Dolichovespula albida (Sladen), a Valid Species, Not a Synonym of D. norwegica (Fabricius) (Hymenoptera: Vespidae, Vespinae)

Entomologica Americana 117(3/4):113–116, 2011

DOLICHOVESPULA ALBIDA (SLADEN), A VALID SPECIES, NOT A SYNONYM OF D. NORWEGICA (FABRICIUS) (HYMENOPTERA: VESPIDAE, VESPINAE) JAMES M. CARPENTER1, LIBOR DVORˇA´K2

AND

KURT M. PICKETT3{

1

Division of Invertebrate Zoology, American Museum of Natural History, New York, New York 10024, USA, email: [email protected] 2 Municipal Museum Maria´nske´ La´zneˇ, Goethovo Na´meˇstı´ 11, CZ-353 01 Maria´nske´ La´zneˇ, Czech Republic, email: [email protected] 3 Department of Biology, University of Vermont, 120A Marsh Life Science Building, 109 Carrigan Drive, Burlington, Vermont 05405, USA Abstract—Study of the male genitalia shows that Dolichovespula albida (Sladen) is not conspecific with D. norwegica (Fabricius). Key words: Vespidae, Vespinae, Dolichovespula, Holarctic species.

INTRODUCTION The world checklist of Vespinae by Carpenter and Kojima (1997) recognized five Holarctic species. Carpenter and Glare (2010) have recently shown that one of these, V. vulgaris (Linnaeus), could be diagnosed as separate Nearctic and Palearctic species by the male genitalia. With regard to the remaining four Holarctic species they stated (Carpenter and Glare, 2010: 6) ‘‘In view of our present findings it would seem that reinvestigation of the status of all these forms is warranted.’’ We here present the results of our study of one of those Holarctic species, Dolichovespula norwegica (Fabricius). We show that the male genitalia of Nearctic and Palearctic specimens differ. The name Dolichovespula albida (Sladen) is applicable to the Nearctic species. TAXONOMIC BACKGROUND Vespa norwegica was described by Fabricius (1781) from ‘‘Norwegia.’’ Kirby (1837) described Vespa marginata from Canada (‘‘Taken in the Route from New York, and again in Lat. 65u’’). De Saussure (1854: 135) treated marginata as a synonym of V. arenaria Fabricius, but de Saussure (1857) treated it as distinct. Vespa marginata was first considered a variety of norwegica by du Buysson (1905), who recorded it from Sikkim, Russia (Siberia, Sakhalin Island), Japan, Peru, Canada (Yukon Territory) and the U.S.A. (Alaska). Sladen (1918) described Vespa albida {

Deceased

from ‘‘Alaska.’’ Birula (1930) treated marginata as a subspecies of norwegicia (as norvegica), which he placed in Vespula. Bequaert (1932) synonymized albida with Vespula norwegica var. marginata. Cockerell (1932) pointed out that Vespa marginata was preoccupied by Gmelin (1790), and Bequaert (1935) used albida as the name of the variety. Miller (1958, 1961) again recognized albida as a distinct species, in Vespula, and Dolichovespula albida was the name used in recent keys in North America (Wagner, 1978; Akre et al., 1981). Miller (1958, 1961) showed that albida was distinct from V. norvegicoides (Sladen), which had also been treated as a variety of norwegica by Bequaert - Miller did not refer to Palearctic norwegica. In the meantime, specialists in Eurasia treated albida as a subspecies of Dolichovespula norwegica (Yamane, 1975; Eck, 1981, 1984; Archer, 1989; Pekkarinen, 1995). The concept of a Holarctic species was based on the failure to find features that could differentiate Nearctic and Palearctic taxa. Du Buysson (1905) used the name marginata for black and white specimens, with the nominotypical variety of norwegica being black and yellow. Following Birula (1930), these two color forms were considered to have a transition zone in Siberia. The white or ivory forms also commonly have reddish spots on the first and second metasomal terga in workers and males in both North America and East Asia. Eck (1981) compared color variation, a few morphometric parameters (body size, length of the oculo-malar space, wing and tibia, and form of the ocellar triangle) and number of tyloids

114

ENTOMOLOGICA AMERICANA

Vol. 117(3/4)

