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Superfamily VESPOIDEA
Family VESPIDAE

The wasps belonging to this family are all social species, the colonies consisting of one or more
fertile queens, workers which are frequently numerous and usually infertile, and, at times, males
and new queens.

Taxonomy: Lewis, 1897. Amer. Ent. Soc, Trans. 24: 169-192, 1 pi. (N. Amer. spp.) — Bohart
and Bechtel, 1957. Calif. Ins. Survey, Bui. 4: 73-102, 8 pis., 34 figs., 7 maps (Calif, spp.).

Biology: Rau, 1929. Canad. Ent. 61: 219-221 (cocoon spinning). —Jeanne, 1975. Quart. Rev.
Biol. 50: 267-287, 9 figs, (adaptiveness of nest architecture).

Morphology: van der Vecht, 1968. K. Nederland. Akad. van Wetensch., Proc, Ser. C, 71:
411-422, 5 figs, (terminal gastral sternite, female, worker).

Subfamily POLISTINAE

These are commonly called paper wasps in North America. The nests consist of a single comb
which is not enclosed in a paper envelope. In most of the Temperate Zone species the nests are
annual, each being founded by one or more overwintering females. One of the females becomes
dominant and is the only one to lay eggs.

The normal larval food consists of dismembered caterpillars. Some species of Polistes exert
considerable predator pressure on economically important insects such as the tobacco hornworm
and cotton bollworm. In an attempt to utilize the wasps as a biological control agent, shelters are
sometimes placed in cultivated fields to afford nesting sites to founding females.

Taxonomy: Yoshikawa, 1962. Jour. Biol. 13: 19-43, 1 map (list of world spp., distribution).

Biology: Yoshikawa, 1962. Jour. Biol. 13: 45-64 (evolution of sociality).

Genus POLISTES Latreille

Polistes Latreille, 1802. Hist. Nat. Crust. Ins., v. 3, p. 363.

Type-species: Vespa gallica Linnaeus. Desig. by Blanchard, 1840.

Richard's (1973) reclassification of the subgenera of Polistes, listed below in Taxonomy, was
received too late to assign the North American species to subgenera in the format used el-
sewhere in the catalog for subgenera. Our species are assigned to the following subgenera.

Polistes subg. Fuscopolistes Richards, 1973, pp. 95, 100 (type- species, Vespa fuscata




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Fabricius, orig. desig.)— apachus, bellicosus, Carolina, dorsalis, flavus, fuscatus, metricus and
perplexus.

Polistes subg. Palisotius Richards, 1973, pp. 95, 100 (type-species, Polistes major Beauvois,
orig. desig.)— ?Hajor.

Polistes subg. Onerarius Richards, 1973, pp. 94, 101 (type-species, Vespa camifex Fabricius,
orig. desig.)— car>n/ex.

Polistes subg. Polistarchus Richards, 1973, pp. 94, 101 (type-species, Vespa canadensis Lin-
naeus, orig. desig.)— awwM/aWs, canadensis, exclamans and iyistabilis.

Polistes subg. Epicnemius Richards, 1973, pp. 96, 102 (type-species, Polistes bicolor Lepeleti-
er, orig. desig.)-paci/i!CMs.

The typical subgenus does not occur in North America.

In the extensive series of papers listed below, Rau consistently used two misidentifications:

Polistes pallipes
***authority mismatch
for the taxon now recognized as metricus, and variatiis for the taxon now recognized as
typical fuscatus; he also used rubiginosus for Carolina, a senior synonym.

Revision: Bequaert, 1937. Arch. Inst. Biol. Veg., Rio de Janeiro 3: 171-205 (New World taxa
of Gallicus and Bicolor species groups). —Bequaert, 1940. N. Y. Ent. Soc, Joiir. 48: 1-31
(N. Amer. taxa). —Bequaert, 1943. Ent. Venezolana, Bol. 2: 107-124 (New World vars. of

Polistes canadensis
***authority mismatch
(L.)). — Snelling, 1955. Amer. Mus. Novitates 1701: 1-9 (subspp. oi major
Beauv. and exclamans Vier.).

Taxonomy: Enteman, 1904. Carnegie Inst. Wash., Pub. 19: 1-88, 6 pis., 27 figs, (coloration).
—Rau, 1942. Ent. Soc. Amer., Ann. 35: 335-338 (behavioral characters of Carolina, f.
fuscatus, metricus). —Richards, 1973. Rev. Ent. Brasil. 17: 85-103, 12 figs, (classification
of subgenera). —Snelling, 1974. Ent. Soc. Wash., Proc. 76: 476-479 (status of some N.
Amer. taxa).

Biology: Rau, 1928. Biol. Bui. 54: 503-519 (honey-gathering by annularis, f fuscatus,
metricus). —Rau, 1928. Psyche 35: 147-150, 4 figs, (modification of nests by Carolina,
annularis, metricus). —Rau, 1928. Brooklyn Ent. Soc, Bui. 23: 230-235 (hibernation,
colony founding by annularis, metricus). — Duncan, 1928. Pan-Pacific Ent. 5: 90 (building
material). —Rau, 1928. Psyche 35: 151-152 (reconstruction of destroyed nests by/.
fuscatus, metricus). —Rau, 1929. Ecology 10: 191-200, 4 figs, (nesting habitat of
annularis, Carolina, f. fuscatus, metricus). —Rau, 1930. Compar. Psychol., Jour. 10:
267-286 (interspecific animosity and tolerance in workers o{ f. fuscatus, metriciis,
annularis). —Rau, 1930. Ent. Soc. Amer., Ann. 23: 461-466 (hibernation behavior of
Carolina, f fuscatus, metricus, annularis). —Rau, 1931. Ecology 12: 690-693 (use of water
by anmdaris, metricus, f. fuscatus, Carolina). —Rau, 1931. Ent. Soc. Amer., Ann. 24:
515-518 (hibernation behavior of annularis, metricus). —Rau, 1939. Compar. Psychol.,
Jour. 27: 259-269 (inter- and intraspecific animosity and tolerance of female annularis, f.
fuscatus, metricus). —Rau, 1939. Ecology 20: 439-442 (population and caste studies of
metricus, f. fuscatus, annularis). —Rau, 1941. Ent. Soc. Amer., Ann. 34: 355-366
(parasites of Carolina, anmdaris, metricus, f. fuscatus). — Rau, 1941. Ent. Soc. Amer.,
Ann. 34: 580-584 (swarming oi annularis, f. fuscatus, metricus). —Rau, 1946. Ent. Soc.
Amer., Ann. 39: 11-27 (nest and cell size off. fuscatus, metricus). — Rabb and Lawson,
1957. Econ. Ent., Jour. 50: 778-784, 2 figs, (predation on tobacco hornworm in N. C. by e.
exclamans, f fuscatus). —Rabb, 1960. Ent. Soc. Amer., Ann. 53: 111-121 (nest founding,
life history, adult and larval food, prey, foraging, parasites in N. C. in f. fuscatus,
annularis, e. exclamans, metricus, d. dorsalis, Carolina). —West, 1967. Science 157:
1584-1585 (female dominancy hierarchies in canadensis, fuscatus). —Nelson, 1968. Ent.
Soc. Amer., Ann. 61: 1528-1539, 3 figs, (parasites, symbionts). — Eberhard, 1969. Mich.
Univ., Mus. ZooL, Misc. Pub. 140: 1-101, 23 figs, (social biology of f fuscatus, canadensis
erythrocephalus Latr., c. infiiscatus Lep.). — Kirkton, 1970. Tall Timbers Conf., Proc. 2:
243-245 (population management in Ark.). — Gillaspy, 1971. Ent. Soc. Amer., Ann. 64:
1357-1361, 4 figs, (population management in Tex., Calif.). —Gillaspy, 1973. Amer. Midland
Nat. 90: 1-12, 5 figs, (behavior of metricus, annularis, major, bellicosus, exclamans,
apachus, Carolina in artificial nesting sites).

