Corresponding author: So Shimizu (
Academic editor: B. Santos
The predominantly tropical ophionine genus
Shimizu S, Broad GR, Maeto K (2020) Integrative taxonomy and analysis of species richness patterns of nocturnal Darwin wasps of the genus
Darwin wasps, the family
The cosmopolitan genus
As well as almost all other genera of
Summary of taxonomic histories of the Japanese species of
New species or names | New records | Deleted species or names | Total species number |
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Species delimitation and taxonomic revisions of poorly known, hyperdiverse groups, such as Darwin wasps, based on traditional morphology-based taxonomy is challenging, but has recently been rapidly improved by advancing integrative approaches that combine multiple perspectives (population genetics, morphometrics, behaviour, host, chemical composition, etc.). A combined morphological and DNA barcoding (partial sequencing of a mitochondrial protein-coding gene, cytochrome c oxidase 1, CO1) approach is the most straightforward method and has been used by many authors for various taxa (e.g.,
Species delineation of ophionines has been considered to be more difficult than many other lineages of insects, including other subfamilies of
There has been much research into patterns of Darwin wasp species richness across latitudinal gradients. This has been summarised fairly recently by
Most ophionines are typical nocturnal koinobiont parasitoids with their centre of species diversity in the (sub-)tropics (e.g.,
The Japanese archipelago is located in a long line between ca. 20–45°N, approximately 3,000 km from south to north, ranging from the southern subtropical to northern and high elevational subarctic zones, containing a high diversity of ecological habitats. Biogeographically, it also includes the Oceanic, Oriental, and Palaearctic regions and is a melting pot of species originating from these regions, so in some ways one of the most interesting biodiversity hotspots (e.g.,
The specimens examined were studied in or borrowed from insect collections, or newly collected for the present study, mainly using High Intensity Discharge (
Field collection
Specimens were observed using a stereoscopic microscope (SMZ1500, Nikon, Tôkyô, Japan). Multi-focus photographs were taken using a single lens reflex camera (α7II, Sony, Tôkyô, Japan) with a micro-lens (LAOWA 25 mm F2.8 2.5–5× ULTRA MACRO, Anhui Changgeng Optics Technology Co., Ltd, Hefei, China and A FE 50mm F2.8 Macro SEL50M28, Sony, Tôkyô, Japan), captured in RAW format, developed using Adobe Lightroom CC v.2.2.1 (Adobe Systems Inc., San Jose, CA, USA), and stacked using Zerene Stacker v.1.04 (Zerene Systems LLC., Richland, WA, USA). The original maps were generated using SimpleMappr (
The morphological terms mainly follow
The abbreviations for specimen repositories used in the present paper (some only in Suppl. material
Morphological terms and measurement characters for head and mandible
Morphological terms for mesosoma
Morphological terms and measurement characters for wings
Morphological terms and measurement characters for metasoma (
Indices used in the present paper.
Indices | Formula | |
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Geno-Orbital Index | maximum breadth of eye in profile [be] / maximum breadth of gena in same line [bg] |
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Alar Index | length of 1m-cu&M between 2m-cu and bulla [cd] / length of 2rs-m [ab] |
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Cubital Index | length of CU between 1m-cu&M and 2cu-a [gf] / length of 2cu-a [fh] |
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Discoidal Index | maximum vertical distance between CU (between 2cu-a and 2m-cu) and 1m-cu&M [k] / length of CU between 2cu-a and 2m-cu [fe] |
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Intercubital Index | length of 2rs-m [ab] / length of M between 2m-cu and 2rs-m [bc] |
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Second Discoidal Index | length of CU between 2cu-a and 2m-cu [ef] / length of CU between M&RS and 1m-cu&M [gi] |
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Sinuousness Index | maximum length between 1m-cu&M and a straight line connecting the intersection of M, 2m-cu, and 1m-cu&M and the intersection of 1m-cu&M and CU [l] / distance between the intersection of M, 2m-cu, and 1m-cu&M [c] and the intersection of 1m-cu&M and CU [g] [j] |
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Second Recurrent Index | length of 2m-cu [ce] / length of CU between 2cu-a and 2m-cu [ef] |
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Nervellar Index | length of CU between M and cu-a [mn] / length of cu-a [no] |
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Dorsal Metasomal Index | length of dorsum of T2 [e] / length of dorsum of T3 [f] |
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Petiolar Index | distance between base of T1 and anterior margin of spiracle [a] / distance between posterior margin of spiracle and apex of T1 [b] |
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Thyridium Index | distance between anterior margin of T2 and anterior margin of thyridium [c] / maximum diameter of thyridium [d] |
Abbreviations for repositories consulted (not all are referred to in the main text and some are only in Suppl. material
Abbreviations | Repositories |
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Entomological Laboratory, Meijô University, Nagoya, Japan |
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Utah State University Insect Collection (= American Entomological Institute: |
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The Parasitic Hymenoptera Collection of the Institute of Beneficial Insect, College of Plant Protection, Fujian Agriculture and Forestry University, Fuzhou, China |
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Fachbereich Zoologie, Luther-Universitat, Halle, Germany |
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Hiwa Museum for Natural History, Shôbara, Japan |
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Museum of Nature and Human Activities, Sanda, Japan |
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Marathwada University Collection, Aurangabad, India |
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Naturhistorisches Museum, Vienna, Australia |
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The Laboratory of Systematic Entomology (= Entomological Institute: EIHU), Hokkaidô University, Sapporo, Japan |
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Tochigi Prefectural Museum, Utsunomiya, Japan |
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There are many published distribution and host records that cannot be verified as we cannot access all voucher specimens or host remains underpinning these literature records;
We used the following order of Japanese Prefectures in the distribution of Japan: [Hokkaidô] Hokkaidô; [Tôhoku] Aomori, Akita, Iwate, Yamagata, Miyagi, Fukushima; [Hokuriku] Niigata, Toyama, Ishikawa, Fukui; [Kantô-Kôshin] Ibaraki, Tochigi, Gunma, Nagano, Yamanashi, Saitama, Tôkyô, Kanagawa, Chiba; [Tôkai] Gifu, Aichi, Shizuoka, Mie; [Kinki] Kyôto, Shiga, Ōsaka, Hyôgo, Nara, Wakayama; [Chûgoku] Tottori, Shimane, Okayama, Hiroshima, Yamaguchi; [Shikoku] Kagawa, Tokushima, Ehime, Kôchi; [Kyûshû] Fukuoka, Saga, Nagasaki, Ōita, Kumamoto, Miyazaki, Kagoshima; [Ryûkyûs] Kagoshima, Okinawa; [Ogasawara] Tôkyô. Honshû consists of Tôhoku, Hokuriku, Kantô-Kôshin, Tôkai, Kinki, and Chûgoku. Prefectures are ordered basically from North to South.
