ISSN 0973-1555(Print) ISSN 2348-7372(Online) HALTERES, Volume 9, 86-110, 2018 © P. GIRISH KUMAR AND JAMES M. CARPENTER doi: 10.5281/zenodo.1258043

A taxonomic review of the genus Thomson (: : Vespinae) from the Indian subcontinent

*P. Girish Kumar1 and James M. Carpenter2

1Western Ghats Regional Centre, Zoological Survey of India, Kozhikode, Kerala–673006, India. 2Division of Invertebrate Zoology, American Museum of Natural History, Central Park West at 79th Street, New York, NY 10024, USA.

(Email: [email protected])

Abstract

The genus Vespula Thomson, 1869, is reviewed from the Indian subcontinent. New synonymy is Vespa orbata var. aurulenta du Buysson, 1905, = Vespula structor (Smith, 1870). The following species are newly recorded from various states of India as follows: V. nursei Archer, 1981, from Delhi, Sikkim and Uttarakhand; V. orbata (du Buysson, 1902) from Nagaland; V. flaviceps (Smith, 1870) from West Bengal; V. germanica (Fabricius, 1793) from Himachal Pradesh; V. structor (Smith, 1870) from Arunachal Pradesh; V. vulgaris (Linnaeus, 1758) from Himachal Pradesh. Parasitic association of Strepsiptera is observed in one worker specimen of V. flaviceps. A modified key to species of the Indian subcontinent is provided.

Keywords: Vespula, review, Indian subcontinent, new record, ethology.

Received: 19 December 2017; Revised: 31 May 2018; Online: 1 June 2018.

Introduction Thomson (1869) erected Vespula as a India. Most of the specimens were properly subgenus of Vespa based on eight species; the preserved and added to the ‗National type species Vespa austriaca Panzer, 1799, Zoological Collections‘ of the Hymenoptera was subsequently designated by Ashmead Section of the Zoological Survey of India, (1902). Members of this social genus are Kolkata (NZC) and some of the specimens at commonly called yellow jackets. They usually Western Ghat Regional Centre of the make underground nests. They are widely Zoological Survey of India, Kozhikode distributed in the Northern Hemisphere. (ZSIK). Currently, 25 species are recorded worldwide List of repositories: Beijing — of which 10 species are present in the Indian Institute of Zoology, Academia Sinica; subcontinent. In this paper, we review the Budapest — Hungarian Natural History Indian subcontinent species. A new synonymy Museum; Dresden — Staatliches Museum für is proposed in this paper, i.e., Vespa orbata Tierkunde; Genève — Museum d‘Histoire var. aurulenta du Buysson, 1905, is Naturelle; Linnean Society — Linnean synonymized under Vespula structor (Smith, Society, London; London — The Natural 1870). The distributional records of many History Museum; New York — American species are augmented in this paper. Parasitic Museum of Natural History; Paris — association of Strepsiptera is observed in one Laboratoire d‘Entomologie, Museum National worker specimen of V. flaviceps. A key to d‘Histoire Naturelle; Stockholm — species of the Indian subcontinent is also Naturhistoriska Riksmuseet (Entomologiska); provided. Taichung — Taiwan Agricultural Research Institute; Washington — U. S. National Materials and Methods Museum of Natural History; Wien — This study is based primarily on the Zoologische Sammlung, Naturhistorisches specimens present in the Zoological Survey of 86

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Museum; Yunnan — Kunming Natural lateral mesoscutal depression without a History Museum of Zoology. backwardly directed depression; Abbreviations used for the terms: HW posterolateral margin of S6 of female = Head width; L = Length; OMS = Oculo- evenly convex; profile of T7 convex in malar space; S = Metasomal sterna; T = male; shape of S7 almost triangular in Metasomal terga; W = Width. male; male genitalia with digitus small, not extending to level of parameral spine; Results and Discussion apex of aedeagus strap-shaped and cuspis Genus Vespula Thomson, 1869 cone-shaped, without dorsal lobe………… Vespula Thomson, 1869: 79, subgenus of rufa species group. ……………..…….. 2 Vespa (eight species). Type species: — T1 usually with long pale hairs; occipital Vespa austriaca Panzer, 1799, by carina usually extending to base of subsequent designation of Ashmead, mandible; lateral mesoscutal depression 1902: 164. with a backwardly directed depression Pseudovespa Schmiedeknecht, 1881: 314, (sometimes indistinct on workers); subgenus of Vespa Linnaeus. Type posterolateral margin of S6 of female with species: Vespa austriaca Panzer, 1799, by a backwardly directed projection; profile monotypy. of T7 concave in male; shape of S7 Paravespula Blüthgen, 1938: 271, subgenus of transversely rectangular in male; male Dolichovespula Rohwer. Type species: genitalia with digitus large, extending Vespa vulgaris Linnaeus, 1758, by nearly as far as or beyond the level of original designation. parameral spine; apex of aedeagus not Allovespula Blüthgen, 1943: 149, subgenus of strap-shaped but either boat-shaped or Paravespula Blüthgen. Type species: spoon-shaped and cuspis pyramid-shaped ―Paravespula rufa (Linné)‖ [= Vespa with a dorsal lobe. ………………………7 rufa Linnaeus, 1758], by monotypy. 2. Hind tibia with long erect hairs; apex of Rugovespula Archer, 1982: 261, 264, clypeus sharply produced in females, subgenus of Vespula Thomson. Type bluntly produced in males. [Worker caste species: Vespa koreensis Radoszkowski, absent]..………..……..austriaca (Panzer) 1887, original designation. — Hind tibia without long erect hairs; apex of Diagnosis: Oculo-malar space short, as short clypeus bluntly produced in females, as or shorter than apical diameter of scape; hardly produced at all in males. [Worker pronotal carina absent or faintly marked caste present]…………………………….3 ventrally; female with outer flange on 3. Large punctures on mesoscutum with mandible; eye almost touching base of distance between punctures usually more mandible; occipital carina present laterally, or less equal or less than the diameter of reaching or not reaching base of mandible, punctures except at middle where sometimes weak; male antenna with long punctures are further apart; large punctures scape, equal to or little shorter than third of two sizes with normally a single smaller flagellar segment, later without tyloids; puncture between larger punctures; branches of aedeagus fused resulting in disc- micropunctures shallow so that surface like or saddle-shaped apical lobe; aedeagus between them appears polished………… longer than parameres. ………………………..…rufa (Linnaeus) — Large punctures on mesoscutum with Distribution: Eurasia, North Africa, North distance between punctures usually greater America, Central America. Adventive in than the diameter of punctures; large Australia, New Zealand, Tahiti, Hawaii, South punctures of two sizes with normally two Africa, Chile, Argentina. smaller punctures between larger punctures; micropunctures deep so that Key to species of the genus Vespula surface between them appears dull……...4 Thomson from the Indian subcontinent 4. Females (queens and workers)....…...…..5 (Modified from Archer, 2012) — Males.………………………………...... 6 1. T1 usually with long black hairs; occipital 5. Oculo-malar space short, about 1/20 to 1/ carina not extending to base of mandible; 25 the width of head; occipital carina long, occupying about 70% of the lateral length

