Corresponding author: Jiang-Li Tan (
Academic editor: N. Johnson
Twenty two species of the paper wasp genus
Tan J-L, van Achterberg K, Chen X-X (2014) Pictorial key to species of the genus
The genus
Some specimens were collected with interception traps (
For the morphological and micro-sculpture terminology used in this paper see
1 | Propodeum with pair of raised vertical carinae anteriorly (a); propodeal orifice narrow, slit-like, acute above (b) | 2 |
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– | Propodeum without pair of raised vertical carinae anteriorly (aa); propodeal orifice relatively broad, more or less rounded above (bb) | 3 |
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2 | Head narrower than mesoscutum just in front of tegulae (a); clypeus slightly higher than wide, shiny and its dorsal half impunctate (b); apical margin of second metasomal tergite armed with a row of spines (c) | |
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– | Head wider than mesoscutum just in front of tegulae (aa); clypeus distinctly wider than high, dull and its dorsal half evenly punctate (bb); apical margin of second tergite simple (cc) | |
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3 | Propodeal valvula large, obscuring most of propodeal orifice in lateral view (a); basal angle of second submarginal cell less than (b) or equal to or slightly greater than 90°(b’) | 4 |
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– | Propodeal valvula medium-sized, most of propodeal orifice visible in lateral view (aa); basal angle of second submarginal cell greater than 90°(bb) | 14 |
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4 | Length between ocellus and eye (OOL) less than twice as long as length between posterior ocelli (POL) (a); first flagellomere of female antenna comparatively short, less than 2.5× as long as its apical width (b); basal angle of second submarginal cell ≥ 90°(c) | 5 |
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– | OOL more than 2.5 × as long as POL (aa); first flagellomere of female antenna long, 3 × longer than its apical width (bb); basal angle of second submarginal cell distinctly less than 90°(cc) | 6 |
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5 | Pronotal carina nearly straight (a); propodeum distinctly obliquely striate (b); propodeal valvula large, nearly circular (c); first metasomal tergite comparatively long, with petiolus parallel-sided, widened part swollen submedially and narrowed near apical margin (d); second metasomal segment oblique apically, with tergite longer than sternite (e); male antenna with distinct tyloids, apical two thirds of apical flagellomere excavated and curved (f) | |
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– | Pronotal carina distinctly sinuate (aa); propodeum largely smooth (bb); propodeal valvula rounded triangular (cc); first tergite comparatively short, petiolus widened basally and apically, and widened part swollen up to apical margin (dd); second metasomal segment vertically cut off apically, with tergite about as long as sternite (ee); male antenna without tyloids and apical flagellomere normal (ff) | |
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6 | Propodeum reticulate-punctate, anteriorly with distinct median depression (a), yellow marks of propodeum absent (b), dorsal side of propodeum reticulate-punctate (c); first metasomal tergite less than twice as long as wide in dorsal view and less than 2.5× as long as high in lateral view (d) | 7 |
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– | Propodeum finely striate, barely depressed antero-medially or if depressed, only as fine median furrow (aa); propodeum with yellow marks confluent basally (bb); dorsal side of propodeum barely punctate, only finely transversely striate (cc); first tergite more than twice as long as wide in dorsal view and more than 2.5× as long as high in lateral view (dd) | 9 |
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7 | Second metasomal tergite as wide as long in dorsal view (a); second segment longer than high in lateral view (b); scutellum black, slightly convex and without median furrow (c) | |
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– | Second tergite wider than long in dorsal view (aa); second segment as long as high in lateral view (bb); scutellum reddish brown, distinctly convex and with a longitudinal median furrow (cc) | 8 |
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8 | Apical flagellomere of male antenna long; carina of the 8th flagellomere (VIII) emarginate (a); second metasomal segment oblique apically, with sternite longer than tergite (b). Male: proximo-ventral margin of penis with a short hook, at most one fourth as long as penis valve (c) | |
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– | Apical flagellomere of male antenna short; carina of the 8th flagellomere (VIII) not emarginate (aa); second metasomal segment vertical apically, with tergite about as long as sternite (bb); male genitalia: proximo-ventral margin of penis valve with a long hook, two thirds as long as penis valve (cc) | |
