Corresponding author: Adam J. Brunke (
Academic editor: J. Klimaszewski
The Oriental species of the relictual genus
Brunke AJ (2017) A revision of the Oriental species of
Species of the genus
Recently, all Neotropical species of
This study is based on 200 specimens that are deposited in the following collections:
Specimens with GIS coordinates on labels were georeferenced using Google Earth. Specimens with only country or ‘country, province/state’ information were not georeferenced. Specimen data were exported to Darwin Core Archive (
All specimens were examined using a Nikon 745T stereomicroscope. To allow for the routine dissection of the terminal abdominal segments (including the aedeagus), distilled water was applied directly to the tip of the abdomen using a fine paintbrush. As a precaution against DNA degradation, specimens examined in the present study were never subjected to high ambient humidity relaxing chambers or entirely submersed in water. This was in direct contrast to the specimens dissected for
Line illustrations were performed in Adobe Illustrator CS6 based on photographs. Photomontage was accomplished using a motorized Nikon SMZ25 microscope and NIS Elements BR v4.5. Photos were processed in Adobe Photoshop CS6. Distribution maps were created using QGIS 2.18 as in
All measurements were made using a live measurement module within NIS Elements BR v4.5. Measurements were taken as listed below, but only proportional (
Greater intraspecific variability in female tergite X and a general lack of male-associated females made it difficult to incorporate female genitalia as fully in the Oriental species concepts compared to those of the Neotropics. Therefore, female genitalia were not illustrated though they were described where possible. Notches in the apex of female tergite X are typical of many Oriental
A specimen level dataset was made available as a Darwin Core Archive and was deposited in GBIF at
A revision of the Oriental
Electus Group
Caesareus Group
Carnifex Group
Lasti Group
Luteus Group
Pictus Group
Vulneratus Group
Loculus Group
Species
1 | Elytra without red, yellow or orange coloration (e.g., Fig. |
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– | Elytra with yellow, red or orange coloration; pronotum with one puncture (marginal) in the dorsal row, anterior angles more sparsely punctate (Fig. |
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2 | Abdomen bicolored red and black, elytra bright metallic green to blue (Fig. |
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– | Abdomen entirely dark, elytra dark, with only faint metallic reflection (Fig. |
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3 | Paramere with constricted stem, exposing median lobe in parameral view (Fig. |
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– | Paramere vaguely constricted, not exposing median lobe in parameral view (Fig. |
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4 | Head with deeply impressed punctures, many punctures confluent, forming rows (Fig. |
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– | Head with regular, non-impressed punctures, most punctures clearly separated (Fig. |
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5 | Paramere with peg setae medially, on projected ridge (Fig. |
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– | Paramere without projected ridge; peg setae with median group extended to no more than just behind level of marginal group (Fig. |
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6 | Hind tibia entirely dark, as dark as darkened portion of femur (Fig. |
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– | Hind tibia in lateral view with at least distal half distinctly lighter than darkened portion of femur (Fig. |
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7 | Antennomeres 7–10 relatively elongate: 6 quadrate and 7 weakly transverse (Fig. |
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– | Antennomeres 7–10 relatively transverse: 6 weakly, and 7 distinctly transverse (Fig. |
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8 | Apex of median lobe in lateral view forming a more elongate triangle (Fig. |
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– | Apex of median lobe in lateral view forming a shorter triangle (Fig. |
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9 | Peg setae absent from broad oval shaped area along middle of paramere (Fig. |
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– | Peg setae absent from only narrow strip along middle of paramere (Fig. |
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10 | Head dark with frons distinctly paler, light orange (Fig. |
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– | Frons similar in color to rest of head (as in Fig. |
|
11 | Pronotal margin greatly expanded, at its widest point, more than four lateral puncture widths wide (Fig. |
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– | Pronotal margin slightly to moderately expanded, at its widest point, no more than three lateral puncture widths wide (Fig. |
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12 | Elytra with distinct black spot near humerus (Fig. |
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– | Elytra without dark markings (Fig. |
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13 | First two visible abdominal tergites without crisp, dark markings on disc; pronotum less transverse; Java (Fig. |
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– | First two visible abdominal tergites with crisp, dark markings on disc; pronotum more transverse; Borneo (Fig. |
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14 | Median lobe in parameral view evenly narrowed to apex (Fig. |
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– | Median lobe in parameral view dilated near apex and then sharply narrowed (Fig. |
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15 | Apical antennomere dark, not contrasting with previous segments (Fig. |
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– | Apical antennomere pale, contrasting with previous segments (Fig. |
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16 | Dorsal surface of head mostly orange with some dark markings (Fig. |
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– | Dorsal surface of head entirely dark (Fig. |
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17 | Elytra with yellow raised marking v-shaped (Fig. |
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– | Elytra with yellow raised marking transverse (Fig. |
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18 | Abdominal tergites IV–V each with strongly impressed and elongate punctures in basal impression (Fig. |
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– | Abdominal tergites IV–V each with typical circular punctures; only apical antennomere distinctly paler than previous segments (Fig. |
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19 | Pronotal margin greatly expanded, at its widest point, more than four lateral puncture widths wide (Figs |
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– | Pronotal margin slightly to moderately expanded, at its widest point, no more than three lateral puncture widths wide (Figs |
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20 | Base of elytra distinctly darker than rest of disc (Fig. |
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– | Disc of elytra entirely reddish (except for pale markings) (as in Fig. |
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21 | Peg setae arranged in one group removed from margin (Fig. |
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– | Peg setae arranged in lateral and medial groups (Fig. |
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22 | Visible abdominal tergites 1–3 with relatively small dark median spots, occupying less than 1/3 of the tergal width (Fig. |
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– | Visible abdominal tergites 1–3 with relatively wide dark median spots, occupying more than 1/3 of the tergal width (Fig. |
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23 | Antennomeres 8–10 quadrate; paramere with apex only slightly narrowed (Fig. |
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– | Antennomeres 8–10 transverse; paramere strongly narrowed to apex (Fig. |
|
24 | Median lobe in lateral view with subapical tooth on distinct carina (Fig. |