To test our assessment of taxonomy based on morphology, we employed a DNA barcoding approach. A total of 168 sequences of CO1 from 41 of 47 Japanese species of
The species, identifiers for the specimens, collection localities, sample codes, and accession numbers for all terminal taxa used in the analyses are listed in Suppl. material
Most specimens for DNA analysis were dried specimens borrowed from collections. Some specimens were newly collected for the present study; these were stored in 80.0–99.9% ethanol and, after DNA extraction, mounted as dried specimens, currently deposited in the respective insect collections. DNA was extracted from a single right mid leg or both right mid and hind legs using the DNeasy Blood and Tissue Kit (Qiagen, Düsseldorf, Germany).
Partial sequences of CO1 were amplified using primers designed by
We conducted multiple sequence alignments in MAFFT v.7.409 (
Analyses were performed with Bayesian Inference (
The latitudinal diversity gradient (
Latitudinal zones used in the latitudinal diversity gradient analysis.
Zones | Latitudinal ranges (LR) | Provinces | Prefectures | Points | Latitudes |
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A | 42 ≤ LR < 45 | Hokkaidô | Hokkaidô | Sapporo | 43°03'51"N |
B | 39 ≤ LR < 42 | Tôhoku | Aomori | Aomori | 40°49'28"N |
Akita | Akita | 39°43'07"N | |||
Iwate | Morioka | 39°42'13"N | |||
C | 36 ≤ LR < 39 | Tôhoku | Yamagata | Yamagata | 38°14'26"N |
Miyagi | Sendai | 38°16'08"N | |||
Fukushima | Fukushima | 37°45'00"N | |||
Hokuriku | Niigata | Niigata | 37°54'08"N | ||
Toyama | Toyama | 36°41'43"N | |||
Ishikawa | Kanazawa | 36°35'40"N | |||
Fukui | Fukui | 36°03'55"N | |||
Kantô-Kôshin | Ibaraki | Mito | 36°20'29"N | ||
Tochigi | Utsunomiya | 36°33'57"N | |||
Gunma | Maebashi | 36°23'28"N | |||
Nagano | Nagano | 36°39'05"N | |||
D | 33 ≤ LR < 36 | Kantô-Kôshin | Yamanashi | Kôfu | 35°39'50"N |
Saitama | Saitama | 35°51'25"N | |||
Tôkyô | Shinjyuku | 35°41'22"N | |||
Kanagawa | Yokohama | 35°26'52"N | |||
Chiba | Chiba | 35°36'17"N | |||
Tôkai | Gifu | Gifu | 35°23'28"N | ||
Aichi | Nagoya | 35°10'49"N | |||
Shizuoka | Shizuoka | 34°58'37"N | |||
Mie | Tsu | 34°43'49"N | |||
Kinki | Kyôto | Kyôto | 35°01'17"N | ||
Shiga | Ōtsu | 35°00'16"N | |||
Ōsaka | Ōsaka | 34°41'11"N | |||
Hyôgo | Kôbe | 34°41'29"N | |||
Nara | Nara | 34°41'07"N | |||
Wakayama | Wakayama | 34°13'34"N | |||
Chûgoku | Tottori | Tottori | 35°30'13"N | ||
Shimane | Matsue | 35°28'20"N | |||
Okayama | Okayama | 34°39'42"N | |||
Hiroshima | Hiroshima | 34°23'47"N | |||
Yamaguchi | Yamaguchi | 34°11'09"N | |||
Shikoku | Kagawa | Takamatsu | 34°20'25"N | ||
Tokushima | Tokushima | 34°03'57"N | |||
Ehime | Matsuyama | 33°50'30"N | |||
Kôchi | Kôchi | 33°33'35"N | |||
Kyûshû | Fukuoka | Fukuoka | 33°36'23"N | ||
Saga | Saga | 33°14'58"N | |||
Ōita | Ōita | 33°14'17"N | |||
E | 30 ≤ LR < 33 | Kyûshû | Nagasaki | Nagasaki | 32°44'41"N |
Kumamoto | Kumamoto | 32°47'23"N | |||
Miyazaki | Miyazaki | 31°54'40"N | |||
Kagoshima | Kagoshima | 31°33'37"N | |||
F | 27 ≤ LR < 30 | Ryûkyûs | Kagoshima | Amami | 28°22'39"N |
Okinawa | Naha | 26°12'45"N | |||
Ogasawara | Tôkyô | Ogasawara | 27°05'39"N |
To understand the regional pattern of sampling biases, numbers of four categories (specimens, collection events, collector, and species) were counted for each area. Each pattern is shown in the heat maps.
To infer the total species richness of the Japanese
A total of 47 morphospecies were recognised in Japan: 32 of which were previously known from the Japanese fauna, eight were new to science, seven were new to Japan, and seven were excluded from the Japanese fauna (Table
Summary of taxonomic results for Japanese species of
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For most Japanese
Bayesian majority-rule consensus tree based on a barcoding gene (BIPP = the Bayesian inference posterior probabilities; MLBS = the maximum likelihood bootstrap percentages).
DNA barcoding analyses sometimes show traditional diagnostic characters to be ineffectual. The confluence or separation of the proximal and distal sclerites of fore wing fenestra has been considered to be an important diagnostic character for
Intraspecific variation of the fore wing sclerite development in
Intraspecific variation of the body colour in
Afrotropical, Australasian, Holarctic, Neotropical, Oceanic, and Oriental regions (
According to the available evidence, species of
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
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
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
Features of mandibles are some of the most important diagnostic characters of
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.
