MAGAZINE THE TAXONOMY OF VESPA CRABRO L. AND ENTOMOLOGIST'S MONTHLY I57 V. DYBOWSKII ANDRE (HYM., VESPINAE) BY MICHAEL E. ARCHER Vespa crabro Linnaeus, 1758 and V. dybowskii Andr6, 1884 are probably a monophyletic group united by the derived character state of the absence of projections from the dorsal part of the male digitus. In a cladistic analysls of Vespa (Archer, unpublished) these two species with the V. tropica species group (I/. orientalis Linnaeus, 177I, V. mandarinia Smiih, 1852, V. soror du Buysson, 1905, V. ducalis Smith, 1852, V. philippinensis de Saussure, 1854, V. tropica (Linnaeus' 1758)) form a monophyletic group characterised by the following derived character state: The apical margin of the seventh gastral sternum of the male has a single shallow emargination, less than a semicircle, and the lateral margins are rounded. This paper reports on the taxonomy and distribution of the two species and also examines the colour subspecies of V. crabro showing polytopic while others show geographical overlap. Contirat some are -suggested the subspecies nomenclature be replaced by sequently it is informal names. It has previously been established (Archer,l99I) that V. walkeri dt Buysson, 1905 is a synonym of V. dybowskii. Characters that separate the species The males of V. crabro and V. dybowskii can be separated from other Vespa species by a genitalia character indicated above. Males of V. crabro can be separated from those of V. dybowskii and other Vespa species by the presence of hairs on the apex of the aedeagus (fig. 1A). Males of V. dybowskii can be separated from those of V. crabro and other Vespa species by the characteristic diamond-shape of, and deep triangulaiemargination in, the apex of the aedeagus (fig. 18). Iniontrast tothe males, the females of V. crabro andV. dybowskii can only be separated from the other species of Vespa by the use of many character states, all of which are primitive or assumed to be primitive: 2. Lateral apical margin of the clypeus semicircular in clypeus without a median tooth between the laterally produced margins. 4. First gastral tergum as seen from above, less than half as long as wide. 5. The posterior ocelli are closer to each other than to the eyes. 6. The clypeus in lateral view appears only weakly convex. 7. Laleral punctures on the second gastral tergum generally with the distances between the punctures greater than the puncture diameters. 8. To separate the females from 7. orientalis the sixth gastral tergum is either yellow (V. crabro) or the third gastral tergum is dark brown or black with at most a very narrow apical yellow band (V. dybowskii). 1. Pretegula carina complete. shape. 3. Apical margin of the Females of V. crabro and V. dybowskii can be separated from each other by colour characteristics. On V. crabro some or all of the second, 31st July, 1992 Vol. 128 (1992) ENTOMOLOGIST'S MONTHLY MAGAZINE Fig. 1. - Dorsal view of the male genitalia: A, Vespa crabro; B, Vespa dybowskii. third, fourth and fifth gastral terga are usually with broad yellow apical bands, but if the bands are narrow then the vertex is yellow or orangeyellow. By contrast on V. dybowsftii the second, third, fourth and fifth gastral terga are dark brown or black with at most a very narrow apical yellow band and the vertex is reddish-brown or dark brown. These colour characteristics will also separate the males of the two species. Vespa crabro The colour forms of V. crabro were last treated by Bequaert (1931) who listed nine colour variations (Table 1). Previously du Buysson (1904-1905) and Birula (1925) had made important contributions. Bequaert (1931) treated the colour forms as varieties, as did Guighlia (1972), but usually the colour forms are considered subspecies (Birula, 1925; van der Vecht, 1959; Edwards, 1980; Matsuura & Yamane, 1e84). After a study of many specimens borrowed from national museums (Archer, 1989) and individuals, particularly N.V. Kurzenko (Ussuriland, U.S.S.R.) and Jeong Gie-Joon (Korea), the following opinions can be given. I. V. crabro crabro is polytopic, as first noted by Birula 1925, with a western distribu- tion in east Europe and an eastern distribution from Lake Baikal to Sakhalin 2. and northern Manchuria. V. crabro germana Christ, 1791 is polytopic, also first noted by Birula 1925, with a western distribution in west Europe and an eastern distribution in Ussuriland of the ENTOMOLOGIST'S MONTHLY 3. MAGAZINE 159 U.S.S.R. Specimens seen from Ussuriland and Europe are similar except possibly the head of the Ussuriland specimens is not quite so red. Intermediates are common between V. crabro birulai Bequaert, 1931 and V. crabro crabroniformis Smith, 1852, as first noted by Bequaert (1931). Intermediates also exist between V. crabro oberthuri du Buysson. 