Based on an examination of 934 specimens from 19 major Asian, European, and North American natural history collections, we documented 25
Within the 300 bp
Maximum likelihood tree of twelve
The East Asian species of
The third tarsomeres are emarginate (e.g., Fig.
Diagnostic morphological characters in
The proportions of funicular antennomere lengths vary between
One species covered here (
Scutella and pronota of various
The Taiwanese species
The morphology of the scutellum is extremely variable and very important in
Variation in the sclerolepidia has been documented in
Sclerolepidia in
Many
Pronotum punctation differs between species in some instances; however, this character is generally not visible as the pronotum is often covered by scales. We used X-ray μCT (as in
Using X-ray μCT, we visualized the metanotum and dorso-longitudinal flight muscles (DLFM), which moderately differ in proportion to the elytra in
Feminine.
Body length 2.0–12.0 mm.
The larvae of
Body length 5.0–6.8 mm. Covered in black and dark gray scales, with longitudinal, black-bordered white stripes across the lateral edges of the pronotum and elytral humeri (compare with
Aedeagi of
This species occurs in far Eastern Russia (Primorsky Krai, Saghalien) and north-central mainland Japan (from Hokkaido south to Gifu Prefecture).
The two
Body length 3.3–4.9 mm. Covered in pale brown scales, with alternating white and black scaled bands on odd elytral intervals. Pronotum with two exposed, bare tubercles medially at the base (one on each side) (Fig.
This species occurs in southern Japan (Okinawa Prefecture, Ryukyu Islands) and South Korea.
On Okinawa Island (Japan) this species is commonly encountered in the spring and is often attracted to residential building lights.
Body length 4.0–5.2 mm. Covered in dark-gray scales, with two V-shaped, transverse black bands across middle of elytra. Pronotum with pale brown and yellow scales dorsally, but dark at base medially. Pronotum approx. sculptured and carinate. Scutellum shiny and bare in examined specimens. Only odd elytral intervals with erect scales across elytral disk (some erect scales on 2nd interval in apical fourth of elytra). Third and fifth elytral intervals with cluster of 3–5 erect scales at elytral base. Sclerolepidia distinct and protruding. Mesosternum with distinctly raised ridge between meso-coxae, highest laterally but dipping shallowly medially (no such ridge or only weakly developed in most other species). Third tarsomeres weakly emarginate.
This species is currently only known from Taiwan.
Body length 5.0–7.0 mm. Easily distinguished from other East Asian
This species is known only from Taiwan.
Many similarly patterned, large species that presumably form a distinct clade within
Body length 5.0 mm. Virtually indistinguishable externally from
This species is known from the type locality (Mau-son, Tongking, which corresponds to modern day northern Vietnam) and apparently also Kuatun, China (Voss’s specimen).
We examined the holotype specimen of
Body length 4.8 mm. This is the only species covered here with costate elytra (on the 2nd, 4th, and 6th intervals). Also, is has erect, elongate scales along the odd-numbered elytral intervals (as in
Body length 4.9–7.0 mm. Covered in dark to pale gray and black scales. Pronotum covered in black scales medially, but with white to yellow scales laterally (compare with
This species is endemic to mainland Japan (excluding Hokkaido), north from Aomori Prefecture and south to Kyushu (Oita Prefecture, Fukuoka Prefecture).
The specific name
This cryptic species was previously assumed to be a pale-color morph of
(Japan: 34).
Body length 4.2–7.0 mm. Covered in black and dark gray scales, with two V-shaped, transverse white and black bands across middle of elytra. Pronotum with weakly patterned gray, black and faint yellow scales. Scutellum covered loosely in yellow scales (denuded in many specimens). Only odd elytral intervals with erect scales across elytral disk (some erect scales on 2nd interval in apical fourth of elytra). Sclerolepidia distinct and protruding. Third tarsomeres distinctly emarginate. Pedon with lateral edges approx. parallel in dorsal view, but converging evenly into a moderately acute point. Internal sac not visible inside aedeagus, but with a non-hooked protruding structure (unmodified in posterior 1/2, with roughened in anterior 1/2) that is clearly longer than pedon (Fig.
This subspecies occurs in mainland Japan (central Japan south to Yakushima Island) and adjunctly on Hachijo Island (Tokyo Metropolis).
Body length 4.8–6.5 mm. Identical to
This subspecies occurs on Amami-Oshima (Kagoshima Prefecture) and Okinawa Island (Okinawa Prefecture).