87 A taxonomic review of the genus Vespula Thomson

of head; clypeus largely yellow with either yellow, brown); lower side of second each lateral yellow stripe broader than submarginal cell on the medial vein before median black stripe or yellow with three the point of contact with the second black spots; metanotum with two yellow medio-cubital or recurrent vein as long as spots. ……………………...nursei Archer or longer than the lower side of second — Oculo-malar space longer, about 1/14 to submarginal cell after the point of contact 1/17 the width of head; occipital carina with the recurrent vein; In male, shaft of short, occupying about 50% of the lateral aedeagus narrow and its apex spoon- length of head; clypeus largely black, with shaped; dorsal process of gonostipes well each lateral yellow stripe narrower than developed and projects laterally; T7 either median black stripe; metanotum without slightly or not at all notched.………… yellow spots………kingdonwardi Archer vulgaris species group. ……………… 13 6. Clypeus more than 50% yellow, black 8. Female (queens and workers)….….…….9 median stripe narrow and/or incomplete; — Males.………………………………… 12 metanotum with two yellow spots; coxa 9. Queens (usually body length greater than with yellow spot.………….nursei Archer 15.5 mm and forewing length greater than — Clypeus less than 50% yellow, black 13 mm).………………………………...10 median stripe broad and complete; — Workers (usually body length less than metanotum without yellow spots; coxa 15.5 mm and forewing length less than 13 without yellow spot………..kingdonwardi mm)..…………………..…………….....11 Archer 10. Propodeum very coarsely rugose all over 7. T1 raised basally with a depression behind its surface and without punctures; ventral this elevation (sometimes indistinct in metapleura anteriorly and ventro-laterally workers); In female, outer basal flange of strongly rugose………………....koreensis mandible nearly straight; In queen, (Radoszkowski) propodeum coarsely rugose all or over — Propodeum less coarsely rugose and major part of its surface; backwardly punctures between and interrupting the directed postero-lateral projection on S6 rugae; ventral metapleura without strong sharply pointed; In worker, either rugae.………...... orbata (du Buysson) propodeum rugose or if not clearly rugose 11. Mesoscutal punctures dense with distances in dorsal aspect, then body tricolored between punctures equal to diameter of (black, yellow, brown); lower side of punctures; propodeum rugose and with second submarginal cell on the medial punctures between and interrupting the vein before the point of contact with rugae; if the rugae indistinct then the body second medio-cubital or recurrent vein in dorsal aspect clearly bicoloured (black, shorter than the lower side of second yellow…..……koreensis (Radoszkowski) submarginal cell after the point of contact — Mesoscutal punctures usually less dense with the recurrent vein; In male, shaft of with distances between punctures greater aedeagus broad and its apex boat-shaped; than diameter of punctures; propodeum dorsal terminal process of gonostipes not rugose, only punctate; body in dorsal weakly developed; T7 deeply notched. aspect tricoloured (black, yellow, brown); ………… koreensis species group ….…8 brown particularly visible on vertex, — T1 not raised basally and without a pronotum and mesoscutum although depression; In female, outer basal flange sometimes restricted in amount…... orbata of mandible strongly curved (sometimes (du Buysson) indistinct in workers); In queen, 12. Propodeum rugose, without punctures; propodeum not coarsely rugose, surface dorsal terminal process almost absent, smooth with punctures and only weakly indicated by a slight undulation on margin rugose around petiole junction; of gonostipes……………………koreensis backwardly directed postero-lateral (Radoszkowski) projection on S6 bluntly rounded; In — Propodeum punctate, without rugae; dorsal worker, propodeum not rugose and in terminal process weakly developed but dorsal aspect body bicoloured (black, with a distinct projection on margin of yellow) except for V. structor and some V. gonostipes..………...orbata (du Buysson) flaviceps which are tricoloured (black, 13. Female (queens and workers)……..….. 14

88 P. Girish Kumar and James M. Carpenter

— Males..………………………………....21 20. Occipital carina incomplete only extending 14. Queens (usually body length greater than just over half way from vertex to 13 mm and forewing length greater than mandible; yellow area of ocular sinus with 11 mm). ………………………………..15 a slightly convex medial margin and — Workers (usually body length less than 13 slightly separated from yellow shield of mm and forewing length less than 11 mm). frons….………………...flaviceps (Smith) ………………………………………….18 — Occipital carina complete, extending from 15. Occipital carina not extending to base of vertex to mandible, but disappearing for a mandible, usually extends for less than ¾ short distance near mandible and then of lateral length of head; light coloured reappearing and reaching to mandible; ocular sinus not deeply penetrated by a yellow area of ocular sinus with a concave medial black projection; black mark on medial margin and widely separated from clypeus, if present, usually not reaching yellow shield of frons……………vulgaris apical margin...... flaviceps (Smith) (Linnaeus) — Occipital carina extending to base of 21. Aedeagus without a small backwardly mandible; other characters partly or directed pointed barb on each side below completely different. …………….…… 16 the apical spoon-shaped region; aedeagus 16. Margin behind third mandibular tooth emarginated apically; aedeagus with a distinctly concave….………….germanica small semicircular projection on each side (Fabricius) below apical spoon-shaped region; dorsal — Margin behind third mandibular tooth terminal process of gonostipes equal to, or straight or at most slightly concave. less than width of shaft of aedeagus. ………………………………………… 17 ………...…………germanica (Fabricius) 17. Bi-coloured (yellow and black); scutellum — Aedeagus with a small backwardly direct- and metanotum black, with yellow spots; ed pointed barb on each side below the clypeus yellow with black median mark; apical spoon-shaped region; aedeagus oculo-malar space shorter, HW/OMS rounded apically; other characters partly or greater than 19………vulgaris (Linnaeus) completely different. ……………...….. 22 — Tri-coloured (yellow, black and brown); 22. Volsella visible in ventral view of brown colour particularly visible on gonostipes; light coloured area of ocular pronotum, T2 and S2-S4; scutellum and sinus not deeply penetrated by a medial metanotum black, without yellow spots, at black projection, medial margin straight or most with brown markings; clypeus convex or continuous with the light yellow to orange without a black median coloured area of frons; mandible without a mark; oculo-malar space longer, triangular black spot at its base; genitalia HW/OMS less than 18.…structor (Smith) short, L/W less than one………...flaviceps 18. Light colour of gastral terga orange (Smith) yellow; dorsal surface of T1 without or — Volsella not visible in ventral view of almost without black marking; metanotum gonostipes; other characters partly or without a yellow mark…..structor (Smith) completely different. …………………..23 — Light colour of gastral terga yellow; dorsal 23. Gastral terga not covered by short brown surface of T1 with a distinct black hairs so that the light coloured areas marking; metanotum with a yellow appear clear yellow; metanotum with mark……………………………………19 yellow spots; light coloured areas of frons 19. Margin behind third mandibular tooth and ocular sinus parallel, or diverging distinctly concave; lateral surface of dorsally, so that they are clearly separated. mesosoma (mesepisternum, mesepimeron, ………………………vulgaris (Linnaeus) dorsal and ventral metapleura) with at — Gastral terga covered by short brown hairs least some long black hairs; if (rarely) no so that the light coloured areas appear black hairs present, then long hairs white. orange-brown; metanotum without yellow …………………...germanica (Fabricius) spots; light coloured areas of frons and — Margin behind third mandibular tooth ocular sinus either continuous or only straight or at most slightly concave; other narrowly separated……...structor (Smith) characters partly or completely different. ………………………………………….20 The rufa species group