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– | Median lobe in lateral view with subapical tooth retracted from outline, not on carina (Fig. |
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25 | Paramere with slender apical portion, peg setae arranged more in rows (Fig. |
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– | Paramere with apical portion only weakly narrowed apicad, peg setae arranged loosely in two circular formations (Fig. |
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26 | Elytra entirely reddish-orange (as in Fig. |
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– | Elytra at least partly dark |
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27 | Pronotum widest at anterior angles; with distinct pronotal protuberance (Fig. |
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– | Pronotum widest at middle or posterior third (Fig. |
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28 | Species distributed in northeastern India |
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– | Species distributed elsewhere |
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29 | In lateral view, discal elytral markings not extending halfway to epipleural margin, therefore epipleuron at most pale at humerus and apex only (as in Fig. |
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– | In lateral view, discal elytral markings extending halfway to epipleural margin, and often continuing laterad in both directions, epipleuron therefore broadly pale in at least basal half (as in Fig. |
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30 | Minute species (3.5–3.7 mm forebody length); paramere without expansion in lateral view; Khasi Hills, Meghalaya and Himalaya of West Bengal, India |
|
– | Average sized species (4.4 mm forebody length); Garo Hills, Meghalaya, India; paramere with subbasal expansion in lateral view (Fig. |
|
31 | Apex of the median lobe in parameral view with single toothed carina (Fig. |
|
– | Apex of median lobe in parameral view with double-toothed carina (Fig. |
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32 | In lateral view, discal elytral markings not extending halfway to epipleural margin, therefore epipleuron at most pale at humerus and apex only (Fig. |
|
– | In lateral view, discal elytral markings extending halfway to epipleural margin, and often continuing laterad in both directions, epipleuron narrowly to broadly pale at middle (Fig. |
|
33 | Pale elytral marking v-shaped or with inner marking slightly triangular (Fig. |
|
– | Pale elytral marking always consisting of oval-shaped inner marking and small circular outer marking; disc of head with moderately impressed and dense punctation; pronotal margin without distinct expansion (Fig. |
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34 | Base of elytron with broad pale area extending to humerus (Figs |
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– | Medial and lateral elytral spots always distinguishable as separate markings, though usually fused to some degree (Figs |
|
35 | Pale basal part of elytra composed of both yellow to orange raised marking and slightly darker non-raised area (Fig. |
|
– | Pale basal part of elytra composed only of yellow to orange raised area (Figs |
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36 | Antennomeres less transverse, 7 weakly transverse (Fig. |
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– | Antennomeres more transverse, 7 distinctly transverse (Fig. |
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37 | Punctures absent or distinctly sparse nearly in posterior half of sutural area (as in Fig. |
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– | Sutural area with moderately dense regularly spaced punctures (Fig. |
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38 | Outer discal spot of elytra thinly connected to inner spot, to form a chevron or ‘v-shaped’ marking (Figs |
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– | Elytra with differently formed markings (as in Figs |
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39 | Epipleuron with distinct dark area in apical 2/3 (Fig. |
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– | Epipleuron broadly pale throughout its length (Fig. |
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40 | Antennomeres 8 and 9 distinctly transverse (Fig. |
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– | Antennomeres 8 and 9 weakly transverse (Fig. |
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41 | Apex of hind femur entirely darkened (Fig. |
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– | Apex of hind femur with, at most, only ventral half darkened (Fig. |
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42 | Median lobe in lateral view strongly narrowed to thin apex (Fig. |
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– | Median lobe in lateral view not strongly narrowed, apex distinctly broader (Fig. |
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43 | Occurring in the Shan Hills of Myanmar; aedeagus as in Fig. |
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– | Occurring east of Myanmar: northern Thailand, Laos and southern Yunnan, China |
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44 | Median lobe in lateral view slender and not distinctly narrowed to apex (Fig. |
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– | Median lobe in lateral view broader and strongly narrowed to hooked apex (Fig. |
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45 | Antennomere 7 distinctly transverse (Fig. |
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– | Antennomere 7 quadrate to weakly transverse (as in Fig. |
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46 | Paramere narrowly bilobed (Fig. |
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– | Paramere not bilobed (Figs |
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47 | Paramere with distinct lateral and medial rows of peg setae, slender (Fig. |
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– | Paramere without distinct rows of peg setae (Fig. |
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48 | Northern Laos; median lobe in lateral view expanded near short apical portion (Fig. |
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– | Southeastern China and Taiwan; median lobe in lateral view expanded much more basally, apical portion longer (Fig. |
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49 | Paramere exceeding apex of median lobe (Fig. |
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– | Paramere shorter than apex of median lobe (Fig. |
|
The Electus Group (
Yulongshan, Yunnan, China.
Among the members of the Electus Group: head punctures clearly separated (Fig.
Measurements ♂ (n = 5):
Measurements ♀ (n = 4):
Coloration: body black, abdominal segments sometimes slightly paler at base and apex, dorsal forebody often with faint metallic reflection; maxillary and labial palpi entirely yellowish-brown, last segment often darkened; antennomeres 1-4 or 1-5 reddish-orange with apices often darkened, 6-11 dark brown, contrasting with previous; legs bicolored: forecoxa yellow with basal fifth (males) or nearly one half dark brown (females), femur yellow and darkened apically, tibia entirely dark brown (very apex sometimes slightly paler), tarsus light brown.
Head distinctly transverse, dorsal surface with moderately dense but clearly separated and asetose punctures, frons distinctly more densely and coarsely punctate.
Pronotum distinctly transverse, posterior puncture in dorsal row occasionally doubled. Elytra weakly to distinctly transverse, shorter than pronotum at middle.
Abdomen with disc of tergites III-V distinctly, VI narrowly impunctate at middle; sternites III-V with basal line projected posteriad (IV-V weakly) at middle.
Median lobe with expansion at apical third in lateral view and narrowed to acute apex, with pair of moderately-sized lateral teeth projecting ventrad (Fig.
Female tergite X triangular with small rounded apex, disc not characteristically flattened.
Figure
The identity of
Longtanghe, Tengchong County, Yunnan, China.
The holotype was not examined of this recently described species but the illustrations in the description (
Among the members of the Electus Group: head punctures clearly separated (Fig.
Figure
This species was collected at a lower elevation (ca. 2080 m) than its closest relative,
In their description of
Phou Pan, Hua Phan, Laos.
China:
Laos:
Among the members of the Electus Group: head punctures clearly separated (Fig.
