The Vespinae of North America (Vespidae, Hymenoptera)

The species of paper wasps in the tribe Vespini, family Vespidae from America North of Mexico are reviewed, including a new identification key to the genera and species, complete synonymy, distribution and biology. This fauna includes six species of Dolichovespula Rohwer, three species of Vespa Linnaeus and 13 species of Vespula Thomson. No Holarctic species are recognized, with the result that Dolichovespula arctica (Rohwer) and Vespula intermedia (du Buysson) are again recognized as species, while Vespula infernalis (de Saussure) is given new status as a species.

The Vespinae of North America (Vespidae, Hymenoptera)

Introduction
Vespinae, or the yellow jackets and hornets, are among the most recognizable wasps in North America.All of the species are either social or are social parasites of other congeners.They construct their nests out of a mixture of plant fibers and salivary secretions, and the nests can range from baseball-sized, with a few thousand cells, to nests with hundreds of thousands of cells.Nests are generally annual but a few species will develop large perennial nests in warm climates.A number of species, such as Vespula alascensis and Vespula germanica, are considered to be pests because of their willingness to build nests in structures and to scavenge a variety of food materials other than live insects, which brings them into frequent contact with humans and because the majority of sting-caused deaths are attributable to yellow jackets.
There have been a number of changes in the North American social vespine fauna since the widely used key published by Akre et al. (1981).Two exotic Vespa affinis Linnaeus and simillima (Smith) have been introduced into North America.One new species has been described, Dolichovespula alpicola Eck (Eck, 1984).In addition, there have been a number of other taxonomic changes based on reevaluation of specimens and study of male genitalia.The most notable of these is the discovery that two species previously considered Holarctic are not in fact found in both North America and the Palearctic.Vespula vulgaris (Linnaeus) and Dolichovespula norwegica (Fabricius) are not conspecific with the species in North America, necessitating name changes for these taxa (Carpenter et al. 2011, Carpenter andGlare 2010): Vespula alascensis (Packard) and Dolichovespula albida (Sladen) are valid species.There are morphological differences now known between North American and Eurasian specimens of another supposed Holarctic species, Vespula austriaca, and the North American species should therefore be known as Vespula infernalis (de Saussure).Another of the supposed Holarctic species, Dolichovespula adulterina (du Buysson) is a social parasite, which does not have the same hosts in Eurasia and North America, hence is probably not conspecific in the Old World and New, and D. arctica (Rohwer) should be treated as a species.Most likely no Holarctic species should be recognized, with the exception of the introduced Vespula germanica, therefore Vespula intermedia (du Buysson) should be used instead of V. rufa (Linnaeus).
Given these changes, and the difficulty of using many of the existing keys, which rely heavily on coloration, we have developed a new key incorporating these taxonomic changes and attempting to place more emphasis on structural features, such as male genitalia.The key by Akre et al. (1981) was also incomplete, not including socially parasitic species, as it was based only on workers.The key below includes all species and castes.

Materials and methods
North America as construed here is America north of Mexico.Thus two species recently described from central Mexico (Vespula inexspectata Eck; Eck 1994) and Guatemala (V.akrei Landolt; Landolt et al. 2010) are not included in the key, nor is Vespa orientalis Linnaeus, recently recorded from Mexico (Dvořák 2006).
Specimens used in the development of this key were from the collections of the American Museum of Natural History, New York; Bohart Museum of Entomology, University of California, Davis; California State Collection of Arthropods, California

1
Head in dorsal view greatly expanded behind eyes, postocular distance more than twice as broad as distance between hindocelli (as in Fig. 5) Distribution.This species occurs in the Hudsonian Zone of North America from Alaska to Maine.
Biology.Dolichovespula albida was split again from D. norwegica (Fabricius) by Carpenter et al. (2012).The nests are generally small and may be subterranean (Bequaert 1932), although Yamane et al. (1980)  Distribution.This is a northern boreal species found in western North America from Alaska to Alberta, and extending as far south as Arizona and New Mexico along the Rocky Mountains (R. Jacobson personal communication).Biology.Dolichovespula alpicola is generally found in mixed hardwood-conifer forests in mountainous regions.There is little information on the nesting biology in this species.Eck (1984) attributed the species to Wagner in a key, thus validly publishing it.Eck (1987)  Distribution.It occurs as far south as California, Arizona and Georgia in North America.Pale markings can be yellow or whitish in D. arctica.Biology.This species is an obligatory social parasite of Dolichovespula arenaria (Wheeler and Taylor 1921) and D. alpicola (Wagner 1978).The name is a replacement for a misidentification of Vespa borealis Kirby by Lewis (1897) Distribution.This species is abundant throughout boreal North America.
Biology.Dolichovespula arenaria feeds on live prey but will occasionally visit carrion.It builds aerial nests, like D. maculata, but its nests are usually built in more sheltered sites, such as within bushes, trees, on houses and outbuildings, and rarely even under rocks.
Distribution.The species occurs throughout North America.Biology.This is the only Dolichovespula in North America with the anterior part of the metasoma completely black, and the pale markings are always whitish.It builds aerial nests, which are usually found in exposed places such as hanging from tree branches.These wasps usually feed on live prey, particularly spiders and flies.Biology.Nests are generally built in above-ground cavities, wall voids, hollow trees, and even in abandoned honey bee hives.Nests are large in size because of the size of the wasps but generally contain only a few thousand cells.These wasps feed on live insects, including honey bees near hives (Akre and Davis 1978) and have been recorded girdling saplings to feed on sap in the spring (Bromley 1931).
Distribution.This East Asian species was introduced into British Columbia, Canada but has apparently not become established.We have included it in this review because of the potential for a reintroduction.
Biology.Nests are built in a wide variety of situations including in bushes, underground cavities, hollow trees, under eaves, in attics, on rock walls, and in wall voids.As with the other Vespa species affinis is predatory on other insects, including honey bees.