89 A taxonomic review of the genus Vespula Thomson

Four species of rufa species group, data (New York). namely, Vespula austriaca (Panzer, 1799), V. rufa (Linnaeus, 1758), V. nursei Archer, 1981 Distribution: India: Jammu & Kashmir; and V. kingdonwardi Archer, 1981, are present Pakistan. Elsewhere: Albania; Austria; in the Indian subcontinent. Belarus; Belgium; Bosnia & Herzegovina; Bulgaria; Canada; China; Croatia; Czech 1. Vespula austriaca (Panzer, 1799) Republic; Denmark; Estonia; Finland; France; Vespa austriaca Panzer, 1799: 2, fig. 2, male, Georgia; Germany; Greece; Hungary; Ireland; ―Viennae Austr.‖ (destroyed). Italy; Japan; Kazakhstan; Korea; Kyrgyzstan; Vespa borealis Smith, 1843: 170, fig., female, Latvia; Liechtenstein; Lithuania; Luxembourg; ―Yorkshire… the north of Scotland‖ Macedonia; Moldova; Mongolia; Netherlands; (London). Junior primary homonym of Norway; Poland; Romania; Russia; Serbia; Vespa borealis Kirby, 1837, and of Slovakia; Slovenia; Spain; Sweden; Vespa borealis Zetterstedt, 1840. Switzerland; Turkey; U. K.; Ukraine. Vespa arborea Smith, 1849: 1x. Replacement name for Vespa borealis Smith. 2. Vespula rufa (Linnaeus, 1758) Vespa biloba Schilling, 1850: 77, [Poland] Vespa rufa Linnaeus, 1758: 572, ―in Europa‖ ―Schlesien‖ (type depository (lectotype female, Linnean Society). unknown). Additional citation: Day, 1979: 71 Vespa arborea ―Ross‖ Dalla Torre, 1894: 138, (designation of lectotype). as a synonym of Vespa austriaca Vespa schrenckii Radoszkowski, 1861: 84, Panzer. Junior primary homonym of figs. 4-5, [Russia] ―Amur‖ (type Vespa arborea Smith, 1849. depository unknown). Diagnosis: Female: Apex of clypeus sharply Vespa sibirica André, 1884: 599, female, produced; occipital carina present only male, [Russia] ―Sibérie‖ (Paris?). laterally; mesoscutum and scutellum dull with Vespa schrenki; Dalla Torre, 1904: 66. sparse shallow punctures; metanotum dull with Incorrect spelling of Vespa schrenckii fine shallow punctures; metasoma dull; body Radoszkowski. covered with long black erect hairs; hind tibia Vespa rufa grahami Archer, 1981: 54, 62, with characteristic long erect hairs; clypeus female, male – ―CHINA, Szechuen, 30 usually with 2-3 black spots; scape yellow miles north of Tatsienku‖ ventrally; ocular sinus yellow at ventral side; (Washington); also from elsewhere in metasoma without reddish areas. Sichuan. Male: Apex of clypeus bluntly produced; Vespula obscura Lee, 1986: 201, 205, 206, fig. genitalia with basal part of aedeagus narrower 5, female – [China] ―Zogang County, than the apical part, the apex evenly rounded Xizang (Tibet) Aut. Reg., 3800 m‖ and without lateral projections, but with (Beijing); also from another locality in subapical constriction; volsella short, Xizang. extending only up to the base of the apical ?Vespula yichunensis Lee, 1986: 202, 203, part; aedeagus entirely light brown. 206, fig. 3, female – [China] ―Yichun Length: Female, 15-18 mm; male, 13-16 mm. City, Heilongjiang Province‖ Ethology: It is an obligatory parasite that does (Beijing). not construct its own nest and breeds in the Vespula yulongensis Dong and Wang, 2002, nests of other Vespula species such as V. in Dong et al.: 396, figs. 1- 5, female - nursei, V. acadica and V. rufa (Williams, ―Yunnan China‖ (Yunnan). 1988; Archer, 2007). It does not produce a Vespula shinanoensis: Takamizawa, 2005, worker caste. The Japanese Social and Bees: 147, 148 (in subgenus Vespula). Material examined: No specimens from the Nomen nudum, described without Indian subcontinent were available for our mentioning repository of the holotype studies. We compared the species description (Articles 13 and 16.4 of the from Das and Gupta (1989) with a queen from International Code of Zoological Suisse - Valais/Grimentz/13 VII - 2 VIII Nomenclature). 1941/J. de Beaumont, and a male from Diagnosis: Female: Large punctures on Germany and Coll. Konow without further mesoscutum with distance between punctures

90 P. Girish Kumar and James M. Carpenter usually more or less equal or less than the ventrally; large punctures on mesoscutum with diameter of punctures except at middle where distance between punctures usually greater punctures further apart; large punctures of two than the diameter of punctures; large punctures sizes with normally a single smaller puncture of two sizes with normally two smaller between larger punctures; micropunctures punctures between larger punctures; shallow so that the surface between them micropunctures deep so that surface between appears polished; metanotum and metasoma them appears dull; propodeum punctate, more shiny with finer micro sculpture and weakly striate ventro-laterally; apical margin more superficial punctation; tibiae with sparse of S6 evenly convex. hairs; a dagger-shaped mark on clypeus; T1 Colour: Head black with yellow marks as and T2 partly red. follows: most of clypeus except linear black Male: Aedeagus not constricted basally, its mark medially; frons with a shield reaching basal part with thick walls; T1 and T2 with red antennal bases; stripe along ocular sinus spots. extending to clypeus; mandible; small spot posteriorly and large linear mark anteriorly on Material examined: No specimens from the gena. Mesosoma black with yellow marks as Indian subcontinent were available for our follows: stripe on pronotum parallel sided; studies. We compared the species description tegula; spot just below hind wing; two spots on from Das and Gupta (1989) with a queen from scutellum; metanotum with two spots Europe without further data, two workers from anteriorly; triangular spot on mesepisternum. Bressoux/Belgium/22/7/52 and Tolumierz/22/ Legs with coxae black or dark brown with X/1990/E. Yildirim, and two males from small yellow spot; femur mainly light brown; Tolumierz/22/X/1990/E. Yildirim and Maele tibia and tarsus brown with yellow dorsal W/Brugge/Belgium (New York). marks. Metasoma with black, reddish brown and pale yellow marks as in figure 6. Body Distribution: Nepal. Elsewhere: Afghanistan; hairs long and black; legs with long black hairs Albania; Austria; Azerbaijan; Belarus; on coxa, trochanter and femur. Belgium; Bosnia & Herzegovina; Bulgaria; Male: Clypeal angles absent, apical margin of China; Croatia; Czech Republic; Denmark; clypeus almost straight; antennal tyloids Estonia; Finland; France; Georgia; Germany; absent; apical margin of S6 deeply concave; Greece; Hungary; Iceland; Ireland; Italy; S7 almost triangular. Colouration similar to Japan; Kazakhstan; Kyrgyzstan; Korea; that of female except as follows: clypeus Latvia; Liechtenstein; Lithuania; Luxembourg; almost completely yellow except narrow black Macedonia; Moldova; Mongolia; Netherlands; median stripe; gena with anterior yellow stripe Norway; Poland; Romania; Russia; Serbia; along its entire length. Slovenia; Slovakia; Spain; Sweden; Length (H+M+T1+T2): Queen, 12.5-13.5 Switzerland; Taiwan; Tajikistan; Turkey; mm; Worker, 9.5-10.5 mm; Male, 9.5-11.5 U.K.; Ukraine; Uzbekistan. mm. Material examined: INDIA: Delhi, Delhi 3. Vespula nursei Archer, 1981 University Ridge, 1 worker, date of collection (Figs. 1-12) and name of collector unknown, NZC Regd. Vespula nursei Archer, 1981: 54, 57, fig. 4A- No. 17587/H3; Delhi University Campus, 1 B, female, male. ―INDIA, Kashmir, 8- male, 27.x.1978, Coll. Meenakshi, NZC Regd. 9,000 ft‖ (holotype female London); No. 17588/H3. Himachal Pradesh, Shimla also from other localities in Kashmir, district, Chini Bungalow, 1 worker, 11.x.1962, Punjab, Himachal Pradesh; China; Coll. V.K. Gupta, NZC Regd. No. 17589/H3; Philippines. Shimla district, Narkanda, 2 worker & 1 male, Diagnosis: Female: Oculo-malar space short, 5.x.1966–13.x.1971, Coll. V.K. Gupta & about 1/20 to 1/25 the width of head; clypeus Party, NZC Regd. Nos. 17591/H3–17593/H3; with coarse punctures separated by more than Shimla, 1 male, 3.x.1966, Coll. Ram, NZC their diameters; lateral angles of clypeus Regd. No. 17590/H3; Chamba district, triangular and projecting outwards; occipital Khajjiar, 1 worker, 27.vi.1965, Coll. Tikar, carina long, occupying about 70% of lateral NZC Regd. No. 17600/H3; Chamba district, length of head; pronotal carina absent; Dainkund, 1 worker, 11.vi.1971, Coll. pronotum punctate and weakly striate Unknown, NZC Regd. No. 17594/H3. Jammu

