ENTOMOLOGIST'S MONTHLY MAGAZINE 43 TAXONOMY, DISTRIBUTION AND NESTING BIOLOGY OF WSPA BINGHAMI, U BASALIS, V UARIABILIS, U FERWDA, U. LUCTUOSA, U MULTIMACULATA AND U BELLICOSA (HYM., VESPINAE) BY MICHAEL E. ARCHER This paper completes the coverage of the remaining seven species of in previous papers (Archer, 1991b, L992, 1993, I994b, 1995,1996). The genus Vespa is considered to contain 23 species (Archer, 1991a). Keys to the species are given in Archer (1989) and a cladistic analysis to show possible phylogenetic relationships in Archer Vespa not considered Q99aa). Van der Vecht (1957,1959) provides important taxonomic papers onVespa species giving much geographical distribution information. Vespa binghami du Buysson, 1905 This species consists of two colour subspecies: the western nominate subspecies with a largely dark brown unicolorous gaster and the eastern V binghami suprunenkoi Birula, 1925 with light and dark brown bands on the gaster. Intermediately coloured females with less extensive light brown gastral bands than those of V binghami suprunenkoi have been found in Sichuan, China. Males may show a greater range in colour variation than the females. A male V binghami suprunenlcoi has been found in northern Thailand where the females are the nominate subspecies. The geographical distribution of V binghami (fig. 1) is: India (northem West Bengal, Meghalaya), northern Burma, Bhutan, northern Thailand, northern Laos, China (Tibet, Sichuan, Yunnan), Russia (Sakhalin Primor'ye ["Maritime Kray']) and Korea. Very little is known about the nesting biology of V bingharni. Information on labels attached to specimens (van der Vecht,I959;Archer, unpublished) indicate that this species is found between the altitudes of 20Om and 273Om, usually over L000m, and is active from March until early November. The life cycle is annual and colonies are synchronised in their development. The queens are larger than the workers and readily separated by size. Vespa basalis Smith, L852 V basalis is widespread in continental Asia extending onto the islands of Sumatra and Taiwan. Its detailed geographical distribution (fig. 2) is: Pakistan (Islamabad), India (Uttar Pradesh, Himachal Pradesh, Meghalaya, Assam, northern West Bengal), Nepal, Burma (Kachin, Shan, Karen, Tenasserim), northern Thailand, Laos, northern Vietnam, China (Sichuan, Yunnan, Fujian), Korea, Sumatra and Taiwan. Further distribution information is given for Nepal by Yamane (L974), Taiwan by Starr (1992), and Korea by Kim et al. (1994). 29th March, 1999 Vol. 135 (1999) 44 ENTOMOLOGIST'S MONTHLY MAGAZINE Fig. 1. -o>=-2? Fig.2.-The world distribution of Vespa basalis. 45 MAGAZINE The nesting biology of V basalis is known to some extent from ENTOMOLOGIST'S MONTHLY Williams (1919), van der Vecht (1957,1959), Yamane, 56. (1977, 1992), Matsuura (1973), Yamane, Sk. (1974), Kuo & Yeh (1985), and Muzaffar & Ahmad (1986). Information on labels attached to specimens has also proved useful (Archer, unpublished). I am grateful to Dr S. Martin for a translation of Kuo & Yeh (1985). Typically, nests are found in mountain forests attached to the branches of trees, sometimes in shrubs and on buildings, in rock crevices and in the ground. Aerial nests in trees are usually at least four metres above the ground. Underground nests are shallow so that the nest is visible from the entrance. Nests which are started in holes are relocated during May or June to aerial sites. In Taiwan, nest densities of ten nests per 150ha have been found. In Nepal, nests are found between 90Om and 1500m although adults have been found between 700m and 1500m. In Taiwan, nests are found from 100m up to 1500m altitude but mainly between 200m and 800m. Generally, throughout the range of V basalis, adults have been found between 1,0m and 2830m, but mainly between 500m and 190Om. Queens overwinter in holes and crevices in walls, trees, and even in old nests. The overwintering period is usually from late December until early March, although adult activity has been recorded during January and February. In Taiwan the queens establish nests during March and April with the first workers appearing