Polistes annularis (Linnaeus)
Conn, to Fla., west to S. Dak. and central Tex. Ecology: Nests

suspended from twigs and branches of shrubs and trees in exposed situations, and




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occasionally in sheltered sites on buildings. Parasite: Xenos pallidus Brues; Sarcophaga
sp.; Apanteles carpatus (Say); Pachysomoides fulvus (Cr.), P. stupidiis (Cr.); Elasmus
polistis Burks. Prey: Halisidota tessellaris (Sm.), Hyphantria cunea (Dru.); Anisota
virginiensis (Dru.), A. senatoria (Sm.), A. stigma (F.), Citheronia regalis (F.);
Geometridae spp.; Limacodidae sp.; Hemerocampa leucostigma (Sm.); Datana spp.,
Heterocampa manteo (Dbldy.), H. spp., Nadata gibbosa (Grt.), Pendea angulosa (A. and
S.), Schizura leptinoides (Grt.), S. spp.; Basilarchia sp.; Psilocorsis sp.; Acronicta
afflicta Grt., A. retardata (Wlkr.), A. spp. Autograpka sp., Catocala sp., Heliothis zea
(Boddie), Hypsoropha homios Hbn., Phosphila turbulente (Hbn.); Tropaea iuna (L.);
Ampeloeca myron (Cram.), Cressonia juglandis (A. and S.), Protoparce sexta (Johan.);
prey is usually obtained in wooded areas. Predator: Dicymolomia pegasalis (Wlkr.),
Chalcoela iphitalis Wlkr.

Vespa annularis Linnaeus, 1763. Centuria Ins. Rar., p. 30.

Vespa cincta Drury, 1773. Illus. Nat. Hist., Index to pt. 1 published with pt. 2.

Biology: Fox, 1896. Ent. News 7: 57 (nest). — Brimley, 1908. Ent. News 19: 107 (male
hibernation). — Rau and Rau, 1918. Wasp Studies Afield, pp. 283-290, fig. 52 (hibernation,
colony founding). —Rau, 1930. Canad. Ent. 62: 81-83 (hibernation mortality). —Rau, 1930.
Canad. Ent. 62: 119-120 (behavior on nest). —Rau, 1931. Brooklyn Ent. Soc, Bui. 26:
116-118, fig. 5 (nest). —Rau, 1940. Ent. Soc. Amer., Ann. 33: 617-620 (cooperative nest
founding). —Rau, 1942. Ent. Soc. Amer., Ann. 35: 94-96 (temperature inducing
hibernation). — Balduf, 1961. Ent. News 72: 259-260 (autumnal swarming). —Hermann and
Dirks, 1974. Ga. Ent. Soc, Jour. 9: 1-8, 3 figs, (smearing on nest by sternal glands).
—Hermann, Gerling and Dirks, 1974. Ga. Ent. Soc, Jour. 9: 203-204 (hibernation, spring
mating). —Hermann and Dirks, 1975. Psyche 82: 97-108 (spring nesting behavior).

Morphology: Snodgrass, 1941. Smithsn. Inst., Misc. Collect. 99 (14): 48-49, figs. 16 I-P (male

genitalia). —Hunt and Hermann, 1971. Ga. Ent. Soc, Jour. 5: 210-216, 2 figs, (poison

apparatus).

Polistes apachus Saussure
Western Kans. and Tex., to southern Colo., introduced into Calif.; Mexico
(Nuevo Leon, San Luis Potosi, Coahuila, Sonora, Durango). Ecology: Nests suspended
from branches, usually at tops of low trees. Parasite: Pachysomoides fulvus Cr.;
Sarcophaga sp.; Xenos peckii Kby. Predator: Chalcoela iphitalis Wlkr. This is commonly
called the Apache wasp in California where it is a pest in fig trees.

Polistes apachus Saussure, 1857. Soc. Ent. France, Ann. (3) 5: 314.

Polistes texanus Cresson, 1872. Amer. Ent. Soc, Trans. 4: 246. ♂, ♀.

Biology: Rau, 1943. Ent. Soc. Amer., Ann. 36: 522, fig. 15 (nest). —Simmons, Fisher and
Tyler, 1948. Ent. Soc. Amer., Ann. 41: 450-455, 1 pi. (nest, life history). — Snelling, 1952.
Pan-Pacific Ent. 28: 177 (hibernation).

Polistes canadensis (Linnaeus)
Southern Ariz, to Peru, Bolivia and Paraguay. Ecology: Nests in trees,
culverts and under eaves of buildings.
Vespa canadensis Linnaeus, 1758. Syst. Nat., ed. 10, v. 1, p. 574.

Biology: Rau, 1943. Ent. Soc. Amer., Ann. 36: 524, fig. 11 (nest).

Polistes carnifex
***authority mismatch
carnifex (Fabricius). South. Tex., Ariz. (Nogales); Mexico to Argentina. Ecology:
Nests under eaves, in culverts and in shrubby trees. Other subspecies occur in South
America.
Vespa carnifex Fabricius, 1775. Syst. Ent., p. 365. ♀.
Polistes onerata Lepeletier, 1836. Hist. Nat. Ins., Hym. 1: 524. ♂.
Polistes valida Say, 1837. Boston Jour. Nat. Hist. 1: 389. ♀, ♂.
Polistes transverso-strigata Spinola, 1851. Accad. Sci. Torino, Mem. (2) 13: 78. ♀.

Biology: Rau, 1943. Ent. Soc Amer., Ann. 36: 521, figs. 3, 4 (nest). —Corn, 1973 (1972).
Psyche 79: 150-157, 2 figs, (nest, behavior).
Carolina (Linnaeus). Eastern U. S., Pa. to Fla., west to Kans. and central Tex. Ecology: Nests
in very sheltered situations, such as in hollow trees or in sidings of houses. Parasite:
Xenos nigrescens Brues, X. rubiginosi (Pierce); Pachysomoides fulvus (Cr.). Predator:
Dicymolomia pegasalis (Wlkr.); Chalcoela iphitalis Wlkr. P. Carolina (L.) has been




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confused with perplexus Cr. in the past. Some of the parasite-predator records and
biological references listed here may actually refer to the latter species.

Vespa Carolina Linnaeus, 1767. Syst. Nat., Ed. 12, v. 1 (pt. 2), p. 948. ♂.

Polistes rubiginosa Lepeletier, 1836. Hist. Nat. Ins., Hym. v. 1, p. 524. ♂.

Biology: Girault, 1907. Canad. Ent. 39: 355-356 (as predator). — Rau and Rau, 1918. Wasp
Studies Afield, pp. 280-283 (prey forage, hibernation). —Rau, 1929. Ent. News 40: 7-13
(behavior female, male at end of season). —Rau, 1931. Brooklyn Ent. Soc, Bui. 26: 115-116
(nest). —Rau, 1931. Psyche 38: 129-144, 6 figs, (nesting habits, pleometrosis). —Wilson,
1933. Fla. Entomologist 17: 12 (as predator). — O'Byrne, 1938. Ent. News 49: 288-289 (nest
site).

Polistes comanchus
***authority mismatch
comanchus Saussure. Southwestern Tex.; Mexico (Coahuila, Durango).

Polistes comanchus Saussure, 1857. Soc. Ent. France, Ann. (3) 5: 314.

Polistes comanchus
***authority mismatch
navajoe Cresson. Western Tex. to Ariz.; Mexico (Sonora, Chihuahua, Durango,
Sinaloa). Ecology: Nests in cavities having very small openings.
Polistes navajoe Cresson, 1868. Amer. Ent. Soc, Trans. 1: 383. ♀.

Polistes dorsalis
***authority mismatch
californicus Bohart. Calif., Ariz.; Mexico (Baja California, Sonora, Sinaloa). Parasite:
Xenos peckii Kby.
Polistes hunteri californicus Bohart, 1949. Pan-Pacific Ent. 25: 101. ♂, ♀.

Polistes dorsalis
***authority mismatch
dorsalis (Fabricius). Va. to Fla., west to Colo., Okla., Tex., N. Mex. south to Honduras.
Ecology: Nests in shrubbery and beneath eaves. Parasite: Xenos peckii Kby.
Vespa dorsalis Fabricius, 1775. Systema Ent., p. 367. ♀.
Polistes fuscatus hunteri Bequaert, 1940. N. Y. Ent. Soc, Jour. 48: 23. ♀, ♂.

Taxonomy: van der Vecht, 1970. K. Nederland. Akad. van Wetensch., Proc, Ser. C, 71: 23
(identity of Fabrician type).

Biology: Spieth, 1948. N. Y. Ent. Soc, Jour. 56: 155-169 (nest, life history). —Hermann,
Gerling and Dirks, 1974. Ga. Ent. Soc, Jour. 9: 203-204 (hibernation).