91 A taxonomic review of the genus Vespula Thomson

& Kashmir, Baramulla district, Gulmarg, 1 apical margin of T7 not notched, convex in worker, 17.vi.1966, Coll. Joseph, NZC Regd. profile; apical margin of S6 concave; S7 No. 17595/H3; Baramulla district, almost triangular; genitalia like that of V. rufa Chandanwari, 1 male, 1.x.1978, Coll. Rina, and V. nursei. Colouration similar to that of NZC Regd. No. 17597/H3; Srinagar, 1 male, female except gena with anterior yellow stripe 6.ix.1978, Coll. A. Kaur, NZC Regd. No. along its entire length. 17596/H3; Anantanag district, Pahalgam, 1 Length: Queen, 17 mm; Worker, 12-13.2 mm; male , 2.x.1978, Coll. Bindu, NZC Regd. No. Male, 13-14.7 mm. 17598/H3. Sikkim, East Sikkim district, Gangtok, 1 worker, 13.v.1966, Coll. Tikar, Distribution: Myanmar; Nepal. Elsewhere: NZC Regd. No. 17599/H3. Uttarakhand, China. Uttarkashi district, Harsil, 5 worker, 9– 13.vi.1977, Coll. V.K. Gupta & Party, NZC Remarks: Specimens of this species was not Regd. Nos. 17601/H3–17605/H3. available for our studies; hence the species Distribution: India: Delhi (new record), diagnosis, colour description and length are Himachal Pradesh, Jammu & Kashmir, taken from Das & Gupta (1989). Punjab, Sikkim (new record), Uttarakhand (new record); Pakistan. Elsewhere: China; The koreensis species group Philippines. This species group consists of only three species, namely, Vespula koreensis 4. Vespula kingdonwardi Archer, 1981 (Radoszkowski, 1887), V. orbata (du Buysson, Vespula kingdonwardi Archer, 1981: 54, 57 1902) and V. arisana (Sonan, 1929) of which (key), 60, fig. 4C-D, female, male – the first two species are present in the Indian [China] ―TIBET, Kashmir, 28.25 N subcontinent. 97.55 E‖ (holotype female London); also from other localities in Xizang; 5. Vespula koreensis (Radoszkowski, Myanmar. 1887) Vespula hirsuta Lee, 1986: 201, 204, 206, fig. Vespa koreensis Radoszkowski, 1887: 432, 4, female – [China] ―Markham male, female. ―Korée‖ (lectotype County, Xizang (Tibet) Aut. Reg., female London). Additional citation: 3300 m‖ (Beijing). Archer, 1982: 263, 267 (designation of Diagnosis: Female: Oculo-malar space longer, lectotype). about 1/14 to 1/17 the width of head; occipital Vespa germanica var. stizoides du Buysson, carina shorter, occupying about 50% of the 1905: 494 (key), 615, female. ―Nord lateral length of head; large punctures on de l‘Inde‖ (holotype London). mesoscutum with distance between punctures Pseudovespa birulai Gussakovskii, 1932: 52, usually greater than the diameter of punctures; female. [Russia] ―Ussuri-Geb.: large punctures of two sizes with normally two Tigrovaja…;Sutshan...Sedanka…Kreis smaller punctures between larger punctures; Chabarowsk…Wladiwostok‖(Stockhol micropunctures deep so that surface between m). them appears dull. Vespula koreensis salebrosa Archer, 1982: Colour: Head black with yellow marks as 261, 267, female. ―CHINA: Szechuen, follows: clypeus largely black with lateral Suifu‖ (London); also from other narrow yellow stripes; frons with a shield localities in China. reaching antennal bases; a stripe along ocular Vespula hainanensis Lee, 1986: 201, 202, 206, sinus not extending to clypeus; Mesosoma fig. 1, female. [China] ―Hainan Island, black with yellow marks as follows: a stripe on Guangdong Province‖ (Beijing). pronotum parallel sided; tegula brown with Diagnosis: T1 raised basally with a depression spots. Metasoma black with reddish brown and behind this elevation (sometimes indistinct in yellow markings; vertical face of T1 reddish workers). Female (queens and workers): outer brown with dorsal and ventral black marks. basal flange of mandible nearly straight. In Body hairs long and black; legs with long queen, propodeum very coarsely rugose all black hairs on coxa, trochanter and femur. over its surface and without punctures; ventral Male: Clypeal angles absent, apical margin of metapleuron anteriorly and ventro-laterally clypeus truncate; antennal tyloids absent; strongly rugose; backwardly directed postero-

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93 A taxonomic review of the genus Vespula Thomson

94 P. Girish Kumar and James M. Carpenter lateral projection on S6 sharply pointed. In queens, propodeum less coarsely rugose and worker, mesoscutal punctures dense with punctures between and interrupting the rugae; distances between punctures equal to diameter ventral metapleuron without strong rugae; of punctures, less dense on centre and denser backwardly directed postero-lateral projection on lateral margins; margin behind inner on S6 sharply pointed. In workers, mesoscutal mandibular tooth distinctly concave; occipital punctures usually less dense with distances carina reaches base of mandible; lower side of between punctures greater than diameter of second submarginal cell on the medial vein punctures, less dense on centre and denser on before the point of contact with the second lateral margins; propodeum not rugose, only medio-cubital or recurrent vein shorter than punctate; margin behind inner mandibular the lower side of second submarginal cell after tooth distinctly concave; occipital carina the point of contact with the recurrent vein. reaches base of mandible; lower side of second Male: Propodeum rugose, without punctures; submarginal cell on the medial vein before the genitalia with dorsal terminal process almost point of contact with the second medio-cubital absent, indicated by a slight undulation on or recurrent vein shorter than the lower side of margin of gonostipes; dorsal terminal process second submarginal cell after the point of of gonostipes weakly developed; parameral contact with the recurrent vein. spine gradually narrowing to an apical point; Male: Propodeum punctate, without rugae; the shaft of aedeagus broad and its apex boat- genitalia with dorsal terminal process weakly shaped; T7 deeply notched. developed but with a distinct projection on Length: Queen, usually greater than 15.5 mm; margin of gonostipes; the shaft of aedeagus Workers, usually less than 15.5 mm. broad and its apex boat-shaped; T7 deeply notched. Material examined: No specimens from the Colour (Worker): Body dorsally tricoloured: Indian subcontinent were available for our black, brown and yellow. Head yellowish studies. We compared the species description brown with vertex darker; mesoscutum brown from Archer (2012) with two queens from except black posteriorly; scutellum yellowish Foochow China/Kellogg coll. and Yen-ping, brown with a black mark in the middle; China/IV.16.1917/Ac. 5148, and two workers metanotum largely black and with a narrow from Yen-ping, China/X.11.1917/Ac. 5148 brownish yellow line across its anterior and VII.10.1917 (New York). margin; mesepisternum and usually the Distribution: India: Meghalaya, Sikkim; mesepimeron with a brown spot; propodeum Myanmar. Elsewhere: China; Korea; Laos; black with two lateral brownish yellow spots; Russia; Thailand; Vietnam. fore leg yellowish brown with coxa, trochanter and femur dorsally blackish brown; middle 6. Vespula orbata (du Buysson, 1902) and hind legs blackish brown with apex of (Figs. 13-18) femur dorsally and tarsus yellowish; T1, T2 Vespa orbata du Buysson, 1902: 140, female – and S2 black or blackish brown with a yellow [India] ―Dardjiling‖ (lectotype Paris); apical band; T3–T5 black basally and with also from Sikkim, China. Additional wider apical yellowish brown bands; T6 citations: Archer, 1982: 262, 268 wholly yellowish brown; S3–S6 largely (designation of lectotype). yellowish brown. Vespa minuta Dover, 1925: 304, fig. 9, female Length (H+M+T1+T2): Worker, 10.5-11.5 – [Myanmar] ―Tenasserim‖ (holotype mm. London). Junior primary homonym of Vespa minuta Linnaeus, 1767. Material examined: INDIA: Nagaland, Vespula nujiangensis Dong and Wang, 2004, Mokokchung, 1 worker, 26.iii.1989, Coll. B.C. in Dong et al., 146, female, figs. 1-5 - Das & Party, Regd. No. ZSI/WGRC/I.R- [China] ―Nujiang Liuku, Yunnan, Alt. INV.8971; Tuensang, 1 worker, 9.viii.2015, 1 100 m. 25°58‘ N. 98°65‘ E.‖ Coll. Lobeno, Regd. No. ZSI/WGRC/I.R- (Yunnan). INV.8972. NEPAL: Kathmandu, Sundarijal, 1 Diagnosis: T1 raised basally with a depression worker, date of collection and name of behind this elevation (sometimes indistinct in collector unknown, Regd. No. workers). Female (queens and workers): outer ZSI/WGRC/I.R-INV.8973. basal flange of mandible nearly straight. In

