Guia de Id de Avispas

1573^9^
iá States
wârtment of
Agriculture
Science and
The Yellowjackets of
Education
Administration
Agriculture
America North of
Handbook
Number 552
Mexico
~^'
United States
Department of The Yellowjackets of America
^""17
Education
North of Mexico
Administration
Agriculture
Handbook
Number 552
For sale by the Superintendent of Documents, U.S. Government Printing Office

Washington, D.C. 20402
ABSTRACT
Akre, R.D., A. Greene, J.F. MacDonald, P.J. Lan- included for rapid identification of species, and a
dolt, and H. G. Davis. 1980. Yellowjackets of glossary defines many of the unusual terms used
America North of Mexico. U.S. Department in the text.
of Agriculture, Agriculture Handbook
No. 552, 102 pp.
Keywords: Hymenoptera, Vespinae, Vespa, Ves-
This volume is a compendium of data on the pula, DolichovespulUy yellowjackets,
pestiferous vespines (hornets and yellowjackets) hornets, yellowjacket biology, econo-
of the United States and Canada. Sections cover mic losses, stings, sting reactions,
taxonomy, biology, economic and medical yellowjacket keys, control, urban
importance, and control. Illustrated keys are problems
This pubhcation reports research involving pesticides. It does

not contain recommendations for their use, nor does it imply that
the uses discussed here have been registered. All uses of
pesticides must be registered by appropriate State and/or Fed-
eral agencies before they can be recommended.
CAUTION: Pesticides can be injurious to humans, domestic

animals, beneficial insects, desirable plants, and fish or other
wildlife—if they are not handled or applied properly. Use all
pesticides selectively and carefully. Follow recommended
practices for the disposal of surplus pesticides and pesticide
containers.
Trade names are used in this publication solely to provide spe-

cific information. Mention of a trade name does not constitute a
warranty or an endorsement of the product by the U.S. Depart-
ment of Agriculture to the exclusion of other products not
mentioned.
Issued June 1981