Polistes exclamans
***authority mismatch
arizonensis Snelling. Ariz., Calif.; Mexico (Sonora).

Polistes exlamans arizonensis Snelling, 1954. Kans. Ent. Soc, Jour. 27: 152. ♀, ♂.

Biology: Hopkins, 1955. Jour. Econ. Ent. 48: 161-163 — Fye, 1972. Envir. Ent. 1: 55-57 (colony

manipulation).

Polistes exclamans
***authority mismatch
bahamensis Bequaert and Salt. La. to Fla.; Bahamas.

Polistes bahamensis Bequaert and Salt, 1931. Ent. Soc Amer., Ann. 34: 794. 9, 5, d.

Polistes exclamans louisianus Bequaert, 1940. N. Y. Ent. Soc, Jour. 48: 14. ♀.

Polistes exclamans
***authority mismatch
exclamans Viereck. N. J. to Fla., west to Nebr., Colo., and Tex.; Mexico (Nuevo

Leon, Coahuila). Ecology: Nests are usually in sheltered sites, frequently under eaves or
roofs. Parasite: Sarcophaga polistensis Hall, S. sp.; Pachysomoides stupidus (Cr.), P.
fulvus (Cr.); Elasmus polistis Burks; Monodontomerus minor (Ratz.). Prey: Arctiidae
sp.; Anisota senatoria (Sm.); Hesperiidae sp.; Heterocampa sp.; Alabama argillacea
(Hbn.), Autographa spp., Feltia subterranea (F.), Heliothis zea (Boddie), Laphygma
frugiperda (Sm.), Plathypena scabra (F.), Prodenia prob. oniithogalli Guen.; Colias
philodice eury theme Bdv.; Loxostege similalis (Guen.); Protoparce sexta (Johan.);
usually obtained from herbaceous vegetation in open fields. Predator: Dicymolomia
pegasalis (Wlkr.), Chalcoela iphitalis Wlkr.

Polistes exclamans Viereck, 1906. Amer. Ent. Soc, Trans. 32: 201. ♀.

Taxonomy: Snelling, 1955. Amer. Mus. Novitates 1701: 5-6 (synonymy). — Eickwort, 1969.
Evolution 23: 391-405 (variation between sexes and castes).

Biology: Rau, 1943. Ent. Soc. Amer., Ann. 36: 525-526, figs. 7-9 (nest). — Hodapp and Bickley,
1959. Ent. Soc. Wash., Proc. 61: 73 (autumnal swarming). —West, 1968. Psyche 75: 118-123
(distribution, nest founding). —Eickwort, 1969. Insectes Sociaux 16: 67-72 (caste separation
female, worker). —Lin, 1972. Ent. Soc. Wash., Proc. 74: 148-155 (male territoriality).

— Hermann, Gerling and Dirks, 1974. Ga. Ent. Soc, Jour. 9: 203-204 (hibernation).

— Hermann and Dirks, 1974. Ga. Ent. Soc, Jour. 9: 1-8 (smearing on nest by sternal
glands). —Hermann, Barron and Dalton, 1975. Ent. News 86: 173-178 (nest founding
behavior in spring).




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Morphology: Crouch and Smith, 1958. Tex. Jour. Sci. 10: 38-59, 20 figs, (sting and associated
glands).

Polistes flavus Cresson
Southern Calif., Nev., Utah, Ariz., N. Mex., western Tex.; Mexico (Sonora,
Chihuahua, Coahuila). Parasite: Xenos peckii Kby.
Poiistes flavus Cresson, 1868. Amer. Ent. Soc, Trans. 1: 383. ♀.

Polistes fuscatus
***authority mismatch
aurifer Saussure. B. C. to Mont., south to Calif, and Nev.; Mexico (Baja California).
Ecology: Nests in sheltered situations.Adventive in Hawaii, Johnston, Niihau and
Society Islands. Parasite: Gordius sp.; Xenos peckii Kby.
Poiistes aurifer Saussure, 1853. Etudes sur la famille des Vespides, v. 2, p. 78. ♀.
Poiistes anaheimensis Provancher, 1888. Addit. Corr. Faune Ent. Canada, Hym., p. 423. ♂.

9.
Poiistes fuscatus var. utahensis Hay ward, 1933. Utah Acad. Sci., Proc. 10: 142. ♂, ♀.
Poiistes fuscatvs connectens Bequaert, 1940. N. Y. Ent. Soc, Jour. 48: 21. ♀, ♂.
Poiistes fuscatus montanus Bequaert, 1940. N. Y. Ent. Soc, Jour. 48: 26. ♀, ♂.

Taxonomy: Snelling, 1954. Kans. Ent. Soc, Jour. 27: 154 (synonymy).

Biology: Duncan, 1928. Pan-Pacific Ent. 5: 90 (nest materials). — Bohart, 1942. Pan-Pacific
Ent. 18: 30 (feeding, hibernation). —Snelling, 1952. Pan-Pacific Ent. 28: 177 (hibernation).
—Snelling, 1954. Kans. Ent. Soc, Jour. 27: 154 (nest site).

Polistes fuscatus
***authority mismatch
bellicosus Cresson. S. C. to Fla., west to Tex.

Poiistes bellicosus Cresson, 1872. Amer. Ent. Soc, Trans. 4: 247. ♂, ♀.

Biology: Rau and Rau, 1918. Wasp Studies Afield, pp. 294-295 (drinking habits).

Polistes fuscatus
***authority mismatch
centralis Hayward. Western Tex., Colo., N. Mex., Ariz., Utah, Calif.; Mexico (Baja
California, Sonora, Chihuahua, Coahuila). Ecology: Nests in sheltered situations.

Poiistes fuscatus var. centralis Hayward, 1933. Utah Acad. Sci., Proc. 10: 143. ♂, ♀.

Polistes fuscatus
***authority mismatch
fuscatus (Fabricius). U. S. east of Rocky Mts. except in extreme southeast and Mich,
where it is replaced by/, bellicosus Cr. and/, laurentianus Beq. respectively. Ecology:
Nests in sheltered situations, frequently beneath eaves and roofs, or in cavities near
ground. Parasite: Xenos peckii Kby.; Pachysomoides fulvus (Cr.); Elasmus polistis
Burks; Dasymutilla castor (BL). Prey: Estgimene acrea (Dru.), Hyphantria cunea
(Dru.); Geometridae spp.; Hesperiidae sp.; Everes comyntes (Godt.); Dasylophia anguina
(A. and S.), Heterocampa manteo (Dbldy.); Euptoieta sp., pupa; Acronicta sp.,
Autographa spp., Feltia subterranea (F.), Heliothis zea (Boddie), H. virescens (F.),
Hypsoropha hornios Hbn., Laphygma frugiperda (Sm.), Peridroma margaritosa (Haw.),
Plathypena scabra (F.), Prodenia prob. omithogalli Guen., Schiriia sp.; Colias philodice
eurytheme Bdv., Phoebis sp.; Loxostege sp.; Protoparce sexta (Johan.); Orthoptera spp.;
usually obtained from herbaceous vegetation in open fields. Predator: Dicymolomia
pegasalis (Wlkr.), Chalcoela iphitalis Wlkr.

Vespa fuscata Fabricius, 1793. Ent. System., v. 2, p. 260.

Vespa nestor Fabricius, 1798. Sup. Ent. System., p. 262.

Poiistes pallipes Lepeletier, 1836. Hist. Nat. Ins. Hym., v. 1, p. 530. ♀.

Poiistes exilis Saussure, 1853. Etudes sur la famille des Vespides, v. 2, p. 85.

Poiistes variatus Cresson, 1872. Amer. Ent. Soc, Trans. 4: 247. ♂, ♀.

Poiistes pallidipes Dalla Torre, 1894. Cat. Hym., v. 9, p. 133. Emend.