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9 | Preapical part of second tergite distinctly swollen (a); lateral profile of second sternite nearly straight anteriorly (b); mesoscutum entirely black (c) | 10 |
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– | Preapical part of second tergite flat (aa); lateral profile of second sternite evenly curved anteriorly (bb); colour of mesoscutum variable (cc) | 11 |
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10 | Female clypeus yellow with a black arrow-shaped mark (a); second tergite with yellow apical band and strongly swollen medio-dorsally in lateral view (b); male genitalia: proximal margin of penis abruptly contracted in dorsal view and with a large hook in ventral view (c) | |
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– | Female clypeus largely black and apically yellow (aa); second tergite largely black and dorsally relatively weakly swollen in lateral view (bb); male genitalia: proximal margin of penis gradually contracted in dorsal view and with a comparatively small hook in ventral view (cc) | |
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11 | Female clypeus reddish or blackish brown and yellow apically (a); gena reticulate-punctate (b); metapleuron black or with reddish brown patch (c); male genitalia: proximo-dorsal margin of penis gradually contracted (d) | 12 |
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– | Female clypeus yellow laterally (aa); gena finely punctate (bb); metapleuron generally with large yellow patch (cc); male genitalia: proximo-dorsal margin of penis abruptly contracted (dd) | 13 |
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12 | Mesoscutum with two separate reddish brown patches (a); tergite and sternite of second metasomal segment fused and suture indistinct except posteriorly (b); male genitalia: proximo-ventral margin of penis valve hardly projecting (c) | |
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– | Mesoscutum entirely black (aa); tergite and sternite of second metasomal segment separated and suture complete (bb); male genitalia: proximo-ventral margin of penis valve with a distinct projection (cc) | |
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13 | Metapleuron ventrally punctate (a); female clypeus reddish brown with two lateral yellow spots (b); first metasomal segment reddish brown baso-laterally, without yellow pattern basally (c); second sternite usually without yellow spots (d); male antenna comparatively robust and serrate (e); male genitalia: proximo-ventral margin of penis valve with a relatively sharp hook-like projection (f) | |
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– | Metapleuron ventrally smooth (aa); female clypeus yellow with a dark arrow-shaped mark medially (bb); first segment partly yellow baso-laterally (cc); second sternite usually with a pair of large yellow spots (dd); male antenna comparatively slender and hardly serrate (ee); proximo-ventral margin of penis valve evenly curved, without a hook-like projection (ff) | |
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14 | Fore wing with two submarginal cells (a); OOL at most twice as long as POL or shorter (b) | |
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– | Fore wing with three submarginal cells (aa); OOL at least 3 × as long as POL or longer (bb) | 15 |
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15 | Length of body 10–11 mm (measured from head to end of second metasomal tergite); pterostigma blackish brown and marginal cell entirely dark brown (a); mesoscutum and second metasomal tergite entirely black (b); mesopleuron (except antero-ventrally) coarsely punctate (c) | 16 |
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– | Length of body 7–8 mm (measured from head to end of second metasomal tergite); pterostigma brown and ventral half of marginal cell subhyaline (aa, in |
18 |
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16 | Female clypeus coarsely and densely punctate (a); propodeum dull and distinctly punctate-striate (b); first metasomal segment black with pair of apical spots (c); second metasomal segment somewhat oblique with sternite shorter than tergite (d) | 17 |
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– | Female clypeus finely punctate (aa); propodeum shiny and smooth, at most very finely striate (bb); first metasomal segment entirely orange (cc); second segment oblique with sternite distinctly larger than tergite (dd) | |
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17 | Ventral half of occipital carina strongly widened, its maximum width nearly one-third genal width (a); pronotum with two small yellow spots medio-dorsally (b); scutellum and metanotum black with yellow pattern (c); propodeum with pair of large yellow patches (d); subapical spots of first tergite yellow and large (e) | |