CONTENTS
Page Page
Introduction 1 Vespula pensylvanica 69
Literature review 3 Vespula vulgaris ,.. 69
Taxonomy 3 Economic importance 74
Introduction 3 Beneficial aspect 74
Identification of genera and species . 7 Economic losses 75
Identification of members of Vespula Losses in agricultural areas 75
species groups 12 Losses in recreational areas 77
Key to Vespinae in North America 12 Losses in urban and
Key to genera 12 suburban areas 77
Key to species 14 Medical importance 78
Vespa 14 Yellowjacket sting apparatus 78
Dolichovespulçi 15 Morphology 78
Vespula 17 Sting function 82
Yellowjacket and hornet worker gaster Venoms 82
patterns drawn to scale (fig. 25) . 23 Yellowjacket stings 82
Vespa 24 Fatal reactions to stings 84
Dolichovespula 24 Symptoms of allergic reaction 85
Vespula 24 Ancillary factors and
Biology 24 sting fatalities 86
General 24 Emergency treatment 86
Hornets— Vespa crabro 32 Long-term treatment 86
Yellowjackets 35 Emergency medical kits 86
Genus Dolichovespula 35 Avoidance of stings 87
Dolichovespula albida 35 Control 87
Dolichovespula árctica 36 Nest destruction 87
Dolichovespula arenaria 38 Subterranean nests 87
Dolichovespula maculata 42 Nests in houses 87
Dolichovespula norvegicoides .. 45 Aerial nests 88
Genus Vespula 46 Use of poison baits 88
Vespula rufa species group 46 Chemical lures 90
Vespula acadica 46 Management of garbage 91
Vespula atropilosa 46 Biological control , 91
Vespula austriaca 51 Control of yellowjacket queens 91
Vespula consobrina 52 Fish or protein bait
Vespula intermedia 53 (no insecticide) traps 91
Vespula vidua 53 Fish/wetting agent/water traps 91
Uncertain status of Vespula Repellent jackets 92
squamosa and V, sulphurea . 55 Control summary 92
Vespula squamosa 56 Acknowledgments 93
Vespula sulphurea 58 Literature cited 94
Vespula vulgaris species group 59 Appendix 100
Vespula flavopilosa 61 Glossary of selected terms used
Vespula germánica 61 in the text lOO
Vespula maculifrons 65 List of chemicals mentioned in this
publication 102
The Yellowjackets of
America North of
Mexico
BY
ROGER D. AKRE, entomologist, Department of Entomology, Washington
State University, Pullman, Wash.
ALBERT GREENE, research associate, Department of Entomology, University
of Maryland, College Park, Md.
JOHN F. MACDONALD, assistant professor^ Department of Entomology,
Purdue University, West Lafayette, Ind.
PETER J. LANDOLT, research assistant, Department of Entomology,
Washington State University, Pullman, Wash.
HARRY G. DAVIS, entomologist, U.S. Department of Agriculture,
Science and Education Administration, Yakima, Wash.
INTRODUCTION
This handbook was written in response to a The most advanced social organization in this
growing need for a general manual on identifica- family is found in the subfamily Vespinae, which
tion, biology, and control of North American includes the yellowjackets and hornets.
yellowjackets. With the continually increasing The term "yellowjacket" seems to be of
popularity of outdoor recreational activities, American origin (Europeans simply call them
more and more people are becoming familiar with wasps). It properly refers to all members of the
these abundant, brightly colored, moderately genera Vespula Thomson and Dolichovespula
large to large wasps, possessing a fiery sting Rohwer, although several of these insects are
seemingly all out of proportion to their size. The white and black rather than yellow and black, and
persistence of some species in partaking of both two northern species are also marked with red.
our food and our garbage, as well as the construc- They are nearly ubiquitous throughout North
tion of many nests in locations with a high prob- America with the exception of the major desert
ability of accidental and unfortunate human areas and the uppermost latitudes with very cool
discovery, often result in a pest status that summers of 6° to 10 °C (43° to 50 °F) during the
ranges from mild nuisance to severe hazard. warmest month (fig. 1). True hornets are closely
All true wasps belong to the order Hymenop- related but are much larger Old World wasps in
tera (along with ants and bees), and have a fully the genus Vespa L.; one introduced species occurs
developed sting. Of the 4,000 wasp species in the in eastern North America, and is included in this
United States (Evans and Eberhard, 1970)' and handbook.
15,000 in the world (Hurd, 1955), most are It is small comfort to know that not only man,
solitary and inoffensive insects that use their but other animals have problems with vespine
sting to paralyze their arthropod prey. Fully wasps, as the following passage from Corbett
social (eusocial) behavior in wasps is Umited
(1955) makes clear: ''On reaching the kill the tiger
almost exclusively to the family Vespidae, whose
started blowing on it. In the Himalayas, and
members use their sting primarily for defense.
especially in the summer, kills attract many hor-
^The year in italic, when it follows the author's name, refers nets, most of which leave as the first light fades
to Literature Cited, p. 94. but those that are too torpid to fly remain, and a
2 AGRICULTURE HANDBOOK 552, U.S. DEPARTMENT OF AGRICULTURE
tiger—possibly after bitter experience—blows off fected, receiving multiple stings about the mouth
the hornets adhering to the exposed portion of and refusing to feed for 4 to 6 hours.
the flesh before starting to feed." During a 1973 What may be the first mentions of human con-
outbreak of yellowjackets in the Pacific North- tact with vespines are references to fear of
west, great masses of the wasps aUghted on both hornets by Middle Eastern peoples in the Old
liuman and animal food in the Portland, Oreg., Testament of the Bible. "Even today, the hornet
zoo. The Uons and tigers were particularly af- in the Middle East is a pest to be feared, their
very cool
summers
major
desert areas
FIGURE 1 ~ Map of North America showing areas of very cool summers with the mean temperature of the warmest month
6** to 10 **C (43° to 50 °F). Yellowjackets seldom occur in these areas. Also shown are desert areas and moun-
tain ranges, which influence vespine occurrence and distribution.
YELLOWJACKETS OF AMERICA NORTH OF MEXICO
ravages of fruit crops and honeybee colonies mak- of a serious allergic reaction to stinging are ex-
ing horticulture and agriculture at times difficult, tremely similar to those of heart attacks and are
at other times virtually impossible" (Spradbery, undoubtedly often mistaken as such.
1973a). As this handbook will demonstrate, yel-
lowjackets in North America are no less an eco- The unsavory reputation of yellowjackets is
nomic problem. In the early 1960's, the U.S. De- therefore, well-deserved, and in many cases
partment of Agriculture (USDA) received 10,000 justifies control measures (although the methods
requests per year for information on these wasps currently available often fail to deal adequately
(Fluno, 1961), and the number today is probably with the problem). This handbook, however, will
far greater. Yellowjackets are responsible for emphasize the paradox that yellowjackets are
dramatic drops in park, campground, and resort also beneficial insects, which appear to be key fac-
attendance, curtailment of logging operations, tors in the natural control of many pestiferous in-
and the impairment of both fruit harvesting and sects. Lastly, it is hoped this work will stimulate
forest fire fighting. Yellowjackets probably cause interest in yellowjackets for their own sake, as
far more than the reported 15 to 20 deaths per some of our most fascinating and dynamic insect
year in the United States, because the symptoms fauna.
LITERATURE REVIEW
A world list of literature useful to researchers and Wilson (1971), The Iwata reference sum-
on yellowjacket biology was published by Akre et marizes information on many species of Far
ál.(1974). The 717 entries were coded as to con- Eastern wasps previously available only in
tents. Other useful references are the books on Japanese.
wasps by Kemper and Dohring (1967) (in
German), Evans and Eberhard (1970), Guiglia Akre and Davis (1978) reviewed the ecology
(1972) (in French), and Spradbery (1973a), The and behavior of the Vespidae, while Davis (1978)
Kemper and Dohring and Guiglia references con- reported on problems caused by yellowjackets in
tain information only on vespids, whereas the the urban environment. MacDonald et al. (1976)
other two references also include information on discussed the status of yellowjacket control pro-
other families of wasps. Three additional authors grams in the United States. All other literature
with chapters or sections on wasps (including on yellowjacket biology is concerned with in-
yellowjackets) are Ebeling (1975), Iwata (1976) dividual species or specific investigations.
TAXONOMY
Introduction human contact is with the social Vespidae. Ves-

