (e.g., sinuous or straight in profile, slender or stout) is useful (Broad and Shaw 2016; Johansson and Cederberg 2019).

Body size. Measuring body length is rather difficult due to the wide range of contraction or expansion of metasomal segments after death, hence fore wing length is a more useful character. However, body size shows a very wide range of variation in many species, such as E. pseudoconspersae, although it is stable in a few species (e.g., E. concentralis and E. nigronotatus). Therefore, this character is occasionally useful for species definition.

Colour. As mentioned above, this character can easily change intraspecifically. For instance, E. nigropectus and E. signativentris show a considerable range of colour variation (body entirely blackish to entirely testaceous). However, this character is stable within some species (e.g., E. acutus sp. nov. and E. nigribasalis, as in Figs 11, 30). Hence, this is also a useful critical character, but we need to be careful when relying on colour pattern.

Total of 19 specimens (all ♀♀): Japan (1♀), Nepal (1♀), Sri Lanka (2♀♀), Taiwan (15♀♀).

Type series: HT ♀ of Ophion semiopacus Matsumura, 1912, Gyochi, TAIWAN, Matsumura leg. (SEHU).

Australasian, Eastern Palaearctic, and Oriental regions (Yu et al. 2016); this is a predominantly Oriental species.

Newly recorded from Nepal.

JAPAN: [Ryûkyûs] Okinawa (Shimizu and Maeto 2016; present study). This species is abundant in Taiwan and in other mountainous areas of the Oriental region, but only one Japanese specimen has been collected from Okinawa-hontô of the Ryûkyûs. This single Japanese individual could have been a wanderer from Taiwan or other southern areas.

Unknown.

The characteristic striking colour pattern of this species (i.e., T1–4 each anteriorly yellow-brown and posteriorly dark brown, as in Fig. 9A) is the most useful diagnostic character among the Japanese species of Enicospilus. Some non-Japanese species, such as E. zebrus Gauld & Mitchell, 1981, have a similar colour pattern, but E. abdominalis is distinguishable by many characters, such as shape and size of fore wing fenestra and sclerites.

This species has been confused with E. nigropectus by many authors (cf. Gauld and Mitchell (1981)) but is easily distinguished from E. nigropectus by presence of central sclerite of fore wing fenestra, as in Fig. 9F (central sclerite absent in E. nigropectus, as in Fig. 33F), larger value of SDI (1.3–1.4 in this species, as in Fig. 9F, but 0.9–1.1 in E. nigropectus, as in Fig. 33F), yellowish interocellar area (Fig. 9B, C) (interocellar area usually blackish in E. nigropectus, as in Fig. 33B, C), etc. Enicospilus abdominalis is also morphologically similar to E. signativentris but can be distinguished by striking colour pattern and absence of distinct posterior transverse carina of propodeum.

Figure 9. 

Enicospilus abdominalis (Szépligeti, 1906) ♀ from Japan A habitus B head, frontal view C head, dorsal view D head, lateral view E mesosoma, lateral view F central part of fore wing.

Total of 54 specimens (29♀♀24♂♂ and 1 unsexed): India (17♀♀5♂♂), Japan (8♀♀17♂♂ and 1 unsexed), Solomon Islands (2♀♀), Sri Lanka (1♀1♂), Taiwan (1♀), Vanuatu (1♂).

Type series: LCT ♀ of Henicospilus okinawensis Matsumura & Uchida, 1926, Okinawa, Ryûkyûs, JAPAN, 1925, S. Sakaguchi leg. (SEHU); HT ♀ of Enicospilus crucis Chiu, 1954, Jûjiro, TAIWAN, 26.IV.1931, T. Shiraki leg. (TARI); HT ♂ of Henicospilus neddenveli Cheesman, 1936, Malekula, New Hebrides [VANUATU], III.1930, L.E. Cheesman leg. (NHMUK, Type 3b.1242).

Australasian, Eastern Palaearctic, Oceanic and Oriental regions (Yu et al. 2016).

JAPAN: [Kyûshû] Fukuoka* and Kagoshima (Momoi 1970; present study); [Ryûkyûs] Kagoshima (Uchida 1956; Momoi 1970; present study) and Okinawa (Matsumura and Uchida 1926; Uchida 1928; present study). *New record. Enicospilus aciculatus is a predominantly (sub-)tropical species and rare in the cooler zone.

Although this is one of the most common species in the Oriental region, there are no host records.

This species is morphologically relatively close to E. laqueatus and E. yonezawanus. However, E. aciculatus is rather easily distinguished from all other species of Japanese Enicospilus by the following combination of character states: mandible with diagonal setose groove, and upper mandibular tooth more than 2.0× as long as lower (Fig. 10B, D); central sclerite of fore wing fenestra moderately sized, ill-defined, very weakly pigmented, and positioned in anterodistal corner of fenestra (Fig. 10F); and proximal angle of proximal sclerite of fore wing fenestra ca. 45° (Fig. 10F). This species is morphologically stable but sometimes exhibits colour variation (i.e., usually the metasoma is entirely orange brown, but rarely the posterior segments are black).

Figure 10. 

Enicospilus aciculatus (Taschenberg, 1875) ♀ from Japan A habitus B head, frontal view C head, dorsal view D head, lateral view E mesosoma, lateral view F central part of fore wing.

The specific name is derived from the characteristic longitudinal acute ridge of the mandibular outer margin.

Total of 87 specimens (66♀♀20♂♂ and 1 unsexed): Japan (65♀♀20♂♂ and 1 unsexed), Taiwan (1♀).

HT: ♀, Ōkuni-rindô (26°43'38.7"N, 128°12'34.4"E, 265 m alt.), Okuma, Kunigami Vil., Kunigami County, Okinawa-hontô, Okinawa Pref., Ryûkyûs, JAPAN, 30.VI–1.VII.2016, S. Shimizu et al. leg. (LT) (MNHA).