Measurements ♂ (n = 3):
Measurements ♀ (n = 1):
Extremely similar to
Figure
The species epithet refers to the previous confusion of this species with
This is the taxon illustrated by
Datang Village, Baoshan City, Tengchong County, Yunnan, China.
MYANMAR:
Among the members of the Electus Group: head punctures clearly separated (Fig.
Measurements ♂ (n = 1):
Extremely similar to
Figure
The holotype was collected at about 1600m in June.
Jianjin Shan, Tianquan County, Sichuan, China.
Among the members of the Electus Group: head punctures clearly separated (Fig.
Measurements ♂ (n = 2):
Measurements ♀ (n = 1):
Extremely similar to
Figure
Collected in June-August at elevations ranging from 1730-2100 m.
Shennongjia Nature Reserve, Pingqian, Hubei, China.
The type series of this recently well illustrated (
Among the members of the Electus Group: many head punctures confluent (Fig.
Figure
The holotype was collected in September but nothing more is known.
Lianhuaping, Leigong Shan, Guizhou, China.
Among the members of the Electus Group: many head punctures confluent (Fig.
Measurements ♂ (n = 1):
Measurements ♀ (n = 1):
Similar to
Figure
The type series was collected in September at an elevation of 1450–1500 m.
Wolong National Nature Reserve, Sichuan, China.
The type series of this recently well-illustrated (
Among the members of the Electus Group: many head punctures confluent (Fig.
Figure
A recently collected specimen was found in August at an unknown elevation.
Bashan, Dabashan, Chengkou Xian, Chongqing, China.
The type series of this recently well illustrated (
Among the members of the Electus Group: many head punctures confluent (Fig.
Measurements ♂ (n = 5):
Measurements ♀ (n = 5):
Similar to
Figure
Specimens have been collected at elevations ranging from 1200-1900 m during April-June and once in August.
The Shaanxi locality reported here is slightly north of the northernmost record of the genus (also for this species) (
One female similar to
The Caesareus Group (
Mt. Matang, Sarawak, Borneo, Malaysia.
Among the members of the Caesareus Group, the yellow-ringed black spot near the humerus of each elytron is unique to
Measurements ♂ (n = 5):
Measurements ♀ (n = 5):
Coloration: body yellowish-red; head black except middle third of frons; elytra with distinct black spot margined with yellow, spot about one third the length of elytra; abdominal tergites III (entirely), IV (basally and medioapically), VI (entirely except very base), VII (entirely except for pale apical one fifth) and VIII (entirely) black; antennomere 1 yellowish except for occasionally darkened apex, 2 reddish with dark apical half, 3-7 dark brown, apical four or apical three (most common) pale yellow to almost white, apical segment asymmetrically dark on one specimen, a few specimens seen with an antennomere half dark and half pale yellow; palpi entirely yellow to dark orange, apices sometimes darkened.
Head distinctly transverse, dorsal surface with moderately dense but clearly separated, asetose punctures, frons nearly impunctate. Antennomeres 6-10 transverse and asymmetrical.
Pronotum variable in shape but always distinctly transverse, convex, pronotal margin greatly expanded to variable degree. Elytra weakly to moderately transverse depending on the degree of lateral dilation, slightly shorter than pronotum at middle, in addition to usual macrosetal rows on disc, scattered punctures bearing setae, nearly all punctures setose on epipleuron of elytron; elytral disc bearing yellow margin of humeral spot raised and impunctate.
Abdomen with disc of tergites III-V distinctly, VI narrowly or not impunctate at middle; sternites III-IV with basal line distinctly, V slightly projected posteriad at middle.
Median lobe in lateral view gradually narrowed toward distinctly to slightly hooked apex, slightly constricted at apical third, without median tooth (Fig.
Female tergite X elongate triangular, constricted at about midlength, with elongate raised discal area that is strongly impressed longitudinally.
Figure
No consistent differences could be found between specimens from mainland Asia and Borneo.
Martapura, South Kalimantan, Borneo, Indonesia.
Borneo, Doherty, 1891 [printed] / C.
Among the members of the Caesareus Group: pronotum not entirely pale; elytra without dark spot; first two visible abdominal segments with distinct darkened area (Fig.
Measurements ♂ (n = 5):
♀ (n = 5).
Similar to
Figure
Like the often co-collected
Although externally indistinguishable from
Among the members of the Caesareus Group: pronotum not entirely pale; elytra without dark spot (Fig.
Measurements ♂ (n = 2):
Measurements ♀ (n = 1):
Almost identical to
Figure
The species epithet recognizes Ulu Temburong National Park in Brunei and its conservation achievements.
Baturaden (“Batoerraden”), Mt. Slamet, Central Java.
In his description of
Among the members of the Caesareus Group: pronotum entirely pale; elytra without dark spot; first two visible abdominal segments without distinct darkened area (Fig.
Measurements ♂ (n = 3):
Measurements ♀ (n = 1):
Similar to
Figure
No additional data accompanies the specimens.
Several female specimens examined are similar to
The true diversity of the Caesareus Group appears to be very poorly known based on the number of female specimens that likely represent additional species. Three females from Borneo, consisting of 2–3 species (
The Carnifex Group (
‘Cambodia’
This specimen in the
Within the Carnifex Group: elytral disc entirely reddish (Fig.
Measurements ♂ (n = 2):
Coloration: head entirely dark; pronotum reddish with median dark, irregular spot; elytra and scutellum reddish, disc with slightly raised yellow v-shaped marking; abdominal tergites III-V reddish with central dark marking slightly more than one-third to two-thirds the tergal width, VI dark, VII-VIII dark with paler base and apex; antennomere 1 yellow, 2-5 reddish, 6-10 dark brown, 11 yellow-brown; palpi brownish orange, apical segment darkened; legs yellowish brown, dorsal surface but not lateral face of mid-femur with darker brown (Fig.
Head distinctly transverse, dorsal surface with moderately dense, clearly separated asetose punctures, frons with only scattered punctures and deep Y-shaped impression. Antennomeres 8-10 slightly transverse and asymmetrical.
Pronotum distinctly transverse, about as wide as head, convex and with very few shallow micropunctures scattered on disc, becoming more distinct on anterior angles. Elytra slightly transverse, suture distinctly shorter than pronotum at middle.