during April and May. Sexuals emerge from October until December. The life cycle is very long: about nine months or 270 days. Queens carry out inter-specific usurpations on Vespa tropica (L.) at the end of April (Kuo & Yeh, 1985), and on V velutina l-epeletier (Yamane, reez). At first, nests in bushes are egg-shaped with a short (2-3cm) ventral tube. After relocation, the nest shape is at first spherical, becoming pearshaped with a conical roof. In one nest the conical roof occupied about I2Vo of. the volume of the nest. The entrance to the nest is at first circular, but later becomes a vertical slit (about Lcm wide and 17cm long). Mature nests usually have three slit-like entrances on thc sides of the envelope. Mature nests vary in height (22-90cm) and width (LG70cm), and in the number of combs (5-16 or more). One large nest was estimated to contain 40,100 cells, of which 27,600 were small cells used to rear the workers and some males, and I2,5O0 large cells used to rear the queens and most of the males. The small cells (diameter 7.1mm) were distinctly smaller than the large cells (diameter 9.4mm). The upper older combs consisted of small cells and the lower combs of large cells. Central combs consisted of small and large cells with the proportion of large cells increasing in the lower combs. In Taiwan, the maximum number of immature brood is reached during October. In a large colony the number of immature brood was estimated to be in excess of 25,000 including sealed brood of 1,848 workers, 2,187 46 ENTOMOLOGIST'S MONTHLY MAGAZINE queens, and2,74}males (about T40insmall cells and 2,000 in lar_g_e,celll)In ttris large colony, meconial counts showed that about 30,000 cells (76.8Vo of Cells) had produced at least one adult: 44.9Vo of cel.ls producing only one adult;'28.5% two adults; 3.3Vo three adults, and 0.1Vo four adults. 'itre developmental period of the brood varied from 4143 days (egg stage 6-7 day3, larva L5 days, sealed brood 2V2I days). It is not clear whlther this developmental period refers to the workers, queens or males. Normally a colony reaches a maximum Population of about 6,000 workers, alihough a lirge colony may have L0,000. In one lq_gt colony, it it would pioduce 37,600 workers, 4,000 males and 3,400 queens. Workers and queens differ in size. was estimated ihat Vespa variabilis du Buysson, L905 'itris sp""ies consist-s of two colour subspecies: the eastern nominate , subspeciei with the first to the fifth gastral terga having alternating black or dark brown and light brown bands, and the western V variabilis fumida van der Vecht, 1959 with the first to the fifth gastral terga black. The geographical distribution of V variabilis (fig. 3) is: India (northern West nEngit, Arunachal Pradesh, Sikkim), Nepal, Burma (Sagaing) and China (Sichuan, Yunnan, Hubei, Fujian). Fig. 3. - The world distribution of Vespa variabilis. Very little is known about the nesting biology of U variabilis. Information on labels attached to specimens (van der Vecht,l959;Archer, unpublished) indicates that this species is found at altitudes between 670m and 3000m but usually over 1000m, and therefore is a montane species. Adults were collected from the end of February until the end of November. The life cycle is annual with the queens overwintering for two to three months. Colonies are synchronous in their development. V fervida Smith, 1859 U fervida and U luctuosa are a species pair united by the synapomorphy of the apical margin of the seventh gastral sternum of the male with two emarginations (Archer, I994a). ENTOMOLOGIST'S MONTHLY MAGAZINE 47 V fervida is restricted to Sulawesi (Indonesia) but is found throughout this island including its southern islands of Salayat and ButangVery little is known about the nesting biology of V fervida except that it is found in forested areas (personal communication, G.R. Else, 1986). Information on labels attached to specimens (van der Vecht, t957, 1959; Archer, unpublished) indicate that this species is found at altitudes from 100m to 1080m so that probably it is restricted to forested areas rather than a particular altitudinal range. The- queens and