PT: 1♀, Kisokoma-kôgen, Kiso County, Nagano Pref., Kantô-Kôshin, JAPAN, 10.VII.1982, Y. Yoshida & S. Yoshimatsu leg. (LT) (NIAES); 4♀♀ and 1 unsexed, Kanaya, Shizuoka Pref., Tôkai, JAPAN, 14.VII.1947 (1♀), 5.VIII.1947 (1♀), 13.VIII.1947 (1♀), 25.IX.1955 (1♀), 30.IX.1955 (1 unsexed), J. Minamikawa leg. (NIAES-JMC); 1♀1♂, Mt. Iwamuro, E. Izu, Shizuoka Pref., Tôkai, JAPAN, 11.X.1966, T. Maenami leg. (NIAES); 4♀♀, Hachikita-kôgen (35°24'2"N, 134°32'5"E, 750 m alt.), Ōsasa, Muraoka, Kami City, Mikata County, Hyôgo Pref., Kinki, JAPAN, 6–7.VIII.2018, K. Sakagami leg. (LT) (2♀♀, NSMT, SEN17–DDBJ-LC484182; 2♀♀, NIAES, SEN16–DDBJ-LC484181); 4♀♀1♂, Kotochi (34.119 136.040, 850 m alt.), Kamikitayama Vil., Nara Pref., Kinki, JAPAN, 16–17.VII.2017, S. Fujie leg. (2♀♀, OMNH; 2♀♀1♂, MNHA); 1♀, Komatsu, Kitayama Vil., Wakayama Pref., Kinki, JAPAN, 12.VIII.2009, Ichikawa leg. (KPMNH); 3♀♀, Mt. Tsurugi-yama (33°51'50.8"N, 134°04'42.4"E, 1,250 m alt.), Higashiyasugeoi, Miyoshi City, Tokushima Pref., Shikoku, JAPAN, 22.VIII.2015, Y. Kitayama leg. (LT) (2♀♀, NIAES; 1♀, SEHU); 1♀, Rainbow Highland, Sugasawa, Matsuyama City, Ehime Pref., Shikoku, JAPAN, 27.VII.2014, R. Okano leg. (LT) (EUM, SEN78–DDBJ-LC484183); 1♀1♂, Tengu-kôgen (33°28'38.7"N, 133°0'19.3"E, 1,360 m alt.), Yoshiunotsu, Tsuno Town, Takaoka County, Kôchi, Shikoku, JAPAN, 10.IX.2016, D. Funamoto leg. (LT) (NIAES); 1♀, Tengu-kôgen, Yoshiunotsu, Tsuno Town, Takaoka County, Kôchi Pref., Shikoku, JAPAN, 13.VIII.2015, Y. Nakatani leg. (EUM); 1♀, Tengu-kôgen, Kôchi, Shikoku, JAPAN, 11.VIII.1998, I. Yamashita leg. (LT) (NIAES); 5♂♂, Mt. Eboshi, Haruno, Kôchi, Shikoku, JAPAN, 10.VII.1991 (1♂), 11.VII.1991 (4♂♂), I. Yamashita leg. (NIAES); 6♀♀, Teragawa, Hongawa Vil., Kôchi Pref., Shikoku, JAPAN, 22.VII.1989 (1♀), 11.IX.1989 (1♀), 25.VII.1991 (3♀♀), 18.VII.1993 (1♀), I. Yamashita leg. (NIAES); 1♀, Jigokudani, Kôchi City, Kôchi Pref., Shikoku, JAPAN, 11.VIII.1992, I. Yamashita leg. (LT) (NIAES); 1♀1♂, Mt. Hikosan, Fukuoka Pref, Kyûshû, JAPAN, 29.VIII.1983, K. Ōhara leg. (LT) (NIAES); 1♀1♂, Mt. Hiko-san, Fukuoka Pref., Kyûshû, JAPAN, 22.IX.1983 (1♀), 20–21.X.1983 (1♂), M.T. Chûjô leg. (LT) (NIAES); 1♀1♂, Mt. Hikosan, Fukuoka Pref., Kyûshû, JAPAN, 5.VII.1983 (1♂), 6.VII.1983 (1♀), R. Noda leg. (LT) (NIAES); 1♂, Hiko-san, Fukuoka Pref., Kyûshû, JAPAN, 1.VIII.1953, S. Kimoto leg. (MNHA-SMCM); 1♀, Hiko-san, Fukuoka Pref., Kyûshû, JAPAN, 5.VIII.1955 (MNHA-SMCM); 1♀, Mt. Hiko-san, Fukuoka Pref., Kyûshû, JAPAN, 18–25.VIII.1989, K. Takeno & M. Sharkey leg. (EMUS); 1♀, Nokonoshima, Fukuoka Pref., Kyûshû, JAPAN, 12.IX.1983, K. Konishi leg. (LT) (NIAES); 1♀, Fukuoka Pref., Kyûshû, JAPAN, 30.VIII.1983, K. Ōhara leg. (LT) (NIAES); 1♀, Mt. Shiratori, Izumi Vil., Kumamoto Pref., Kyûshû, JAPAN, 6.VII.1991, R. Noda leg. (NIAES); 1♀, Shiiya pass (1,400 m alt.), Kumamoto Pref., Kyûshû, JAPAN, 15.VI.1985, K. Konishi et al. (LT) (NIAES); 1♀, Mt. Kurodake, Mts. Kujû, Ōita Pref., Kyûshû, JAPAN, 3.X.1983, N. Kôda & R. Noda leg. (LT) (NIAES); 1♀, Mt. Kurodake, Mts. Kujû, Ōita Pref., Kyûshû, JAPAN, 15.IX.1985, K. Konishi leg. (LT) (NIAES); 1♂, Mt. Hoyoshi-dake, Kouyama Town, Kagoshima Pref., Kyûshû, JAPAN, 12–14.VII.1991, R. Noda leg. (LT) (NIAES); 3♀♀, Anbô-rindô (350 m alt.), Yakushima, Kagoshima Pref., Kyûshû, JAPAN, 22.VII.1982, K. Konishi & S. Yoshimatsu leg. (LT) (NIAES); 1♂, Shiratani-unsuikyô, Yakushima, Kagoshima Pref., Kyûshû, JAPAN, 25.VII.1982, K. Konishi & S. Yoshimatsu leg. (LT) (NIAES); 1♀, Chûô-rindô, Amami-ôshima, Kagoshima Pref., Ryûkyûs, JAPAN, 19.IX.1993, M. Yoshida leg. (LT) (NIAES); 1♀, Mt. Yui-dake, Amami-ôshima, Kagoshima Pref., Ryûkyûs, JAPAN, 14.X.2004, H. Makihara leg. (MsT) (KPMNH); 1♀, Kamiya, Amami-ôshima, Kagoshima Pref., Ryûkyûs, JAPAN, 30.VI.1992, R. Noda leg. (NIAES); 1♀, Higashinakama, Amami-ôshima, Kagoshima Pref., Ryûkyûs, JAPAN, 13.V.1983, S. Yoshimatsu & S. Nomura leg. (LT) (NIAES); 1♀, Mt. Yuidake, Amami-ôshima, Kagoshima Pref., Ryûkyûs, JAPAN, 1.IX.1983, R. Noda & K. Hirano leg. (LT) (NIAES); 1♂, Hiji agricultural road (26°43'16.8"N, 128°10'43.4"E, 85 m alt.), Hiji, Kunigami Vil., Kunigami County, Okinawa-hontô, Okinawa Pref., Ryûkyûs, JAPAN, 3–4.VII.2016, S. Shimizu et al. (LT) (NIAES); 2♀♀, Ōkuni-rindô (26°43'38.7"N, 128°12'34.4"E, 265 m alt.), Okuma, Kunigami Vil., Kunigami County, Okinawa-hontô, Okinawa Pref., Ryûkyûs, JAPAN, 30.VI–1.VII.2016, S. Shimizu et al. (LT) (SEHU); 1♀, Ōkuni bridge (26.704179 128.197087, 200 m alt.), Ōkuni-rindô, Hama, Kunigami Vil., Kunigami County, Okinawa-hontô, Okinawa Pref., Ryûkyûs, JAPAN, 6.IV.2019, S. Shimizu leg. (LT) (TARI); 1♀2♂♂, Oku-nigô-rindô (26.815735 128.297288, 200 m alt.), Sosu, Kunigami Vil., Kunigami County, Okinawa-hontô, Okinawa Pref., Ryûkyûs, JAPAN, 7.V.2019, S. Shimizu & T. Tokuhira leg. (LT) (1♀1♂, EMUS; 1♂, CNC); 9♀♀, Ōkuni bridge (26.704179 128.197087, 200 m alt.), Ōkuni-rindô, Hama, Kunigami Vil., Kunigami County, Okinawa-hontô, Okinawa Pref., Ryûkyûs, JAPAN, 29–30.III.2019, S. Shimizu leg. (LT) (2♀♀, NSMT; 5♀♀, KPMNH; 2♀♀, NIAES); 1♀, Okuyona-rindô (26.783608 128.282954, 240 m alt.), Ada, Kunigami Vil., Kunigami County, Okinawa-hontô, Okinawa Pref., Ryûkyûs, JAPAN, 7.IV.2019, S. Shimizu leg. (LT) (NIAES); 2♀♀1♂, Sate (26.774333 128.234803, 230 m alt.), Kunigami Vil., Kunigami County, Okinawa-hontô, Okinawa Pref., Ryûkyûs, JAPAN, 5–6.V.2019, S. Shimizu et al. (LT) (1♀1♂, MNHA; 1♀, NHMUK); 1♂, Janagusuku-rindô (26.688693 128.172478, 200 m alt.), Janagusuku, Ōgimi Vil., Kunigami County, Okinawa-hontô, Okinawa Pref., Ryûkyûs, JAPAN, 8.V.2019, S. Shimizu et al. (LT) (CNC); 1♂, Hama-ichigô-rindô (26.708926 128.181583, 200 m alt.), Hama, Kunigami Vil., Kunigami County, Okinawa-hontô, Okinawa Pref., Ryûkyûs, JAPAN, 7.V.2019, S. Shimizu & T. Tokuhira leg. (LT) (NHMUK); 1♀, Pingtung County (22°25'3"N, 120°43'16"E, 920 m alt.), Chunri Township, TAIWAN, 3.X.2015, S. Shimizu & M. Ito leg. (LT) (TARI, SEN130–DDBJ-LC484184).