Abdomen with disc of tergites III-V distinctly impunctate; sternites III-IV with basal line distinctly, V weakly projected posteriad at middle.
Median lobe in lateral view strongly constricted in apical fourth, apical fourth deflexed ventrad, apex knob-like without distinct median tooth, with weakly formed pair of subapical teeth (Fig.
Female unknown.
Figure
One specimen has been collected in July.
Ban Thongvay, Muang Paxong, Bolaven Plateau, Champasak, Laos.
Within the Carnifex Group: elytral disc entirely reddish (Fig.
Measurements ♂ (n = 2):
Extremely similar to
Figure
The holotype was collected by pyrethrum fogging of fungusy logs in a disturbed primary rainforest (selective logging) at 1000 m during June. The paratype was collected in a malaise trap during October.
This species is named after Jan Pedersen (
Bao Lac (town, likely in vicinity of), Cao Bang, Vietnam..
Within the Carnifex Group: elytral disc entirely reddish (Fig.
Measurements ♂ (n = 1):
Measurements ♀ (n = 1):
Extremely similar to
Figure
One specimen was collected in an FIT in a rainforest during May, at a relatively low (but unspecified elevation).
Manfei, Nabanhe Nature Reserve, Xishuangbanna, Yunnan, China.
Within the Carnifex Group: elytral disc entirely reddish (Fig.
Measurements ♂ (n = 4):
Measurements ♀ (n = 1):
Extremely similar to
Figure
Specimens have been collected in January, April and May at elevations ranging from 810-1500 m.
Mon Lake, Phu Khieo Wildlife Sanctuary, Chaiyaphum, Thailand
Within the Carnifex Group: elytral disc entirely reddish (Fig.
Measurements ♂ (n = 1):
Extremely similar to
Figure
The holotype was collected in a pitfall trap in January at an elevation of 1000 m.
In recognition of Phu Khieo Wildlife Sanctuary, which encompasses the type locality: a remarkable bowl-like plateau raised out of the surrounding lowland landscape to approximately 800-1000 m. The sustainable conservation of Phu Khieo is the subject of a collaborative project between Thailand and the European Union.
Fifth District, Ledong County, Hainan, China
The type series of this recently described, and well-illustrated species was not examined (
Within the Carnifex Group: elytral disc entirely reddish (Fig.
Figure
This species has been collected at elevations ranging from 525-978 m, in November and December. One specimen was collected by ‘beating the shrubs’ (
Nokrek Biosphere Reserve, Garo Hills, Meghalaya, India.
Within the Carnifex Group,
Measurements ♂ (n = 1):
Measurements ♀ (n = 3):
Extremely similar to
Figure
This species is named in recognition of the Nokrek UNESCO Biosphere Reserve in the Western Garo Hills of Meghalaya, India, where all known specimens were collected.
Two females, (Khasi Hills Meghalaya (
The members of the Lasti Group share the following character states: a deeply bilobed paramere, unique among Oriental
Cinchona [=Cinchona Rd. between Makkanduru and Hwy 88?], Karnataka, India.
This species can be distinguished by the following character states: head entirely dark; elytral disc dark with pale markings not extending onto epipleuron (Fig.
Measurements ♂ (n = 3):
Measurements ♀ (n = 3):
Coloration: head, pronotum and abdomen entirely dark; elytra with pale yellow, raised markings, inner marking oval shaped and larger than outer circular marking; antennomeres 1-3 dark brown, 4-5 reddish, 6-10 dark brown, 11 distinctly paler, light brown to yellowish; palpi dark brown, apical segment paler; legs yellow, forecoxae dark brown, femur with dark band in apical half, tibia entirely dark with ventral face sometimes paler.
Head distinctly transverse; dorsal surface with moderately dense, clearly separated asetose punctures, frons with only scattered punctures and poorly impressed. Antennomere 6 slightly, 7-10 distinctly transverse and more or less symmetrical.
Pronotum distinctly (males) to slightly transverse, about as wide as head, with shallow micropunctures scattered on disc, becoming more distinct on anterior angles. Elytra slightly transverse, suture slightly to distinctly shorter than pronotum at middle.
Abdomen with disc of tergites III-VI distinctly and broadly impunctate; sternites III-IV with basal line distinctly projected posteriad at middle.
Median lobe in lateral view with apical portion projected ventrad, apex deflexed dorsad (Fig.
Female with tergite VIII entire (female paratype) to emarginate, emargination minute to small; tergite X elongate shield-shaped, with broadly rounded apex, disc slightly raised and without depressions or strong ridges.
Figure
10 km SW Munnar, Idukki District, Kerala, India.
Measurements ♂ (n = 1):
♀ (n = 1).
Coloration: body orange to reddish-orange (darkened paratype); head orange with central darkened area larger in female; disc of pronotum orange with central darkened area, a pair of dark spots laterally, a darkened areas along apex and base; elytra orange to reddish-orange (darkened paratype), with darkened area around scutellum; abdominal tergite III mostly dark with lateroapical areas orange, IV-V orange, dark basally and in narrow median stripe, VI entirely dark on disc, VII-VIII orange; antennomeres 1-5 brownish orange, 6-10 dark brown, 11 slightly paler than previous, lighter brown;
palpi brownish orange; legs brownish orange, with dorsal and lateral surfaces of mid and hind femur darkened, outer faces of tibia darker.
Head distinctly transverse; dorsal surface with moderately dense, clearly separated asetose punctures, frons with only scattered punctures. Antennomeres 8-10 transverse and asymmetrical.
Pronotum distinctly transverse, about as wide as head, convex and with shallow micropunctures scattered on disc, becoming more distinct on anterior angles. Elytra slightly transverse, suture slightly shorter than pronotum at middle; disc of elytron with slightly raised yellow v-shaped marking.
Abdomen with disc of tergites III-V distinctly impunctate; sternites III-IV with basal line distinctly projected posteriad at middle.
Median lobe in lateral view with apical portion projected ventrad, apex deflexed dorsad, with pair of basal teeth at level of apical fourth (Fig.
Female with tergite VIII entire in single specimen studied, tergite X elongate triangular, with acute apex, raised disc of similar shape with broad median depression.
Figure
This taxon shares its specific epithet with the Bengal Tiger (
The members of the Luteus Group share the following character states: head and pronotum mostly orange to reddish-orange; disc of elytra with yellow, raised marking elongate and transverse (Fig.
