Corresponding author: Hongbin Liang (
Academic editor: Terry Erwin
Three new species of genus
To date, nine species of the subgenus have been described from Asia, namely,
To date, no key including all known species of
We measured all available specimens for each species except for those with plentiful specimens, for which five males and five females were measured, including smallest and largest specimens for each sex (determined by visual inspection of the assembled samples). Measurements were made with the aid of a Nikon SMZ1500 stereoscopic dissecting microscope with a micrometer. Body length (BL) was measured as the linear distance along the midline from the apex of the longer mandible to the apex of the longer elytron. Other measurements, and abbreviations used for them in this paper, are as follows: HW = maximum head width including the eyes; EYL = eye length measured along the longitudinal diameter of the eye (dorsal–lateral view); PL = length of pronotum measured along median line; PW = pronotum width at its widest point; EL = elytron length from base to apex; EW = width across both elytra at widest point (equal to body width).
Wherever we refer to abdominal ventral plates, we use the numbering system that recognizes the generally accepted segmental homologies in
All photographs were taken through a Nikon stereoscopic dissecting microscope fitted with a Canon 450D camera, and were edited by Helicon Focus and Photoshop software.
Specimens examined in the course of this study were deposited at the following collections:
Natural History Museum, London, U.K.
California Academy of Sciences, San Francisco, U.S.A.
Private Collection of Changchin Chen, Tianjin, China
Museum of Hebei University, Baoding, China
National Zoological Museum of China, Institute of Zoology, Beijing, China
Museum National d'Histoire Naturelle, Paris, France
Narodni Muzeum, Prirodovedecke Muzeum, Prague, Czech Republic
Osaka Museum of Natural History, Osaka, Japan
Institute of Plant Physiology & Ecology, Shanghai Institutes for Biological Sciences, Chinese Academy of Sciences, Shanghai, China
Zoological Institute, Russian Academy of Sciences, St. Petersburg, Russia
Antennomere 3 distinctly longer than antennomeres 1 and 2 combined (
Length 12.0–18.0 mm, width 4.6–6.4 mm. Head and pronotum black, with green, blue or coppery metallic luster; elytra black, with blue or coppery luster in a few species; ventral surface black; legs black, brown or yellow; antennae yellow, brown, or dark brown.
Head with vertex nearly glabrous, or sparsely and coarsely punctate behind posterior level of eye; eyes moderately prominent; genae pubescent; antennae long, antennomere 1 (scape) coniform (
Pronotum cordate (
Elytra moderately convex, oblong, with distinct isodiametric microsculpture at least laterally; scutellar striae long, with basal setiferous pore present (
Abdominal sterna densely pubescent laterally, sparsely pubescent or glabrous medially (
Pro- and mesosterna and pro-, mes-, and metepisterna densely pubescent; metasternum pubescent laterally, nearly glabrous medially; prosternal intercoxal process punctate and bordered at apex; scutellum triangular, glabrous.
Anterior femora without tooth; tarsomeres short, sparsely setose dorsally; anterior tarsomere 4 short and triangular, deeply emarginate (more so in male), with two rows of long setae ventrally (
Male genitalia with aedeagus tubular, simple; ejaculatory orifice long (
Character states of
Habitus of
Habitus of
China (Heilongjiang, Liaoning, Shaanxi, Gansu, Henan, Hubei, Hunan, Zhejiang, Fujian, Taiwan, Jiangxi, Guangdong, Hainan, Guangxi, Guizhou, Sichuan, Yunnan, Tibet), North Korea, Japan, Russia (Far East), Afghanistan, Pakistan, India. The known localities of
Members of this subgenus are typically collected on sandy beaches of rivers or streams (
Based on the metallic body surface, single supraorbital seta, antennomere 3 longest and antennomeres 4–11 densely pubescent,
As presently conceived, the subgenus can be divided into two species groups: 1) the
In his treatment of the North American species of genus