Eastern Palaearctic and Oriental regions.

JAPAN: [Kantô-Kôshin] Nagano; [Tôkai] Shizuoka; [Kinki] Hyôgo, Nara, and Wakayama; [Shikoku] Tokushima, Ehime, and Kôchi; [Kyûshû] Fukuoka, Ōita, Kumamoto, and Kagoshima; [Ryûkyûs] Kagoshima and Okinawa. This is a rather common species in western and southern Japan.

Unknown.

This species resembles E. maai in colour pattern but can be readily distinguished by the following characters: mandible strongly twisted ca. 85° so that outer margin forming acute median longitudinal ridge between near centro-proximal part of mandible and base of mandibular apical teeth (Fig. 11B, D) (mandible more or less weakly twisted less than 35° so that outer mandibular surface without any ridge or groove in E. maai); fore wing with AI more than 1.0 (Fig. 11F) (AI less than 0.7 in E. maai); fore wing fenestra with strongly pigmented central sclerite (Fig. 11F) (central sclerite absent in E. maai); and mesopleuron closely striate to punctostriate (Fig. 11E) (mesopleuron coarsely striate in E. maai). This species is uniquely distinctive and easily distinguishable within the Japanese Enicospilus on account of the mandibular morphology and colour pattern.

Female (n = 66). Body length 23.5–26.0 (HT: ca. 24.0) mm.

Head with GOI = 2.7–3.3 (HT: 2.8) (Fig. 11D). Lower face 0.7× as wide as high, strongly shiny, finely longitudinally striate centrally, and finely punctate with setae laterally (Fig. 11B). Clypeus 1.5–1.7× (HT: 1.5) as wide as high, sparsely finely punctate with setae, weakly convex in profile, and ventral margin acute (Fig. 11B, D). Malar space 0.4–0.5× (HT: 0.4) as long as basal mandibular width (Fig. 11B, D). Mandible strongly twisted by 80–85° (HT: ca. 85°), moderately long, evenly narrowed, outer surface without diagonal setose groove but with longitudinal acute ridge between near centroproximal part of mandible and base of mandibular teeth (Fig. 11B, D). Mandible teeth equal length, but upper tooth stouter than lower (Fig. 11B). Frons, vertex and gena strongly shiny with fine setae (Fig. 11B–D). Posterior ocellus close to eye (Fig. 11B–D). Ventral end of occipital carina not joining oral carina. Antenna with 54–60 (HT: 56) flagellomeres; first flagellomere 1.6–1.9× (HT: 1.6) as long as second; 20^th flagellomere 1.8–2.0× (HT: 1.8) as long as wide.