First, the number, shape, and position of sclerites of the fore wing fenestra is usually very useful in
Second, the shape and setosity of the fore wing fenestra is sometimes a very useful character. For example, among Japanese species, the fenestra of
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.,
1 | Fore wing fenestra lacking both sclerites and quadra (Fig. |
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– | Fore wing fenestra with more or less distinct sclerites (e.g., Figs |
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2 (1) | Fore wing fenestra without proximal and central sclerites and only with rather strongly pigmented and thick distal sclerite (Fig. |
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– | Fore wing fenestra with weakly to strongly pigmented proximal sclerite and also sometimes with central and/or distal sclerite (if distal sclerite present, it is never thickened) (e.g., Figs |
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3 (2) | Mandible very strongly twisted by ca. 85°, therefore outer margin forming acute median longitudinal ridge between centroproximal part of mandible and base of apical teeth (this ridge is the ventral margin of the mandible) (Fig. |
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– | Mandibular torsion various, not very strongly twisted and outer margin never forming acute median longitudinal ridge as above. Colour of mesoscutum various. |
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4 (3) | Clypeus nasute, strongly convex, anterior margin acute and strongly projecting, and ventral margin strongly and abruptly impressed (Figs |
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– | Clypeus not nasute, flat to moderately convex, anterior margin obtuse and rounded if convex, and ventral margin impressed to acute (e.g., Figs |
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5 (4) | Proximal margin of proximal sclerite of fore wing fenestra distinctly separated from proximal margin of fenestra by more than width of proximal sclerite (e.g., Figs |
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– | Proximal margin of proximal sclerite of fore wing fenestra joining or close to proximal margin of fenestra, if separated then by less than half width of proximal sclerite (e.g., Figs |
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6 (5) | Interocellar area entirely black (Fig. |
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– | Interocellar area entirely yellow- to red-brown (e.g., Fig. |
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7 (6) | Proximal sclerite of fore wing fenestra narrow and more or less linear (e.g., Fig. |
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– | Proximal sclerite of fore wing fenestra various but rather broad (triangular, circular, comma-shaped, etc.) (e.g., Figs |
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8 (7) | Fore wing fenestra with linear central sclerite (Fig. |
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– | Fore wing fenestra without central sclerite (Figs |
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9 (8) | Fore wing vein 1m-cu&M evenly curved (Fig. |
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– | Fore wing vein 1m-cu&M weakly to moderately sinuous (Fig. |
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10 (7) | Fore wing with both |
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– | Fore wing with either or both |
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11 (10) | Proximal part of marginal cell of fore wing widely glabrous (Fig. |
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– | Marginal cell of fore wing uniformly setose (Figs |
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12 (11) | Fore wing fenestra with central sclerite (Fig. |
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– | Fore wing fenestra without central sclerite (Figs |
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13 (12) | Proximal and distal sclerites of fore wing fenestra strongly confluent and distal sclerite strongly sclerotised (Fig. |
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– | Proximal sclerite of fore wing fenestra isolated and distal sclerite absent or vestigial (Figs |
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14 (13) | Proximal sclerite of fore wing fenestra usually entirely weakly pigmented and half moon-shaped (Fig. |
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– | Proximal sclerite of fore wing fenestra partly to entirely strongly pigmented and drop-shaped (Fig. |
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15 (5) | Outer mandibular surface usually with strong diagonal groove between dorsoproximal corner and medio- to ventrobasal part of apical teeth; groove usually bearing moderate to very dense setae (e.g., Figs |
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– | Outer mandibular surface smooth; sometimes with punctures or setae but never forming diagonal line and never dense (e.g., Figs |
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16 (15) | Clypeus flat, ventral margin acute and projecting (Fig. |
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– | Clypeus flat to convex, and ventral margin acute to impressed but never projecting (e.g., Figs |
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17 (16) | Fore wing fenestra without central sclerite (Fig. |
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– | Fore wing fenestra with central sclerite (e.g., Figs |
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18 (17) | Proximal part of marginal cell of fore wing adjacent to vein 2r&RS with distinct and wide glabrous area (Fig. |
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– | Marginal cell of fore wing uniformly setose, never with wide glabrous area (e.g., Figs |
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19 (18) | Face broad; lower face more than 0.82× as wide as high; clypeus more than 1.8× as wide as high (e.g., Figs |
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– | Face moderately broad; lower face less than 0.81× as wide as high; clypeus less than 1.8× as wide as high (e.g., Figs |
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20 (19) | Meso- and metapleuron entirely very densely punctate so that matt or submatt, punctures of metapleuron contiguous or separated by less than diameter of puncture, thus very weakly shiny or not (Fig. |
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– | Meso- and metapleuron finely to moderately punctate to punctostriate, strongly shiny, and never matt (e.g., Figs |
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21 (20) | Posterior area of propodeum entirely punctate and strongly shiny. Meso- and metapleuron entirely punctate and strongly shiny (Fig. |
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– | Posterior area of propodeum with fine to coarse sculpture (e.g., reticulation, rugae, striae), never entirely punctate, and usually weakly shiny. Meso- and metapleuron punctate to striate (e.g., Figs |
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22 (21) | Posterior ocellus distinctly separated from eye (Fig. |
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– | Posterior ocellus close to eye but not touching (e.g., Figs |
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23 (22) | Mandible very long and slender; apical 0.7 parallel-sided; proximal outer surface with very wide subtriangular concavity (Fig. |
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– | Mandible moderately long; apical 0.3–0.5 (sub-)parallel-sided; proximal outer surface usually with shallow and narrow crescent-shaped concavity (Figs |
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24 (23) | Mesosoma and terminal segments of metasoma blackish (Figs |
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– | Body entirely orange brown, except for terminal segments of metasoma sometimes infuscate or black (Figs |
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25 (24) | Proximal and distal sclerites of fore wing fenestra not confluent (Fig. |
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– | Proximal and distal sclerites of fore wing fenestra confluent (Fig. |
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26 (24) | T6–8 black (metasoma entirely testaceous in very rare cases) (Fig. |
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– | Metasoma always uniformly orange-brown, with posterior segments never black (Fig. |
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27 (19) | Central sclerite of fore wing fenestra rather large, but ill-defined and never strongly sclerotised (Fig. |
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– | Central sclerite of fore wing fenestra variously sized, well defined, and strongly sclerotised (e.g., Figs |
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28 (27) | Proximal and distal sclerites of fore wing fenestra confluent (e.g., Figs |
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– | Proximal and distal sclerites of fore wing fenestra not confluent (e.g., Figs |
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29 (28) | Body entirely yellow-brown (Fig. |
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– | Mesosoma and posterior segments of metasoma dark brown to black (Figs |