1 | All intervals punctate, pubescent, slightly convex ( |
|
– | At least basal half of intervals 1–5 smooth and glabrous medially, distinctively convex ( |
3 |
2 | Intervals dull, with dense and regular punctures ( |
|
– | Intervals shining, with sparse and irregular punctures (large and small punctures mixed) ( |
|
3 | Antennomere 1 with apical end much thicker than basal end, coniform ( |
4 |
– | Antennomere 1 with apical end as thick as basal end, elongate-ovoid ( |
8 |
4 | Intervals 1–7 glabrous medially, with a row of pubescence laterally (near striae) ( |
|
– | Intervals 6–7 pubescent, intervals 1–5 glabrous medially, with a row of pubescence laterally (near striae) ( |
5 |
5 | Tibiae and femora bi-colored, with tibiae black or dark brown ( |
|
– | Tibiae and femora concolorous, yellow or brown ( |
6 |
6 | All antennomeres brown or dark brown ( |
|
– | At least antennomere 1 yellow or brown, paler than the rest ( |
7 |
7 | Antennomeres 1–3 brown, color paler than antennomeres 4–11 ( |
|
– | Only antennomere 1 yellow or brown, color paler than antennomeres 2–11 ( |
|
8 | Intervals 1–7 glabrous medially ( |
|
– | At most intervals 1–5 glabrous medially ( |
9 |
9 | Male with media lobe of aedeagus gradually bent near base ( |
|
– | Male with media lobe of aedeagus abruptly bent near the base ( |
Holotype: male (IZCAS), "China, Guizhou, Xishui County, Dabaitang, 600m”/ "IOZ & Guizhou Univ. Joint Expedition, 2000.9.24, Liang H.B” / ""Holotype,
Antennomere 1 coniform (
Total length = 16.0–17.0 mm (mean = 16.8), width = 5.60–6.13 mm (mean = 5.86); HW = 2.60–2.95 mm (mean = 2.82), EYL = 0.95–1.10 mm (mean = 1.03), ratio Ant3/Ant1 = 1.56–1.72 (mean = 1.64), PL/PW = 0.76–0.84 (mean = 0.81), EL/EW = 1.68–1.91 (mean = 1.79), EL/ PL = 1.37–1.49 (mean = 1.42).
Head and pronotum black with green or coppery metallic luster; elytra black; ventral surface black; mandibles and trochanters dark brown; antennomere 1, femora, tibiae yellow to brown; antennomeres 2–11, palpi and tarsi dark brown to nearly black.
Head with vertex smooth medially, coarsely punctate behind eyes; labrum slightly emarginate at apex; antennomere 1 coniform (
The Latinized name
Mensural data cited in the description were obtained from the holotype and all paratypes.
Members of this species are most similar to those of
They are also similar to
Holotype: male (IZCAS), "Fujian, Jianyang, Huangkeng, Guiling, 270–340m""/ ""1960.III.26, Zhang Yiran collector""/ ""Holotype,
Antennomere 1 coniform (
Total length = 14.50–16.50 mm (mean = 15.71), width = 5.33–5.87 mm (mean = 5.60); HW = 2.40–2.85 mm (mean = 2.71), EYL = 1.00–1.15 mm (mean = 1.05), ratio Ant3/Ant1= 1.37–1.61 (mean = 1.50), PL/PW = 0.82–0.90 (mean = 0.86), EL/EW = 1.67–1.81 (mean = 1.76), EL/ PL = 1.27–1.41 (mean = 1.35).
Head and pronotum black, with green or coppery metallic luster; elytra black; ventral surface black; antennae, palpi, femora, and tibiae brown; mandibles, trochanters, and tarsi dark brown.
Head with vertex smooth medially, coarsely punctate and sparsely pubescent behind eye; labrum slightly emarginate at apex; antennomere 1 strongly coniform (
The Latinized name
Mensural data cited in the description were obtained from the holotype and all paratypes.
Male members of this species are similar to those of
Holotype: male (IZCAS),"Southwest Yunnan, Gongshan, Dulongjiang, 0.5km N of Dizhengdang,
Antennomere 1 elongate ovoid (
Total length = 15.50–17.00 mm (mean = 15.71), width = 5.60–6.13 mm (mean = 5.81); HW = 2.75–2.95 mm (mean = 2.82), EYL = 1.00–1.15 mm (mean = 1.09), ratio Ant3/Ant1 = 1.73–1.93 (mean = 1.81), PL/PW = 0.78–0.85 (mean = 0.83), EL/EW = 1.62–1.79 (mean = 1.73), EL/ PL = 1.29–1.45 (mean = 1.36).
Head and pronotum black with green or coppery metallic luster; elytra black; ventral surface black, with some slight metallic reflection; coxae almost black; mandibles dark brown; antennomeres 4–11, palpomere, trochanters, tibiae at both ends, and tarsomeres reddish or brown; antennomeres 1–3, femora and middle portions of tibiae yellow or yellowish-brown.