Mesosoma entirely weakly to moderately shiny with setae (Fig. 11E). Pronotum finely punctate or diagonally wrinkled (Fig. 11E). Mesoscutum 1.4–1.5× (HT: 1.4) as long as maximum width, rather strongly shiny and finely punctate to smooth with setae, and evenly rounded in profile (Fig. 11E). Notauli absent (Fig. 11E). Scutellum moderately convex, smooth with some irregular rugae, with lateral longitudinal carinae along entire length of scutellum. Epicnemium densely punctate with setae. Epicnemial carina strong, straight or very slightly curved, inclined to anterior, dorsal end not reaching anterior margin of mesopleuron (Fig. 11E). Mesopleuron entirely longitudinally striate to punctostriate (Fig. 11E). Submetapleural carina broadened anteriorly (Fig. 11E). Metapleuron coarsely rugose (Fig. 11E). Propodeum evenly rounded in profile; anterior transverse carina complete centrally and not joining pleural carina laterally; anterior area longitudinally striate; spiracular area smooth; posterior area rather coarsely rugose; propodeal spiracle elliptical and joining pleural carina by ridge (Fig. 11E).

Wings (Fig. 11F). Fore wing length 16.5–18.0 (HT: 16.5) mm with AI = 1.0–1.4 (HT: 1.4), CI = 0.4, DI = 0.3, ICI = 0.5, SDI = 1.3–1.4 (HT: 1.4), SI = 0.1–0.2 (HT: 0.1), SRI = 0.3; vein 1m-cu&M sinuate; vein 2r&RS slightly sinuate; vein RS rather evenly curved; fenestra and sclerites of discosubmarginal cell as in Fig. 11F; proximal sclerite triangular, not confluent with distal sclerite, strongly pigmented; central sclerite suboval, pigmented, positioned in centrodistal part of fenestra; distal sclerite pigmented; proximal corner of marginal cell evenly setose; posterodistal corner of second discal cell 90–100° (HT: ca. 100°) and of subbasal cell 80–85° (HT: ca. 80°); vein 1cu-a antefurcal to M&RS by 0.1–0.2× (HT: 0.2) length of 1cu-a. Hind wing with NI = 2.9–3.5 (HT: 2.9); vein RS straight; vein RA with 7–9 (HT: 8) uniform hamuli.

Legs. Hind legs with coxa in profile 1.7–1.8× (HT: 1.8) as long as deep; basitarsus 1.9–2.0× (HT: 2.0) as long as second tarsomere; fourth tarsomere 3.3–3.5× (HT: 3.5) as long as wide; tarsal claw simply pectinate.

Metasoma with DMI = 1.2–1.3 (HT: 1.3), PI = 3.3–3.5 (HT: 3.3), THI = 3.0–3.4 (HT: 3.3); thyridium oval; ovipositor sheath not longer than posterior depth of metasoma.

Colour (Fig. 11). Entirely light to rather dark brown except for apex of mandible, posterior part of mesoscutum, and central part of frons infuscate. Wings moderately infuscate; sclerites pigmented and amber; veins dark reddish brown.

Male (n = 20). Very similar to female.

Figure 11. 

Enicospilus acutus Shimizu, sp. nov. ♀ from Japan (PT) A habitus B head, frontal view C head, dorsal view D head, lateral view E mesosoma, lateral view F central part of fore wing.

Total of 112 specimens (66♀♀42♂♂ and 4 unsexed): Japan (1♀), India (57♀♀41♂♂), Kenya (2♀♀1♂ and 1 unsexed), Madagascar (1♀ and 1 unsexed), Malaysia (1♀), Saudi Arabia (1 unsexed), South Africa (1♀), Uganda (2♀♀ and 1 unsexed), Zimbabwe (1♀).

Type series: HT ♂ of Henicospilus yanagiharai Sonan, 1940, Kitadaitô-jima, Okinawa Pref., Ryûkyûs, JAPAN, 18.III.1939, M. Yanagihara leg. (TARI) (Fig. 12); HT ♀ of Enicospilus fossatus Chiu, 1954, Jahore, MALAYSIA, 1.X.1916, J. Sonan leg. (TARI); HT ♀ of Henicospilus euxoae Wilkinson, 1928, Salisbury, ZIMBABWE, 31.XII.1927, J.I. Roberts leg. (from Euxoa) (NHMUK, Type 3b.1289).

Afrotropical, Australasian, Oceanic, and Oriental regions (Yu et al. 2016).

JAPAN: [Ryûkyûs] Okinawa (Sonan 1940; present study).

This species has a very wide distribution from South East Asia to South Africa. According to Gauld and Mitchell (1981), this distribution pattern is hardly surprising when considering many of their host moths are also widely distributed throughout the Old World tropics. Enicospilus capensis is frequently encountered as a parasitoid of economically important noctuid moths; however, only a single specimen has been collected in Japan.

Recorded from various Lepidoptera hosts, but reliable records are mainly from Noctuidae (e.g., Gauld and Mitchell 1981; Nikam and Gaikwad 1989; Nikam 1990). No host records from Japan.

The Japanese specimen of E. capensis is very easily distinguishable from all other Japanese Enicospilus specimens on account of very wide face (i.e., lower face 1.2× as wide as high, as in Fig. 12B) and long mandible. This species is morphologically similar to E. ramidulus, but distinguishable by the following combination of morphological characters: metapleuron matt (Fig. 12E) (metapleuron evenly moderately punctate and never matt in E. ramidulus, as in Fig. 39E); metasoma usually entirely orange-brown (Fig. 12A) (posterior metasomal segments usually strongly infuscate in E. ramidulus, as in Fig. 39A). This species is usually morphologically rather uniform, but the Japanese specimen has a much broader lower face than others (Fig. 12B). However, there does not seem to be enough of a difference to justify a separate species, as Gauld and Mitchell (1981) also concluded.

Figure 12. 

Enicospilus capensis (Thunberg, 1824) (HT ♀ of E. yanagiharai (Sonan, 1940)) A habitus B head, frontal view C head, dorsal view D head, lateral view E mesosoma, lateral view F central part of fore wing.

Total of 54 specimens (39♀♀14♂♂ and 1 unsexed): England (18♀♀2♂♂), Japan (19♀♀10♂♂ and 1 unsexed), Russia (1♂), unknown (2♀♀1♂).

Oriental and Palaearctic regions (Yu et al. 2016); this is a predominantly Palaearctic species that may be restricted to there, although Lee et al. (2011) reported this species from the Oriental region, probably based on a misidentification.