Biology: Girault, 1911. Wis. Nat. Hist. Soc, Bui. 9: 49-63 (observations on female). —Davis,
1919. Brooklyn Ent. Soc, Bui. 14: 122-123 (nest). —Bequaert, 1923. Brooklyn Ent. Soc,
Bui. 18: 73-80 (vestigial pleometrosis). —Davis, 1924. Staten Island Inst. Arts and Science
2: 137-138 (nest). — Bugbee, 1936. Ent. Soc. Amer., Ann. 29: 614-616 (nest). —Gaul, 1940. N.
Y. Ent. Soc, Jour. 48: 391-393 (artificial brood rearing). — Beall, 1942. Canad. Field-Nat.
56: 64-67 (autumnal swarming). — Eberhard, 1969. Mich. Univ., Mus. Zool., Misc. Pub. 140:
1-101 (social biology). — Gibo, 1972. N. Y. Ent. Soc, Jour. 80: 105-108 (hibernation sites,
temperature tolerance). —Hermann and Dirks, 1974. Ga. Ent. Soc, Jour. 9: 1-8 (smearing
on nest by sternal glands). —Hermann, Gerling and Dirks, 1974. Ga. Ent. Soc, Jour. 9:
203-204 (hibernation). —Gibo, 1974. Canad. Ent. 106: 101-106 (colony founding in lab.). Rau
published many notes on this taxon as variatus; they are cited under the generic heading.




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Morphology: Marshall and Dernehl, 1905. Ztschr. f. Wiss. Biol. 80: 122-154, 2 pis.
(embryology). —Marshall, 1907. Ztschr. f. Wiss. Biol. 86: 173-213, 3 pis. (embryology).

Polistes fuscatus
***authority mismatch
laurentianus Bequaert. Que., Ont., Mich.

Polistes fuscatus laiirentianus Bequaert, 1942. Canad. Ent. 74: 159. ♂, ♀.

Polistes instabilis Saussure
South. Tex.; Mexico to Costa Rica. Ecology: Nests in trees, in culverts,
under bridges and overhanging rocks. Predator: Chalcoela iphitalis Wlkr.
Polistes instabilis Saussure, 1853. Etudes sur la famille des Vespides, v. 2, p. 91, pi. 10, fig.
2, pi. 11, fig. 1. ♂.

Biology: Rau, 1943. Ent. Soc. Amer., Ann. 36: 524-525, figs. 5, 6, 13 (nest).

Polistes kaibabensis
***authority mismatch
Hayward. Ariz. (Grand Canyon), southern Utah.

Polistes cmiadensis var. kaibabensis Hayward, 1932. Utah Acad. Sci., Proc. 9: 89. ♂, ♀.

Polistes major
***authority mismatch
castaneicolor Bequaert. Southern N. Mex. and Ariz.; Mexico (Baja California, Sonera,
Chihuahua, Jalisco), Colombia.
Polistes major var. castaneicolor Bequaert, 1936. Ent. News 47: 12. ♀, ♂.

Polistes major
***authority mismatch
major Beauvois. Southern Fla., Tex., N. Mex. and Ariz.; Mexico south to Brazil; Antilles.
Ecology: Nests in shrubbery, in culverts and beneath eaves. Parasite: Elasmus polistis
Burks. Predator: Chalcoela iphitalis Wlkr.
Polistes Major Beauvois, 1818. Ins. Afr., Amer., p. 206.
Polistes major bakeri Bequaert, 1940. N. Y. Ent. Soc, Jour. 48: 15. ♂, ♀.

Biology: Rau, 1943. Ent. Soc. Amer., Ann. 36: 522, fig. 12 (nest).

Polistes major
***authority mismatch
palmarum Bequaert. Southern Calif.; Mexico (Baja California).

Polistes major var. palmanim Bequaert, 1936. Ent. News 47: 11. ♀.

Polistes metricus Say
Eastern U. S. from N. Y. to Fla., west to Nebr., Okla. and Tex. Ecology: Nests
in shrubbery. Parasite: Xenos peckii Kby.; Sarcophaga polistensis Hall, S. sp.;
Sphecophaga vesparum hurra (Cr.); Pachysomoides fulvus (Cr.). Predator:
Dicymolomia pegasalis (Wlkr.), Chalcoela iphitalis Wlkr.
Polistes metrica Say, 1831. Indiana: School Press, New Harmony, p. 15.

Taxonomy: Rau, 1929. Psyche 36: 34-36 (color variation).

Biology: Turner, 1912. Psyche 19: 184-190 (orphan nest, feeding, homing, hunting). — Pellett,
1916. Iowa Acad. Sci., Proc. 23: 275-284 (life history, behavior in lab.). —Rau and Rau,
1918. Wasp Studies Afield, pp. 244-280, 291-294 (homing experiments with female, worker;
nest sites). —Rau, 1928. Psyche 35: 153-156 (trophallaxis). —Rau, 1929. Canad. Ent. 61:
25-30 (feeding experiments). —Rau, 1929. Ent. News 40: 226-232, 256-259 (orphan nests).
—Rau, 1930. Canad. Ent. 62: 143-147, 2 figs, (life history, nest destruction). —Rau, 1931.
Brooklyn Ent. Soc, Bui. 26: 111-113 (nest, nest site). —Rau, 1935. Ent. News 46: 25-27
(female duties). —Madden and Chamberlain, 1938. Jour. Econ. Ent. 31: 705 (as predator).
—Hermann, Gerling and Dirks, 1974. Ga. Ent. Soc, Jour. 9: 203-204 (hibernation). In these
papers Rau cited metncus as pallipes.

Polistes pacificus
***authority mismatch
pacificus Fabricius. South. Tex. to South America. Other subspp. occur in Central
and South America.
Polistes pacifica Fabricius, 1805. Systema Piezatorum, p. 274. ♀.

Taxonomy: Bequaert, 1937. Inst. Biol. Veget. Arch. 3: 196.

Polistes perplexus Cresson
Md. to Ga., west to southern 111., Kans., Okla. and Tex. P. perplexus Cr. has
been confused with Carolina (L.) in the past. Some of the parasite-predator records and
biological references listed under the latter species may actually refer to perplexus.
Polistes perplexus Cresson, 1870. Amer. Ent. Soc, Trans. 4: 245. ♂.
Polistes generosus Cresson, 1870. Amer. Ent. Soc, Trans. 4: 246. ♂.

Unplaced Taxon of Polistes Latreille

Polistes nigripennis (DeGeer)
Pa.

Vespa nigripennis Degeer, 1773. Mem. Serv. Hist, des Ins., v. 3, p. 582, pi. 29, fig. 10.
Bequaert suggested that this was possibly the same as rubiginosus {^Carolina).




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Subfamily POLYBIINAE

Revision: Bequaert, 1933. Ent. Amer. (n. s.) 13: 87-150, 3 pis. (Nearctic taxa).

Genus MISCHOCYTTARUS Saussure

The nests are similar to those of Polistes, but average somewhat smaller. They consist of a
single free comb of hexagonal paper cells and are attached by a short pedicel in North American
forms to a natural or man-made support usually some distance from the ground. Each colony is
generally started by a single fertile female or queen.

Revision: Richards, 1945. Roy. Ent. Soc, London, Trans. 95: 295-462, 4 pis., 119 figs. (New
World spp.).

Biology: Jeanne, 1972. Mus. Comp. Zool., Bui. 144: 63-150, 46 figs., 4 pis. (biology, social
behavior of M. drewseni Sauss.).

Genus MISCHOCYTTARUS Subgenus MISCHOCYTTARUS Saussure

Mischocy thorns (!) Saussure, 1853. Etudes sur la famille des Vespides, v. 1, Introduct. p.

VIII. Nom. nud.
Mischocy ttarns Saussure, 1853. Etudes sur la famille des Vespides, v. 2, p. 19.
Type-species: Zethus labiatus Fabricius. Desig. by Ashmead, 1902.

The species occur in Mexico, Central America, and South America.

Genus MISCHOCYTTARUS Subgenus KAPPA Saussure

Kappa Saussure, 1854. Etudes sur la famille des Vespides, v. 2, p. 200.

Type-species: Polybia injucunda Saussure. Desig. by Bequaert, 1933.

Mischocyttarus cubensis
***authority mismatch
cubcnsis (Saussure). Ga., Fla., Ala.; Bahamas, Cuba.

Polybia cube7isis Saussure, 1854. Etudes sur la famille des Vespides, v. 2, p. 202. ♂, ♀.

Polybia phthisica Cresson, 1865. Ent. Soc. Phila., Proc. 4: 167. ♂, ♀.

Mischocyttarus cubensis
***authority mismatch
mexicanus (Saussure). Tex. (Brownsville); Mexico, Central America.