Head rugose near eyes and occiput, vertex sparsely rugose; eyes prominent; antennomere 1 elongate–ovoid (
Pronotum cordate, moderately convex; lateral margins bordered; disk almost smooth, sparsely rugose and setose near base and lateral margins; basal foveae moderately deep.
Elytra widest at mid-length; humeral angle obtuse (
Abdominal sterna bordered, densely pubescent laterally; sterna IV–VI sparsely pubescent medially; sternum VII rugose–pubescent, with apex narrowly rounded or subobtuse in males (
Apical lamella of aedeagus moderately triangular (
The Latinized name
Mensural data cited in the description were based on measurements obtained from 5 males and 5 females selected for maximum variation.
Specimens of
This species is clearly very closely related to
Total 15 specimens.
Paratype: 2 males (MNHN), "J. Klapperich, Asmar, 900m, Kunartal, 3.4.53, O-Afghanistan”/ "
Antennomere 1 elongate ovoid (
Total length = 15.50–16.00 mm (mean = 15.50), width = 5.60–6.10 mm (mean = 5.73); HW = 2.75–2.95 mm (mean = 2.82), EYL = 1.00–1.15 mm (mean =1.00), ratio Ant3/Ant1 = 1.67–1.83 (mean = 1.71), PL/PW = 0.84–0.85 (mean = 0.85), EL/EW = 1.63–1.73 (mean = 1.71), EYL/ PL = 1.15–1.44 (mean = 1.34). Mensural data cited in the description were obtained from type and cotype specimens examined.
Head, pronotum and elytra black, with green or blue metallic luster; ventral surface black; mandibles and trochanters dark brown; palpomeres, femora, and tarsomeres brown to dark brown; antennae and tibiae yellow to brown (
Head with vertex smooth or very sparsely punctate behind eyes; antennomere 1 elongate ovoid (
Labels of
Antennomere 1 of
Humeral region of elytra of
Elytral intervals of
Abdominal sternum VII (ventral aspect) of
Aedeagi of
Aedeagi of
Color of antennae, palpomeres, femora, and tibiae varied from medium brown to dark brown or even black among different individuals (
Mensural data cited in the description were obtained from type and cotype specimens examined. The name
Total 24 specimens:
Antennomere 1 cylindrical (
Total length = 16.0–17.0 mm (mean = 16.45), width = 5.87–6.40 mm (mean = 5.99); HW = 2.90–3.00 mm (mean = 2.96), EYL = 1.10–1.15 mm (mean = 1.12), ratio Ant3/Ant1 = 1.71–1.88 (mean = 1.76), PL/PW = 0.81–0.88 (mean = 0.85), EL/EW = 1.68–1.86 (mean = 1.76), EL/ PL = 1.32–1.48 (mean = 1.39).
Head and pronotum black, with green or coppery metallic luster; elytra black; ventral surface black; mandibles, trochanters, and tarsomeres dark brown; antennae, palpomeres, femora, and tibiae yellow to brown.
Head with vertex nearly glabrous, very sparsely punctate and pubescent near eyes; Antennomere 1 cylindrical (
Mensural data cited in the description were based on measurements obtained from 5 males and 5 females selected for maximum variation.
This species was first described as a subspecies of
Total 14 specimens:
Antennomere 1 coniform (
Total length = 15.00–18.00 mm (mean = 16.50), width = 5.33–6.13 mm (mean = 5.78); HW = 2.55–3.00 mm (mean = 2.79), EYL = 0.95–1.10 mm (mean = 1.04), ratio Ant3/Ant1 = 1.53–1.73 (mean = 1.62), PL/PW= 0.80–0.89 (mean = 0.86), EL/EW = 1.64–1.89 (mean = 1.76), EL/ PL = 1.26–1.46 (mean = 1.35).
Head and pronotum black, with green or coppery metallic luster; elytra black; ventral surface black; antennomeres 2–11, palpomeres, trochanters, tibiae, and tarsomeres dark brown to black; antennae and femora yellow to light brown.
Head with vertex coarsely punctate and pubescent behind eyes; antennomere 1 coniform (
Photographs of habitats for
Photographs of habitats for
Mensural data cited in the description were obtained from all cited specimens.