workers clearly differ in size. Queens are found throughout the year so it is possible that colonies are asynchronous in their development. V luctuosa de Saussure, 1854 The nomenclatural history of V luctuosa is complex. Bequaert (193a) recognised five colour varieties of V luctuosa: 1, The nominate colour variety; 2, semperi du Buysson, L905; 3, malayana, nom. nov- for annulata Smith, 1858; 4, bellicosa de Saussure, 1854; and 5, luzonensis, nova var. Van der Vecht (1957) raised bellicosa to species rank and recognised malayana as a synonym of the subspecies V multimaculata multimaculata Perez,19L0. The other three colour forms of V luctuosa were considered to be subspecies. Kojima & Tano (1985) described V luctuosa negrosensls based on two queens, which could not be distinguished from V luctuosa luctuosa, and fbur workers, with very minor colour differences from V. luctuosa luctuosa. With further study of a larger number of specimens this new colour variety might be considered a microgeographic variation but not a macrogeographic variation or subspecies. As such V luctuosa negrosensis should be treated as a syn. n. of V luctuosa luctuosa. V luctuosa is restricted to the Philippine Islands. V luctuosa semperi (southern colour form) has been found on the island of Mindanao and is distinguished from the other colour forms by its yellow sixth gastral tergum. The other two colour forms have a black sixth gastral tergum. V luctuosa luctuosa (northern-central colour form) has been found on the islands of Luzon, Mindoro, Panay, Negros and Cebu. It is recognised by the narrow yellow apical band on the first to the fourth or fifth gastral terga. The width of the yellow band on the second gastral tergum extends about one sixth of the way across the tergum. V. luctuosa luzonensis (eastern colour form) has been found on the islands of Leyte, Biliron, Panaon and Samar. It is recognised by the wide yellow apical band on the first to the fourth or fifth gastral terga. These yellow bands extend at least one quarter of the way across the tergg. Kojima (1982) and Kojima & Reyes (1936) give keys and distribution notes on the colour forms of V luctuosa. Alittle information is available on the nesting biology of V luctuoss in Williams (1919), Kojima & Yamane (1980), Kojima (1982) and Starr & 48 ENTOMOLOGIST,S MONTHLY MAGAZINE Jacobson (1990). Information on labels attached to specimens. and the specimens'themselves (van der Vecht, 1957 ;Archer, unpublished) provide information on female caste differences, altitude distribution, and months of activity. Nests are usually found in forested areas attached to the branches of trees from Lm to 6m, but sometimes up to 2Om, above the ground. Nests also have been found in bushes, and one was attached to a ceiling of an abandoned house. queen nest of V luctuosa luctuosa consisted of a single comb completely covered by an envelope with a ventral entrance extended into a ,rrty short (about 3mm long) cylindrical tube. Two queen nests- of V luctuosa luzonensis had bell-shaped envelopes so that from a ventral view A the single comb was clearly visible. Later nests are . spherical and completely covered by envelopes except fo1 a ventral circular or oval entrince situated to one side. Mature nests elongate vertically but lack a roof cone. Probably up to six combs are present in mature nests with about'l',250 cells, alttrougtr larger nests have been seen. Small and large cells are s"putit"d. ftre oldest comb, at the top of the nest, consists of small readily-occisionally with a few peripheral large cells, while the other combs cells, consist of large c-ells, sometimes with small cells in the second comb. Cells in the oldest comb have up to five meconia per cell, indicating that a cell has been used to rear five adults. The second comb may have up to four meconia per cell, while younger combs have progressively fewer meconia per cell. Adulti have been found from 2OOm to 150Om, and to be active in all months except February. Males have been found during January, March, May, September, and October. Queen, small and large nests all have been found during March. Thus it is