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Distribution: India: Arunachal Pradesh, subgenus Vespula). [Korea] ―Za-mo Nagaland (new record), Sikkim, Uttarakhand, san, 60 km NE from Pyongyan‖ West Bengal; Nepal: Sundarijal; Myanmar. (Budapest); also from other localities Elsewhere: China; Thailand; Laos; Vietnam. in Korea. This species was previously recorded by other Vespa vulgaris var. flavitor; Das & Gupta, workers from ―Uttar Pradesh: Mussoorie‖ and 1984: 449. Incorrect spelling of Vespa from ―Assam: Mishmi Hills‖. The locality vulgaris var. flavior Stolfa. Mussoorie presently belongs to Uttarakhand Vespula gracilia Lee, 1986: 201, 203, 206, fig. state and the locality Mishmi Hills presently 2, female. [China] ―Zhoushan, belongs to Arunachal Pradesh. Zhejiang Province‖ (Beijing); also from Fujian. The vulgaris species group Diagnosis: Queens: occipital carina not Four species of this species group, extending to base of mandible, usually extends namely, Vespa flaviceps (Smith, 1870), V. for less than ¾ of lateral length of head; light germanica (Fabricius, 1793), V. structor coloured ocular sinus not deeply penetrated by (Smith, 1870) and V. vulgaris (Linnaeus, a medial black projection; black mark on 1758) are present in the Indian subcontinent. clypeus, if present, usually not reach apical margin. Workers: pale markings ivory-white; 7. Vespula flaviceps (Smith, 1870) light coloured ocular sinus not deeply (Figs. 19-36) penetrated by a medial black projection; black Vespa japonica de Saussure, 1858: 261, mark on clypeus rarely reaches its apical female - ―Japon.‖ (Genève). Junior margin. Males: volsella visible in ventral view primary homonym of Vespa japonica of gonostipes; light coloured area of ocular Radoszkowski, 1857. sinus not deeply penetrated by a medial black Vespa flaviceps Smith, 1870: 174, 191, pl. projection, medial margin straight or convex XXI figs. 10, 11, female. ―Binsur, or continuous with the light coloured area of Kumaon, North-west Provinces of frons; mandible without a triangular black spot India‖ (London). at its base; genitalia short, L/W less than one. “Vespa Lewisii, Sauss. Guêpes Soc. (MS.)‖ Colour: Body markings highly variable in Smith, 1873: 198. Nomen nudum. queen and worker. Yellow usually paler, Vespa “Lewisii, Sauss. MS.‖ Cameron, 1903: approaching ivory white. Head yellowish 280, as a synonym of Vespa japonica except frons and vertex black; clypeus usually de Saussure, 1858. Available under yellow, rarely with a basal black line, or this Art. 11(e) of the International Code of line divided in to three spots; genal band Zoological Nomenclature. complete; antenna black with scape yellow to Vespa Saussurei Schulz, 1906: 231. yellowish brown ventrally; mesosoma black Replacement name for Vespa japonica except following yellow markings: posterior de Saussure. margin of pronotum; spot under wing base; Vespa karenkona Sonan, 1929: 137 (key), 148, anterior half of scutellum; anterior margin of female. [Taiwan] ―Karenkô (3000- metanotum; a pair of spots on propodeum; 6500 ft.)‖ (Taichung). gaster black with yellow bands that are darker Vespa 4-maculata Sonan, 1929: 137 (key), than mesosomal markings; legs largely 148, female. [Taiwan] ―Arisan‖ (coll. yellowish. Male similar to worker in M. Maki). colouration. Vespa vulgaris var. flavior Stolfa, 1934: 49, female. ―India: Bagarkote 8000 piedi. Length: Queen, 13-15 mm. Worker, 8-9 mm. Kumaon W. Himalaya… Dal, sopra Male, 13-14 mm. Dharmsala, 5500 piedi Punjab… Pindi Point, Murree, 7242 piedi. Material examined: INDIA: Himachal Punjab…Rive del fiume Deo Gad e Pradesh, Chamba dist., Dalhousie, Bakrota Foresta S. O. di Pinath, 7 miglia da Hills, 1 queen, v-vi.1927, Coll. S.L. Hora, Kausani Distr. Almora‖ (type NZC Regd. No. 2280/H3; Shimla, Matiana, 2 depository unknown). queen, 28-30.iv.1907, Coll. N. Annandale, Vespula japonica pionganensis Giordani Regd. Nos. ZSI/WGRC/I.R-INV.8974 & Soika, 1976: 287, 290, female (in 8975; Ahla, 1 queen, vii.1971, Collector

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99 A taxonomic review of the genus Vespula Thomson