JAPAN: [Hokkaidô] (Uchida 1928, 1935; present study); [Honshû] (Uchida 1935; present study); [Tôhoku] Aomori (Uchida 1928; present study), Yamagata*, and Fukushima (Uchida 1928; present study); [Hokuriku] Niigata (Uchida 1928; present study); [Kantô-Kôshin] Tochigi (Uchida 1928; present study), Nagano*, and Tôkyô (Uchida 1928, 1930); [Tôkai] Gifu (Uchida 1928); [Kinki] Kyôto* and Hyôgo*; [Chûgoku] Hiroshima*; [Shikoku] (Uchida 1935); [Kyûshû] (Uchida 1935), Fukuoka*. *New records.

Reared from one species of Noctuidae in Japan: Trachea tokiensis (Butler, 1884) (Uchida 1928, 1930). Notodontid and noctuid moths are recorded as hosts, but reliable records are only from Noctuidae of the subfamily Hadeninae (e.g., Broad and Shaw 2016).

This species is usually very easily distinguished from all other Palaearctic Enicospilus species by the black mesosoma, thyridium, and posterior segments of metasoma, as in Fig. 13A. Enicospilus combustus has sometimes been confused with E. multidens stat. rev., E. ramidulus, and E. shikokuensis; moreover, some authors have treated E. combustus and E. ramidulus as a single species (e.g., Viktorov 1957; Townes et al. 1965; Gauld and Mitchell 1981). However, E. combustus is easily separated from E. shikokuensis by the separated proximal and distal sclerites of fore wing fenestra, as in Fig. 13F (proximal and distal sclerites usually obviously confluent in E. shikokuensis, as in Fig. 44F), from E. multidens stat. rev. and E. ramidulus by the entirely more or less blackish mesosoma, as in Fig. 13A, E (mesosoma entirely orange-brown in E. multidens stat. rev. and E. ramidulus, as in Figs 29A, E and 39A, E respectively). Moreover, this species is similar to E. sharkeyi sp. nov. in colour pattern (Figs 13, 43), however, E. combustus can be readily distinguished from it by many characters, such as separated proximal and distal sclerites of fore wing fenestra, as in Fig. 13F (proximal and distal sclerites confluent in E. sharkeyi sp. nov., as in Fig. 43F), larger central sclerite of fore wing fenestra, as in Fig. 13F (central sclerite smaller in E. sharkeyi sp. nov., as in Fig. 43F), wider lower face, as in Fig. 13B (narrower in E. sharkeyi sp. nov., as in Fig. 43B), etc.

Figure 13. 

Enicospilus combustus (Gravenhorst, 1829) ♀ from Japan A habitus B head, frontal view C head, dorsal view D head, lateral view E mesosoma, lateral view F central part of fore wing.

Total of 59 specimens (51♀♀6♂♂ and 2 unsexed): Brunei (27♀♀2♂♂), Japan (22♀♀2♂♂ and 1 unsexed), Taiwan (2♀♀2♂♂ and 1 unsexed).

Australasian, Eastern Palaearctic, and Oriental regions (Yu et al. 2016).

JAPAN: [Kantô-Kôshin] Tôkyô* and Kanagawa*; [Kinki] Hyôgo*; [Chûgoku] Hiroshima (Konishi and Nakamura 2010; present study); [Shikoku] Tokushima*, Ehime (Konishi and Yamamoto 2000; present study), and Kôchi*; [Kyûshû] Fukuoka (Konishi 1993; present study), Nagasaki (Konishi 1993; present study), Ōita (Konishi 1993; present study), and Kumamoto (Konishi 1993; present study). *New records. Enicospilus concentralis is restricted to the warmer Pacific coast in Japan although has not been recorded from Ryûkyûs.

Hosts unknown.

Although adult wasps are most active during summer, it is also relatively easily encountered in winter: hibernating adults are often found on the underside of leaves of evergreen plants (such as Aucubaceae shrubs).

Wing characters of this species (e.g., fore wing with proximal part of marginal cell widely glabrous, CI = 0.1–0.3, ICI = 0.2–0.3, and central sclerite of fore wing fenestra linear and parallel to distal margin of fenestra, as in Fig. 14F) are unique within the genus; hence, this species can very easily be distinguished from all other species of Enicospilus.

Figure 14. 

Enicospilus concentralis Cushman, 1937 ♀ from Japan A habitus B head, frontal view C head, dorsal view D head, lateral view E mesosoma, lateral view F central part of fore wing.

Total of 49 specimens (44♀♀5♂♂): Japan (8♀♀3♂♂), Sri Lanka (2♀♀), Taiwan (34♀♀2♂♂).

Type series: HT ♀ of Eniscospilus (sic) dasychirae Cameron, 1905, Pundaluoya, SRI LANKA, I.1899, P. Cameron leg. (NHMUK, Type 3b.1267); HT ♀ of Eniscospilus (sic) horsfieldi Cameron, 1905, Peradeniya, SRI LANKA, IX.1902, P. Cameron leg. (NHMUK, Type 3b.1265).

Australasian, Eastern Palaearctic, and Oriental regions (Yu et al. 2016); this is a predominantly Oriental species.

JAPAN: [Ryûkyûs] Okinawa (Chiu 1954; Gauld and Mitchell 1981; present study). Though this species is widely distributed in Southeast Asia, it is restricted to Okinawa in Japan.

Recorded as a parasitoid of several species of Erebidae (subfamily Lymantriinae) (Gauld and Mitchell 1981; Chiu et al. 1984; Chen et al. 2009) and Noctuidae (Tang 1990), although there are no host records from Japan.

This species is readily distinguishable from all other Enicospilus species by the unique small drop-shaped and isolated proximal sclerite of fore wing fenestra, as in Fig. 15F.

Figure 15. 

Enicospilus dasychirae Cameron, 1905 ♀ from Japan A habitus B head, frontal view C head, dorsal view D head, lateral view E mesosoma, lateral view F central part of fore wing.

Total of 14 specimens (12♀♀2♂♂): Japan (2♀♀1♂), Malaysia (6♀♀), Philippines (1♀), Sri Lanka (3♀♀), Taiwan (1♂).

Type series: HT ♀ of Pleuroneurophion erythrocerus Cameron, 1905, Peradeniya, SRI LANKA (NHMUK, Type 3b.1214); LCT ♀ of Allocamptus orientalis Uchida, 1928, Okinawa Pref., Ryûkyûs, JAPAN, VII.1926, Hirayama leg. (SEHU).