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– | Occipital carina hardly or not widened (aa); pronotum reddish brown medio-dorsally (bb); scutellum and metanotum reddish brown (cc); propodeum black (dd); apical spots of first tergite reddish brown and large (ee) | |
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18 | Head reddish brown and with yellow patches near its occipital carina, and temple distinctly shorter than eye in dorsal view (a); pterostigma pale brown and basal half of marginal cell subhyaline (b); first metasomal tergite strongly widened in lateral view, its length less than 1.5× its height (c); male: flagellum more or less serrate ventrally and terminal flagellomere weakly curved (d) | |
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– | Head black with yellow spots and temple as long as eye in dorsal view (aa); pterostigma dark brown and posterior half of marginal cell subhyaline (bb); first metasomal tergite weakly widened in lateral view, its length more than 2.2× its height (cc); male: flagellum slightly serrate and terminal flagellomere bullet-shaped (dd) | 19 |
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19 | Vertex, frons, mesoscutum and scutellum shiny, impunctate (a); mesopleuron largely yellow and without distinct coarse punctures (b); second metasomal sternum yellow on basal two-thirds (c) | |
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– | Vertex, frons, mesoscutum and scutellum dull, finely punctulate with rather large, shallow, flat-bottomed punctures (aa); mesopleuron black with less extensive yellow pattern (bb, bb’) and with distinct coarse punctures (bb, bb’); second metasomal sternite black with small yellow spots (cc), but more extensively yellow in |
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20 | Occipital carina sinuate near middle level of eye (a); clypeus yellow with an isolated black medial spot (b); apical part of first metasomal tergite comparatively wide (beyond spiracle slightly wider than long) and spiracle invisible in dorsal view (c); basal slender part of first metasomal tergite rather short in lateral and ventral view, rising directly from posterior end of reception of propodeal suspensory ligament (d) | |
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- | Occipital carina evenly curved near middle level of eye (aa); clypeus entirely yellow (bb) or a large black spot (bb’); apical part of first metasomal tergite comparatively slender (beyond spiracle longer than wide or quadrate) and spiracle visible in dorsal view (cc); basal slender part of first metasomal tergite longer in lateral and ventral view, rising further away from propodeal suspensory ligament (dd) | 21 |
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21 | Gena yellow, except upper half of occipital carina black (a); clypeus entirely yellow (b); mesopleuron with extensive yellow pattern (c) as second sternite (d) | |
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- | Gena mainly black anteriorly and posteriorly, but medially yellow (aa); clypeus with a large triangular black medial spot (bb); mesopleuron with restricted yellow pattern (cc); second sternite black except a narrow yellow band apically (dd) | |
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CHINA: 3 ♀ (NWUM), Yaoqu, Mengla, Yunnan, 28–30.ix.2010, JL Tan; 1 ♀ (NWUM), Longmen, Shangyong, Yunnan, 26.ix.2010, JL Tan; 1 ♀ (ZJUH), Jinghong Forestry Park, Yunnan, 31.vii.2003, J Lu (no. 20045916); 5 ♀ (ZJUH), Fugong, Yunnan, 24.viii.2003, Q Li (no. 20046662, -4, -6, -8, -70); 1 ♀ (ZJUH), Mt. Youle, Yunnan,17.iv.1981, JH He (no. 811876); 1 ♀ (ZJUH), Shuangjiang, Yunnan, 21.iv.1981, coll. JH He (no. 813609); 2 ♀ + 8 ♂ (ZJUH), Tongzhong, Shiwandashan, Guangxi, 3.xii.2001, ZF Xu (no. 20029971–4, -77–79, -80–82). INDONESIA: 2 ♀ (RMNH), W. Java: Djampang Tengah, 1634, M Walsh; 1 ♂ (RMNH), Udjung Kulon, Tjidaon, Java, 29.xi.1951, AMR Wegner. VIETNAM: 1 ♀ (RMNH), S. Vietnam: Dak Lak, Chu Yang Sin N.P., 50 km S. Buon Ma Thuot, Krong K’Mar, 550–600 m, 23–25.x.2005, near rainforest, C van Achterberg & R de Vries; 1 ♀ (RMNH), id., but 590–840 m, 22–26.x.2005, Malaise traps 13–23; 1 ♂ (RMNH), N. Vietnam: Viet Try, Thuong Cuu, near Thanh Son,
Some specimens have the apical margin of the second metasomal tergite flattened, not convex as in typical specimens. In the species
China (Yunnan, Guangxi); Borneo; Java; Myanmar; Malay Peninsula; Sumatra; Vietnam (
1 ♀ (RMNH), syntype of
Among the specimens from China, there are two colour forms,
China (Yunnan); Borneo; India; Malaysia; Myanmar; Thailand; Vietnam. (
1 ♀, Luang Prabang, Hat Thoun, Siam, 10. xi.1917, RV De Salvaza (holotype, BMNH) 1♂, Mergui, Tenasserim, May 1890, CT Bingham (allotype, BMNH. “This specimen bears a label in Bingham’s handwriting: “
No specimens from China were available, but this species is included in the key because it has been listed for China by
China (Guangdong); Myanmar; Thailand. (
MYANMAR (specimens collected by R. Malaise are paratypes of
China (Guizhou, Fujian, Zhejiang), Myanmar. (
INDONESIA: 1 ♀ (RMNH), Java, Malang, 11.iv.1930, JG Betrem. CHINA: 1 ♀ (NWAM), Xishuangbanna, Mengla, Yunnan, 20.iv.1982, JR Zhou & SM Wang.