100 P. Girish Kumar and James M. Carpenter unknown, Regd. No. ZSI/WGRC/I.R- worker & 1 male, 30.iii.1999, Coll. B. Mitra & INV.8976; Solan dist., Kandaghat, 1 queen, Party, Regd. Nos. ZSI/WGRC/I.R-INV.9007 6.iv.2011, Coll. D. Banerjee & Party, Regd. & 9008; Uttarkashi Dist., Jaspur, 1 worker, No. ZSI/WGRC/I.R-INV.8977; Chamba dist., 28.viii.2006, Coll. B. Mitra & Party, Regd. Khajjiar, 1 queen, 24.iv.2012, Coll. S.K. No. ZSI/WGRC/I.R-INV.9009. West Bengal, Ghosh & Party, Regd. No. ZSI/WGRC/I.R- Jalpaiguri Dist., Bangrakote, 1 queen, INV.8978; Kullu dist., Manali, 1 queen, 2.v.1914, Coll. Tytler, NZC Regd. No. 4.vi.2012, Coll. B. Mitra & Party, Regd. No. 1099/H3. ZSI/WGRC/I.R-INV.8979; Shimla, 1 worker, 26.ix.1918, Coll. Brunetti, Regd. No. Ethology: Parasitic association of Strepsiptera ZSI/WGRC/I.R-INV.8980; Shimla, 1 worker, was observed in one worker specimen 11.x.1974, Coll. L.K. Ghosh & Party, Regd. collected in Himachal Pradesh. The female No. ZSI/WGRC/I.R-INV.8981; Solan, 1 strepsipteran extruded out from the third worker, 14-17.vi.1979, Coll. V.K. Gupta & tergite of the wasps as seen in the figure 30. Party, Regd. No. ZSI/WGRC/I.R-INV.8982; Chamba dist., Dalhousie, 2 workers, Distribution: India: Assam, Himachal 10.xi.2010, Coll. G.P. Mandal & Party, Regd. Pradesh, Jammu & Kashmir, Manipur, Nos. ZSI/WGRC/I.R-INV.8983 & 8984. Meghalaya, Punjab, Sikkim, Uttarakhand, Jammu & Kashmir, Srinagar, 1 male, West Bengal (new record); Myanmar; Nepal; 20.x.1976, Coll. S.K. Gupta, Regd. No. Pakistan. Elsewhere: China; Japan (including ZSI/WGRC/I.R-INV.9084; Shopian district, Amami Is. In Ryukyu Is.); Korea; Laos; Heff village, apple orchards, 1 worker, Russia; Taiwan; Thailand; Vietnam. 9.ix.2017, Coll. Altaf Hussain, Regd. No. ZSI/WGRC/I.R-INV.10018. Manipur, Imphal 8. Vespula germanica (Fabricius, 1793) Valley, 3 workers, 24.xii.1945, Coll. M.L. (Figs. 37-48) Roonwal, Regd. Nos. ZSI/WGRC/I.R- ?Vespa maculata Scopoli, 1763, 312, INV.9085–9087. Uttarakhand, Nainital, 1 [Slovenia] ―Carnioliae‖ (destroyed?). male, 1908, Coll. R.E. Lloyd, Regd. No. Junior primary homonym of Vespa ZSI/WGRC/I.R-INV.8985; Chamoli dist., maculata Linnaeus, 1763. Lohba, 1 male, 17.iv.1914, Coll. Tytler, Regd. ?Vespa macularis Olivier, 1792: 695. No. ZSI/WGRC/I.R-INV.8986; Dehradun Unjustified emendation of Vespa dist., Mussoorie, 1 worker, 15.vi.1930, Coll. maculata Scopoli. B.N. Chopra, Regd. No. ZSI/WGRC/I.R- Vespa germanica Fabricius, 1793: 256. INV.8987; Pauri Garhwal dist., Kiwarsh, 1 [Germany] ―Kiliae‖ (type depository worker, 17.i.1966, Coll. T.D. Soota, Regd. No. unknown). ZSI/WGRC/I.R-INV.8988; Solan dist., Diagnosis: Queen and Worker: Margin behind Dagshai, 1 worker, 10.xi.1970, Coll. Kailash third mandibular tooth distinctly concave; Chandra & Party, Regd. No. ZSI/WGRC/I.R- outer basal flange of mandible strongly INV.8989; Nainital, 4 workers, 2-10.x.1978, curved; occipital carina reaching mandibular Coll. V.K. Gupta, Regd. Nos. ZSI/WGRC/I.R- base; apical margin of clypeus concave and INV.8990–8993; Uttarkashi Dist., Pilang, 5 depressed medially; mesosomal punctures workers, 18.xi.1999, Coll. B.C. Das & Party, indistinct; propodeum not rugose. Regd. Nos. ZSI/WGRC/I.R-INV.8994–8998; Colour: Body black with yellow markings as Uttarkashi Dist., Bhatwari, 6 workers, follows: mandible except teeth brown; 19.xi.1999, Coll. B.C. Das & Party, Regd. clypeus; shield on lower frons; ocular sinus (in Nos. ZSI/WGRC/I.R-INV.8999–9004; queen, yellow mark on frons connected to that Uttarkashi Dist., Bhatwari, 6 workers, of ocular sinus); area between antennal toruli 19.xi.1999, Coll. B.C. Das & Party, Regd. and inner eye margin; temple; broad mark on Nos. ZSI/WGRC/I.R-INV.8999–9004; pronotum; two marks on either sides of Rudraprayag Dist., Chopta, 1 worker, scutellum and metanotum; triangular mark on 21.xi.1999, Coll. B.C. Das & Party, Regd. No. mesopleuron near to forewing; small spots on ZSI/WGRC/I.R-INV.9005; Almora Dist., either sides of propodeum; tegula; mark Ranikhet, 1 worker, 27.xi.1999, Coll. B.C. Das behind base of hindwing; mark on mid and & Party, Regd. No. ZSI/WGRC/I.R- hind coxa; apical half of all femora; all tibia INV.9006; Pithoragarh Dist., Berinag, 1 and tarsus entirely; dorsal side of T1 except

101 A taxonomic review of the genus Vespula Thomson extended black mark at middle (large) and at Tunisia; Turkey; Turkmenistan; U.K.; sides (small); T2-T5 except at base black Ukraine; U.S.A. (Introduced); Uzbekistan. which triangularly extends towards apex at middle and separate small spots on either 9. Vespula structor (Smith, 1870) sides; S2-S5 except basal black area which (Figs. 49-60) extends to apex at middle and both sides; T6 Vespa structor Smith, 1870, in Horne and and S6 entirely. Smith, 191, pl. XXI fig. 12, female – Length: Queen, 14.5 mm; Worker, 8-12 mm. ―Binsur, Kumaon, North-west Provinces of India‖ (London). Material examined: INDIA: Himachal Vespa orbata var. aurulenta du Buysson, Pradesh, Lahaul Spiti district, Chango, 1 1905: 579, female, ―Chine, Mou-pin ... worker, 14.ix.2011, Coll. G. Srinivasan & ; Amur‖ (Paris, Wien?). NEW Party, Regd. No. ZSI/WGRC/I.R-INV.9553; SYNONYMY.1 Lahaul Spiti district, Pao, 1 worker, Vespa structrix Schulz, 1906: 231. Unjustified 21.ix.2013, Coll. V.D. Hegde & Party, Regd. emendation of Vespa structor Smith. No. ZSI/WGRC/I.R-INV.9554. Jammu & Vespula gongshanensis Dong 2005, in Kashmir, Ladakh, Kargil, 1 worker, Dong et al.: 65, female, male – 15.vi.1973, Coll. V.K. Gupta & Party, NZC ―China, Gongshan, Yunnan Province, Regd. No. 16574/H3; Ladakh, Leh, 98 alt. 2,950‖ (holotype female Yunnan). workers, 19-23.vi.1973, Coll. V.K. Gupta & Diagnosis: Queen: Tri-coloured (yellow, black Party, NZC Regd. Nos. 16575/H3–16672/H3; and brown); brown colour particularly visible Ladakh, Chusul, 1 worker, 7.ix.2008, Coll. on pronotum, T2 and S2-S4; scutellum and A.K. Sanyal & Party, Regd. No. metanotum black, without yellow spots, at ZSI/WGRC/I.R-INV.9555; Ladakh, Nider, 1 most with brown markings; clypeus yellow to worker, 8.ix.2008, Coll. A.K. Sanyal & Party, orange without a black median mark; oculo- Regd. No. ZSI/WGRC/I.R-INV.9556; Kargil malar space longer, HW/OMS less than 18; district, Panikhar, 2 workers, 20.ix.2008, Coll. margin behind third mandibular tooth straight A.K. Sanyal & Party, Regd. Nos. or at most slightly concave; occipital carina ZSI/WGRC/I.R-INV.9557 & 9558. extend to base of mandible. Workers: Light PAKISTAN: Chitral, 1 queen & 1 worker, colour of gastral terga orange yellow; dorsal 9.ix.1929, Coll. B.N. Chopra, Regd. No. surface of T1 without or almost without black ZSI/WGRC/I.R-INV.9559 (queen) & NZC marking; metanotum without a yellow mark. Regd. No. 1096/H3 (worker). Material examined: INDIA: Arunachal Distribution: India: Himachal Pradesh (new Pradesh, West Kameng district, Bomdila, 1 record), Jammu & Kashmir; Nepal; Pakistan. worker, 30.viii.1998, Coll. A.R. Lahiri & Elsewhere: Afghanistan; Albania; Algeria; Party, Regd. No. ZSI/WGRC/I.R-INV.10234; Armenia; Argentina (Introduced); Ascension Jammu & Kashmir, Srinagar, 1 worker, Island (Introduced); Australia (Introduced); 29.ix.1978, Coll. Bindu, Regd. No. Austria; Azerbaijan; Belarus; Belgium; Bosnia ZSI/WGRC/I.R-INV.10235; Sikkim, exact & Herzegovina; Bulgaria; Canada collection locality and date of collection (Introduced); Chile (Introduced); China; unknown, 2 workers, Coll. A.V. Knyvett, Croatia; Czech Republic; Denmark; Estonia; Regd. Nos. ZSI/WGRC/I.R-INV.10236 & Finland; France; Georgia; Germany; Greece; 10237; Uttarakhand, Uttar Kashi district, Hungary; Iceland (Introduced); Iran; Iraq; Ireland; Israel; Italy; Jordan; Kazakhstan; 1 Korea; Kyrgyzstan; Lebanon; Liechtenstein; One of us (JMC) has seen a specimen from Mou- Lithuania; Luxembourg; Macedonia; Malta; pin in Paris, with a determination label by du Mayotte; Moldova; Mongolia; Morocco; Buysson, a label reading “Type/examined by/Dr. J. Netherlands; New Zealand (Introduced); v. d. Vecht 1956” and a determination label as Norway; Poland; Portugal; Romania; Russia; Paravespula structor Sm. ab. by I. H. Yarrow. 1958. The determination appears to be correct, which Sakhalin Island; Scotland; Serbia; Slovakia; represents revised synonymy for this taxon. JMC Slovenia; South Africa (Introduced); Spain checked for material in Vienna, but only found an (including Canary Islands); Sweden; empty labeled space in the collection. Switzerland; Syria; Taiwan; Tajikistan;