highly likely that colonies are asynchrononJitr their development. Queens and workers clearly differ in size. V multimaculata Perez, 1910 U multimaculata and U bellicosa are synapomorphy just a species pair although a for these two species has not been found (Archer, 1994a). Van der Vecht (L957) described the western colour form, V multi- maculsta pendleburyi, with wider yellow gastral telgal bands, e.g. on the second teigum, at short distance from the middle of the tergum, the yellow band extends about one quarter across the tergum. In the eastem colour form, the nominate subspecies, it only extends about one sixth across the tergum. I have seen specimens of V. multimaculata pendleburyi and V multiiaculata multimacilata together at the same site both in the range of the westem colour form (in Peninsular Malay-sia) Td the eastern i colSur form maintained. (in Sabah). Th" two subspecies therefore should not be ENTOMOLOGIST'S MONTHLY The geographical distribution of U MAGAZINE multimaculata is: 49 Singapore, Malaysia (Peninsular, Sarawak, Sabah), Brunei, and Indonesia (Sumatra, Kalimantan). Very little is known about the nesting biology of V multimaculata. Martin (1995) found that it nested in the ground, especially under the roots of trees with nests from 20 to 30cm in diameter. Nests are associated with both lowland and montane forests (van der Vecht, 1957; Martin, 1995). Information on labels attached to specimens (van der Vecht, 1957, I959;Archer, unpublished) indicate that this species is found at altitudes between 50m and 21,40m. Adults have been found throughout the year except January. Queens have been found active away from the nest during February, April, July and October, and males have been found during February, March, April and July. Thus it is highly possible that colonies are asynchronous in their development. Queens and workers clearly differ in size. V bellicosa de Saussure, 1853 The geographical distribution of V bellicosa is: Indonesia (Sumatra, Kalimantan) and Malaysia (Sarawak). Very little is known about the nesting biology of V. bellicosa. It is with lowland forests (van der Vecht, t957; Martin, 1995). Information on labels attached to specimens (van der Vecht, 1957, I959;Archer, unpublished) indicate that this species is found at altitudes between 75m and I25m. Adults have been found throughout the year except for March and December so that it is highly likely that colonies are asynchronous in their development. Queens and workers probably differ associated in size. REFERENCES Archer, M.8., 1989, A key to the world species of the Vespinae (Hymenoptera). Part 1, Key, Checklist and Distribution: 147 Part 2, (72 figs): t1-34), Research Monograph of the College of Ripon and York St John, No. 2; l99la, The number of species that can be recognised within the genus Vespa (Hy-., Vespinae), Entomologistb mon. Mag., 127; 16l-164;1991b, Taxonomy and bionomics of the Vespa tropica group (Hym., Vespinae), ibid., 127: 2?5-232; 1992, T\e taxonomy of Vespa crabro L. and V. dybowskii Andre (Hym., Vespinae), ibid.,l28 157-163;1993, The life history and colonial characteristics of the Hornet, Vespa crabro L. (Hym., Vespinae), ibid., 129 1,51-163; 1994a, A phylogenetic study of the species of the genus Vespa (Hymenoptera: Vespinae), Ent. scand.,24:469478; 1994b, Thxonomy, distribution and nesting biology of the Vespa bicolor group (Hym., Vespinae), Entomologist's mon. Mog., 130: 14L158; 1995, Thxonomy, _distribution and nesting biology of the Vespa mandarinia group (Hym., Vespinae), ibi.d., l3l: 47-53;1996, Taxonomy, distribution and nesting biology of Vespa affinis and Vespa mocsaryann du Buysson (Hym., Vespinae), ibid., 133l. 27-38. Bequaert, J., 1930, On the generic and subgeneric divisions of the Vespinae (Hymenoptera), Bull. Brooklyn ent. Soc.,25: 59-70; L934, I-es races de coloration de Vespa luctuosa de Saussure et de Polistes tenebricosus r Lrpeletier, Bull. Mus. Hist. nat. Belg.,10: 1-11. Kim, J.K., Moon, T.