Banks of Khwae Noi River, near Sai Yok, Kanchanaburi, Thailand.
This species is easily distinguished by the combination of the orange head and pronotum, transverse and raised yellow marking on the elytra (Fig.
Measurements ♂ (n = 1):
Coloration: body orange, head with small medial darkened spot, pronotum with single median and pair of lateral dark spots, elytra each with large lateral and triangular marking that extends onto epipleuron only apically, elytra each with transverse and raised yellow marking (Fig.
Head distinctly transverse; dorsal surface with moderately dense, clearly separated asetose punctures, frons with only scattered punctures. Antennomeres 6-7 slightly, 8-10 distinctly transverse, 10 slightly asymmetrical.
Pronotum distinctly transverse, about as wide as head, with very few shallow micropunctures scattered on disc, becoming more distinct on anterior angles. Elytra slightly transverse, suture slightly shorter than pronotum at middle; disc of elytron with slightly raised yellow transverse marking.
Abdomen with disc of tergites III-IV distinctly, V narrowly impunctate.
Median lobe in lateral view with apical portion sinuate due to expansion, projected ventrally, with large median tooth arising from median carina (Fig.
Female unknown.
Figure
The specific epithet refers to the warm, reddish-orange body coloration.
The holotype was collected in March along the banks of a large river with several other species of
Sepilok Nature Resort, Sabah, Borneo, Indonesia.
This species is easily distinguished by the combination of the orange head and pronotum, transverse and raised yellow marking on the elytra (as in Figs
Measurements ♂ (n = 1):
Similar to
in lateral view, with pair of broad basal teeth (Fig.
Figure
In recognition of the Kabili-Sepilok Forest Preserve, which adjoins the type locality and preserves a significant remnant of virgin lowland Dipterocarp forest in northern Borneo.
The members of the Pictus Group (
Puwen, Xishuangbanna, Yunnan, China.
The holotype of this species is a female and this initially caused doubt about the identity of
This species may be recognized by the following: head entirely dark; pronotum widest in posterior third; elytra partly dark, discal markings of elytra extending onto epipleuron, not forming v-shape (Fig.
Measurements ♂ (n = 5):
Measurements ♀ (n = 2):
Coloration: color dimorphic, dark morph with dark pronotum and dark abdomen; bicolored morph with red pronotum and abdomen with tergites III-V entirely and base of VI reddish. Head entirely dark; elytra with raised median marking oval-shaped and larger than raised lateral marking, markings always connected, pale area extended onto epipleuron, epipleuron varying from pale at midlength to nearly entirely pale; scutellum dark; antennomere 1 yellow with darkened apex, 2-5 reddish, 6-10 dark brown, 11 either dark brown or vaguely paler; palpi yellow with apical segment slightly darker; legs yellow, fore and midfemur with small ventroapical dark marking, hind femur with small apical band.
Head distinctly transverse, dorsal surface with dense, clearly separated asetose punctures, frons with very few scattered punctures and deep, coarse Y-shaped impression. Antennomeres 6-7 quadrate to slightly transverse, 8-10 distinctly transverse and slightly asymmetrical, 10 longer than 9.
Pronotum distinctly transverse, about as wide as head, with a few shallow micropunctures on disc, becoming more distinct on anterior angles. Elytra slightly transverse, suture slightly shorter than pronotum at middle.
Abdomen with disc of tergites III-VI distinctly impunctate medially, some specimens with VI only narrowly impunctate; sternites III-IV with basal line distinctly projected posteriad at middle.
Median lobe in lateral view sinuate, apex pointed and sharply deflexed ventrad, basal teeth appearing at ventral face (Fig.
Female tergite VIII with median emargination deep and elongate; female tergite X shield-shaped, disc with distinct and entirely rounded dorsal expansion in apical half, apex truncate with narrow and rounded median projection.
Figure
This species has been collected in March, May and July, at elevations ranging from 600-1500 m.
Mintaingbin Forest Camp, ca. 35 km north of Aungban, Shan State, Myanmar.
This species may be recognized by the following: head entirely dark; pronotum widest in posterior third; elytra partly dark, discal markings of elytra extending onto epipleuron, not forming v-shape, medial and lateral markings of different sizes and never entirely fused (Fig.
Measurements ♂ (n = 5):
Measurements ♀ (n = 2):
Extremely similar to
Figure
This species has been collected in March, May and June at 900–1320 m. Specimens have been collected from rotten wood and from within fungi.
The species is named in honor of Harald (Harry) Schillhammer (
Banks of Khwae Noi River, near Sai Yok, Kanchanaburi, Thailand.
This species may be recognized by the following: head entirely dark; pronotum widest in posterior third; elytra partly dark, discal markings of elytra extending onto epipleuron, not forming v-shape, medial and lateral markings of different sizes and never entirely fused (Fig.
Measurements ♂ (n = 2):
Extremely similar to
Figure
The type series was collected in March along the banks of a large river, with many other beetles including
This species is named in honor of Mr. Guillaume de Rougemont (United Kingdom), the sole collector of this and many other Oriental
At present,
Menglun Nature Reserve, Xishuangbanna, Mengla County, Yunnan, China.
The type specimen of this recently described, and well-illustrated species was not examined (
This species may be recognized by the following: head entirely dark; pronotum widest in posterior third; elytra partly dark, discal markings of elytra extending onto epipleuron, not forming v-shape, medial and lateral markings of different sizes and never entirely fused (Fig.
Measurements ♂ (n = 3):
Measurements ♀ (n = 1):
Extremely similar to
Figure
The acute apex of the median lobe was observed to be longer in specimens south of the type locality in Yunnan, China. This is attributed to intraspecific variation.
Nokrek National Park, West Garo Hills, Meghalaya, India.
This species may be recognized by the following: head entirely dark; pronotum widest in posterior third; elytra partly dark, discal markings of elytra extending onto epipleuron, not forming v-shape, medial and lateral markings of different sizes and never entirely fused (Fig.
Measurements ♂ (n = 1):
Extremely similar to
Figure
The holotype was collected at approximately 1100 m in May.
The species epithet refers to the apex of the median lobe, which is uniquely thin and concave in lateral view.
In species of the Vulneratus Group (
Cochinchina [=Southeast and Mekong River Delta regions], Vietnam.