pidae and Eumenidae are distributed transcon-
The family Vespidae formerly included the tinentally, whereas Masaridae are found in the
subfamilies Masarinae, Eumeninae, and Ves- Western United States and southwestern
pinae (Bequaert, 1918\ Bradley, 1922), which Canada.
have since been elevated to family rank (Broth- Most Masaridae are solitary, yellow and black
ers, 1975\ Richards, 1962). They now constitute wasps that provision their mud cells with pollen
the three families of the Vespoidea. Vespoid and nectar rather than arthropod prey. They are
wasps are recognized by the elongate discoidal easily separated from other Vespoidea by clubbed
cell in the forewing and the longitudinal folding antennae and the presence of two, rather than
of the wings at rest (with the exception of most three, submarginal cells in the forewing. Masa-
Masaridae). Members of all three vespoid fami- rids are relatively scarce in most localities and
lies are found in North America, although most rather innocuous.
AGRICULTURE HANDBOOK 552, U.S. DEPARTMENT OF AGRICULTURE
Eumenidae, also solitary and harmless to family includes the large and diverse tribes,
humans, are predaceous and make a variety of Ropalidiini and Polybiini, and the monotypic
nest types in soil, in twig cavities, or on various tribe that contains the cosmopolitan genus Pol-
surfaces. They are recognized by a combination istes Latreille (Polistini). Richards (1978) revised
of characteristics: Middle tibiae with one apical the American species and subspecies of Polistinae
spur, toothed or bifid tarsal claws, filiform and summarized information on their biology.
antennae, and three submarginal cells in the The Ropalidiini are found only in Africa and
forewing. Australasia (fig. 3). The Polybiini are primarily
The Vespidae, consisting of the subfamilies Neotropical (23 genera), with three additional
Stenogastrinae, Polistinae, and Vespinae, have genera found in Africa and Asia, and four (or pos-
two apical spurs on the middle tibiae, simple tar- sibly six) species distributed into temperate lati-
sal claws, and three submarginal cells in the tudes in North America (fig. 4).
forewing. The stenogastrines are found only in Mischocyttarus flavitarsis (Saussure) is found
Southeast Asia (fig. 2). Their small colonies are in the Western United States and into southern
found in deep forests, and most species are pre- British Columbia. It is similar in appearance to
social (Spradbery, 1975), Van der Vecht (1977) dis- Polis tes but can be readily distinguished by the
cussed the phylogeny and taxonomic status of narrow, stalklike first gastral segment. Nests
the Stenogastrinae and tentatively concluded are small and similar in architecture to Polis tes
they should either be a separate family or a sub- nests. Like Polistes, these wasps may be found
family of Eumenidae. hibernating in large numbers in buildings and
The Polistinae are eusocial wasps. This sub- attics of houses. A related species, M cubensis
FIGURE 2 — Distribution of Stenogastrinae (drawn by G. Shinn).