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Syana Chatti, 4 queens & 1 worker, Distribution: India: Arunachal Pradesh (new 14.xi.2003, Coll. S.K. Chakraborty & Party, record), Assam, Jammu & Kashmir, Himachal Regd. Nos. ZSI/WGRC/I.R-INV.10238– Pradesh, Sikkim, Uttarakhand, West Bengal; 10242. Nepal; Myanmar. Elsewhere: China; Laos;

107 A taxonomic review of the genus Vespula Thomson

Thailand. irregular zigzag margin; apex of S2-S5 with almost regular band, narrowed towards sides; 10. Vespula vulgaris (Linnaeus, 1758) outer margin of T6; visible part of S6 almost (Figs. 61-66) entirely. Vespa vulgaris Linnaeus, 1758: 572, ―in Length: worker, 12. 5- 13.5 mm. Europa‖ (Lectotype female, Linnaean Society, London). Additional citation: Material examined: INDIA: Himachal Day, 1979: 77 (designation of Pradesh, Kinnaur district, Sangla, 1 worker, lectotype). 17.x.2006, Coll. R. Babu & Party, Regd. No. Vespa sexcincta Panzer, 1799: 1, fig. 1, male – ZSI/WGRC/I.R-INV.9560; Jammu & ―Viennae Austr.‖ (destroyed). Kashmir, Pahalgaon, 2 workers, 1-2.x.1978, Vespa vulgaris var. pseudogermanica Stolfa, Coll. Bindu, Regd. Nos. ZSI/WGRC/I.R- 1932: 26 (key), female – [Italy] ―Lago INV.9561 & 9562. superior di Weissenfels (Tarvisio …. Senosecchia‖ (type depository Distribution: India: Himachal Pradesh (new unknown). record), Jammu & Kashmir; Pakistan. Vespa mixta ―Schenck‖ Edwards, 1980: 363. Elsewhere: Albania; Argentina (Introduced); Nomen nudum. Australia (Introduced); Austria; Azerbaijan; Vespula rufosignata Eck, 1998: 299, female, Belarus; Belgium; Bosnia & Herzegovina; figs. 1A, C, 2 - ―India: Kashmir, Bulgaria; Chile (Introduced); China; Croatia; Gulmarg, 2900 m‖ (type depository Czech Republic; Denmark; Estonia; Finland; not stated; London); also from two France; French Polynesia (Introduced); other localities. Georgia; Germany; Greece; Hungary; Iceland Vespula vulgaris vetus Eck, 1999: 309, 329, (Introduced); Iran; Ireland; Israel; Italy; Japan; female, male - ―China, Quinghai Kazakhstan; Korea; Kurile Islands; Prov., Datong-Tal, 120 km NE Kyrgyzstan; Latvia; Liechtenstein; Lithuania; Xinging, 2360 m‖ (holotype female Luxemburg, Macedonia; Mexico; Moldova; Dresden); also from five other Mongolia; Netherlands; New Zealand localities. (Introduced); Norway; Poland; Portugal; Vespula yunlongensis Dong and Wang, 2003, Romania; Russia; Serbia; Slovakia; Slovenia; in Dong et al.: 212, fig. 1, female [?] - South Africa (Introduced); Spain; Sweden; ―Yunnan, China‖ (Yunnan). Switzerland; Syria; Turkey; U.K.; Ukraine. Diagnosis: Worker. Occipital carina complete, extending from vertex to mandible, but Acknowledgements disappear for a short distance near to mandible The authors are grateful to Dr. Kailash and then reappear and reaching to mandible; Chandra, Director, Zoological Survey of India, yellow area of ocular sinus with a concave Kolkata, and Dr. P.M. Sureshan, Officer-in- medial margin and widely separated from Charge, Western Ghats Regional Centre, yellow shield of frons; margin behind third Zoological Survey of India, Kozhikode, for mandibular tooth almost straight. providing facilities and encouragement. Colour: Body black with yellow markings as follows: mandible except teeth brown; clypeus References except an anchor-shaped black mark; shield on André, E. 1884. Species des Hyménoptères lower frons; ocular sinus; area between d‘Europe et d‘ Algerie, 2, Vespinae. antennal toruli and inner eye margin; temple Beaune. except wide black interruption; narrow mark Archer, M.E. 1981. The Euro-Asian species of on pronotum; two marks on either sides of the Vespula rufa Group (Hymenoptera, scutellum and metanotum; triangular mark on Vespidae) with descriptions of two new mesopleuron; spots on either sides of species and one new subspecies. Kontyû propodeum; tegula; small mark behind base of 49: 54-64. hindwing; mark on mid and hind coxa; apical Archer, M.E. 1982. A revision of the subgenus half of all femora; all tibia and tarsus entirely; Rugovespula nov. of the genus Vespula apical half of dorsal side of T1, band medially (Hymenoptera, Vespidae). Kontyû 59: invaginated; apex of T2-T5, anterior side with 261-269.