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30 (29) | Metasoma entirely yellow-brown with posterior segments always black (Fig. |
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– | Metasoma uniformly yellow-brown and posterior segments never infuscate (Fig. |
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31 (29) | Coxae, T1–2 and T5–8 dark-brown to black (Fig. |
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– | Coxae and T1–2 testaceous, never coloured (Fig. |
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32 (28) | Distal part of fore wing fenestra between central and distal sclerites covered with setae (Fig. |
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– | Fore wing fenestra without any setae (e.g., Figs |
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33 (32) | Outer mandibular surface with deep basal concavity and dense punctures and stout setae, without diagonal groove but punctures and setae sometimes forming diagonal line (Fig. |
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– | Outer mandibular surface with shallow basal concavity, with conspicuous diagonal deep setose groove (e.g., Figs |
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34 (33) | Central sclerite of fore wing fenestra well delimited, D-shaped (Figs |
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– | Central sclerite of fore wing fenestra poorly delimited, oval to linear (Figs |
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35 (34) | Central sclerite of fore wing fenestra positioned in anterodistal part of fenestra, smaller, moderately sclerotised (Fig. |
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– | Central sclerite of fore wing fenestra positioned in centrodistal part of fenestra, larger, strongly sclerotised (Fig. |
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36 (34) | Fore wing vein 1cu-a more or less curved or angulate (Fig. |
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– | Fore wing vein 1cu-a straight (Fig. |
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37 (15) | Interocellar area entirely black (Figs |
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– | Interocellar area entirely yellow- to red-brown (e.g., Figs |
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38 (37) | Fore wing fenestra without central sclerite or quadra (Fig. |
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– | Fore wing fenestra widely covered with very weakly pigmented quadra (Fig. |
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39 (37) | Fore wing fenestra usually without central sclerite (Figs |
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– | Fore wing fenestra with conspicuously pigmented central sclerite (e.g., Figs |
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40 (39) | Fore wing fenestra widely covered with unpigmented to very weakly pigmented quadra (Fig. |
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– | Fore wing fenestra without quadra (Figs |
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41 (40) | Fore wing fenestra with bullet-shaped proximal sclerite (Fig. |
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– | Fore wing fenestra with triangular proximal sclerite (Figs |
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42 (41) | Fore wing fenestra wider and anterodistal corner antefurcal to RS by less than length of 2rs-m (Fig. |
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– | Fore wing fenestra smaller and anterodistal corner antefurcal to RS by more than length of 2rs-m (Fig. |
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43 (39) | Fore wing fenestra long and anterodistal corner (sub)interstitial or antefurcal to RS by less than 0.3× length of 2rs-m (Figs |
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– | Fore wing fenestra moderately long and anterodistal corner antefurcal to RS by more than 0.4× length of 2rs-m (e.g., Figs |
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44 (43) | Mesoscutum, ventral part of T3–4, and entire of T5–8 infumate to black, otherwise pale brown (Fig. |
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– | Mesoscutum and metasoma entirely orange-brown (Fig. |
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45 (43) | Central sclerite of fore wing fenestra linear and parallel with distal margin of fenestra (Fig. |
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– | Central sclerite of fore wing fenestra various (Fig. |
|
46 (45) | Propodeum with distinct posterior transverse carina arising from pleural carina (Fig. |
|
– | Propodeum without posterior transverse carinae (e.g., Figs |
|
47 (46) | Metasoma with striking colour pattern, i.e., anterior parts of T1–4 yellow-brown and posterior parts dark brown to black (Fig. |
. |
– | Metasoma entirely orange-brown or brown (e.g., Figs |
|
48 (47) | Fore wing fenestra with larger circular to D-shaped, very strongly sclerotised central sclerite (Fig. |
|
– | Fore wing fenestra with smaller linear to elongate suboval, weakly to strongly sclerotised central sclerite (Figs |
|
49 (48) | Fore wing with central sclerite strongly sclerotised and pigmented; setae and veins darker brown (Fig. |
|
– | Fore wing with central sclerite weakly to moderately sclerotised and pigmented; setae and veins brighter brown (Fig. |
|
Total of 19 specimens (all ♀♀): Japan (1♀), Nepal (1♀), Sri Lanka (2♀♀), Taiwan (15♀♀).
Type series:
Australasian, Eastern Palaearctic, and Oriental regions (
Newly recorded from Nepal.
JAPAN: [Ryûkyûs] Okinawa (
Unknown.
The characteristic striking colour pattern of this species (i.e., T1–4 each anteriorly yellow-brown and posteriorly dark brown, as in Fig.
This species has been confused with
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:
Australasian, Eastern Palaearctic, Oceanic and Oriental regions (
JAPAN: [Kyûshû] Fukuoka* and Kagoshima (
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
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♀).
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
Female (n = 66). Body length 23.5–26.0 (
Head with
Mesosoma entirely weakly to moderately shiny with setae (Fig.
Wings (Fig.
Legs. Hind legs with coxa in profile 1.7–1.8× (
Metasoma with
Colour (Fig.
Male (n = 20). Very similar to female.
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:
Afrotropical, Australasian, Oceanic, and Oriental regions (
JAPAN: [Ryûkyûs] Okinawa (
This species has a very wide distribution from South East Asia to South Africa. According to
Recorded from various
The Japanese specimen of
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 (
JAPAN: [Hokkaidô] (
Reared from one species of
This species is usually very easily distinguished from all other Palaearctic
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 (
JAPAN: [Kantô-Kôshin] Tôkyô* and Kanagawa*; [Kinki] Hyôgo*; [Chûgoku] Hiroshima (
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
Wing characters of this species (e.g., fore wing with proximal part of marginal cell widely glabrous,
Total of 49 specimens (44♀♀5♂♂): Japan (8♀♀3♂♂), Sri Lanka (2♀♀), Taiwan (34♀♀2♂♂).
Type series:
Australasian, Eastern Palaearctic, and Oriental regions (
JAPAN: [Ryûkyûs] Okinawa (
Recorded as a parasitoid of several species of
This species is readily distinguishable from all other
Total of 14 specimens (12♀♀2♂♂): Japan (2♀♀1♂), Malaysia (6♀♀), Philippines (1♀), Sri Lanka (3♀♀), Taiwan (1♂).
Type series:
Australasian, Eastern Palaearctic, and Oriental regions (
JAPAN: [Ryûkyûs] Okinawa (
Recorded from
This species especially resembles
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:
Australasian, Oceanic, and Oriental regions (
Newly recorded from Laos.
JAPAN: [Kyûshû] Kagoshima*; [Ryûkyûs] Kagoshima (
Recorded as a parasitoid of three species of
This species is easily distinguishable from all other
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:
Eastern Palaearctic and Oriental regions (
Newly recorded from Laos and Malaysia.
JAPAN: [Kantô-Kôshin] Tôkyô*; [Tôkai] Mie*; [Kinki] Ōsaka (
Unknown.
This species is easily distinguishable by the wide face (Fig.
Total of 36 specimens (35♀♀1♂): Brunei (1♀), China (1♀), Japan (28♀♀1♂), Taiwan (5♀♀).
Type series:
Australasian, Eastern Palaearctic, and Oriental regions (
Newly recorded from Taiwan.
JAPAN: [Kyûshû] Kagoshima*; [Ryûkyûs] Kagoshima* and Okinawa*. * New records.
Unknown.
This species is closest to
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:
Australasian, Eastern Palaearctic, and Oriental regions (
JAPAN: [Kantô-Kôshin] Kanagawa (
Unknown.
This species is very easily distinguishable from any other Japanese
Total of 3 specimens (2♀♀1♂): Japan (2♀♀1♂).
Eastern Palaearctic region (
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
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
Total of 6 specimens (5♀♀1♂): Japan (5♀♀1♂).
Type series:
Eastern Palaearctic region (
JAPAN: [Hokuriku] Fukui*; [Kantô-Kôshin] Tochigi* and Saitama (
Unknown.
As in Fig.
The specific name is derived from the type locality.
A holotype female only.
Oriental region.
JAPAN: [Ryûkyûs] Okinawa.
Unknown.
This species is similar to
Female (
Head with
Mesosoma entirely strongly shiny with setae (Fig.
Wings (Fig.
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
Colour (Fig.
Male. Unknown.