Australasian, Eastern Palaearctic, and Oriental regions (Yu et al. 2016); this is a predominantly Oriental species.

JAPAN: [Ryûkyûs] Okinawa (Uchida 1928; present study).

Recorded from Erebidae by Sharma (1985).

This species especially resembles E. grandis (Cameron, 1905) and E. plicatus (Brullé, 1846), but is distinguishable by the smaller size, shorter antennae, and more matt and uniformly punctate meso- and metapleuron (Fig. 16A, E). Other than this species, all Japanese Enicospilus species have at least one fore wing sclerite; hence, it is fortunately very easily identifiable.

Allocamptus orientalis was described based on two females and one male from Okinawa and Taiwan (Uchida 1928). Although the Taiwanese specimen was designated as the lectotype by Gauld and Mitchell (1981), the lectotype label was attached to a Japanese specimen.

Figure 16. 

Enicospilus erythrocerus (Cameron, 1905) ♀ from Japan A habitus B head, frontal view C head, dorsal view D head, lateral view E mesosoma, lateral view F central part of fore wing.

Total of 57 specimens (34♀♀22♂♂ and 1 unsexed): Australia (5♀♀2♂♂ and 1 unsexed), Brunei (2♀♀1♂), Japan (21♀♀13♂♂), Laos (1♀), Singapore (1♀), Taiwan (4♀♀6♂♂).

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

Australasian, Oceanic, and Oriental regions (Yu et al. 2016); new to the Eastern Palaearctic region.

Newly recorded from Laos.

JAPAN: [Kyûshû] Kagoshima*; [Ryûkyûs] Kagoshima (Uchida 1956; present study) and Okinawa (Matsumura and Uchida 1926; Uchida 1928; Sonan 1940; present study). *New record.

Recorded as a parasitoid of three species of Euproctis (Erebidae: Lymantriinae) (Corbett and Miller 1928; Sonan 1944; Chen et al. 2009). Other host records seem less likely, and there are no host records from Japan.

This species is easily distinguishable from all other Enicospilus species by the angulate fore wing vein 1m-cu&M, large value of CI (i.e., 0.6–0.8), and characteristic sclerites of fore wing fenestra (Fig. 17F). Enicospilus flavocephalus is very similar to E. xanthocephalus in colour pattern, body size, and body profile, as in Figs 17 and 52, but these species are easily distinguishable from each other by the colour of interocellar area (entirely yellow-brown in E. flavocephalus, as in Fig. 17C, but entirely black in E. xanthocephalus, as in Fig. 52C), shape of fore wing vein 1m-cu&M (angulate in E. flavocephalus, as in Fig. 17F, but evenly curved in E. xanthocephalus, as in Fig. 52F), etc.

Figure 17. 

Enicospilus flavocephalus (Kirby, 1900) ♀ from Japan A habitus B head, frontal view C head, dorsal view D head, lateral view E mesosoma, lateral view F central part of fore wing.

Total of 55 specimens (19♀♀35♂♂ and 1 unsexed): Brunei (1♂), India (1 unsexed), Japan (18♀♀32♂♂), Laos (1♂), Taiwan (1♀), unknown (1♂).

Type series: HT ♀ of Henicospilus formosensis Uchida, 1928, Baibara [= Meiyuan], TAIWAN, 15.VI.1926, Y. Saito & Kikuchi leg. (SEHU); HT ♀ of Enicospilus vacuus Gauld & Mitchell, 1981, Chizuka, Okinawa, Ryûkyûs, JAPAN, VII–IX, G.E. Bohart & C.L. Harnage leg. (EMUS); PT ♀ of E. vacuus, same data and repository as HT; HT ♀ of Enicospilus saepis Chiu, 1954, Nara, Kinki, JAPAN, 17.VIII.1918, J. Sonan leg. (TARI).

Eastern Palaearctic and Oriental regions (Yu et al. 2016).

Newly recorded from Laos and Malaysia.

JAPAN: [Kantô-Kôshin] Tôkyô*; [Tôkai] Mie*; [Kinki] Ōsaka (Chiu 1954; present study), Nara (Chiu 1954; Shimizu 2020; present study), and Wakayama*; [Chûgoku] Hiroshima (Konishi and Nakamura 2005, 2010; present study) and Yamaguchi*; [Shikoku] Ehime (Konishi and Yamamoto 2000; present study) and Kôchi*; [Kyûshû] Fukuoka*, Saga (Chiu 1954; present study), Nagasaki*, Kumamoto (Chiu 1954; present study), and Kagoshima*; [Ryûkyûs] Kagoshima* and Okinawa (Gauld and Mitchell 1981; present study). *New records.

Unknown.

This species is easily distinguishable by the wide face (Fig. 18B), unique conspicuous long line of setae of fore wing discosubmarginal cell (Fig. 18F), shape of central sclerite of fore wing fenestra (Fig. 18F), quadrate scutellum, and large size.

Gauld and Mitchell (1981) had separated E. formosensis and E. vacuus based on differences of value of CI. However, the CI of these ‘species’ are continuous and no other morphological differences could be recognised. Tang (1990) also suggested that these names represented the same species. Hence, E. vacuus is newly synonymised under E. formosensis in the present paper.

Figure 18. 

Enicospilus formosensis (Uchida, 1928) ♀ from Japan A habitus B head, frontal view C head, dorsal view D head, lateral view E mesosoma, lateral view F central part of fore wing.

Total of 36 specimens (35♀♀1♂): Brunei (1♀), China (1♀), Japan (28♀♀1♂), Taiwan (5♀♀).

Type series: HT ♀ of Enicospilus zyzzus Chiu, 1954, Foochow, Fukien, CHINA, 13.II.1948, H.F. Chao leg. (TARI); PT ♀ of Enicospilus zyzzus, Oshima, Kyûshû, JAPAN, V.1930, Takahashi leg. (TARI).

Australasian, Eastern Palaearctic, and Oriental regions (Yu et al. 2016).

Newly recorded from Taiwan.

JAPAN: [Kyûshû] Kagoshima*; [Ryûkyûs] Kagoshima* and Okinawa*. * New records. Chiu (1954) has recorded this species from Japan, but lacking collection locality data. Most Japanese specimens were collected in Yakushima by Malaise traps.

Unknown.