We continue to recognize this subspecies as it is significantly isolated from the mainland populations and therefore presumably does represent a distinct intraspecific lineage.
Body length 2.1–3.0 mm. Covered in pale brown, white, and gray scales, with two white-scaled spots in the apical 1/2 of the elytra. Scutellum covered in yellow scales. Only odd elytral intervals with erect scales. Sclerolepidia small but protruding. Third tarsomeres distinctly emarginate. Pedon with lateral edges approx. parallel in dorsal view, but converging evenly into a rounded point. Internal sac without tuberculate or roughened musculature inside the aedeagus, but with a non-hooked protruding structure (unmodified in posterior 2/3, sclerotized in anterior 1/3) that reaches the tip of the temo (Fig.
This small species occurs throughout the Ryukyu Islands (southern Kagoshima Prefecture and Okinawa Prefecture, Japan) as well as Taiwan.
This species appears to be closely allied with the similarly minute, long hind-legged Australian species
Body length 4.5–7.0 mm. Covered in pale and dark brown scales, with two obliquely shaped pale brownish, black-bordered spots across middle of elytra. Second funicular antennomere longer than funicular antennomeres 3 + 4 (Fig.
This distinctive species occurs in Japan (mainland, throughout the Ryukyu Islands) and Taiwan.
(Japan: 3).
Body length 4.5–7.0 mm. Covered in pale and dark brown scales, with two obliquely shaped pale brownish, black-bordered spots across middle of elytra. Second funicular antennomere shorter than funicular antennomeres 3 + 4 (Fig.
This rare species is only known from two localities in mainland Japan, Mt. Takao (Tokyo Metropolis) and Mt. Otoh (Wakayama Prefecture).
The specific name
This cryptic species was clearly overlooked due to its remarkable similarity with
Body length 4.1–5.8 mm. Covered in pale brown scales, with alternating white and black scaled bands on odd elytral intervals. Scutellum covered in yellow to brown scales. Only odd elytral intervals with erect scales. Sclerolepidia distinct and protruding. Posterior edge of prosternum with prominent projections contiguous with the fore-coxae (Fig.
We only encountered this species from Taiwan. The records from Japan and the Kuril Islands (Russia / Japan) noted in
This species is superficially quite similar to
Body length 4.0–5.4 mm. Covered in pale brown scales, with two obliquely shaped tan-colored, black-bordered spots across middle of elytra. Pronotum lacking impunctate longitudinal midline (present in
This species is found in mainland Japan (except Hokkaido) and has also been reported from South Korea (
Body length 2.6–5.0 mm. Covered in pale and dark brown scales, with two obliquely shaped pale brownish, black-bordered spots across middle of elytra. Pronotum lacking impunctate longitudinal midline (present in
This species occurs broadly throughout Japan (mainland Japan and Ryukyu Islands) and Taiwan and is also recorded from China by
Body length 4.1–5.2 mm. Covered in dark gray and black scales, with a pale-yellow scale, V-shaped, transverse band across middle of elytra. Basal patch of black scales surrounding scutellum approx. V-shaped and extending prominently onto the second elytral intervals (as opposed to the similar
This species is currently only known from Taiwan.
Body length 4.9–6.0 mm. Covered in dark gray and black scales, with a white-scaled, V-shaped, transverse band across middle of elytra. Elytra with elongate I-shaped patch of black scales on the first elytral interval (surrounding the scutellum) that barely extends onto the second elytral interval. Pronotum lacking impunctate longitudinal midline (present in
This species is apparently endemic to mainland Japan and occurs from Hokkaido south to Kyushu (Oita Prefecture).
Body length 6.1–7.0 mm. Covered in pale brown, dark gray, and black scales, with a tan-colored, V-shaped, transverse band across middle of elytra. Pronotum lacking impunctate longitudinal midline (present in
This species occurs in Japan (mainland) and China, and has also been reported from South Korea (
An examination of the holotype of
• Kosempo, “VII–19”, H. Sauter,
Body length 3.2–5.2 mm. Covered in orange, white, dark and pale gray scales. Elytra with inner five or six intervals black and gray; outer intervals with pale brown to orange scales; with V-shaped band of white scales along middle three or four intervals in center of elytra. Pronotum lacking impunctate longitudinal midline (present in
This species occurs commonly throughout southern Japan (Ryukyu Islands) and Taiwan.