This species may be recognized by the following: head entirely dark; pronotum widest in posterior third; elytra partly dark, discal markings of elytra extending onto epipleuron, not forming v-shape, medial and lateral markings not entirely fused (Fig.
Measurements ♂ (n = 1):
Measurements ♀ (n = 1):
Coloration: head, pronotum and abdomen entirely dark; elytra with raised median marking oval-shaped and larger than raised lateral marking, markings connected, pale area extended onto epipleuron, epipleuron pale, pale area varying from near humerus to basal half; scutellum dark; antennomere 1 yellow with darkened apex, 2-5 reddish, 6-10 dark brown, 11 either dark brown or vaguely paler; palpi yellow with apical segment slightly darker; legs yellow, hind femur with small ventroapical dark marking.
Head strongly transverse, dorsal surface with dense, clearly separated asetose punctures, frons with few scattered punctures and deep, coarse Y-shaped impression. Antennomeres 6-10 distinctly transverse and 8-10 slightly asymmetrical.
Pronotum distinctly transverse, slightly wider than head, center of disc almost entirely without micropunctures, micropunctures becoming more distinct on anterior angles. Elytra slightly transverse, suture distinctly shorter than pronotum at middle.
Abdomen with disc of tergites III-VI distinctly impunctate medially, VI slightly more narrowly than others.
Median lobe in lateral view projected ventrad, with hooked apex, without basal teeth (Fig.
Female tergite VIII with median emargination broad triangular, moderately deep; female tergite X elongate shield-shaped, with the disc evenly convex and lacking distinct raised areas, apex acuminate with apex projected.
Figure
One specimen was collected in March at 100 m.
Manfei, Nabanhe Reserve, Xishuangbanna, Yunnan, China.
This species may be recognized by the following: head entirely dark; pronotum widest in posterior third; elytra partly dark, base of elytra with broad pale area composed entirely of pale, raised yellow marking extending onto epipleuron (Fig.
Measurements ♂ (n = 5):
Measurements ♀ (n = 5):
Similar to
Figure
As far as known,
Near Lushan, Nantou County, Taiwan.
This species may be recognized by the following: head entirely dark; pronotum widest in posterior third; elytra partly dark, discal markings of elytra extending onto epipleuron, not forming v-shape, medial and lateral markings of different sizes and never entirely fused (Fig.
Measurements ♂ (n = 5):
Measurements ♀ (n = 5):
Similar to
Figure
Specimens have been collected in January, March-May and July, at elevations ranging from 400-2000 m. Some specimens have been sifted from litter.
Nanling National Forest Park, Ruyuan County, Guangdong, China.
The type series of this recently described, and well-illustrated species was not examined (
This species may be recognized by the following: head entirely dark; pronotum widest in posterior third; elytra partly dark, discal markings of elytra extending onto epipleuron, not forming v-shape, medial and lateral markings of different sizes and never entirely fused (Fig.
Figure
The holotype was collected in July.
Although
Mt. Phu Phan, Hua Phan, Laos.
This species may be recognized by the following: head entirely dark; pronotum widest in posterior third; elytra partly dark, discal markings of elytra extending onto epipleuron, not forming v-shape, medial and lateral markings of different sizes and never entirely fused (Fig.
Measurements ♂ (n = 3):
Similar to
Figure
Specimens were collected at an elevational range of 1500-1900 m in May and June.
The specific epithet refers to the swollen, preapical portion of the median lobe.
This species is most similar to
Taitung, Taitung County, Taiwan.
Taiwan:
This species may be recognized by the following: head entirely dark; pronotum widest in posterior third; elytra partly dark, with broadly pale basal area composed of yellow raised marking and slightly darker non-raised area (Fig.
Measurements ♀ (n = 1):
Similar to
Figure
This species has been observed in April-July and October at an approximate elevation of 1200 m. One pair was found
The photo records of this species by D. Huang and L. Wei-Ren represent the only detailed data available as the type series is only reported from the general area of Taitung city.
Kuatun [=Gua Dun], Fujian, China.
This species may be recognized by the following: head entirely dark; pronotum widest in posterior third; elytra partly dark, with broadly pale basal area composed of yellow raised marking and slightly darker non-raised area; antennomere 7 distinctly transverse; female tergite X with disc raised medioapically.
Measurements ♀ (n = 5):
Similar to
Figure
Specimens have been collected in April-July at elevations ranging from 900–1100 m.
There have been concerns that this species was conspecific with
The diverse members of the Loculus Group (
Menglun Nature Reserve, Xishuangbanna, Yunnan, China.
The type series of this recently described, and well-illustrated species was not examined (
This species may be recognized by the following: head entirely dark; pronotum widest in posterior third; elytra partly dark, discal markings of elytra extending onto epipleuron, not forming v-shape, forming broadly pale basal area composed entirely of yellow raised marking (similar to Fig.
Figure
The type series was collected in February at elevations ranging from 560-860 m.
Mt. Yingge Ling [base of], Hainan, China.
The type series of this recently described, and well-illustrated species was not examined (
This species may be recognized by the following: head entirely dark; pronotum widest in posterior third; elytra partly dark, discal markings of elytra extending onto epipleuron, not forming v-shape, medial and lateral markings of different sizes and never entirely fused (Fig.
Figure
Specimens have been collected in November and December at elevations ranging from 450-666 m.
Except for one, much larger, member of the Carnifex Group,
Ban Thongvay, Muang Paxon, Bolaven Plateau, Champasak, Laos.
This species may be recognized by the following: head entirely dark; pronotum widest in posterior third; elytra partly dark, discal markings of elytra extending onto epipleuron, forming v-shape (Fig.
Measurements ♂ (n = 1):
Measurements ♀ (n = 3):
Coloration: head entirely dark; pronotum entirely reddish; scutellum reddish, at least at base, disc of elytra dark with yellow v-shaped, raised marking, base sometimes broadly reddish; epipleuron not entirely pale, with subapical darkened area; abdominal tergites III-V entirely reddish, VI varying from dark with reddish base to entirely reddish; antennomere 1 yellow with darkened apex, 2-5 light brownish-red with darkened apex, 6-10 dark brown, apical segment brownish to dark brown; palpi yellow-brown with apical segment slightly darker; legs yellow, midfemur with ventroapical darkened area, hind femur with apical area of femur darkened.