This species is closest to E. pallidistigma Cushman, 1937, but distinguished from it by the smaller fore wing fenestra and weaker central sclerite (= quadra) (Fig. 19F). According to Gauld and Mitchell (1981), this species also resembles E. maai and E. rogus, but is easily distinguishable from them by the mesoscutum colour (i.e., mesoscutum with longitudinal vittae or rarely entirely infuscate in E. insinuator, but only posterior part of mesoscutum strongly infuscate in E. maai and E. rogus). Furthermore, it differs from E. maai by the shape of fore wing fenestra (i.e., fenestra moderate length and anterodistal corner of fenestra antefurcal to RS by more than 0.7× length of 2rs-m in E. insinuator, as in Fig. 19F; but fenestra very long and anterodistal corner of fenestra interstitial to RS in E. maai) and from E. rogus by the torsion of mandible (i.e., mandible twisted by 15–35° in E. insinuator, as in Fig. 19B, D; but 75–80° in E. rogus).

Figure 19. 

Enicospilus insinuator (Smith, 1860) ♀ from Japan A habitus B head, frontal view C head, dorsal view D head, lateral view E mesosoma, lateral view F central part of fore wing.

Total of 122 specimens (103♀♀19♂♂): Brunei (30♀♀2♂♂), India (2♀♀), Japan (64♀♀16♂♂), Papua New Guinea (4♀♀), Singapore (1♀), Sri Lanka (2♀♀), Taiwan (1♂).

Type series: HT ♀ of Enicospilus gephyrus Chiu, 1954, Tokusa, Yamaguchi Pref., Chûgoku, JAPAN, 1.VIII.1922, T. Shiraki leg. (TARI).

Australasian, Eastern Palaearctic, and Oriental regions (Yu et al. 2016).

JAPAN: [Kantô-Kôshin] Kanagawa (Watanabe et al. 2016); [Tôkai] Shizuoka* and Mie*; [Kinki] Kyôto*, Ōsaka*, and Hyôgo*; [Chûgoku] Hiroshima* and Yamaguchi (Chiu 1954; present study); [Shikoku] Tokushima*, Ehime (Konishi and Yamamoto 2000; present study), and Kôchi*; [Kyûshû] Fukuoka* and Kumamoto*; [Ryûkyûs] Kagoshima*. *New records. It is restricted to the warmer Pacific coast in Japan.

Unknown.

This species is very easily distinguishable from any other Japanese Enicospilus by its characteristic fenestra and sclerites of fore wing (i.e., fenestra widened proximally, and proximal and distal sclerites confluent and shaped like a letter ‘P’, as in Fig. 20F).

Enicospilus javanus exhibits a very wide range of variation in the shape of fore wing sclerites, body size, and colour, as mentioned by Gauld and Mitchell (1981: 262). We examined 24 female specimens from the same locality and date collected by I. D. Gauld in Brunei, and these specimens indeed show an extremely wide range of morphological variation. However, the variation is more or less continuous. This variation suggests that this species includes some cryptic species and integrative approaches are needed to reveal species boundaries. Regarding the Japanese population, it is very stable, suggesting that there are no cryptic species included.

Figure 20. 

Enicospilus javanus (Szépligeti, 1910) ♀ from Japan A habitus B head, frontal view C head, dorsal view D head, lateral view E mesosoma, lateral view F central part of fore wing.

Total of 3 specimens (2♀♀1♂): Japan (2♀♀1♂).

Eastern Palaearctic region (Tang 1990; Yu et al. 2016).

Newly recorded from Japan.

JAPAN: [Kantô-Kôshin] Tôkyô and Chiba. This species is collected only from a big city in Japan.

Unknown.

All Japanese specimens had been misidentified as E. yonezawanus, because they lack the central sclerite of fore wing fenestra and have a strongly pigmented triangular proximal sclerite, as in Figs 21F, 54F. However, E. jilinensis can be distinguished from E. yonezawanus by many characters, such as the smooth outer mandibular surface, smaller fore wing fenestra, entirely punctate mesopleuron, and larger value of ICI.

Some morphological features (e.g., flattened clypeus, moderately long and slender mandible, and absence of central sclerite of fore wing fenestra) suggest a relation to E. shinkanus, which should be tested when fresh specimens are available.

Figure 21. 

Enicospilus jilinensis Tang, 1990 ♀ from Japan A habitus B head, frontal view C head, dorsal view D head, lateral view E mesosoma, lateral view F central part of fore wing.

Total of 6 specimens (5♀♀1♂): Japan (5♀♀1♂).

Type series: HT ♀ of Enicospilus kikuchii Shimizu, 2017, Kawamata, Chichibu, Saitama Pref., Kantô-Kôshin, JAPAN, 28–30.VIII.2012, N. Kikuchi leg. (NIAES); PT ♂ of Enicospilus kikuchii, Eboshi-dake, Kagoshima Pref., Kyûshû, JAPAN, 4.V.1969, K. Kusigemati leg. (SEHU-KUSIG).

Eastern Palaearctic region (Shimizu 2017; present study).

JAPAN: [Hokuriku] Fukui*; [Kantô-Kôshin] Tochigi* and Saitama (Shimizu 2017; present study); [Shikoku] Ehime* and Kôchi*; [Kyûshû] Kagoshima (Shimizu 2017; present study). *New records.

Unknown.

As in Fig. 22, the characteristic colour pattern easily distinguishes this species from other Enicospilus species. Morphologically, E. kikuchii resembles E. melanocarpus, but is distinguishable by the following combination of character states: mesosoma, T1, T2, and T5–8 black (Fig. 22A, E) (usually most of body yellow-brown with black T5–8 in E. melanocarpus, as in Fig. 28A, E); and metapleuron roughly diagonally punctostrigose (Fig. 22E) (uniformly punctate or finely diagonally punctostriate in E. melanocarpus, as in Fig. 28E).

Figure 22. 

Enicospilus kikuchii Shimizu, 2017 ♀ from Japan A habitus B head, frontal view C head, dorsal view D head, lateral view E mesosoma, lateral view F central part of fore wing.

The specific name is derived from the type locality.

A holotype female only.

HT: ♀, Benoki, Kunigami Vil., Kunigami County, Okinawa-hontô, Okinawa Pref., Ryûkyûs, JAPAN, 9.IV.1996, M. Aoyagi leg. (LT) (NIAES).

Oriental region.

JAPAN: [Ryûkyûs] Okinawa.

Unknown.