of the male antenna, and concluded that D. norwegica and albida were conspecific. Pekkarinen (1995) compared measurement data for seven parameters (oculo-malar length, distance between compound eyes on vertex, postocellar width, distance between posterior ocellus and occipital margin, clypeal width, clypeal length and pronotal width), along with features of the male genitalia (upper inner margin of the paramere and tip of the aedeagus) and likewise concluded that D. norwegica and albida were conspecific. He concluded this despite considering (Pekkarinen, 1995: 66) that ‘‘The the tip of aedeagus is more slender in D. albida.’’ However, his figures do not show much of a difference. MATERIAL AND METHODS Male specimens of D. albida and D. norwegica in the collection of the American Museum of Natural History were dissected, the genital capsule extracted, cleared slightly in warm lactophenol, and examined in glycerin. For photography (Figs. 1–2) the paramere was detached. Males of D. albida and D. norwegica in the collections of Leopoldo Castro, Libor Dvorˇa´k and Bob Jacobson were also dissected, as well as one male D. albida borrowed from the Royal Alberta Museum. Nearctic specimens were from localities in Alaska (N 5 5), Alberta (N 5 1) and the Northwest Territories (N 5 1). Palearctic specimens were from localities in the Czech Republic (N 5 10), Finland (N 5 1), France (N 5 2), Italy (N 5 2), Slovakia (N 5 2), Spain (N 5 2), China (Xinjiang; N 5 2), Mongolia (N 5 2), Russia (Bolshoi Shantar Island in the Sea of Okhotsk; N 5 1) and ‘‘Europe’’ (N 5 1). RESULTS The parameres of North American and Eurasian males are shaped very differently (cf. Figs. 1 vs. 2). In lateral view, North American males have the dorsal margin (left side in Figs. 1–2) distinctly bowed out or bulging; it is close to straight in Eurasian males. The shape difference shows little variability: from almost straight to a little sinuate apically in norwegica, but with the basal part never as wide as in albida, never with such a strongly projecting bulbous aspect. Although previous investigators have compared male genitalia of D. norwegica and D. albida, their published observations have been limited to the

Fig. 1. Paramere of Dolichovespula norwegica in lateral view.

upper inner margin of the paramere and tip of the aedeagus (Eck, 1984; Pekkarinen, 1995). In both D. norwegica and D. albida the upper inner margin of the paramere is produced as a lobe; there is some variation in shape, but not an obvious, consistent difference. The tip of the aedeagus may be more slender in D. albida than in D. norwegica, as concluded by Pekkarinen (1995), but this is at best a subtle difference. The difference in paramere shape has been overlooked. This is immediately obvious, and is a species-specific sort of difference. TAXONOMY The Nearctic and Palearctic forms of Dolichovespula norwegica are readily diagnosed by the male genitalia, hence should be treated as distinct

2011

DOLICHOVESPULA ALBIDA (SLADEN)

115

material of the worker caste collected during the Novara expedition. It was synonymized with Vespa marginata by du Buysson (1905), who saw a specimen from de Saussure. The whereabouts of that specimen, or of any type material from de Saussure’s ‘‘Reise der Novara’’ monograph, are unknown; they are not in the Naturhistorisches Museum Wien, as stated by Bequaert (1935) for peruana, and verified by the senior author, nor at the Muse´um National d’Histoire Naturelle. The type locality of peruana is an obvious error, as noted by Bequaert (1935: 128), but as he stated: ‘‘While this locality ‘Quito’ was undoubtedly due to some error, it is by no means clear how the Novara Expedition could have obtained workers of ‘marginata Kirby,’ a wasp peculiar to Alaska and Labrador. Moreover, de Saussure’s description does not quite agree with the boreal form of norwegica; he does not mention the rufous spot which is almost always present on the sides of the second tergite in the worker of ‘marginata Kirby.’’’

Fig. 2. Paramere lateral view.

of

Dolichovespula

albida

in

species. The name D. albida is available for the Nearctic species, and we so recognize it. Dolichovespula albida (Sladen), revised status Vespa marginata Kirby, 1837: 265, pl. VI fig. 2, female—[Canada] ‘‘Taken in the Route from New York, and again in Lat. 65u’’ (type depository unknown). Junior primary homonym of Vespa marginata Gmelin, 1790. Vespa albida Sladen, 1918: 71, male, female—[U. S. A.] ‘‘Alaska’’ (lectotype male Ottawa). One other detail remains to be discussed: the status of Vespa peruana de Saussure, 1867. This was described from ‘‘Peru altior; Quito’’ based on

Therefore Bequaert (1935) did not use the name peruana when rejecting marginata due to homonymy, instead using albida, whose identity he regarded as ‘‘beyond dispute.’’ Of course, du Buysson’s concept of marginata was simply white-marked specimens of his norwegica, and as already noted he included specimens from several localities in the Palearctic. Bequaert (1932: 121) considered that the specimens from Siberia and Sakhalin were ‘‘undoubtedly’’ V. norwegica pacifica Birula. And it is true that workers of D. albida generally have reddish markings on the second metasomal tergum. Given these considerations, Vespa peruana is better left unrecognized, and like Bequaert we use D. albida (Sladen) as the substitute name for Vespa marginata Kirby. DISCUSSION The number of Holarctic species in Vespinae continues to decline. With our present results, there are just three recognized: Vespula austriaca, V. rufa and D. adulterina. Pekkarinen (1995) recognized six. Yet for one of these, D. norvegicoides, he confirmed a difference in the sculpture between the gena and clypeus that allows diagnosis of two species, the Nearctic D. norvegicoides and the Palearctic D. pacifica. Carpenter and

116

ENTOMOLOGICA AMERICANA

Glare (2010) demonstrated diagnostic differences in the male genitalia between the Nearctic V. alascensis and the Palearctic V. vulgaris, and now we have done so for the Nearctic D. albida and the Palearctic D. norwegica. Two of the three remaining Holarctic species are social parasites, inquilines, which have different hosts in the Nearctic and Palearctic Regions. This difference in itself may indicate that a single species is not involved in each case. The same sort of study of the male genitalia undertaken here and by Carpenter and Glare (2010) should be pursued. ACKNOWLEDGMENTS We thank Matthias Buck, Leopoldo Castro and Bob Jacobson for supplying specimens, Adrien Perrard and Dominique Zimmermann for information, and Leopoldo Castro for comments on the manuscript. This work was partially supported by National Science Foundation Grant DEB-0843505 to JMC and KMP.