Males of this species (
Total 47 specimens:
Antennomere 1 coniform (
Total length = 14.50–16.00 mm (mean = 15.33), width = 5.07–5.60 mm (mean = 5.33); HW = 2.50–2.90 mm (mean = 2.67), EYL = 1.00–1.10 mm (mean = 1.02), ratio Ant3/Ant1 = 1.56–1.88 (mean = 1.72), PL/PW = 0.83–0.90 (mean = 0.86), EL/EW = 1.75–1.91 (mean = 1.80), EL/ PL = 1.29–1.40 (mean = 1.35).
Head and pronotum black with strong green or bluish green metallic luster; elytra black with very weak blue luster; ventral surface black; legs, antennae, mandibles and palpomere dark brown.
Head with vertex coarsely punctate behind eyes; antennomere 1 coniform (
Mensural data cited in the description were based on measurements obtained from 5 males and 5 females selected for maximum variation.
This species may well be endemic to Japan, and its reported occurrence in Korea (
Total 41 specimens.
Antennomere 1 coniform (
Total length = 16.00–17.00 mm (mean = 16.4), width = 5.33–6.13 mm (mean = 5.68); HW = 2.70–2.95 mm (mean = 2.80), EYL = 1.00–1.05 mm (mean =1.01), ratio Ant3/Ant1 = 1.42–1.65 (mean = 1.52), PL/PW = 0.78–0.80 (mean = 0.79), EL/EW = 1.74–1.90 (mean = 1.80), EL/ PL= 1.37–1.47 (mean = 1.41).
Head and pronotum black with green and coppery metallic luster; elytra black; ventral surface black; antennomeres 4–11, trochanters, palpomeres, and tarsomeres dark brown; antennomeres 1–3, femora, tibiae yellow to brown.
Head with vertex sparsely pubescent and coarsely punctate behind eyes; antennomere 1 coniform (
Mensural data cited in the description were based on measurements obtained from 5 males and 5 females selected for maximum variation.
Members of
Total 603 specimens.
Antennomere 1 cylindrical (
Total length = 14.00–17.00 mm (mean = 15.10), width = 5.33–5.87 mm (mean = 5.60); HW = 2.40–2.85 mm (mean = 2.59), EYL = 0.95–1.00 mm (mean = 0.97), ratio Ant3/Ant1= 1.94–2.21 (mean = 2.08), PL/PW = 0.86–0.90 (mean = 0.88), EL/EW = 1.62–1.74 (mean = 1.68), EL/ PL = 1.30–1.39 (mean = 1.34).
Head, pronotum and elytra black, with blue or greenish blue metallic luster; ventral surface black; antennomeres 4–11, mandibles, palpomeres, trochanters, and tarsomeres brown; antennomeres 1–3, femora, and tibiae yellow.
Head convex, vertex punctate and pubescent behind eyes; antennomere 1 cylindrical (
Mensural data cited in the description were based on measurements obtained from 5 males and 5 females selected for maximum variation.
Members of this species are very similar to
Total 66 specimens.
Antennomere 1 elongate-ovoid (
Total length = 14.00–17.50 mm (mean = 15.60), width = 4.80–6.00 mm (mean = 5.60); HW = 2.40–2.95 mm (mean = 2.63), EYL = 0.95–1.00 mm (mean = 0.96), ratio Ant3/Ant1 = 1.96–2.10 (mean = 2.04), PL/PW = 0.86–0.90 (mean =0.87), EL/EW = 1.65–1.73 (mean = 1.68), EYL/ PL = 1.29–1.40 (mean = 1.35).
Head and pronotum black, with green or coppery metallic luster; elytra black in general, but with slight coppery luster in a few specimens; ventral surface black; antennomeres 4–11, mandibles, palpomeres, trochanters, and tarsomeres brown; antennomeres 1–3, femora and tibiae yellow.
Head convex, vertex punctate and pubescent behind eyes; antennomere 1 elongate–ovoid (
Map showing the known geographical distributions of
Mensural data cited in the description were based on measurements obtained from 5 males and 5 females selected for maximum variation.
We compared a photograph of the holotype (type lost according
We thank Mr. Changchin Chen, Mr. Wensin Lin, Dr. Shigehiko Shiyaki and Dr. Seiji Morita for the loan of specimens in their collections, and Dr. Christine Taylor (BMNH) for loan of the type specimens of