Examination of the
Body length 3.2–5.0 mm. Covered in dark gray, black, and bright yellow scales, with a black-bordered, yellow spot at the center of each elytron. Apex of elytra covered in yellow scales, and contrasting with overall dark dorsal surface of elytra. Pronotum lacking impunctate longitudinal midline (present in
This species is known only from Okinawa Prefecture, Japan (Yambaru National Park (Okinawa Island) and forested regions of Ishigaki Island.
This species is very distinct from other Japanese
Body length 4.9–6.2 mm. Covered in pale and dark brown scales, with two obliquely shaped white, black-bordered spots across middle of elytra. Pronotum lacking impunctate longitudinal midline (present in
This species occurs in southern Japan (Kagoshima Prefecture) and Taiwan, and has also been recorded from South Korea (
This species is closely allied with
Body length 3.0–3.8 mm. Covered in dark and pale brown, white, and black scales, with a white-scaled, V-shaped band in the center of the elytron. Pronotum lacking distinct longitudinal, impunctate midline (present in
This species occurs in Japan (mainland, Ryukyu Islands south to Amami-Oshima), South Korea, and Taiwan.
Body length 5.1–7.0 mm. Covered in pale brown, dark gray, and black scales, with a tan-colored, V-shaped, transverse band across middle of elytra. Pronotum lacking distinct longitudinal, impunctate midline (present in
This species is endemic to Okinawa Prefecture, Japan, and is currently only known from Ishigaki, Iriomote, and apparently also Okinawa Island.
This species is quite similar in size and color to
Body length 4.1–5.0 mm. Covered in pale and dark brown scales, with bright X-shaped band of white scales across apical 1/2 of elytra, which extends up to the elytra humeri. Distinctive bright white scale bands also present on hind femora. Pronotum lacking distinct longitudinal, impunctate midline (present in
This widely distributed species occurs in Vietnam, Thailand, Myanmar, and Taiwan.
One of K.M. Heller’s specimen of this species in the
Body length 3.0–4.4 mm. Covered in pale yellow, white, gray, and black scales. Pronotum with central longitudinal band of dark gray to black scales, bordered laterally by pale brown scales. Elytra with first intervals alternating black (at base of scutellum) and yellow; second to sixth intervals gray in posterior 1/2 of elytra, humeri with yellow to pale scales; apical 1/2 of elytra with transverse white V-shaped band extending from first to fifth intervals; apex of elytra largely with pale to yellow scales. Pronotum lacking impunctate longitudinal midline (present in
This species is only known from mainland Japan, from Hokkaido south to Kyushu (Kumamoto Prefecture).
The specific name
See
X-ray μCT generated 3D models of two
Body length 4.0–4.7 mm. Covered in pale brown, dark and pale gray, pale yellow, and black scales, with two small white spots on the elytra (one on each side) in the apical 1/2 of the elytra. Pronotum lacking impunctate longitudinal carina (present in
This distinctive species occurs in Japan (mainland), Taiwan, and has also been reported from South Korea and the Kuril Islands (Russia) (
| 1 | Third tarsomeres truncate or weakly emarginate (Fig. |
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| – | Third tarsomeres strongly emarginate (Fig. |
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| 2 | Second and fourth elytral intervals with erect scales |
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| – | Second and fourth elytral intervals without erect scales (may have small erect scales at elytral apex) |
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| 3 | Erect scales on odd elytral intervals elongate, hair-like (distinctly longer than antennal club) |
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| – | Erect scales on odd elytral intervals stubby, round (shorter than antennal club) |
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| 4 | Sclerolepidia inconspicuous, low and rounded; third tarsomeres truncate; scales covering body predominantly brown |
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| – | Sclerolepidia distinct and protruding; third tarsomeres weakly emarginate; scales covering body predominantly gray |
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| 5 | Second elytral interval with erect scales |
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| – | Second elytral interval lacking erect scales (may have small erect scales at elytral apex) |
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| 6 | Scutellum distinctly large, wide, and lacking scales (Fig. |
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| – | Scutellum slender and largely covered in scales dorsally (e.g., Fig. |
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| 7 | Internal sac of aedeagus protruding but with roughened musculature proximal to pedon (Fig. |
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| – | Internal sac of aedeagus protruding but with roughened musculature located at distance from pedon (Fig. |
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| 8 | Even-numbered elytral intervals costate |
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| – | Elytral intervals not costate |
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| 9 | Posterior edge of prosternum with prominent projections contiguous with the fore-coxae (Fig. |
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| – | Posterior edge of prosternum lacking prominent projections contiguous with the fore-coxae (e.g., Fig. |