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:
Afrotropical and Oriental regions (
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
Comparison of diagnostic characters of
Characters | Species | ||
---|---|---|---|
|
|
|
|
Lower margin of clypeus | acute | blunt and impressed | blunt |
Torsion of mandible | 10–15° | 10–12° | 60–80° |
Outer surface of mandible | with a diagonal groove | covered with dence and long setae | smooth |
Basal concavity of mandibular outer surface | shallow | deep | shallow |
Discosubmarginal cell | rather short | rather short | long |
|
0.4–0.5 | 0.5–0.7 | 0.6–0.9 |
This species probably prefers lakes. Hence, the specific name is derived from the Greek
Total of 37 specimens (18♀♀19♂♂): Japan (18♀♀19♂♂).
Eastern Palaearctic region.
JAPAN: [Hokkaidô]; [Hokuriku] Fukui.
All specimens have been collected from marshes or lakes of rather cooler regions, suggesting that it is restricted to hosts that inhabit open, aquatic conditions. However, some factors, such as a progression of plant succession, isolation of habitats, and increasingly dry conditions, have led many wetland insects to become endangered in Japan (e.g.,
The distally setose fore wing fenestra is unique to this species within the Asian
Female (n = 18). Body length 15.0–18.0 (
Head with
Mesosoma entirely weakly to moderately shiny with setae (Fig.
Wings (Fig.
Legs. Hind leg with coxa in profile 1.7–1.9× (
Metasoma with
Colour (Fig.
Male (n = 19). Very similar to female.
This is a fairly morphologically uniform species, although
Total of 51 specimens (42♀♀9♂♂): India (2♀♀), Japan (40♀♀8♂♂), Philippines (1♂).
Type series:
Eastern Palaearctic region (
Newly recorded from India and Philippines.
JAPAN: [Hokkaidô] (
Unknown.
This species resembles
Comparison of diagnostic characters of
Characters | Species | |||
---|---|---|---|---|
|
|
|
|
|
Sculpture of meso- and metasoma | roughly punctate to punctostriate | punctostriate | punctate | striate |
Fore wing vein 1m-cu&M | evenly curved | weakly sinuous | evenly curved | weakly sinuous |
|
<0.5 | ≥ 0.5 | > 0.5 | ≥0.5 |
Pectinae of hind tarsal claw | complete | complete | lacking proximally | complete |
The specific name is derived from one of the greatest Japanese entomologists, Shônen Matsumura.
Total of 12 specimens (8♀♀4♂♂): Japan (8♀♀4♂♂).
Eastern Palaearctic region.
JAPAN: [Shikoku] Ehime and Kôchi; [Kyûshû] Fukuoka, Nagasaki and Ōita.
Unknown.
This species is similar to
Female (n = 8). Body length 28.0–30.5 (
Head with
Mesosoma entirely weakly to moderately shiny with setae (Fig.
Wings (Fig.
Legs. Hind leg with coxa in profile 1.7–1.8× (
Metasoma with
Colour (Fig.
Male (n = 4). Very similar to female except body length 22.0–30.5 mm; antennae with 55–63 flagellomeres.
Total of 217 specimens (166♀♀41♂♂ and 10 unsexed): Australia (1♀), China (1♀), India (26♀♀), Indonesia (4♀♀2♂♂ and 1 unsexed), Japan (26♀♀12♂♂ and 1 unsexed), Malaysia (1♀), Maldives (1♂), Papua New Guinea (7♀♀1♂), Philippines (7♀♀), Singapore (1 unsexed), Sri Lanka (8♀♀), Taiwan (85♀♀25♂♂ and 7 unsexed).
Type series:
Australasian, Eastern Palaearctic, Oceanic, and Oriental regions (
JAPAN: [Ryûkyûs] Kagoshima* and Okinawa (
No host records from Japan.
This species is morphologically most similar to
In Japanese collections, they are sometimes confused with
Comparison of diagnostic characters of
Characters | Species | |||
---|---|---|---|---|
|
|
|
|
|
Lower face (width / height) | 0.7–0.8 | 0.8–0.9 | 0.7–0.8 | 1.0 |
Length of tooth (upper teeth / lower teeth) | 1.2–1.5 | 1.9–2.2 | 1.2–1.3 | 1.7–2.0 |
Posterior ocelli and orbit | very close | touching or very close | very close | distinctly separated |
Sculpture of meso- and metapleuron | punctate to punctostriate | entirely punctate | punctate to punctostriate | punctate to punctostriate |
Marginal cell of fore wing | entirely setose | entirely setose | with a glabrous | entirely setose |
Shape of central sclerite | oval | oval | linear | comma-shaped |
Proximal and distal sclerites | confluent | separated | confluent | separated |
Total of 30 specimens (14♀♀16♂♂): Japan (14♀♀16♂♂).
Type series:
Eastern Palaearctic region (
JAPAN: [Hokkaidô]*; [Tôhoku] Aomori*; [Hokuriku] Niigata*; [Kantô-Kôshin] Gunma*, Nagano*, Yamanashi*, and Saitama*; [Tôkai] Shizuoka* and Mie*; [Kinki] Ōsaka (
Unknown.
In some ichneumonid collections, this species has been confused with
This species is also similar to
Total of 35 specimens (26♀♀9♂♂): India (5♀♀), Japan (14♀♀8♂♂), Sri Lanka (1♀), Taiwan (6♀♀1♂).
Type series:
Eastern Palaearctic and Oriental regions (
JAPAN: [Kantô-Kôshin] Kanagawa (
No host records from Japan. Recorded as a parasitoid of
This species is very easily distinguishable from any other
Total of 26 specimens (19♀♀7♂♂): Japan (18♀♀7♂♂), Taiwan (1♀).
Oriental region (
JAPAN: [Ryûkyûs] Kagoshima (
Unknown.
This species is one of the largest Japanese ophionines, along with
Total of 18 specimens (11♀♀4♂♂ and 3 unsexed): Brunei (2♀♀2♂♂ and 1 unsexed), Indonesia (2♀♀), Japan (1♀2♂♂), Singapore (1♀), Sri Lanka (2♀♀ and 2 unsexed), Taiwan (2♀♀), unknown (1♀).
Type series:
Oriental region (
Newly recorded from Indonesia.
JAPAN: [Ryûkyûs] Okinawa (
Unknown.
This species is an extremely large insect, as is
This species is easily distinguishable from all other Japanese
Total of 225 specimens (123♀♀100♂♂ and 2 unsexed): Brunei (3♀♀2♂♂), China (1♀1♂), Indonesia (1♀), Japan (99♀♀81♂♂ and 2 unsexed), Korea (1♀), Malaysia (1♀2♂♂), Sri Lanka (1♀), Taiwan (15♀♀13♂♂), Thailand (1♀), unknown (1♂).
Type series:
Australasian, Eastern Palaearctic, and Oriental regions (
JAPAN: [Tôhoku] Aomori (
This is one of the most frequently encountered wasps in Japan as well as in southeast Asia, especially Sundaland.