Polybia mexicana Saussure, 1854. Etudes sur la famille des Vespides, v. 2, p. 203. ♀.

Mischocyttarus flavitarsis
***authority mismatch
centralis Bequaert. Utah, Colo., Nebr., N. Mex., western Tex.

Mischocyttarus flavitarsis var. centralis Bequaert, 1933. Ent. Amer. (n. s.) 13: 129. ♀, ♂.

Mischocyttarus flavitarsis
***authority mismatch
flavitarsis (Saussure). Calif., Oreg., Utah, Colo., Nebr., Ariz.

Polybia flavitarsis Saussure, 1854. Etudes sur la famille des Vespides, v. 2, p. 199. ♀.

Biology: Smith, 1944. Pan-Pacific Ent. 20: 80 (hibernation site). — Snelling, 1953. Kans. Ent.

Soc, Jour. 26: 143-145 (hibernation, nest size and site).

Mischocyttarus flavitarsis
***authority mismatch
idahoensis Bequaert. B. C, Wash., Greg., Idaho, Mont., Nev., Utah, Colo.

Mischocyttarus flavitarsis var. idahoensis Bequaert, 1933. Ent. Amer. (n. s.) 13: 133. ♀, ♂.

Mischocyttarus flavitarsis
***authority mismatch
kaibabensis Bequaert. Ariz. (Grand Canyon, Kayenta).

Mischocyttams flavitarsis var. kaibabensis Bequaert, 1933. Ent. Amer. (n. s.) 13: 133. ♀.

Mischocyttarus flavitarsis
***authority mismatch
navajo Bequaert. Ariz.; Mexico (Chihuahua).

Mischocyttarus flavitarsis var. navajo Bequaert, 1933. Ent. Amer. (n. s.) 13: 131. ♀, ♂.

Genus BRACHYGASTRA Perty

Brachygastra Perty, 1833. Delect. Anim. Articul. Brasil, p. 145.

Type-species: Brachygastra analis Perty Desig. by Bequaert, 1933 {=Polistes
lecheguana Latreille).
Nectarina Swainson and Shuckard, 1840. On the history and natural arrangement of

insects, p. 183. N. name for Brachygastra.
Melissaia Shuckard, 1841. hi White, Ann. and Mag. Nat. Hist. 7: 320. N. name for

Nectariyia.
Caba Ihering, 1904. Rev. Mus. Pauhsta 6: 103. N. name for Nectarina.

The genus is essentially Neotropical but one species extends its range into southern United
States. The perennial paper nests are attached to branches of low trees and are begun by small




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swarms of fertile queens and workers. An average nest is subglobular, is enclosed in a single
sheet of papery material, is about 1 foot in diameter, contains about 50,000 cells arranged in
layers, has several entrances, and has spiral ramps connecting the tiers of cells. The adults col-
lect and store honey but do not cap the cells.

Revision: Buysson, 1905. Soc. Ent. France, Ann. 74: 537-566, 6 pis. — Naumann, 1968. Kans.
Univ. Sci. Bui. 47: 929-1003, 69 figs. (New World spp.).

Brachygastra mellifica (Say)
Southern Tex. and Ariz, to Costa Rica and Panama (?).

Polistes mellifica Say, 1837. Boston Jour. Nat. Hist. 1 (pt. 4): 390. ♂, ♀.

Nectarina mellifera Dalla Torre, 1904. Gen. Ins., fasc. 19, p. 86. Emend.

Chartergus aztecus Cameron, 1906. Invertebrata Pacifica 1: 154. ♀. Preocc.

Chartergus arizonaensis Cameron, 1907. Invertebrata Pacifica 1: 181. ♀.

Chartergus centralis Cameron, 1907. Invertebrata Pacifica 1: 181. ♀.

Nectarina cameroni Meade-Waldo, 1911. Ann. and Mag. Nat. Hist. (8) 7: 112. N. name.

Biology: Barber, 1905. Ent. Soc. Wash., Proc. 7: 25 (nest). — Schwarz, 1929. Nat. Hist. 29:
421-426, 5 figs, (nest, honey production).

Genus POLYBIA Lepeletier

Polybia Lepeletier, 1836. Hist. Nat. Ins. Hym., v. 1, p. 533.

Type-species: Polistes liliacea Fabricius Desig. by Ashmead, 1902.
Myrapetra White, 1841. Ann. and Mag. Nat. Hist. 7: 320.

Type-species: Myrapetra scutellaris White. Monotypic.
Myraptera Curtis, 1844. Linn. Soc. London, Proc, p. 188. Emend.
Alpha Saussure, 1853. Etudes sur la famille des Vespides, v. 2, p. 167.

Type-species: Polybia bifasciata Saussure. Desig. by Bequaert, 1944.
Eupolybia Dalla Torre, 1904. In Wytsman, Gen. Ins., fasc. 19, p. 76.

Type-species: Polistes liliacea Fabricius. Present desig. by Richards.

The species of Polybia build phragmocyttarous nests which consist of one to many combs at-
tached to a branch, leaf or rock, surrounded by an envelope of carton or of mud containing an
exit hole at the side or bottom of the comb. A number of species enlarge the nest by adding suc-
cessive combs, each covered by the envelope, so that access to the upper combs is through the
exit-holes of earlier stages of the nest. Some of the taxa store large quantities of honey, or sex-
ual castes of ants or termites, some of which are fed to larvae and some of which maintain the
adults during hibernation. Larvae may also be fed on partially masticated insects.

Taxonomy: Richards, 1951. In Richards and Richards, Royal Ent. Soc. London, Trans. 102:
129-150, figs. 11-16 (taxa allied to occidentalis (Oliv.) with key including the two taxa
occurring in America north of Mexico).

Polybia diguetana Buysson
Ariz. (Nogales); Mexico, Central America, Peru. Ecology: Buysson noted
that the nest was like that of typical occidentalis which consists of as many as 6 combs
surrounded by a carton envelope.
Polybia occidentalis var. diguetana Buysson, 1905. Soc. Ent. France, Bui, p. 9. 9, 5 , 6.
occidentalis nigrateila Buysson. Ariz. (Nogales); Mexico, Guatemala, Honduras. Typical
occidentalis (OHv.) occurs in South America.
Polybia occidentalis var. nigrateila Buysson, 1905. Soc. Ent. France, Bui., p. 9. ♀, ♂.
Polybia occidentalis nausicaa Richards, 1951. In Richards and Richards, Royal Ent. Soc.
London, Trans. 102: 140. 9, cJ. N. syn. (0. W. Richards).

Subfamily VESPINAE

This is morphologically the most specialized subfamily of the social wasps. Component species
are commonly called hornets (those nesting above ground in North America) and yellow jackets
(those nesting usually subterraneously). The nests consist of several to many combs of hex-
agonal cells composed of paper; cells constructed early in the year may be used for several lar-
vae in succession. The combs are usually enclosed in a paper envelope. The nests are annual, new
queens and males being produced late in the summer or early in the fall; the newly fertilized
queens overwinter and begin new nests in the spring. There are relatively few precise records




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of the insects used to feed vespine larvae; apparently dismembered and masticated adult Dip-
tera and honeybees are commonly used; the wasps may also obtain bits of flesh from fresh and
decaying carcasses. Adult vespines feed on liquid foods, primarily nectar or honey; some species
are known to prey upon adult honeybees, which they kill and extract nectar from the crop.

Revision: Bequaert, 1932. Ent. Amer. (n. s.) 12: 71-138, 6 figs. (N. Amer. taxa).

Taxonomy: McFarland, 1888. Amer. Ent. Soc, Trans. 15: 297 (key). —Bequaert, 1930.

Brooklyn Ent. Soc, Bui. 25: 59-70 (generic and subgeneric classification). —Bequaert, 1935.
Brooklyn Ent. Soc, Bui. 30: 119-124 (check list, correction, additions). —Duncan, 1939.
Stanford Univ., Pubs., Univ. Ser. Biol. Sci. 8 (1): 85-97 (genera). —Bequaert, 1941.
Brooklyn Ent. Soc, Bui. 36: 111-117 (corrections, additions).

Biology: Duncan, 1939. Stanford Univ., Pubs., Univ. Ser. Biol. Sci. 8 (1): 98-176 (composite
summary for American spp.).