China (Yunnan); Bali; India; Java; Lombok; Nepal; Malay Peninsula; Myanmar; Philippine Islands; Sumba, Sulawesi, Sri Lanka; Thailand; Vietnam. (
1 ♂ (ZMUC), with three handwritten labels “Lectotype, J.v.d.Vecht, 1957”, “
China (Fujian, Guangdong, Hainan, Hong Kong, Taiwan, Yunnan, Zhejiang); Bali; Bangka; Borneo; Flores; India; Java; Kariman Djava I.; Myanmar; Nepal; Nias; Malay Peninsula; Sumatra; Timor; Palawan; Ryukyu Islands; Philippines (Palawan); Sri Lanka; Thailand; Vietnam. (
MALAYSIA: 1 ♂ (RMNH), S. Sabah, near Long Pa Sia, c 1010 m, 25.xi.–8.xii.1987, Mal. trap, C van Achterberg; 1 ♂ (RMNH), S. Sabah, Beaufort,
No specimens from China were available in the present study, but this species was reported from China by
China (Hong Kong); Borneo; Malay Peninsula; Sumatra; Vietnam. (
CHINA: 1 ♂ (BMNH), Hummocks, Hainan, 24.v.1936, G. Ros, (allotype of
China (Guangdong, Hainan, Hong Kong, Jiangxi, Yunnan); Bangka; India; Java; Myanmar; Vietnam. (
MYANMAR: 4 ♂ (RMNH), N. East Burma: Sadon, 1200 m, 28.vi–5.vii.1934, R. Malaise (paratypes of
The yellow stripes of the mesoscutum are lacking in some individuals.
China (Yunnan), Myanmar. (
VIETNAM: 1 ♀ (RMNH), S. Vietnam: Dak Lak, Chu Yang Sin N.P. 50 km S. Buon Ma Thuot, Krong K’Mar, 550–600 m, 23–25.x.2005, near rainforest, C van Achterberg & R de Vries; 1 ♂ (RMNH), S. Vietnam: Dông Nai, Cát Tien N.P., Head Quarters, 3.x.2005, at light, C van Achterberg & R de Vries. INDIA: 1 ♀ (RMNH), S. India, Madras State, Coimbatore, 1400 ft., v.1960, P Susai Nathan; 1 ♀ (RMNH), Madras, India; 1 ♂ (RMNH), India, Ajanda, 13.vii.1978, J Timor. INDONESIA: 1 ♀ + 1 ♂ (RMNH), Java, Malang, Kawi, iv.1933, JG Betrem. SRI LANKA: 1 ♂ (RMNH), Col. Dist., Colombo, 50 ft., museum gardens, 15.i.1977, KV Krombein & P Fernando.