1902 and V. crabro crabroniformis and both colour forms have been found at the same locality. V. crabro birulai andV. crabro oberthuri would seem to be colour variations of V. crabro crabroniformis and can hardly be maintained. 4. Some specimens of V. crabro gribodoi Bequaert, 1931 and V. crabro crabroniformis can only be separated with difficulty with perhaps the pronotum being more extensively red on V. crabro gribodoi. Bequaert (1931) noted the similarity between V. crabro gribodoi and V. crabro birulai and separated the two colour forms by the yellow blotches on the second gastral tergum of V. crabro birulai. In practice this difference is not always present. In many ways a polytopic colour form is represented in the west by V. crabro gribodoi from England and Wales and in the east by V. crabro crabroniformis from Himalayan, southern and eastern China and Taiwan. 5. V. crabro caspicaPerez,l9l0 is found in Transcaucasia and northern Iran and differs from the west European V. crabro gerrnana by the presence of a transverse dirty to pale yellow blotch on the second gastral tergum. However, specimens from the geographical range of V. crabro caspica may lack the second gastral tergal blotch while west European specimens from Corsica, Italy and Sicily may show the second gastral tergal blotch. These observations were also noted by Birula (1925) for V. crabro caspica and by Guighlia (1972) concerning V. crabro germana. It is probable thaL V. crabro caspica is a colour variety of V. crabro germana although the two colour forms still differ in that the third gastral tergum of V. crabro caspica is largely dark coloured while that of V. crabro germana is largely yellow coloured. 6. V. crabro flavofasciata Cameron, 1903 is usually considered to be the characteristic colour form of Japan. However, specimens indistinguishable from V. crabro flavofasciata have been found in Korea and Ussuriland of the U.S.S.R. Giordana-Soika (1976, 1982) also found V. crabro flavofasciata in Korea. Thus V. crabro flavofasciata is not restricted to Japan. THE TNFORMAL AND FORMAL (SUBSPECIES) COLOUR FORM 1. - YESP, CRABRO NAMES FOR TABLE Informal Names Subspecies Subspecies (this paper) (based on Bequaert, 1931) altaica Southern Siberian East European Baikal to Manchurian West European Far east Asian altaica crabro crabro germana germana British gribodoi crabroniformis gribodoi birulai, crab roniformis, oberthuri flavofasciata flavofasciata Chinese Japanese crabro, caspica crabro Sermana The above opinions would reduce the number of colour subspecies from nine to six (Table 1). However, the subspecies V. crabro crabro, V. crabro germana and V. crabro gribodoi with part of. V. crabro crabroniformis would be polytopic. With the exception of V. crabro gribodoi, intermediate coloured specimens are found in the geographi- 160 ENTOMOLOGIST'S MONTHLY MAGAZINE cal areas between the other colour forms. Because of these complications it is suggested the subspecies nomenclature be replaced by 'informal' names following the arguments of Wilson & Brown (1953). These informal names are given in Table 1. Separation of the informal colour forms of V. crabro The eight informal colour forms can be separated by reference mainly to the colour of the vertex and gena (black, orange, yellow), ocellar region (black, not black), pronotum (red or yellow in part), mesoscutum (black or red in part), the third gastral tergum (largely yellow or largely dark coloured) and the presence or absence of light coloured blotches on the second gastral tergum. The southern Siberian colour form is characterised by a yellow vertex with a black or red spot on the gena of the female, black ocellar region, pronoturn and mesoscutum and the third gastral tergum about equally yellow and dark coloured. To the west of the southern Siberian colour form is found the east E,uropean colour form on which the vertex and gena become red or sometimes the vertex is orange, the pronotum is often red and the third gastral tergum is largely yellow. Further west is found the west European colour form on which the ocellar region is usually red, sometimes with traces of black, the pronotum and mesoscutum showing red and the third gastral tergum largely yellow except in the near eastern region where it is mainly dark, the second gastral tergum may have a transverse yellow basal blotch varying in colour intensity but particularly distinct in southern and near eastern Europe. In England and Wales is found the British colour form which differs from the near eastern-west European colour form in having