ROPALIDIINI Ropalidia M
FIGURE 3 - Distribution of Ropalidüni (G. Shinn).
FIGURE 4 - Distribution of Polybüni (G. Shinn).

(Saussure), occurs from Florida to South Caro- Colony size is small with usually less than 200
lina, whereas M mexicanas (Saussure) occurs in adults reared through the season. More detailed
southern Texas. (Richards, 1978, raised M caben- accounts were given by Evans and Eberhard
sis mexicanas to species level.)
(1970), West Eberhard (1969), and Wilson (1971).
Brachygastra mellifera (Say) has a Neotropical Paper wasps are generally not pests unless the
distribution but also extends north into south- nest is disturbed or large numbers of females,
ern Texas and Arizona. Their large and populous looking for suitable overwintering sites, find
paper nests are comprised of concentric combs their way into buildings in autumn. Their preda-
covered by a single paper envelope (Bequaert, tory habits, especially on caterpillars, make
1933), They are known for their honey-storing them beneficial to gardeners and farmers.
behavior. In addition, two species of Polybia The Vespinae (Provespa Ashmead, Vespa.
have been collected (or intercepted in quaran- Vespula, Dolichovespala) occur throughout the
tine) at Nogales, Ariz., on the U.S. border. Northern Hemisphere and in Southeast Asia
Polistes (paper wasps) occur throughout most (fig. 6). The three species of Provespa are found
of the world (fig. 5). They make a single, usually only in Southeast Asia and are nocturnal (van
horizontal, comb with no surrounding envelope. der Vecht, 1957). Vespa is primarily eastern
Colonies are founded in spring by one or more Asian with two species, Vespa orientalis Fab.
overwintered females. One female becomes the and V. crabro L., found in the western Palearctic.
dominant queen and egg layer while the re- K crabro has also been introduced into the East-
mainder function as subordinate workers (Pardi, ern United States. In this handbook, the yellow-
1948), Emerging females are workers until later jackets are considered to belong to two distinct
in the season when reproductives are produced. genera, the generally ground-nesting Vespula
POLISTINI Polistes
FIGURE 5 - Distribution of Polistini (G. Shinn).