No specimens from China were available in the present study, but this species was reported from China by
China (Guangdong, Hong Kong); Australia (Thursday Islands, Queensland); Bali; Bangka; Borneo; India; Java; Kariman Djava I.; Lombok; Malay Peninsula; Mariana Islands; Myanmar; New Britain; New Guinea; Pakistan; Palau Islands; Philippine Islands; Sri Lanka; Sulawesi; Sumatra; Sumba; Sumbawa; Talud Islands; Tukang Besi Islands; Vietnam; Volcano Islands. (
INDONESIA: 1 ♀ (RMNH), W. Flores, Rana Mese, 1300 m, 21.xi.1949, Dr. Bühler & Dr. Sutter (labeled as
No specimens from China were available in the present study, but the species was reported from China (Hong Kong) by
?China (Hong Kong), India, Thailand, Vietnam, Bangka, Sumatra, Java, Kariman Djava Isl., Bali, Lombok, Sumbawa, Flores, Sumba, Timor, Sulawesi. (
CHINA: 3 ♀ (NWUM), Mengla, Yaoqu, Yunnan, 27–30.ix.2010, JL Tan; 1 ♀ (NWUM), Longmen, Shangyong, Yunnan, 26.ix.2010, JL Tan; 1 ♀ (NWUM), Banna, Menglun, Yunnan, 5.x.2010, JL Tan. THAILAND: 1 ♀ (RMNH), 50 km SW Loei (
China (Yunnan), Thailand. (
MALAYSIA: 1 ♀ (RMNH), Penang Hill, Penang, 2500 ft, 27.i.1959, H.T. Pagden (paratype of
No specimens from China were available in the present study, but this species was reported from China by
China (Yunnan); Borneo; Malay Peninsula. (
CHINA: 1 ♀ (CATAS), Nada, Danzhou, Hainan,
The clypeus is completely yellow, but sometimes with a small black spot; the occipital carina is variable, sometimes the carina is bent anteriorly as in
China (Hainan); Cambodia; India, Malay Peninsula; Myanmar; Thailand; Vietnam. (
♀ (NWUM), CHINA: Longmen, Shangyong, Yunnan, 27.ix.2010, JL Tan.
The new species is similar to
♀, length of body (head + mesosoma + metasomal segments I–II) 11–12 mm, fore wing 10.5–12 mm.
The species name is derived from “para” (Latin for “near”) and “artifex”, because it is similar to
China (Yunnan); Thailand.
CHINA: 1 ♀ (RMNH), Tibet, Khamba Jong, F.A.M. INDIA: 1 ♀ + 1 ♂ (RMNH), Khasia Hills, Assam; 1 ♀ (RMNH), British Sikkim, 200 m, valley at Tista Bridge, 8–15.xii.1934, R Malaise; 1 ♀ (RMNH), E. Himalayas, Pashok, 2000 ft, Darjiling district, 11.vi.16, LC Hartless; 1 ♀ (RMNH), India, Lushai Hills, Miso district, Thingsat, 2000–3500 ft, 9–10.ix.1960, F Schmid. MYANMAR: 1 ♂ (RMNH), Burma: Washaung, 20 km, East of Myitkyina, c 200 m, 14.vii.1934, R Malaise; 1 ♀ (RMNH), Nord East Burma: Punkatang, road Sadon-Myitkyina, 8.vii.1934, R Malaise.
Occipital carina is curved forward ventrally and the gena narrow ventrally; the propodeum has the yellow stripes merged, but are sometimes lacking.
China (Tibet); India; Laos; Myanmar; Thailand; Vietnam. (
INDIA: 1 ♀ (RMNH), Sikkim, Kambur, 3280 ft, 15.viii.1959, F. Schmid; 1 ♀ (RMNH), Assam, Div. Kamens, Bokhar, 2200–2500 ft, 28.v.1961, F Schmid (both paratypes labelled “
The specimens from China agree well with the examined paratypes from India.
China (Tibet); India. (
INDIA: 1 ♂ (RMNH), India, Kulara, Kumaon, 12000 ft, 4.viii.1958, F Schmid. MYANMAR: 1 ♀ (RMNH), Carin Chebá, Karen Hills, Kayin State, 900–1000 m, 5.viii.1988, L Fea. CHINA: 5 ♀ (ZJUH), Ruili, Yunnan, 1.v.1981, no. 812465, JH He.