Genus VESPA Linnaeus

Vespa Linnaeus, 1758. Syst. Nat., ed. 10, v. 1, p. 343.

Type-species: Vespa crabro Linnaeus. Desig. by Lamarck, J801.
Macrovespa Dalla Torre, 1904. In Wytsman, Gen. Ins., fasc. 19, p. 64.

Type-species: Vespa crabro Linnaeus. Desig. by Bequaert, 1980.

The giant hornets are conspicuous members of the wasp fauna in the Palaearctic and Oriental
Regions; one taxon is adventive in North America. The species of Vespa are quite diverse in
nesting habits; some construct aerial nests in trees, others nest in sheltered sites above ground,
such as in hollow trees, and still others are subterranean. The nests, especially in exposed situa-
tions, are usually enclosed in a paper envelope.

crabro germana Christ. Ont., Mass., R. I., Conn., N. Y., N. J., Pa., Del., Md., D. C, Va., N. C, S.
C, Ga., Ala., W. Va., Ohio, Ind., Ky., Tenn., Wis., N. Dak., S. Dak.; central and western
Europe. Ecology: Nests in very sheltered sites above ground, frequently in hollow trees,
attics, porches, sheds, and rarely in underground cavities; exposed nests have a
complete paper envelope, but in quite sheltered sites the envelope may be only partial
or completely lacking. Adventive from Europe in mid-1800's. The official common name
is the giant hornet; in some rural areas of the mid-Atlantic states it is known as the
hybrid bee. Typical crabro L. and other subspecies occur in the Palaearctic Region.

Vespa crabro germana Christ, 1791. Naturgesch. Insekt. Bienen, Wespen u.
Ameisengeschl., p. 215.

Vespa crabro vulgata Birula, 1925. Arch. f. Naturgesch. 90, Abt. A, H. 12, p. 100. 9,^,6.

Taxonomy: Bequaert, 1931. Konowia 10: 101-109 (color forms of crabro L. with key).

Biology: Beutenmuller, 1898. N. Y. Ent. Soc, Jour. 6: 199, pis. 9-10 (nest). —Felt, 1915. N. Y.
State Mus., Bui. 180: 71-73, pi. 2 (nest). —Cory, 1931. Jour. Econ. Ent. 24: 50-52 (nest,
injury to lilac, apples). — Ikan et al., 1969. Jour. Insect Physiol. 15: 1703-1712 (queen
pheromones). — Islay and Schaudinschky, 1973. Acoust. Soc Amer., Jour. 53: 640-649
(acoustical communication).

Morphology: Snodgrass, 1941. Smithsn. Inst., Misc. Coll. 99 (14): 49-50, pi. 17, figs. I-Q, pi. 18,
figs. A-D (male genitalia).

Genus VESPULA Thomson

These smaller social vespines are restricted to the Holarctic Region.

Revision: Miller, 1961. Canad. Ent., Sup. 22, v. 93: 1-52, 84 figs., 15 maps.

Taxonomy: Sladen, 1918. Ottawa Nat. 32: 71-72 (key to Canadian spp.). —Miller, 1958 (1956).
Tenth Internatl. Cong. Ent., Proc. 1: 257-264 (nomenclature and distribution). — Fluno,
1973. Ent. Soc. Wash., Proc. 75: 80-83, 1 fig. (differentiation of subgenera by attraction to
chemical).

Biology: Taylor, 1939. Ent. Soc. Amer., Ann. 32: 304-315 (origin and evolution of social
parasitism). —Gaul, 1941. Psyche 48: 16-19 (artificial housing of colonies). —Gaul, 1942.




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Brooklyn Ent. Soc, Bui. 37: 57-61 (brood rearing). —Morse and Gary, 1961. Bee World 42:
179-181 (predation on honeybees). — Akre, Hill and MacDonald, 1973. Jour.Econ. Ent. 66:
803-805, 4 figs, (artificial housing of colonies).

Genus VESPULA Subgenus VESPULA Thomson

Vespula Thomson, 1869. Opusc. Ent., v. 1, p. 79.

Type-species: Vespa austriaca Panzer. Desig. by Ashmead, 1902.
Pseudovespa Schmiedeknecht, 1881. Ent. Nachr. 7: 314.

Type-species: Vespa austriaca Panzer. Monotypic.
Paravesptila Bluethgen, 1938. Konowia 16: 271.

Type-species: Vespa vulgaris Linnaeus. Orig. desig.
Allovespula Bluethgen, 1943. Stettin. Ent. Ztg. 104: 149.

Type-species: Vespa rufa Linnaeus. Orig. desig.

Nests are usually underground or close to it in hollow logs or stumps. Combs, except for a
small conical area around the main suspensorium, are usually essentially plane. Members of this
subgenus are commonly called yellow jackets.

Vespula acadica (Sladen)
Transcontinental in Canadian Zone from Newfoundland to Yukon Terr, and
Alaska, Maine to Minn., west of 100th meridian south to N. Mex. and Calif. Ecology:
Location of nests variable, sometimes aerial, subterranean or in rotten logs.

Vespa rufa var. americana Buysson, 1905. Soc. Ent. France, Ann. 63: 499. ♂. Preocc.

Vespa acadica Sladen, 1918. Ottawa Nat. 32: 72. 6, 9, 5.

Vespida rufa sladeni Bequaert, 1931. Ent. Amer. (n. s.) 12: 102. 6, 9, 5.

Biology: Sladen, 1918. Ottawa Nat. 32: 72 (nest site). —MacDonald, Akre and Hill, 1975.
Kans. Ent. Soc, Jour. 48: 114-121 (nest location and structure).

Vespula atropilosa (Sladen)
Canada and U. S. west of 100th meridian mostly in Canadian and

Transition Zones; Mexico (Baja California). Ecology: Most nests are subterranean and
usually in rodent burrows. Parasite: Sphecophaga vesparum hurra (Cr.); Triphleba
lugubris (Meig.).
Vespa atropilosa Sladen, 1918. Ottawa Nat. 22: 72. 9, 0.

Biology: MacDonald, Akre and Hill, 1974. Melanderia 18: 1-93, 13 figs, (comparative biology

and behavior). —MacDonald, Akre and Hill, 1975. Kans. Ent. Soc, Jour. 48: 53-63 (nest

associates). —MacDonald, Akre and Hill, 1975. Kans. Ent. Soc, Jour. 48: 114-121, 2 figs.

(nest location and structure).

Vespula austriaca (Panzer)
Que., Ont., Man., Sask., Alta., B. C, N. W. T., Alaska, Maine, N. Y., N. J.,
Colo., N. Mex., Utah, Calif., Oreg., Wash., Idaho; Holarctic Host: Inquiline in nests of
rufa (L.) in Europe; North American host(s) unknown. The worker caste is absent.

Vespa austriaca Panzer, 1799. Faunae Ins. German., h. 63, pi. 2. ♂.

Vespa borealis Smith, 1843. Zoologist 1: 170. ♀. Preocc.

Vespa arborea Smith, 1849. Zoologist 7, App., p. Ix. N. name.

Vespa infemalis Saussure, 1853. Etudes sur la famille des Vespides, v. 2, p. 139. ♀.

Vespa tripunctata Packard, 1870. Chicago Acad. Sci., Trans. 2: 26. ♀.

Vespula consobrina (Saussure)
Transcontinental in Canada and northern U. S. in Canadian and
Transition Zones, W. Va., N. C, Ga., Colo., N. Mex., Ariz., Calif. Ecology: Nests are
usually subterranean, but occasionally are in shrubs just above the ground surface.

Vespa consobrina Saussure, 1864. Etudes sur la famille des Vespides, v. 2, p. 141. ♂.

Vespa scelesta McFarland, 1888. Amer. Ent. Soc, Trans. 15: 298. 6, 9, 5.

Vespa sulcata Howard, 1901. Insect Book, pi. 6, fig. 18. ♂.

Biology: Dow, 1930. Boston Soc. Nat. Hist., Bui. 56: 12, 1 fig. (nest).

Vespula germanica (Fabricius)
N. Y., N. J., Pa., Del, Md.; Europe. Ecology: Nests in sheltered

situations, frequently in structures. Adventive from Europe, apparently m mid-20th
century. References to taxonomy, biology and morphology of the Palaearctic population
will be found in Guigha, 1973 (1972), Les Guepes Sociales d'Europe Occidental et
Septentrionale, vol. 6 of Faune de I'Europe et du Bassin Mediterraneen.
Vespa Germanica Fabricius, 1793. Ent. System., v. 2, p. 256.