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Archer, M.E. 2007. Taxonomy, distribution Dong, D., Wang, Y., He, Y. and Wang, R. and nesting biology of species of the genus 2002. A New Species of Vespula Vespula or the Vespula rufa species group (Hymenoptera; Vespidae) from Yunnan, (Hymenoptera, Vespidae). Entomologist‘s China. Journal of the Southwest Monthly Magazine 143: 35-48. Agricultural University 24(5): 396-397. Archer, M.E. 2012. Vespine wasps of the Dong, D., He, Y., Wang, Y., and Wang, R. World. Behaviour, Ecology & Taxonomy 2003. A New Species of Vespula of the Vespinae. Monograph series, (Hymenoptera: Vespidae) from Yunnan, volume 4, 352 pp. Published by Siri China. Journal of the Southwest Scientific Press, Manchester, UK. Agricultural University 25(3): 212-213. Ashmead, W.H. 1902. Classification of the Dong, D., Wang, Y. He, Y. and Wang, R. fossorial, predaceous and parasitic wasps, 2004. A New Species of Vespula or the Superfamily Vespoidea. Paper no. 6. (Hymenoptera; Vespidae) from Nujiang Family XXVIII. Vespidae. Canadian of Yunnan Province. Journal of Southwest Entomologist 34: 163-166. Agricultural University (Natural Science) Blüthgen, P. 1938. Systematisches Verzeichnis 26(2): 146-147. der Faltenvespen Mitteleuropas, Dong, D., Liang, X., Wang, Y. and He, Y. Skandinaviens und Englands. Konowia 16: 2005. A New Species of Vespula from 270-295. Gongshan, Yunnan, China. Blüthgen, P. 1943. Taxonomische und Entotaxonomia 27(1): 65-68. biologische Natizen über paläarktische Dover, C. 1925. Further notes on the Indian Faltenwespen (Hym., Ves.). Stettiner Diplopterous wasps. The Journal of the Entomologische Zeitung 104: 149-158. Asiatic Society of Bengal, new series 20: Buysson, R. du. 1902. Descriptions de trois 289-305. guêpes nouvelles. Bulletin de la Société Eck, R. 1998. Eine nueu Art der Vespula entomologique de France: 140-141. vulgaris gruppe aus Asien (Insecta; Buysson, R. du. 1905. Monographie des Hymenoptera; Vespidae). Reichenbachia guêpes ou Vespa. Annales de la Société 32(43): 299-302. entomologique de France 73: 485- 556., Eck, R. 1999. Zur Populationsgleirung und 565- 634. Zoogeographie von Vespula vulgaris Cameron, P. 1903. Descriptions of four Factoren der intraspecifichen species of Vespa from Japan. Merkmalsanderungen (Insecta: Entomologist 36: 278-281. Hymenoptera: Vespidae). Entomologische Dalla Torre, K.W. von. 1894. Catalogus Abhandlungen, Staatliches Museum fur Hymenopterorum 9, Vespidae Tierkunde, Dresden 58: 309-342. (Diploptera), Leipzig, 181 pp. Edwards, R. 1980. Social Wasps. Rentokil Dalla Torre, K. W. von. 1904. Vespidae, Limited, East Grinstead, 385 pp. Genera Insectorum 19: 1-108. Fabricius, J.C.F. 1793. Entomologia Das, B.P. and Gupta, V.K. 1984. A catalogue Systematica Emendate et acuta. of the families Stenogastridae and Secundum, Classes, Ordines, Genera, Vespidae from the Indian Subregion Species, Adiectis Synonymis, Locis, (Hymenoptera: Vespoidea). Oriental Observationibus, Descriptionibus 2. 17: 395-464. Hafniae, viii+519pp. http://dx.doi.org/10.1080/00305316.1983. Giordani Soika, A. 1976. Vespidi ed Eumenidi 10433698 raccolti in Corea (Hymenoptera). Annales Das, B.P. and Gupta, V.K. 1989. The social d‘Histoire Nationale de Hungary 68: 287- wasps of India and the adjacent countries 293. (Hymenoptera: Vespidae). Oriental Insects Griffin, F.J. 1939. On the dates of publication Monograph 11: 1-292. of de Saussure (H. de): Etudes sur la Day, M.C. 1979. The species of Hymenoptera famille des Vespides 1-3. 1852-1858. described by LINNAEUS in the genera Journal of the Society for the Bibliography Sphex, Chrysis, Vespa, Apis and Mutilla. of natural History 1: 211-212. Biological Journal of the Linnean Society Gussakovskii, V. 1932. Verzeichnis der von 12: 45-84. Herrn Dr. Malaise im Ussuri und Kamtshatka gesammelten aculeaten

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Hymenoptera. Arkiv för Zoologi 24 Schulz, W.A. 1906. Spolia A(10):1-66. Hymenopterologica. Insel Creta. 355 pp. Horne, C. and Smith, F. 1870. Notes on the Paderborn. habits of some hymenopterous insects Scopoli, G.A. 1763. Entomologia Carniolica from the north-west provinces of India. exhibens insecta Carnioliae indigena et With an appendix, containing descriptions distributa in ordines, genera, species, of some new species of Apidae and varietates. Vienna: Johan Thomas von Vespidae collected by Mr. Horne: by Trattner. Frederick Smith, of the British Museum. Smith, F. 1843. Descriptions of the British Transactions of the Zoological Society of wasps. Zoologist 1: 161-171. London 7: 161-196, pls. 19-22. Smith, F. 1849. Descriptions of new species of Lee, T.S. 1986. Notes on the genus Vespula British bees. Zoologist 7, App.: lvii-lx. from China (Hymenoptera: Vespidae). Smith, F. 1873. Descriptions of aculeate Sinozoologica 10(4): 201-206. Hymenoptera of Japan, collected by Mr. Linnaeus, C. 1758. Systema Naturae, (10th George Lewis at Nagasaki and Hiogo. Ed.) Vol. 1. (Laurentii Salvii, Holmiae). Transactions of the Entomological Society Olivier, G.A. 1791-92. Encyclopedie of London 1873: 181-206. methodique histoire naturelle. Insectes 6. Sonan, J. 1929. On Vespa from Formosa. Paris (Panckoucke): 704 pp. Transactions of the Natural History Panzer, G.W.F. 1799. Faunae Insectorum Society of Formosa 19: 136-149. Germanicae, Heft 63: 1-2. Stolfa, E. 1932. Il genere Vespa nella Venezia Radoszkowski, O. 1861. Beschreibung einiger Giulia. Memorie della Societa neuen Hymenopteren-Arten. Horae Entomologica Italiana 9(1): 24-29. Societatis Entomologicae Rossicae 1: 79- Stolfa, D.E. 1934. Due nuovi Vespidi indiani. 86, pl. II. Bollettino del Museo civico di Storia Radoszkowski, O. 1887. Hyménoptères de Naturale di Venezia 1: 47-49. Koree. Horae Societatis Entomologicae Takamizawa, K. 2005. Description of a New Rossicae 21: 428-436. Species of Vespula (Subgenus Vespula) Saussure, H. de. 1858. Note sur la famille des From Nagano Pref., Central Honshu, Vespides. Revue et magasin de zoologie Japan (Hymenoptera: Vespidae): P. 148, 2(10): 259-261. (In: Takamizawa, K., Ed, The Japanese Schilling, P.S. 1850. Die in Schlesien und der Social Wasps and Bees), The Shinano Graftschaft Glatz gesammeleten Arten der Manichi Shimban, Nagano. Gattung Vespa. Jahresbericht der Thomson, C.G. 1869. Ofversigt of Sveriges Schlesischen Gesellschaft für Vespariae. Opuscula Entomologica 1: 78- Vaterländische Cultur 1850(1): 76-78. 82. Schmiedeknecht, O. 1881. Über einige Williams, P.H. 1988. Social wasps (Hym., deutsche Vespa-Arten. Entomologische Vespidae) from the Kashmir Himalaya. Nachrichten 7: 313-318. Entomologist‘s Monthly Magazine 124: 149-152.

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