China (Yunnan); India; Myanmar; Thailand. (
CHINA: 1 ♀ (ZJUH), Guishan, Heyuan, Guangdong, 18.v.2002, no. 20028494, ZF Xu; 1 ♀ (NWUM), Luofu, Huizhou, Guangdong, 250 m, 12.vii.2004, CT Zhang; 1 ♀ (ZJUH), Dongzhong, Fangcheng, Guangxi, 8.xii.2001, ZQ He; 1 ♀ (ZJUH), Baise, Guangxi, 2.vi.1982, no. 822126, JH He; 9 ♀ + 2 ♂ (NWUM), Yaoqu, Mengla, Yunnan, 27–30.ix.2010, JL Tan; 7 ♀ + 2 ♂ (NWUM), Longmen, Shangyong, Yunnan, 26–27.ix.2010, JL Tan; 1 ♀ (GSFA) Xishuangbanna, Wangtianshu, Yunnan, 610 m, 23.iv. 2002, SP Sun; 1 ♀ (NWAM), Yaoqu, Mengla, Yunnan, 640 m, 6.v.1991, GC Liu & WZ Cai; 12 ♀ + 1 ♂ (NWUM), Beibeng, Motuo, Tibet, 10.viii.2011, HL Yang & JB Wang; 1 ♀ (NWUM), Motuo, Tibet, (
Specimens from Tibet are slightly darker than those from Yunnan and the basal yellow marks of the second sternite are very small or lacking. Chinese specimens have the gena slightly wider and the first tergite longer than other specimens, 3.0–3.5 times as long as high with small yellow patches. Specimens from Vietnam and Malaysia have the first tergite shorter, about 2.9 times as long as high with large yellow patches.
China (Hainan, Guangdong, Guangxi, Yunnan, Tibet); Bali; Borneo; India; Java; Malay Peninsula; Myanmar; Nepal; Philippine Islands; Sri Lanka; Sumatra; Thailand; Vietnam. (
CHINA: 1 ♀ (NWAM), Menglun, Yunnan,19.v.1991, YL Wang & RG Tian; 1 ♀ (ZJUH), id., but 12.iv.1981, JH He, no. 811796; 2 ♀ (ZJUH), Sanchahe, Yunnan, 10.iv.1981, JH He, no. 811812; 1 ♀ (ZJUH), Mt. Youle, Yunnan,17.iv.1981, JH He, no. 811873. MALAYSIA: 2 ♂ (RMNH), Sabah, Brumas, NBT, 23–27.i.1973, K.M. Guichard; 1 ♂ (RMNH), Borneo, Sarawak, trail Bario-Pa Lungan,
China (Yunnan); Bangka; Borneo; India; Malaysia (Sabah, Sarawak) (rec. n.); Myanmar; Sumatra; Thailand; Vietnam. (
CHINA: Type material of
China (Jiangxi, Fujian, Taiwan); Myanmar. (
INDIA: 1 ♀, Type, 60-15, E.I.C., with a handwritten label “
No specimens from China were available, but the species is included in the key because it has been listed for China.
China; India; Malay Peninsula; Myanmar; Nepal; Pakistan; Sulawesi. (
We thank the two anonymous referees, Dr JM Carpenter (American Museum of Natural History, New York), and Prof. Baozhen Hua (Northwest A&F University, China) for their important comments. We are in debt to Prof. Maoling Sheng (Shenyang), Dr Yihai Zhong (Hainan), Prof. C. Villemant (Paris), Dr G. Broad (London) and Dr F. Gusenleitner (Austria) for their kindness to loan specimens. Open access to this paper was supported by the Encyclopedia of Life (EOL) Open Access Support Project (EOASP). The first author was supported jointly by the Foundation for Key University Teachers by the Ministry of Education (No. 2011697505), the National Natural Science Foundation of China (NSFC, No. 31201732), the foundation of Shaanxi Educational Committee (No. 11JK0614), the Research Fund for the Doctoral Program of Higher Education of China (No. 20116101120001) and the Opening Foundation of Key Laboratory of Resource Biology and Biotechnology in western China (Northwest University), Ministry of Education.