This species is similar to E. matsumurai sp. nov. but easily distinguishable by the number of antennal flagellomeres (i.e., antenna with 54 flagellomeres in E. kunigamiensis sp. nov. but 59–60 in E. matsumurai sp. nov.), value of ICI (i.e., ICI = 0.4 in E. kunigamiensis sp. nov. but 0.7–0.8 in E. matsumurai sp. nov.), length of lateral longitudinal carinae of scutellum (i.e., lateral longitudinal carinae along anterior 0.2 of scutellum in E. kunigamiensis sp. nov. but more than 0.8 in E. matsumurai sp. nov.), body size (body length ca. 23.0 mm in E. kunigamiensis sp. nov. but 28.0–30.5 mm in E. matsumurai sp. nov.), etc. However, the detailed affinities of E. kunigamiensis sp. nov. are unknown.

Female (HT). Body length ca. 23.0 mm.

Head with GOI = 2.5 (Fig. 23D). Lower face 0.7× as wide as high, strongly shiny, and smooth with fine setae (Fig. 23B). Clypeus 1.4× as wide as high, smooth, moderately convex in profile, and ventral margin acute (Fig. 23B, D). Malar space 0.2× as long as basal mandibular width (Fig. 23B, D). Mandible twisted by ca. 30°, moderately long, evenly rather strongly narrowed, outer surface smooth (Fig. 23B, D). Upper tooth of mandible 1.4× as long as lower. Frons, vertex and gena strongly shiny with fine setae (Fig. 23B–D). Posterior ocellus almost touching eye (Fig. 23B–D). Ventral end of occipital carina not joining oral carina. Antennae with 54 flagellomeres; first flagellomere 1.9× as long as second; 20^th flagellomere 1.7× as long as wide.

Mesosoma entirely strongly shiny with setae (Fig. 23E). Pronotum finely diagonally striate (Fig. 23E). Mesoscutum 1.4× as long as maximum width, strongly shiny and finely punctate with setae, and evenly rounded in profile (Fig. 23E). Notauli absent (Fig. 23E). Scutellum moderately convex, rather weakly shiny, punctate, with lateral longitudinal carinae along anterior 0.2 of scutellum. Epicnemium densely punctate with setae. Epicnemial carina present, almost straight and inclined to anterior, dorsal end not reaching anterior margin of mesopleuron (Fig. 23E). Dorsal part of mesopleuron smooth to finely punctate and ventral part longitudinally finely punctostriate (Fig. 23E). Submetapleural carina broadened anteriorly (Fig. 23E). Metapleuron moderately diagonally striate (Fig. 23E). Propodeum declivous in profile; anterior transverse carina complete; anterior area longitudinally striate; spiracular area smooth; posterior area moderately rugose; propodeal spiracle elliptical and not joining pleural carina by ridge (Fig. 23E).

Wings (Fig. 23F). Fore wing length ca. 16.0 mm with AI = 0.7, CI = 0.3, DI = 0.3, ICI = 0.4, SDI = 1.1, SI = 0.1, SRI = 0.3; vein 1m-cu&M almost evenly curved; vein 2r&RS almost straight; vein RS rather evenly curved; fenestra and sclerites of discosubmarginal cell as in Fig. 23F; proximal sclerite triangular, faintly confluent with distal sclerite, strongly pigmented; central sclerite not well-delimited and pigmented part suboval, positioned in anterodistal part of fenestra; distal sclerite vestigial; proximal corner of marginal cell evenly setose; posterodistal corner of second discal cell ca. 100° and of subbasal cell ca. 70°; vein 1cu-a antefurcal to M&RS by 0.1× length of 1cu-a. Hind wing with NI = 2.3; vein RS straight; vein RA with 7 uniform hamuli.

Legs. Hind leg with coxa in profile 1.8× as long as deep; basitarsus 1.9× as long as second tarsomere; fourth tarsomere 3.1× as long as wide; tarsal claw simply pectinate.

Metasoma with DMI = 1.4, PI = 3.2, THI = 3.5; thyridium oval; ovipositor sheath not longer than posterior depth of metasoma (Fig. 23A).

Colour (Fig. 23). Longitudinal vittae of mesoscutum and metasoma red-brown; apex of mandible black, otherwise whitish yellow. Wings weakly infuscate; sclerites pigmented and amber; vein dark reddish brown.

Male. Unknown.

Figure 23. 

Enicospilus kunigamiensis Shimizu, sp. nov. ♀ (HT) from Japan A habitus B head, frontal view C head, dorsal view D head lateral view E mesosoma, lateral view F central part of fore wing.

Total of 76 specimens (48♀♀26♂♂ and 2 unsexed): India (2♀♀1♂), Japan (22♀♀23♂♂), Taiwan (23♀♀2♂♂ and 2 unsexed), Zambia (1♀).

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

Afrotropical and Oriental regions (Yu et al. 2016); new to the Eastern Palaearctic (Hachijô-jima, Tôkyô, Kantô-Kôshin, Japan) and Oceanic (Nishi-jima, Tôkyô, Ogasawara, Japan) regions (cf. Suppl. material 1: Table S1); this is a predominantly (sub-)tropical species. This species is widely distributed from the Afrotropical to Oriental regions and Gauld (1982) suggested that it has possibly been introduced from Asia to Africa, although there is no reliable evidence to support or refute his hypotheses.

Newly recorded from Japan.

JAPAN: [Kantô-Kôshin] Tôkyô; [Ryûkyûs] Kagoshima and Okinawa; [Ogasawara] Tôkyô. This species is restricted to southern regions of Japan.

Unknown.

This species resembles E. pseudantennatus, E. vestigator, and E. tripartitus on the shape of fore wing fenestra, sclerites, and venation. However, E. laqueatus is easily distinguishable from them by the outer mandibular surface morphology (i.e., outer mandibular surface with diagonal setose groove between dorsoproximal corner and base of apical teeth in E. laqueatus, but smooth or just densely punctate with stout setae in the other three species, as summarised in Table 6). In addition, this species morphologically resembles E. aciculatus and E. yonezawanus but is distinguished from them by its strongly pigmented central sclerite of fore wing fenestra (Fig. 24F) (central sclerite very weakly pigmented or vestigial in E. aciculatus, as in Fig. 10F, and completely lacking in E. yonezawanus, as in Fig. 54F).

Figure 24. 

Enicospilus laqueatus (Enderlein, 1921) ♀ from Japan A habitus B head, frontal view C head, dorsal view D head, lateral view E mesosoma, lateral view F central part of fore wing.