Head distinctly transverse, dorsal surface with moderately dense, clearly separated asetose punctures, frons with few scattered punctures and distinct Y-shaped impression. Antennomeres 6 quadrate, 7 slightly and 8-10 distinctly transverse, 10 more elongate than 9, 8-10 slightly asymmetrical.
Pronotum slightly transverse to quadrate, about as wide as head, with sparse moderately impressed micropunctation on disc, becoming more distinct on anterior angles. Elytra distinctly transverse, suture distinctly shorter than pronotum at middle.
Abdomen with disc of tergites III-V distinctly, VI narrowly impunctate medially; sternites III-IV with basal line distinctly projected posteriad at middle.
Median lobe in lateral view with narrowed apical portion nearly straight, apex with tooth (Fig.
Female tergite VIII with median emargination moderately deep and varying from triangular to U-shaped; female tergite X elongate shield-shaped, with moderately projecting and rounded apex, disc distinctly raised, this area longitudinally impressed at middle.
Figure
The type series was collected during May-June at 800-1200 m elevation. Specimens were collected using flight intercept traps and from tree fungi.
This species is named in honor of Alexey Solodovnikov (
Karen Hills, Kayin State, Myanmar.
This species may be recognized by the following: head entirely dark; pronotum widest in posterior third; elytra partly dark, discal markings of elytra extending onto epipleuron, forming v-shape (as in Fig.
Measurements ♂ (n = 1):
Similar to
Figure
The holotype was collected in May at 900–1100 m.
This species is named after Leonardo Fea (1852–1903), an Italian explorer and naturalist who collected a rich variety of
Mt. Mulayit, Kayin State, Myanmar.
This species may be recognized by the following: head entirely dark; pronotum widest in posterior third; elytra partly dark, discal markings of elytra extending onto epipleuron, forming v-shape (as Figs
Measurements ♂ (n = 1):
Measurements ♀ (n = 1):
Similar to
Figure
The type series was collected in March at a relatively high elevation (1800–1900 m).
The species is named after Mulayit Taung, a mountain in southern Myanmar. The type series was collected near its summit.
Banks of Khwae Noi River, near Sai Yok, Kanchanaburi, Thailand.
This species can be recognized by the following combination of characters: orange frons contrasting with dark head (Fig.
Measurements ♂ (n = 2):
Coloration: head dark with orange frons; pronotum reddish orange with hourglass-shaped dark marking medially; elytral disc dark brown to dark brownish-red, with orange to yellow marking at middle and along suture, apical angles and humeri paler; scutellum dark; epipleuron pale along its length; abdominal tergites III-VI ranging from almost entirely dark, to reddish with central dark marking, VII entirely dark; antennomere 1 yellow, 2-5 reddish with dark apices, 6-10 dark brown, 11 vaguely paler; palpi yellowish; legs yellowish with small dark marking at ventral apex of femora.
Head distinctly transverse, dorsal surface with moderately dense but clearly separated, asetose punctures, frons with few scattered punctures. Antennomeres 8-10 slightly transverse and asymmetrical.
Pronotum distinctly transverse, center of disc with very few micropunctures, about as wide as head. Elytra moderately transverse, suture shorter than pronotum at middle; elytral disc bearing a pair of small, raised, oval and light yellow markings near the center, and an additional, smaller pair laterad.
Abdomen with disc of tergites III-V distinctly, impunctate at middle.
Median lobe in lateral view evenly narrowed to apex, ventral face flat, with minute pair of median teeth formed from median carina (Fig.
Female unknown.
Figure
Specimens were collected in March and May at both lowland and montane elevations (900–1100 m). It is unlikely that
This charming species is named in honor of Aleš Smetana (
Mawsynram, Meghalaya, Khasi Hills, India.
This species can be distinguished by the following character states: head entirely dark; elytral disc dark with pale markings not extending halfway to epipleural margin (Fig.
Measurements ♂ (n = 1):
Measurements ♀ (n = 1):
Coloration: head, pronotum and abdomen entirely dark; elytra with yellow, raised markings, medial marking thinly connected to smaller lateral marking that does not extend halfway to epipleural margin; antennomere 1 yellow with darkened apex, 2-5 reddish with darkened apices, 6-11 dark brown; palpi yellow with apical segment slightly darkened; legs yellow, mid and hind tibia with darkened ventroapical area.
Head distinctly transverse, dorsal surface with moderately dense, clearly separated asetose punctures, frons with only scattered punctures and distinctly impressed. Antennomere 6 slightly, 7-10 distinctly transverse and 8-10 asymmetrical.
Pronotum distinctly transverse, about as wide as head to slightly wider, with a few shallow micropunctures on disc, becoming more distinct on anterior angles. Elytra slightly transverse, suture shorter than pronotum at middle.
Abdomen with disc of tergites III-VI distinctly impunctate medially.
Median lobe in lateral view strongly constricted in apical portion, with apical tooth formed from median carina, basal teeth absent, ventral face more or less straight (similar to Fig.
Female with tergite VIII entire; tergite X elongate shield-shaped, with broadly rounded but projected apex, disc distinctly raised and without depressions or strong ridges.
Figure
The type series was collected in June at approximately 800 m. The area around the type locality is also considered to be the wettest terrestrial place on earth, receiving record annual rainfall levels.
The species epithet refers to the type locality in the Khasi Hills of Meghalaya, India, and recognizes this global biodiversity hotspot.
Sevoke, Darjeerling, West Bengal, India.
This species can be distinguished by the following character states: head entirely dark; elytral disc dark with pale markings extending halfway to epipleural margin (Fig.
Measurements ♂ (n = 1):
Extremely similar to
Figure
The holotype was collected in October at 200 m and was sifted from leaves in a forest ravine (
The species epithet refers to its distribution in the Himalayan mountains.
This is the species
Mawsynram-Balat Road, Khasi Hills, Meghalaya, India.
This species can be distinguished by the following character states: head entirely dark; elytral disc dark with pale markings extending halfway to epipleural margin (Fig.
Measurements ♂ (n = 1):
Extremely similar to
Figure
The holotype was collected in October at 1000 m.
The species epithet refers to the fact that this species is the smallest known in the Oriental fauna at 3.5 millimeters in forebody length.