Unknown.
This species is very easily distinguished from all other Japanese
This species exhibits a wide range of colour variation, but the Japanese specimens are morphologically stable. Japanese specimens have a slightly wider mandible than specimens from other regions, as mentioned by
Total of 21 specimens (12♀♀9♂♂): China (1♀1♂), Japan (10♀♀6♂♂), Nepal (1♂), Taiwan (1♀1♂).
Type series:
Eastern Palaearctic and Oriental regions (
JAPAN: [Kantô-Kôshin] Saitama*; [Tôkai] Shizuoka*; [Kinki] Hyôgo*; [Chûgoku] Shimane* and Hiroshima (
No host records from Japan. Described as a parasitoid of
This species can be very easily distinguished from all other
This species exhibits a wide range of morphological variation in size and colour pattern. The proximal sclerite is usually weakly pigmented but is strongly pigmented in the holotype of
This species is very close to
Total of 8 specimens (5♀♀3♂♂): Japan (5♀♀3♂♂).
Eastern Palaearctic region.
JAPAN: [Hokuriku] Niigata; [Kantô-Kôshin] Ibaraki; [Tôkai] Aichi; [Kinki] Ōsaka.
Unknown.
Although this species is very close to
Female (n = 5). Body length 20.0–22.0 (
Head with
Mesosoma entirely very strongly shiny with setae (Fig.
Wings (Fig.
Legs. Hind leg with coxa in profile 1.9–2.0× (
Metasoma with
Colour (Fig.
Male (n = 3). Very similar to female.
Total of 124 specimens (85♀♀37♂♂ and 2 unsexed): Brunei (3♀♀), India (1♀), Japan (81♀♀37♂♂ and 2 unsexed).
Type series:
Eastern Palaearctic and Oriental regions (
JAPAN: [Hokkaidô] (
Reared from
This species resembles
Total of 31 specimens (24♀♀7♂♂): Japan (22♀♀6♂♂), Taiwan (2♀♀1♂).
Oriental region (
Newly recorded from Japan.
JAPAN: [Hokkaidô]; [Hokuriku] Niigata; [Kantô-Kôshin] Tochigi, Nagano, Yamanashi, and Saitama; [Kinki] Hyôgo; [Shikoku] Kôchi; [Kyûshû] Fukuoka.
Unknown.
This species is most similar to
Specimens with various shapes of sclerites of the fore wing fenestra but which are very similar in sculpture are identified as this species by the key provided by
Total of 174 specimens (143♀♀25♂♂ and 6 unsexed): Australia (1♀), Brunei (2♀♀), India (37♀♀9♂♂ and 1 unsexed), Japan (54♀♀3♂♂), Nepal (4♀♀), Papua New Guinea (2♀♀), Sri Lanka (1♀), Taiwan (41♀♀13♂♂ and 5 unsexed), unknown (1♀).
Type series:
Australasian, Eastern Palaearctic, Oceanic, and Oriental regions (
Newly recorded from Australia, Bhutan, Brunei, Indonesia, Laos, Malaysia, Nepal, New Caledonia, Papua New Guinea, Philippines, Solomon Islands, Sri Lanka, Tajikistan, and Taiwan.
JAPAN: [Hokkaidô] (
No host records from Japan. A variety of hosts have been reported in the literature (e.g.,
This species is easily distinguished from all other Japanese species of
According to
The treatment of
Total of 144 specimens (93♀♀50♂♂ and 1 unsexed): England (4♀♀1♂), Germany (1♀), Italy (1♀), Japan (71♀♀47♂♂ and 1 unsexed), Korea (1♀), Mallorca (6♀♀), Russia (1♀), Scotland (1♀), Spain (1♀), Sweden (2♀♀1♂), Switzerland (1♂), unknown (4♀♀).
Afrotropical, Oriental, and trans-Palaearctic regions (
JAPAN: [Hokkaidô] (
Recorded from a wide variety of hosts, but some records are undoubtedly the result of misidentifications of the ichneumonid. Reliable rearings are from species of
This species is sometimes confused with
Total of 29 specimens (20♀♀7♂♂ and 2 unsexed): Indonesia (1♀), Japan (16♀♀5♂♂ and 1 unsexed), Madagascar (1 unsexed), Malaysia (2♀♀), Taiwan (1♀2♂♂).
Type series:
Afrotropical, Australasian, Eastern Palaearctic, Oceanic and Oriental regions (
Newly recorded from Indonesia.
JAPAN: [Kantô-Kôshin] Tochigi (
Unknown.
This species is one of the most easily identified
The central sclerite of the Madagascan specimens is smaller than others. The colour of the interocellar area is usually a useful character for identification of
The nasute clypeus of
Total of 141 specimens (94♀♀29♂♂ and 18 unsexed): Japan (45♀♀13♂♂), Taiwan (49♀♀16♂♂ and 18 unsexed).
Type series:
Eastern Palaearctic and Oriental regions (
Newly recorded from Indonesia.
JAPAN: [Tôhoku] Aomori*; [Kantô-Kôshin] Tochigi* and Kanagawa*; [Tôkai] Shizuoka*; [Kyûshû] Kagoshima (
Reared from two species of
This species is very easily distinguished from all other species of
We could find no morphological differences between
Total of 9 specimens (6♀♀3♂♂): Laos (4♀♀1♂), Taiwan (2♀♀2♂♂). No Japanese specimens available.
Type series:
Eastern Palaearctic and Oriental regions (
JAPAN: [Ryûkyûs] Okinawa (
Unknown.
This species resembles
The specific name is dedicated to Dr Michael Sharkey (University of Kentucky Lexington, Kentucky, USA) who collected many of the type series.
Total of 25 specimens (15♀♀10♂♂): Japan (15♀♀10♂♂).
Eastern Palaearctic region.
JAPAN: [Hokkaidô]; [Hokuriku] Toyama; [Kantô-Kôshin] Tochigi and Gunma; [Tôkai] Mie; [Kinki] Kyôto and Wakayama; [Shikoku] Tokushima and Kôchi; [Kyûshû] Fukuoka, Ōita, and Kagoshima.
Unknown.
This species has a very similar colour pattern to
Female (n = 15). Body length 19.0–22.5 (
Head with
Mesosoma entirely shiny with setae (Fig.
Wings (Fig.
Legs. Hind leg with coxa in profile 1.7–1.8× (
Metasoma with
Colour (Fig.
Male (n = 10). Very similar to female.
This species is morphologically very stable except that the mesosoma varies from entirely dark to reddish.
Total of 93 specimens (36♀♀55♂♂ and 2 unsexed): Japan (35♀♀54♂♂ and 2 unsexed), Korea (1♀1♂).
Type series:
Eastern Palaearctic and Oriental regions (
JAPAN: [Hokkaidô]*; [Hokuriku] Niigata (
No host records from Japan.