Distribution. Widespread in North America
Biology.Vespula alascensis nests are usually built in subterranean cavities or in structures.The nests can be huge and may become perennial in warmer climates.This species feeds on live prey but will scavenge any source of protein or sugar.It has been introduced into Hawaii, but is not now established (Carpenter 2008, under vulgaris).
This North American species was usually known as Vespula vulgaris (Linnaeus), based on the similarity between the American and European populations.Carpenter and Glare (2010) discovered that the European and American populations are not conspecific based on features of the male genitalia and mitochondrial DNA evidence.Biology.This predatory species preys on live insects and usually nests in cavities, using sites in logs, wall voids and rodent burrows.Distribution.The distribution of Vespula austriaca closely resembles that of acadica, occurring in subarctic Alaska and Canada, and southward in the western mountain ranges.Biology.This is an obligatory social parasite of Vespula acadica (Reed et al. 1979).This species has been known as Vespula austriaca, a Palearctic species, since Bequaert (1916), but there are sculptural differences in queens between Palearctic and Nearctic specimens.Distribution.These wasps occur in the far northern Nearctic Region.

Vespula flavopilosa
Biology.The biology of North American intermedia has not been studied.Vespula rufa in the Palearctic usually nests below ground, in cavities or under eaves.They prey on live insects.
Although treated as a synonym of Vespula rufa (Linnaeus), which is thus a Holarctic species, the North American population is probably distinct and should be treated as a separate species, V. intermedia (du Buysson).It differs in coloration from V. rufa, with the pale markings being whitish and the metasomal terga I-II with reddish markings.The pale markings are either yellow in V. rufa (European specimens), or the terga lack reddish markings (Eastern Palearctic), or there are whitish spots in addition to reddish markings (Eastern Palearctic).Distribution.This species is the most common Vespula occurring east of the 100 th meridian.
Biology.It builds subterranean nests in a wide variety of situations, even in abandoned vehicles.These yellow jackets prey on live insects and are also scavengers of sources of protein and sugar.
Distribution.This is the most abundant pest species of Vespula on the West Coast and much of the interior west of North America.Biology.Nests are usually built in cavities, which may be in the ground or in structures, such as attics, wall voids and even basements.The nests can become huge and often become perennial in warmer climates.These wasps are general scavengers, and will feed on live prey or any other source of protein or sugar including garbage.It is adventive in Hawaii.Distribution.This is an eastern species, occurring east of the 100 th meridian and south to Honduras.Biology.It is a facultative social parasite of Vespula maculifrons.The two striped scutum is a distinctive feature of both Vespula squamosa and Vespula sulphurea.Queens of squamosa are quite different in color from workers and males, with their extensive orange-brown coloration, particularly on the metasoma.
Distribution.Vespula sulphurea is abundant in mid elevation and wildland areas in western North America.
Biology.This is the yellowest of the North American species, with a two yellowstriped scutum much like that seen in Vespula squamosa, but sulphurea occurs west of the 100 th meridian.They build small, subterranean nests, generally feed on live prey; in unusual circumstances they might scavenge food.
Distribution.Vespula vidua occurs in the Transition and Upper Austral Zones of eastern North America.
Biology.Most nests are subterranean but Vpl.vidua will also build nests in hollow logs.

Figure 1 .
Figure 1.Vespula germanica worker a side view b dorsal view; Vespula pensylvanica worker c side view d dorsal view.
reported an aerial nest just above ground.
later published a fuller description and designated a lectotype.
Distribution.The central European color form of this Palearctic species was introduced into the New York area in the mid-1800's (de Saussure 1898).It now occurs throughout the eastern United States, east of the Mississippi River.

Vespula infernalis (de Saussure), stat. n.
(Menke and Snelling 1975)s in the eastern United States as far south as Georgia (R.Jacobson, personal communication).Biology.It builds subterranean nests.This species feeds on live insects, as well as other sources of sugar and protein, and is a scavenger much like Vespula germanica, V. pensylvanica and V. alascensis.Vespa maculata Scopoli 1763:312.Type destroyed; "Carnioliae" (Slovenia).Nec Vespa maculata Linnaeus 1763.?Vespa macularis Olivier 1792:695.Unjustified emendation of Vespa maculata Scopoli.Vespa germanica Fabricius 1793:256.Type unknown; "Kiliae" (Germany) (repository unknown).Vespula germanica has been unintentionally introduced into temperate regions worldwide.It apparently first appeared in Montreal in the 1960's and other parts of eastern North America in the 1970's, although there is a record of the species collected in Ithaca, New York in 1891(Menke and Snelling 1975).The species reached California by 1989.Biology.The nests are usually built in structures or less commonly in the ground.Nests can be huge and may become perennial in warmer climates.These wasps feed on live prey or scavenge any source of protein or sugar.Vespa infernalis deSaussure 1854:139.Holotype female; "L'Amérique du Nord, Philadelphie (TURIN?).Vespa tripunctata Packard 1870:26, pl.II fig.11.Holotype female: "Kutleet", USA (repository unknown).Nec Vespa tripunctata Fabricius 1787 and Vespa tripunctata Schenck 1861.