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| 10 | Cuticle distinctly naked (lacking scales) across much of body; pronotum roughly sculptured and carinate |
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| – | Cuticle largely covered in scales (sometimes specimens are denuded); pronotum punctate but not roughly sculptured or carinate (except at base of pronotum in |
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| 11 | Pronotum with two exposed, bare, red tubercles medially at the base (Fig. |
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| – | Pronotum lacking tubercles at base (Fig. |
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| 12 | Scutellum distinctly shiny and bare dorsally (Fig. |
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| – | Not with the above combination of characters |
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| 13 | Pedon distinctly tapered apically into a fine point (Fig. |
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| – | Not with the above combination of characters |
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| 14 | Small species (3.0–3.8 mm) with brown scaling and distinct white scaled V-shaped band in posterior 1/2 of elytra; pedon tube-shaped and internal sac with a small, sclerotized structure basally, and 2 prong-like structures anteriorly at its apex (Fig. |
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| – | Size and color variable ; aedeagus not as above |
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| 15 | Small species (3.0–4.4 mm) with pale and dark brown scales, and shallow, white-scaled, V-shaped band in posterior 1/2 of elytra; internal sac of aedeagus distinctly scaly in appearance and protruding; apex of internal sac ending in two scaly feather-like tips; apex of pedon with distinct sinuation and pointed (Fig. |
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| – | Size and color variable; aedeagus not as above |
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| 16 | Large species (5.1–7.0 mm) with pale brown, dark gray, and black scales, and a tan-colored, V-shaped, transverse band across middle of elytra; scutellum covered in erect scales (Fig. |
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| – | Size and color variable; aedeagus not as above |
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| 17 | Exceptionally small species (2.1–3.0 mm), covered in pale brown, white, and gray scales, with 2 white-scaled spots in the apical 1/2 of the elytra; pedon with lateral edges approx. parallel in dorsal view, but converging evenly into a blunt, rounded point; internal sac without tuberculate or roughened musculature inside the aedeagus, but with a non-hooked protruding structure (unmodified in posterior 2/3, sclerotized in anterior 1/3) that reaches the tip of the temo (Fig. |
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| – | Larger species (3.2–7.0 mm), variable in color; aedeagus not as above |
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| 18 | Elytra with distinct white-scaled, X-shaped band across elytra (extremely bright in some specimens); pedon with lateral edges approx. parallel in dorsal view, but sinuating in distal 1/2 into a distinct point; internal sac protruding and clear across basal 2/3; with two bean-shaped and U-shaped sclerotized structure anteriorly at its apex (Fig. |
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| – | Elytra with white-scaled V-shaped or transverse band, but never an X-shaped band; aedeagus not as above |
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| 19 | Brightly colored species, with dull orange, white, and brown scales; pedon with lateral edges approx. parallel in dorsal view, but converging evenly into a blunt, rounded point; internal sac not visible inside aedeagus, but protruding basally and with 2 sclerotized cylinder-shaped structures posteriorly at the base (Fig. |
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| – | Dark scaled species, with black and gray scales covering most of body; aedeagus not as above |
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| 20 | Internal sac with tuberculate (roughened) musculature, visible inside of aedeagus; internal sac protruding from aedeagus and weakly hooked anteriorly (Fig. |
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| – | internal sac not with tuberculate musculature visible inside of aedeagus (only in protruding portion of sac in some species); body size variable |
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| 21 | internal sac with hook-shaped protruding structure and roughened musculature at the base (Fig. |
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| – | Internal sac with modified protruding structure but not with roughened musculature and weakly hooked shape; pronotum never with impunctate longitudinal midline; size variable |
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| 22 | Internal sac without club-shaped structure anteriorly at its apex (Fig. |
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| – | Internal sac with club-shaped structure anteriorly at its apex; pronotum nearly uniform in color, with black and dark gray scales (no prominent lateral bands) |
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| 23 | Club of internal sac surrounded closely by roughened musculature (Fig. |
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| – | Club of internal sac not surrounded closely by roughened musculature (Fig. |
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