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Taxonomy: Menke and Snelling, 1975. U. S. Dept. Agr., Coop. Ins. Rpt. 25: 193-200, 33 figs,
(characters to distinguish from native North American spp.).
intermedia (Buysson). Newfoundland, Que., Man., Sask., Alta., N. W. T., Yukon Terr., Alaska in
Hudsonian Zone.
Vespa rufa var. intermedia Buysson, 1905. Soc. Ent. France, Ann. 73: 591. ♂, ♀.

Vespula maculifrons (Buysson)
Southern Canada and U. S. east of 100th meridian in Transition and

Austral Zones. Ecology: Nests are usually subterranean, occasionally in decayed stumps
or logs; nest usually contains 4-8 combs, rarely only a single comb. It may live as an
inquiline in nests of vulgaris (L.) and interbreed with it. Parasite: Vespula squamosa
(Dru.). V. maculifrons was frequently misidentified as gerynanica in publications prior
to 1931.
Vespa maculifrons Buysson, 1905. Soc. Ent. France, Ann. 63: 608.
Vespa communis \a.r.flavida Sladen, 1918. Ottawa Nat. 32: 71. ♀.

Biology: Couper, 1870. Canad. Ent. 2: 49-53 (nest). — Marlatt, 1891. Ent. Soc. Wash., Proc. 2:
80-83 (nest). — Rau and Rau, 1918. Wasp Studies Afield, pp. 295-297, fig. 53 (foraging,
nest). — Beamer, 1925. Ent. News 36: 309-310 (hibernation female). —Rau, 1930. Ent.
News 41: 185-190, pi. 19 (life history, nest). —Rau, 1931. Brooklyn Ent. Soc, Bui. 26: 85-89,
pi. 4 (nest, colonization). —Bromley, 1931. N. Y. Ent. Soc, Jour. 39: 129 (foraging,
predation). —Gaul, 1948. Brooklyn Ent. Soc, Bui. 43: 37-41 (interspecific tolerance).
— Haviland, 1962. Ent. Soc. Wash., Proc. 64: 181-183 (colony size). — Kurczewski, 1968. N.
Y. Ent. Soc, Jour. 76: 84-86 (predation). — Balduf, 1968. Ent. Soc. Wash., Proc. 70: 336-338,
1 fig. (inquilinism and interbreeding with vulgaris). —Simon and Benton, 1968. Ent. Soc.
Amer., Ann. 61: 542 (winter activity in aerial nest). —Green, Heckman, Benton and Coon,
1970. Ent. Soc Amer., Ann. 63: 1197-1198, 2 figs, (exposed aerial nest). — MacDonald and
Matthews, 1975. Science 190: 1003-1004, 1 fig. (parasite).

Vespula pensylvanica (Saussure)
Canada and western U. S. west of 100th meridian in Canadian and

Transition Zones; Mexico (Michoacan, Mexico). Ecology: Nests are subterranean.

Parasite: Bareogonalos canadensis (Harr.); Sphecophaga vesparuni burra (Cr.);

Triphleba lugubris (Meig.).
Vespa pensylvanica Saussure, 1857. Stettin. Ent. Ztg. 18: 117. ♀.
Vespa occidentalis Cresson, 1874. Amer. Ent. Soc, Trans. 5: 100. ♀, ♂. Preocc.

Biology: Snow, 1882. Psyche 3: 339 (predation). — Metzmain, 1903. Entomologist 36: 137
(female hibernation). —Duncan, 1939. Stanford Univ., Pubs., Univ. Ser. Biol. Sci. 8 (1):
98-173 (nest, life history). —Smith, 1956. Ent. News 67: 141-146, 1 pi. (nest). —Chapman,
1963. Ecology 44: 766-767 (predation). —MacDonald, Akre and Hill, 1974. Melanderia 18:
1-93, 13 figs, (comparative biology and behavior).

Morphology: Duncan, 1939. Stanford Univ., Pubs., Univ. Ser. Biol. Sci. 8 (1): 13-84, pis.

I-XXVII (female, male).

Vespula squamosa (Drury)
Ont., N. Y. to Wis. and Iowa, south to Fla. and Tex., most common in
Austral Zones; Mexico, Guatemala. Host: Vespula maculifrons (Buyss.), V. vidua
(Sauss.); the squamosa queen is a temporary, apparently facultative social parasite.

Vespa squamosa Drury, 1773. Illus. Nat. Hist., Index to pt. 1 published with pt. 2.

Vespa lineata Fabricius, 1775. Systema Ent., p. 365.

Vespa conchacea Christ, 1791. Naturgesch. Insekt. Bienen, Wespen u. Ameisengeschl, p.
259.

Vespa cuneata Fabricius, 1804. Systema Piezatorum, p. 258.

Vespa cruciata Lepeletier, 1836. Hist. Nat. Ins. Hym., v. 1, p. 513. Emend, of cuneata.

Vespa bistriata MacFarland, 1888. Amer. Ent. Soc, Trans. 15: 298. ♀. Preocc.

Vespa macfarlandi Lewis, 1897. Amer. Ent. Soc, Trans. 24: 172. 9. N. name.

Vespula squamosa var. (or subsp.) michoacana Bequaert, 1941. Ent. News 52: 249. ♂.

Taxonomy: Turner, 1908. Psyche 15: 1-3, 1 pi. (coloration worker, male).

Biology: Taylor, 1939. Ent. Soc. Amer., Ann. 32: 310-313 (temporary parasitism by female in
nest of vidua). —Gaul, 1947. Brooklyn Ent. Soc, Bui. 42: 87-96 (behavior, nest). —Gaul,
1948. Brooklyn Ent. Soc, Bui. 43: 37-41 (interspecific tolerance). — Tissot and Robinson,
1954. Fla. Ent. 37: 73-92 (aerial and subterranean nests). —MacDonald and Matthews,
1975. Science 190: 1003-1004, 1 fig. (host, nest).




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Vespula sulphurea (Saussure)
Oreg., Calif., Nev., Ariz.; Mexico (Baja California). Ecology: Nests are
subterranean.
Vespa sulphurea Saussure, 1854. Etudes sur la famille des Vespides 2: 137. ♀.

Biology: Bequaert, 1931. Ent. Amer. (n. s.) 12: 112 (nest).

Vespula vidua (Saussure)
N. S., N. B., Ont., in northeastern U. S. from Maine to N. Dak., south to Va.
and Iowa, south in Appalachian Mts. to N. C, Ga., mostly in Transition and U. Austral
Zones. Ecology: Nests are subterranean. Parasite: Sphecophaga vespai-um burra (Cr.);
Vespula squamosa (Dru.), the queen of which is a temporary social parasite.
Vespa vidua Saussure, 1854. Etudes sur la famille des Vespides, v. 2, p. 136.

Biology: Taylor, 1939. Ent. Soc. Amer., Ann. 32: 310-313 (nest, parasitism by squamosa).

Vespula vulgaris (Linnaeus)
Alaska, Canada, U. S. south to Calif., Ariz., N. Mex., S. Dak., Iowa, Ind.,
111., Ohio, N. C, mostly in Canadian and Transition Zones; Mexico (Michoacan, Mexico);
widely distributed in Palaearctic Region; Ecology: Nests are usually subterranean or in
stumps with paper envelope present or absent; aerial nests are close to ground and
always with envelope, adventive in New Zealand. Parasite: Sphecophaga vesparum
burra (Cr.). May interbreed with maculifrons (Buyss.).

Vespa vulgaris Linnaeus, 1758. Syst. Nat., ed. 10, v. 1, p. 572. ♀.

Vespa communis Saussure, 1857. Stettin. Ent. Ztg. 18: 117. ♀.

Vespa alasce7isis Packard, 1870. Chicago Acad. Sci., Trans. 2: 27. ♀.

Vespa westwoodii Shipp, 1893. Psyche 6: 450.

Biology: Bequaert, 1931. Ent. Amer. (n. s.) 12: 92-93 (nest). — Balduf, 1968. Ent. Soc. Wash.,
Proc. 70: 332-336, 1 fig. (nest, life history).