LITERATURE CITED Akre, R. D., A. Greene, J. F. MacDonald, P. J. Landolt and H. G. Davis. 1981. The yellowjackets of America North of Mexico. U.S.D.A. Agric. Handb. 552: 1–102. Archer, M. E. 1989. A key to the world species of the Vespinae (Hymenoptera). Res. Monogr. College of Ripon & York St. John 2. York. Bequaert, J. 1932. A tentative synopsis of the hornets and yellow-jackets (Vespinae; Hymenoptera) of America. Entomol. Am. (n. s.) 12(2): 71–138. Bequaert, J. 1935. Additions and corrections to the revision of North American Vespinae (Entomologica Americana, 1931). Bull. Brooklyn Entomol. Soc. 30: 119–124. ¨ ber die russischen Wespen und ihre Birula, A. 1930. U geographische Verbreitung. (Dritter Beitrag). Ann. Mus. Zool. Acad. Sci. U.R.S.S. 31(2): 291–339. Buysson, R. du. 1905 (1904). Monographie des gueˆpes ou Vespa (suite*). Ann. Soc. Entomol. Fr. 73(3): 485–556, 565–634. Carpenter, J. M. and T. R. Glare. 2010. Misidentification of Vespula alascensis as V. vulgaris in North America (Hymenoptera: Vespidae; Vespinae). Am. Mus. Novitat. 3690: 1–7.

Vol. 117(3/4)

Carpenter, J. M. and J. Kojima. 1997. Checklist of the species in the subfamily Vespinae (Insecta: Hymenoptera: Vespidae). Nat. Hist. Bull. Ibaraki Univ. 1: 51–92. Cockerell, T. D. A. 1932. The Vespula marginata of Kirby. Pan-Pacific Entomol. 8(4): 161. Eck, R. 1981. Zur Verbreitung und Variabilita¨t von Dolichovespula norwegica (Hymenoptera, Vespidae). Entomol. Abh. Staat. Mus. Tierk. Dresden 44: 133–152. Eck, R. 1984. Bestimmungsschlu¨ssel fu¨r die Arten der Gattung Dolichovespula ROHWER, 1916 (Hymenoptera; Vespidae). Entomol. Abh. Staat. Mus. Tierk. Dresden 48(4): 35–44. Fabricius, J. C. 1781. Species Insectorum. I. C. E. Bohni, Hamburg. Gmelin, J. F. 1790. Caroli a Linne´ Systema naturae. ed. XIIIa, aucta & reformata. I, pars V: 2225–3020 Lipsiae. Kirby, W. 1837. Fauna Boreali-Americana. Part IV. The Insects, Norwich, xxxix + 325 pp., 8 pls. Miller, C. D. F. 1958. Distributional and nomenclatorial problems in some forms of Vespula in North America (Hymenoptera: Vespidae). Proc. Tenth Intl. Congr. Entomol., Montreal, 1956, 1: 257–264. Miller, C. D. F. 1961. Taxonomy and distribution of Nearctic Vespula. Can. Entomol. 93, Suppl. 22: 1–52. Pekkarinen, A. 1995. Geographic variation and taxonomy of the species of Dolichovespula in the boreal zone of the Holarctic region (Hymenoptera, Vespinae). Acta Zool. Fenn. 199: 61–70. Saussure, H. de. 1853–58. Monographie des Gueˆpes Sociales ou de la Tribu des Vespiens. Masson, Paris, and J. Cherbuliez, Gene`ve. Saussure, H. de. 1857. Bemerkungen u¨ber die Gattung Vespa, besonders u¨ber die amerikanischen Arten. Stettin. Ent. Ztg. 18: 114–117. Saussure, H. de. 1867. Hymenoptera. Reise der Oesterreichischen Fregatte Novara um die Erde in den Jahren 1857–1859. Zool. Theil II. pt 1: 1–38, pls. I–IV., Wien. Sladen, F. W. L. 1918. The genus Vespa in Canada. Ottawa Nat. 32: 71–72. Wagner, R. E. 1978. The genus Dolichovespula and an addition to its known species of North America (Hymenoptera: Vespidae). Pan-Pacif. Entomol. 54: 131–142. Yamane, S. 1975. Taxonomic notes on the subgenus Boreovespula Blu¨thgen (Hymenoptera, Vespidae) of Japan, with notes on specimens from Sakhalin. Kontyuˆ 43(3): 343–355.