As mentioned in the diagnosis of
This species exhibits a wide range of colour variation from entirely testaceous to dark brown or black. Paler individuals have the proximal and distal sclerites separated and the central sclerite weak, so it is likely that the degree of melanisation has an effect on the sclerite development as well as the colour. The holotype of
Total of 87 specimens (64♀♀21♂♂ and 2 unsexed): Chagos archipelago (2♀♀10♂♂ and 2 unsexed), India (40♀♀5♂♂), Japan (18♀♀3♂♂), Solomon Islands (1♀1♂), Taiwan (2♀♀2♂♂), Vanuatu (1♀).
Type series:
Australasian, Eastern Palaearctic, Oceanic, and Oriental regions (
JAPAN: [Hokkaidô]*; [Hokuriku] Niigata*; [Kantô-Kôshin] Tôkyô*; [Tôkai] Shizuoka*; [Chûgoku] Hiroshima*; [Kyûshû] Kagoshima*; [Ryûkyûs] Kagoshima* and Okinawa (
No host records from Japan. Reported as a parasitoid of
According to
The holotype of
Total of 208 specimens (154♀♀46♂♂ and 8 unsexed): India (39♀♀17♂♂), Indonesia (7♀♀5♂♂ and 3 unsexed), Japan (95♀♀23♂♂), Sri Lanka (1♀), Taiwan (12♀♀1♂ and 5 unsexed).
Type series:
Australasian, Eastern Palaearctic, Oceanic, and Oriental regions (
JAPAN: [Kantô-Kôshin] Tôkyô (
Japanese host records are from several species of plusiine
This species is morphologically close to
Total of 7 specimens (5♀♀1♂ and 1 unsexed): Japan (2♀♀1♂), Sri Lanka (2♀♀ and 1 unsexed), Taiwan (1♀).
Oriental region (
Newly recorded from Japan.
JAPAN: [Ryûkyûs] Okinawa.
Unknown.
The specific name is derived from the name of the collector of the holotype specimen, a famous Japanese insect taxonomist, Masatoshi Takakuwa.
A holotype male only.
Eastern Palaearctic region.
JAPAN: [Kantô-Kôshin] Tôkyô (Miyake-jima Island).
Unknown.
This species is more or less similar to
Male (
Head with
Mesosoma entirely rather weakly shiny with fine setae (Fig.
Wings (Fig.
Legs. Hind leg with coxa in profile 1.7× as long as deep; basitarsus 1.9× as long as second tarsomere; fourth tarsomere 3.3× as long as wide; tarsal claw simply pectinate.
Metasoma with
Colour (Fig.
Female. Unknown.
Total of 67 specimens (39♀♀26♂♂ and 2 unsexed): China (1♀), India (1♂), Japan (12♀♀14♂♂ and 1 unsexed), Nepal (24♀♀8♂♂ and 1 unsexed), Taiwan (2♀♀), unknown (3♂♂).
Type series:
Eastern Palaearctic and Oriental regions (
JAPAN: [Tôhoku] Miyagi*; [Hokuriku] Niigata* and Ishikawa*; [Kantô-Kôshin] Kanagawa*; [Tôkai] Shizuoka*, Aichi*, and Mie*; [Chûgoku] Shimane* and Hiroshima (
Unknown.
This species resembles
The specific name is derived from the Latin
The holotype female only.
Eastern Palaearctic region.
JAPAN: [Shikoku] Ehime.
Unknown.
Some characters (e.g., wide face, long and slender mandible, shape of sclerites) suggest that
Female (
Head with
Mesosoma entirely strongly shiny with fine setae (Fig.
Wings (Fig.
Legs. Hind leg with coxa in profile 1.9× as long as deep; basitarsus 2.1× as long as second tarsomere; fourth tarsomere 3.7× as long as wide; tarsal claw simply pectinate.
Metasoma with
Colour (Fig.
Male. Unknown.
Total of 23 specimens (19♀♀4♂♂): Brunei (2♀♀), India (11♀♀1♂), Japan (5♀♀1♂), Sri Lanka (1♂), Taiwan (1♀1♂).
Type series:
Australasian, Oceanic, and Oriental regions (
Newly recorded from Japan.
JAPAN: [Kyûshû] Nagasaki; [Ryûkyûs] Okinawa.
Unknown.
This species resembles
Although Japanese specimens do not vary significantly, some wing characters of the holotype of
Total of 123 specimens (103♀♀19♂♂ and 1 unsexed): Brunei (3♀♀), India (83♀♀5♂♂), Japan (7♀♀9♂♂), Philippines (1♀), Sri Lanka (3♀♀), Taiwan (6♀♀5♂♂ and 1 unsexed).
Type series:
Australasian and Oriental regions (
JAPAN: [Ryûkyûs] Kagoshima (
No Japanese rearings. A range of hosts have been recorded in the literature, with some looking more reliable than others.
This species is sometimes confused with
Total of 31 specimens (18♀♀12♂♂ and 1 unsexed): Japan (16♀♀12♂♂ and 1 unsexed), South Korea (1♀), unknown (1♀).
Type series:
Eastern Palaearctic region (
Newly recorded from South Korea.
JAPAN: [Hokkaidô] (
Unknown.
This species is similar to
This species is rather morphologically stable. The holotype of
Total of 303 specimens (196♀♀103♂♂ and 4 unsexed): India (12♀♀ and 2 unsexed), Indonesia (1♀), Japan (166♀♀101♂♂), Laos (11♀♀1♂), Malaysia (5♀♀ and 2 unsexed), Papua New Guinea (1♀), Taiwan (1♂).
Type series:
Australasian, Eastern Palaearctic, and Oriental regions (
JAPAN: [Tôhoku] Akita*, Yamagata (
Unknown.
There is some variation in the shape of the proximal sclerite, but in Japanese specimens it is usually very stable.
Total of 5 specimens (3♀♀2♂♂): Japan (1♂), Taiwan (3♀♀1♂).
Type series:
Oriental region (
Newly recorded from Japan.
JAPAN: [Ryûkyûs] Okinawa.
Unknown.
This species can very easily be distinguished from all other
Both
The following species have been recorded from Japan in error so were not included in the present study.
A total of 12, 8, 19, 31, 25, and 33 species were observed from the latitudinal zones A to F respectively, and saturation species richness was estimated at 13.98, 8.65, 20.00, 36.99, 32.11 and 56.46 in each zone (Figs
Individual-based extrapolated species accumulation curve, comparing each zone rarefied to 2,000 individuals (LR = latitudinal ranges).
Latitudinal pattern of
Regional patterns of the four categories (i.e., number of specimens, collection events, collectors, and species) are visualised as heat maps in Fig.