Genus VESPULA Subgenus DOLICHOVESPULA Rohwer

Dolichovespula Rohwer, 1916. Conn. State Geol. and Nat. Hist. Survey, Bui. 22: 642.

Type-species: Vespa maculata Linnaeus. Orig. desig.
P seudovespula Bischoft, 1931. Gesell. Naturf. Freunde Sitzber. (1930), p. 346.

Type-species: Vespa noiivegica var. adulterina Buysson. Orig. desig.
Boreovespula Bluethgen, 1943. Stettin. Ent. Ztg. 104: 149.

Type-species: Vespa norwegica Fabricius. Orig. desig.
Metavespula Bluethgen, 1943. Stettin. Ent. Ztg. 104: 149.

Type-species: Vespa silvestris ScopoH. Orig. desig.

Nests are usually aerial, suspended at some height or in bushes near the ground. Combs are
usually turned up at the margins so as to be concave above.

Vespula albida (Sladen)
Labrador, Newfoundland, Que., Maine, Man., Alta., B. C, N. W. T., Yukon
Terr., Alaska, chiefly in Hudsonian Zone. Ecology: Nests are small and subterranean.
Vespa marginata Kirby, 1837. Fauna Bor.-Amer., v. 4, p. 265. ♀. Preocc.
Vespa albida Sladen, 1918. Ottawa Nat. 32: 71. c?, ?.

Biology: Sladen, 1919. Rpt. Canad. Arctic Exped. 1913-18, v. 3, p. 26g (nest).

Vespula arctica Rohwer
Newfoundland to Yukon Terr., Alaska, in lower U. S. chiefly in Canadian
Zone. Host: Vespula arenaria {¥.). The worker caste is absent.
Vespa borealis Lewis, 1897. Amer. Ent. Soc, Trans. 24: 171. ♂, ♀. Preocc.
Vespula arctica Rohwer, 1916. Conn. State Geol. and Nat. Hist. Survey, Bui. 22: 642. N.
name.

Biology: Wheeler and Taylor, 1921. Psyche 28: 135-144, 3 figs, (permanent social parasitism

of arctica in nests of arenaria). —Wheeler, 1939. Ent. Soc. Amer., Ann. 32: 305-310 (social

parasitism in arenaria nests).

Vespula arenaria (Fabricius)
In Canada from Newfoundland to Yukon Terr., Alaska, in lower U. S.
transcontinental chiefly in Canadian and Transition ones. Ecology: Nests are usually
aerial but placed close to ground in shrubbery, occasionally under eaves or roofs, rarely
the nests may be attached to tree roots and only partly exposed. Parasite: Sphecophaga
vesparum burra (Cr.); Vespula arctica Roh.

Vespa arenaria Fabricius, 1775. Systema Ent., p. 365.

Vespa borealis Kirby, 1837. Fauna Bor.-Amer., v. 4, p. 264. Preocc.



1522 Hymenoptera in America Nort^h of Mexico

Vespa diabolica Saussure, 1854. Etudes sur la famille des Vespides, v. 2, p. 138. 2,5.
Vespa fernaldi Lewis, 1897. Amer. Ent. Soc, Trans. 24: 171. ♀, ♂.

Biology: Wheeler and Taylor, 1921. Psyche 28: 135-144, 3 figs, (nest, parasite). — Hungerford,
1930. Ent. News 41: 329-330, 1 pi. (nest). —Taylor, 1939. Ent. Soc. Amer., Ann. 32: 305-310
(nest, parasite). —Gaul, 1941. N. Y. Ent. Soc, Jour. 49: 367-369 (taste sensitivity of adults,
larvae). — Gaul, 1941. Psyche 48: 16-19 (colony housing, interspecific tolerance). — Gaul,
1942. Brooklyn Ent. Soc., Bui. 37: 57-61 (brood rearing, mating). —Gaul, 1948. Brooklyn
Ent. Soc, Bui. 43: 73-79 (distribution of labor). —Sailer, 1950. Kans. Ent. Soc, Jour. 23:
134-137, 3 figs, (nest temperature). —Gaul, 1952. N. Y. Ent. Soc, Jour. 60: 17-20 (flight in
stormy weather). —Gaul, 1952. Brooklyn Ent. Soc, Bui. 47: 79-92 (temperature regulation
in nest). — Gibo, 1972. N. Y. Ent. Soc, Jour. 80: 105-108 (hibernation sites, temperature
tolerance). —Gibo, Yarascavitch and Dew, 1974. Canad. Ent. 106: 503-507, 1 fig. (colony
thermoregulation). —Gibo, Dew and Hajduk, 1974. Canad. Ent. 106: 873-879, 2 figs,
(relation of colony biomass to calorie production).

Vespula maculata (Linnaeus)
In Canada from Nova Scotia to Yukon Terr., Alaska, in lower U. S. from
Maine to Fla., west to 100th meridian in Canadian, Transition, U. and L. Austral Zones,
west of 100th meridian chiefly in Canadian, Transition and U. Sonoran Zones. Ecology:
Nests are always aerial and are usually suspended from limbs or beneath eaves and
roofs. Parasite: Sphecophaga vesparum burra (Cr.). The official common name is the
bald-faced hornet; sometimes it is also caUed the white-faced hornet.
Vespa maculata Linnaeus, 1763. Centuria Ins. Rar., p. 30.

Vespa maculata am,ericana Christ, 1791. Naturgesch. Insekt. Bienen, Wespen u.
Ameisengeschl., p. 239.

Biology: Couper, 1870. Canad. Ent. 2: 49-53 (nest). — Fyles, 1903. 33rd Ann. Rpt., Ent. Soc.
Ontario, pp. 69-70, figs. 47-48 (nest). —Howard, 1915. Ent. Soc. Wash., Proc 17: 148 (nest
color). — Rau and Rau, 1918. Wasp Studies Afield, pp. 297-298 (predation). —Davis, 1919.
Brooklyn Ent. Soc, Bui. 14: 119-123, 1 pi. (nest). — parker, 1928. Ent. Soc. Wash., Proc. 30:
14 (succession of brood in one cell). —Rau, 1929. Ent. Soc. Amer., Ann. 22: 659-675, 10 figs,
(nesting habits). —Bromley, 1931. N. Y. Ent. Soc, Jour. 39: 126-128 (predation). — Betz,
1932. Quart. Rev. Biol. 8: 197-209, 4 figs, (colony size). —Rau, 1934. Brooklyn Ent. Soc,
Bui. 19: 170 (hibernating female). —Rau, 1934. Brooklyn Ent. Soc, Bui. 19: 171 (predation).
— Balduf, 1936. Canad. Ent. 68: 138-139 (colony size). —Gaul, 1941. Psyche 48: 16-19
(colony housing, interspecific tolerance). —Gaul, 1952. N. Y. Ent. Soc, Jour. 60: 21-24
(metabolic cycles and flight). —Balduf, 1954. Ent. Soc. Amer., Ann. 47: 445-458, 3 pis. (nest
structure and growth, foraging, life history, population size). — Gibo, 1972. N. Y. Ent. Soc,
Jour. 80: 105-108 (hibernation sites, temperature tolerance). —Howell, 1973. Ent. News 84:
141-142 (predation on Vespula maculifrons (Buyss.)). — Gibo, Yarascavitch and Dew, 1974.
Canad. Ent. 106: 503-507, 1 fig. (colony thermoregulation). —Gibo, Dew and Hajduk, 1974.
Canad. Ent. 106: 873-879, 2 figs, (relation of colony biomass to calorie production).

Morphology: Bequaert, 1932. Ent. Amer. (n. s.) 12: 73-75, fig. 1 (male genitalia). — Snodgrass,
1941. Smithsn. Inst., Misc Collect. 99 (14): 49, pi. 17, figs. A-H (male genitalia). —Hermann
and Krispyn, 1975. Ga. Ent. Soc, Jour. 10: 307-313, 8 figs, (venom apparatus).

Vespula norvegicoides (Sladen)
In Canada from Newfoimdland to Yukon Terr., Alaska, in lower U. S.
transcontinental chiefly in Canadian Zone. Ecology: Nests in low shrubbery.
Vespa norvegicoides Sladen, 1918. Ottawa Nat. 32: 71. ♂, ♀.

Biology: Bequaert, 1932. Ent. Amer. (n. s.) 12: 119 (nest).



1523




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