-Y. & Yoon, I.8., 1994, Systematics of vespine wasps from Korea, I. Genus Vespa Linnaeus (Vespidae, Hymenoptera), KoreanJ. Entom.,24:1'07-11'5. Kojima, J.,1,992? The genusVespa in the ftritipplnes (Hymenoptera, Vespidae), KontyA, 50: 4344M. Kojimar_J. & Reye^s, S.G., t986,?tritippine Vespidae, I, Genus Vespa, Philipp. Ent.,6:445454. Kojima, J. & Thno' 50 ENToMoLocrsls MoNTHLy MAGAZINE 1985, A new subspecies of Vespa luctuosa from Negros Island, ibid.,6: 43H37. Kojima, J. & Yamane, Slc, 1980, Biological notes on Vespa luctuosa luzonensis from lryte Island, The Philippines, with descriptions of adults and larvae (Hymenoptera: Vespidae), Insecta matsum.,19:79-87. Kuo, M.C. & Yeh, W.H., 1985, Studies of Vespidae T., in Taiwan. II. Ecological studies of Vespa basalis Smith, Vespa velutina flavitarsis Sonan and Vespa tropica pseudosoror Vecht, J. Natn. Chinyi lrut. Agric., 11: 95-106. Martin, S.J., 1995. Hornets (Hymenoptera: Vespinae) of Malaysia. Malayan Nature J., 49:71-82. Matsuura, M., 1973. Nesting habits of several species of the genus Vespa in Formosa, Kontyfi,4l:28G293. Matsuura, M. & Yamane, Slc, 1990, Biology of the vespine wasps, Springer-Verlag, Berlin. Muzaffa4 N. & Ahmad, R., 1986, Studies on hornets attacking honeybees in Pakistan, Pak J. agric. Res., 7: 5943. Staq C.K., 1992, The social wasps (Hymenoptera: Vespidae) of Thiwan, Bull. Nat. Mus. Nat..Scr.,3:9!138. Star, C.K. & Jacobson, R.S., 1990, Nest structure in Philippine Hornets (Hymenoptera, Vespidae,Vespa spp.),Jap.J. Ent.,58:125-143. VechtJ. van der,l95T,TheVespinaeof the Indo-Malayan and Papuan areas (Hymenoptera, Vespidae), Zool. Verh. Leiden,34: 1-83; 1959, Notes on Oriental Vespidae, including some species from China and Japan (Hymenoptera, Vespidae), Zool. Meded.,36:205-232. Williams, EX., 1919, Philippine wasp studies, 2, Descriptions of new species and life history studies, Bull. Hawaiian Sug. Pltrs' Ass. Srn. (Ent. Ser.), 14: 119-186. Yamane, Sk, 1974, On the genus Vespa (Hymenoptera, Vespidae) from Nepal, KontyL, 42: 29-39. Yamane, 56., 1977, On the collecting of vespine nests, based on practices through a survey in Thiwan from 1972to 1974 (Hymenoptera: Vespidae), Seibutsu Ky6zai 12: 42-59; 1992, A huge nest of Vespa basalis collected in Taiwan (Hymenoptera: Vespidae), Chinese J. Entom.,12: 1-11. The University College of Ripon and York St John, l,ord Mayor's Walk, York, YO3 7EX, England, U.K. November 13th,1996. Platyspathius sp. (Hym., Braconidae, Doryctinae) in S.W France, new to the Palaearclic Region A single male specimen of a species of Platyspathius Yiereck, 191 I was collected in a Malaise Trap operated at Bernac, l-ot-et-Garonne, France by R.R. Askew between 29 June and 31 July 1995. The trap was situated at the edge of ancient mature woodland (Quercus, Juniperus, Cornus, Ace4 Prunas, etc.) and collected several rare insects characteristic of such sites (e.g. Shaw, 1996, Entomologist\ mon. Mag.,132:204). Platyspathius is a relatively small genus, previously known only from the Afrotropical and Indo-Australian regions. As far as is known they are parasitoids of insects living in dead wood. The French specimen runs smoothly to the type species of the genus, P pictipennis Viereck, in Nixon's key (1943, Trans. R. ent. Soc. Lond,93: 173456) by virtue of the distinctive structure of its hind leg and (while being near the edge of the range of variation in its relatively light reddish coloration, rather weak sculpture, and slender build) it agrees well with the seven male specimens of this species from Uganda and Sudan preserved in the BMNH. However, more specimens, and especially females, would be needed to decide its specific identity. The record is brought forward now because the Askews have moved on from their former house at Bernac, substantially reducing the hope that more specimens can be obtained, not only to demonstrate presence of a breeding population but also to enable identification to species. I am grateful to Dick Askew for giving me the ichneumonoid portions of Malaise Trap catches, to Sergei Belokobylskij for his opinion on the specimen, and to Mike Fitton for access to the BMNH collection. The French specimen is deposited in the National Museums of Scotland. M.R. SHAW National Museums of Scotland, Chambers Street, Edinburgh, EHl lJF: January 28th,1998. - -