a yellow vertex and gena, the third gastral tergum always largely yellow and the basal yellow blotch on the second gastral tergum absent or faint. To the east of the southern Siberian colour form is found the BaikalManchurian colour form which is similar to the east European colour form. Further east is found the far east Asian colour form which is similar to the west European colour form except possibly the head is not quite so extensively red, the red colour being replaced apically by orange. The Chinese colour form is similar to the British colour form except in Taiwan and the north and east of its distribution where the vertex and gena can be orange, the pronotum often red and the second gastral yellow basal blotches more distinct. The Japanese colour form has an orange vertex and gena, usually a black ocellar region, red markings on the pronotum, black mesoscutum and the third gastral tergum either mainly dark coloured or about equally yellow and dark coloured. The geographical distributions of these eight informal colour forms are given in Table 2. ENTOMOLOGIST'S MONTHLY MAGAZINE t67 -a :J b q \v li a o B () F I c.i bb t! r62 ENTOMOLOGIST'S MONTHLY MAGAZINE Vespa dybowskii It has previously been established (Archer, 199I) that the colour differences between V. dybowskii and V. walkeri used by du Buysson (1904, 1905) were not reliable and V. walkeri is now considered a synonym of V. dybowskii. Distribution of V. crabro and V. dybowskii The distribution of V. crabro (tig, 2) is throughout Europe to about 64" north, U.S.S.R. to about 56'north and east to Sakhalin; western Turkey, northern Iran, western Mongolia, north-eastern central, eastern and southern China, Korea and Japan. The west European colour form has been introduced into eastern U.S.A. and south eastern Canada. "o*u Fig. 3. - The world distribution of Vespa dybowskii. ENTOMOLOGIST'S MONTHLY MAGAZINE 163 The distribution of V. dybowskii (tig.3) is north Burma, eastern Tibet, central and eastern China, U.S.S.R. (Ussuriland) and Japan. The range of V. dybowskii is probably entirely within the range of V. crabro. COLOUR FORMS OF VESPA CRABRO WITH THEIR TABLE 2. -THE NAMES AND GEOGRAPHICAL DISTRIBUTIONS INFORMAL 1. Southern Siberian South western and central Siberia from the Ural Mountains to Lake Baikal and southwards to Altai Mountains and western Mongolia. Scandinavia to about 64" north. Central and east European 2. East European U.S.S.R. 3. Baikal to Manchurian U.S.S.R. east of the Yenisei river to about 56' north. Sakhalin. Manchuria. France to west European U.S.S.R. Portugal to western Turkey. Transcaucasia and northern Iran by the Caspian Sea. Introduced to eastern U.S.A. and south eastern Canada. 4. West European 5. Far Eastern Ussuriland of the U.S.S.R. 6. British England and Wales. 7. Chinese Himalayan, southern and eastern China. Taiwan. 8. Japanese Japan. Korea. Ussuriland of U.S.S.R. ACKNOWLEDGEMENT AND DEDICATION dedicate this paper to the late Dr Ian Yarrow (Obituary, 1990, Entomologist's mon. Mog., 126 253-257) who, before he died, kindly gave me his extensive notes on the Vespinae which included some analysis of the colour forms of Vespa crabro. I REFERENCES Archer, M.E., 1989, A Key to the World Species of the Vespinae (Hymenoptera). Parts 1 &2. Res. Monog. College Ripon &York StJohn, no.2'. 1-41, figs 1-72;7991, The number of species that can be recognised within the genus Vespa (Hy-., Vespinae), Entomologist's mon. Mag., 127:167-164. Bequaert, J., 1931, The colour forms of the common hornet, Vespa crabro Linnaeus, Konowia l0: 101-109. Birula, A., 1925, Ueber die russischen Wespen und ihre geographische Verbreitung, Arch. Naturgesch. 90A(12): 88-102. Buysson, R. du, 1904-5. Monographie des gudpes ou Vespa, Annls Soc. ent. Fr., (1904) 72:260-288; (1905) 73 486-556,565-634. Edwards, R., 1980, Social Wasps. Their biology and control, The Rentokil Library, East Grinstead, England. GiordaniSoika, A,,1.976, Vespidi ed Eumenidi raccolti in Corea (Hymenoptera). Ann. Hist. Nat. Mus. Natl. Hung., 68:287-293;7982, Vespidi ed Eumenidi raccolti in Corea (Hymenoptera) II. Folia Ent. Hung., 43:39-41. Guiglia, D.,1972, Les Guepes Sociales (Hymenoptera: Vespidae) d'Europe Occidentale et Septentrionale. Masson, Paris. Matsuura, M. & Yamane, Sk., 1990, Biology of the Vespine Wasps. Springer-Verlag, Berlin. Vecht, J. van der,'1959, Notes on Oriental Vespidae, including some species from China and Japan (Hymenoptera, Vespidae). Zool. Meded., 36:205-232. Wilson, E.O. & Brown, W.L., 1953, The subspecies concept and its taxonomic appiication. Syst. Zool.,2:99-111- The College of Ripon & York St John, Lord Mayor's Walk, York YO3 7EX. September 6th, 1991