Individual-based observed and rarefaction numbers of
Heat maps of regional patterns
Individual-based species accumulation curve, comparing the observed and estimated numbers of
Regional patterns of the number of specimens, collection events, collectors, and species. Bold indicates especially small numbers (fewer than 5).
Provinces | Prefectures | Specimens | Collection events | Collectors | Species |
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Hokkaidô | Hokkaidô | 81 | 53 | 33 | 12 |
Tôhoku | Aomori | 34 | 30 | 8 | 8 |
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Yamagata | 24 | 17 | 7 | 6 | |
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Fukushima | 27 | 15 | 10 | 6 | |
Hokuriku | Niigata | 52 | 36 | 27 | 13 |
Toyama | 11 | 7 |
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Fukui | 38 | 9 |
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Kantô-Kôshin |
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Tochigi | 37 | 22 | 17 | 12 | |
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Nagano | 21 | 15 | 13 | 9 | |
Yamanashi | 6 | 6 | 6 |
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Saitama | 19 | 19 | 13 | 7 | |
Tôkyô | 48 | 36 | 28 | 15 | |
Kanagawa | 34 | 31 | 17 | 12 | |
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Tôkai |
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Shizuoka | 159 | 94 | 33 | 15 | |
Mie | 16 | 16 | 14 | 13 | |
Kinki | Kyôto | 11 | 7 | 7 | 9 |
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Ōsaka | 8 | 6 | 6 | 6 | |
Hyôgo | 52 | 42 | 37 | 17 | |
Nara | 9 |
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Wakayama | 26 | 16 | 12 | 9 | |
Chûgoku |
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Shimane | 12 | 12 | 9 | 9 | |
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Hiroshima | 108 | 101 | 34 | 17 | |
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Shikoku |
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Tokushima | 19 | 13 | 11 | 8 | |
Ehime | 76 | 43 | 23 | 16 | |
Kôchi | 62 | 52 | 32 | 20 | |
Kyûshû | Fukuoka | 105 | 69 | 44 | 18 |
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Nagasaki | 12 | 10 | 8 | 8 | |
Ōita | 10 | 7 | 6 | 6 | |
Kumamoto | 20 | 15 | 15 | 11 | |
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Kagoshima | 112 | 62 | 37 | 18 | |
Ryûkyûs | Kagoshima | 285 | 112 | 69 | 20 |
Okinawa | 242 | 181 | 125 | 31 | |
Ogasawara | Tôkyô | 38 |
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Rarefaction of
Individuals | Observed number of species | Estimated number of species | |
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ACE Mean | Chao 1 Mean | ||
50 | 22.12 | 30.84 | 32.94 |
100 | 29.14 | 33.50 | 34.77 |
150 | 32.64 | 36.58 | 38.07 |
200 | 34.68 | 37.75 | 39.18 |
250 | 36.02 | 39.78 | 42.21 |
300 | 36.99 | 40.83 | 43.55 |
350 | 37.74 | 41.34 | 44.66 |
400 | 38.35 | 41.60 | 43.88 |
450 | 38.88 | 42.15 | 44.75 |
500 | 39.34 | 42.69 | 45.43 |
550 | 39.75 | 43.40 | 45.70 |
600 | 40.12 | 43.61 | 46.69 |
650 | 40.46 | 43.83 | 46.93 |
700 | 40.78 | 44.21 | 47.63 |
750 | 41.07 | 44.65 | 48.16 |
800 | 41.34 | 44.93 | 48.40 |
850 | 41.60 | 45.11 | 48.52 |
900 | 41.84 | 45.88 | 49.08 |
950 | 42.07 | 45.83 | 49.10 |
1000 | 42.29 | 46.02 | 49.07 |
1050 | 42.49 | 46.29 | 49.33 |
1100 | 42.69 | 46.31 | 49.12 |
1150 | 42.88 | 46.42 | 49.32 |
1200 | 43.06 | 46.72 | 50.08 |
1250 | 43.23 | 47.11 | 50.63 |
1300 | 43.40 | 47.25 | 50.05 |
1350 | 43.56 | 47.38 | 50.37 |
1400 | 43.72 | 47.70 | 50.79 |
1450 | 43.87 | 47.89 | 50.67 |
1500 | 44.02 | 48.04 | 51.12 |
1550 | 44.16 | 48.38 | 52.16 |
1600 | 44.30 | 49.07 | 52.50 |
1650 | 44.44 | 49.61 | 53.15 |
1700 | 44.58 | 50.77 | 53.98 |
1750 | 44.72 | 51.97 | 54.66 |
1800 | 44.85 | 53.02 | 54.83 |
1850 | 44.99 | 54.79 | 55.02 |
We revised the Japanese species of
Species richness of
An overwhelmingly larger number of species in zone F (= Ryûkyûs) is worthy of special mention, especially as the diversity of habitats in Ryûkyûs is apparently narrower than the other zones; these suggest that Ryûkyûs is one of the biodiversity hotspots of
Nine of 47 (19%) Japanese
In the present study, a total of 47 species are recognised in Japan. However, some species, such as
Sampling of
Regional sampling biases also seem to be related to the distribution of universities with traditional entomological laboratories or of active and large entomological societies.
In regions which are difficult to access and/or far from large cities (and which are not famous collection sites, nor near entomological laboratories and societies),
These regional sampling biases affect not only analyses of species richness of
More comprehensive sampling can be achieved through a combination of professional study and citizen science, amateur collecting. Either alone will be insufficient. In Japan, there are many people who enjoy entomology, especially field collecting (like hunting) and making private collections. This hobby, and feeling insects to be very special, is often called “Mushi-ya” (
The study of a large specimen base has been vital for resolving the taxonomic confusion that has surrounded Japanese
We would like to express our sincere thanks to the following museum curators, researchers, and collectors who made specimens available to us and/or loaned valuable material: Akihiko Shinohara (
This research is partially supported by the Grants-in-Aid for JSPS KAKENHI (Grant numbers 19H00942, 24405028 and 26840134) and the Grant-in-Aid for JSPS Fellows (Grant Number 18J20333) to SS from the Japan Society for the Promotion of Science. The JSPS Overseas Challenge Program for Young Researchers enabled SS to carry out research at
Table S1. Specimens examined
specimen data
Abbreviations for repositories are as follows:
Table S2. Specimen data used in the DNA barcoding analyses
molecular data
Abbreviations for identifiers: AB, Andrew Bennett; DJ, Daniel Janzen; DQ, Donald L.J. Quicke; PR, Pascal Rousse; and SS, So Shimizu. Abbreviations for countries: BEN, Benin; BLZ, Belize; CAN, Canada; CRI, Costa Rica; ENG, England; ESP, Spain; GUF, French Guiana; JPN, Japan; LAO, Laos; MDG, Madagascar; MYS, Malaysia; PYF, French Polynesia; THA, Thailand; TWN, Taiwan; and ZAF, South Africa.
Checklist and nomenclatural summary of the Japanese species of
species list