BaBe National Park, Bac Kan Province, Vietnam.
This species can be distinguished by the following character states: head entirely dark; elytral disc dark with pale markings extending halfway to epipleural margin (Fig.
Measurements ♂ (n = 1):
Similar to
Figure
The holotype was collected in May using an FIT in lowland forest (180 m).
This species is named in honor of Stewart Peck (Carleton University, Ottawa, Canada), the collector of its holotype and many other rare taxa belonging to
A single female specimen from Kachin State, Myanmar (NHMW) was studied that is similar to
Baturaden (“Batoerraden”), Mt. Slamet, Central Java.
This species can be recognized by the following combination of characters: orange frons contrasting with dark head (Fig.
Measurements ♂ (n = 2):
Measurements ♀ (n = 4):
Coloration: head dark with orange frons; pronotum varying from entirely reddish-orange, to with median and pair of darkened areas; elytra dark with humeri, apical angles and v-shaped , raised discal marking light yellow, base and area along suture orange, these pale areas often connecting to leave only outer apical areas dark; scutellum dark to reddish with dark border; antennomere 1 yellow with darkened apex, 2-5 light brownish-red, 6-10 dark brown, apical segment distinctly paler, light brown to yellow; palpi yellow-brown; legs yellow, on ventral face, very apex of femur darkened.
Head distinctly transverse, dorsal surface with moderately dense, clearly separated asetose punctures, frons with very few scattered punctures and shallow Y-shaped impression. Antennomeres 6 slightly, 7-10 distinctly transverse, 10 more elongate than 9, 7-10 slightly asymmetrical.
Pronotum slightly transverse to quadrate, about as wide as head, with scattered shallow micropunctures on disc, becoming more distinct on anterior angles. Elytra slightly transverse, suture distinctly shorter than pronotum at middle, punctation relatively sparse, with most punctures separated by a distance not less than their diameter.
Abdomen with disc of tergites III-VI distinctly impunctate medially, basal impressions with elongate punctures; sternites III-IV with basal line distinctly projected posteriad at middle.
Median lobe in lateral view with apical fourth more strongly narrowed to apex, which is deflexed ventrad (Fig.
Female tergite VIII with median emargination varying from small and triangular to moderately deep and elongate triangular; female tergite X elongate, with acute, acuminate apex; disc only weakly raised in apical half, not separated by ridges.
Figure
Specimens were collected in March and July.
Mount Kendang, West Java.
This species can be distinguished by the following character states: head entirely dark; elytral disc dark with pale markings not extending onto epipleuron (Fig.
Measurements ♂ (n = 4):
Measurements ♀ (n = 5):
Similar to
Figure
Specimens have been collected in January and June-August, at elevations ranging from 1000-1700 m.
Mt. Muria (“Gn. Moeria”), Central Java.
Measurements ♂ (n = 1):
Coloration: body dark; elytra reddish; abdominal tergite III mostly dark with lateroapical areas reddish, IV with middle dark and remaining area reddish, V entirely reddish, VI-VII dark with base and apex reddish, VIII dark; antennomere 1 yellow, II-V reddish, 6-10 dark brown, 11 pale yellow; palpi yellowish, apical segment darkened; legs yellowish, femora with dorsal surface darkened.
Head distinctly transverse; dorsal surface with moderately dense, clearly separated asetose punctures, frons with coarse depressions in addition to usual frontal impressions and with scattered punctures, frons bearing a pair of setose punctures between anterior frontal punctures. Antennomeres 7-10 transverse and asymmetrical.
Pronotum distinctly transverse, convex and with micropunctures scattered on disc, becoming rather dense on anterior angles. Elytra slightly transverse, slightly shorter than pronotum at middle, in addition to usual macrosetal rows on disc, scattered punctures bearing setae, nearly all punctures setose on epipleuron of elytron.
Abdomen with disc of tergites III-V distinctly impunctate; sternites III-IV with basal line distinctly projected posteriad at middle.
Median lobe in lateral view gradually narrowed apicad to level of single median, apical tooth formed from projection of apical carina, apex of median lobe slightly constricted apicad of tooth, median lobe with pair of basal teeth at about midlength (Fig.
Female unknown.
Figure
The holotype was collected in February at 700-1000 m.
Dikoya, Central Province, Sri Lanka.
Measurements ♂ (n = 1):
Measurements ♀ (n = 1):
Coloration: body dark; head, pronotum and abdomen entirely dark, elytra dark, each with yellow v-shaped marking; antennomere 1 yellow with darkened apex, 2-5 reddish-orange with darkened apices, 6-10 dark brown, 11 pale yellow; palpi yellowish with apical segment darkened; forecoxae yellow with basal fifth dark brown in both sexes, legs yellow, forefemur with dorsal surface dark brown, lateral face and dorsal surface of mid and hind tibia with subapical dark marking, tibia with lateral face darker.
Head strongly transverse, more so in male; dorsal surface including frons glossy with sparse, small and poorly impressed punctures; in male, lateral part of head beneath eye expanded ventrad. Antennomeres 8-10 distinctly transverse but not asymmetrical.
Pronotum transverse and widest at anterior angles, strikingly more strongly transverse in males than females, with lateral portions explanate in males, protuberance moderate (male) or distinct (female); medial part of disc almost entirely without micropunctures, anterior angles with many impressed micropunctures. Elytra slightly transverse, suture distinctly to slightly shorter than pronotum at middle, nearly all punctures of epipleuron setose.
Abdomen with disc of tergites III-VI distinctly impunctate.
Median lobe in lateral view evenly converging to apex, ventral face nearly straight except for very apex flexed ventrad, without teeth (Fig.
Female with tergite VIII bearing large, circular emargination (Fig.
Figure
Specimens were collected in September and December-January at 1160–1280 m. One specimen was collected by a malaise trap.
Two female specimens from Java are similar to
Dorsal habitus of
Dorsal habitus of
Pronotum, lateral:
Antenna:
Abdominal tergite III:
Elytra:
Female abdominal apex:
Distribution of:
Distribution of:
Distribution of:
Distribution of:
This project has received funding in the form of fellowships from the NSERC PRP program and from the European Union’s Horizon 2020 research and innovation program under the Marie Sklodowska-Curie grant agreement no. 642241 (BIG4). The author would like to thank A. Solodovnikov (