Vespula/Dolichovespula M
Vespa WMi
VESPINAE Provespa lililí
FIGURE 6 — Distribution of Vespinae (G. Shinn).
and the generally aerial-nesting Dolichovespula. Although the Vespinae are a well-defined and
Both are Holarctic. In addition, Vespula ger- easily recognized group, they are often confused
mánica (Fab.), a European species, has been with paper wasps in the genus Polistes. Paper
introduced into New Zealand, Australia, South wasps can be readily distinguished by their elon-
Africa, and Chile, as well as the Eastern United gate legs and larger, more slender bodies (fig.
States (Edwards, 1976), Vespula vulgaris (L.), a 76). Their nests always consist of a single, ex-
Holarctic species, has been introduced into posed comb (fig. 8a, 6), suspended by a narrow
Australia (Spradbery, 1973b). stalk and often located under eaves, porches, or
other manmade structures. Yellowjacket nests,
even very young ones, have a globular or oval
Identification of Genera and Species
paper envelope that usually encloses multiple
Worker yellowjackets are fairly small wasps combs (fig. 9).
The following pictorial key will serve to identi-
about 10 to 14 mm (1/2 inch) long (fig. la) and
are strikingly marked with black and yellow or fy workers of the 16 species of free-living yellow-
black and white patterns. In the United States, jackets and the single species of hornet found in
the term **hornet" is generally applied to two North America. (Two parasitic yellowjackets,
much larger species in the same subfamily (Dol- Dolichovespula árctica (Rohwer) and Vespula
ichovespula maculata (L.), Vespa crabro), and austriaca (Panzer), do not have a worker caste,
sometimes to all aerial-nesting yellowjackets. and utilize workers of the host species to care
Technically, however, only species in the genus for their own brood). Figure 10 illustrates those
Vespa (of which V. crabro is the only North structures referred to in the key. Except for the
American example) are hornets. recognition of Vespula and Dolichovespula as
3
=3
o
ï:3
Oí
O
identification in eastern North America is compli-

cated by the fairly close resemblance of four spe-
cies, all of which may occur in some areas: Vespu-
la vulgaris, V. maculifrons (Buysson), V. germán-
ica, and V. flavopilosa Jacobson.
Until recently, only the first three species were
recognized, with the differences between them
elaborated in detail in Menke and Snelling {1975);
however, morphological, biological, and electro-
phorectic studies have now revealed a previously
unrecognized form, V. flavopilosa, which has long
been confused with the closely related V. vulgaris
and V. maculifrons (Jacobson et al., 1978; figs. 10,
11, and 16 of Menke and Snelling, 1975, labeled V.
maculifrons, are V. flavopilosa). Much of this con-
fusion was due to the large amount of color pat-
tern dimorphism that exists between V. flavo-
PN-6527
pilosa queens and workers, making asssociation
of individually collected females difficult. For in-
stance, V. flavopilosa queens sometimes closely
resemble those of V. vulgaris, whereas the
gasters of highly xanthic V. flavopilosa workers
may be practically indistinguishable from those
of melanic V. germánica. Examination of V.
flavopilosa from over 25 nests, however, reveal-
ed stable color patterns among females and
males both within and among all colonies exam-
ined from Georgia, North Carolina, Pennsylvan-
ia, and New York. In addition, comparative elec-
trophoretic studies showed consistent enzymatic
differences between V. flavopilosa and the three
FIGURE 8 — a, Polistes queen on her nest; b, Polistes nest

showing single, exposed comb (J. Krispyn).
separate genera, the taxonomic treatment of

Miller (1961) has been followed. Most species are
quite distinct and often can be determined on
the basis of gaster color pattern alone; however,
since these markings always vary to some degree
and may appear similar in some closely related
species, identifications should always be made on
a series of specimens when possible. Two sources
of confusion are commonly encountered: (1) The
color patterns of worker Vespula acadica (Sladen)
and V. atropilosa (Sladen) vary widely and often
bear a superficial resemblance to each other. A
complete range of patterns may be found within
one colony, although certain colonies tend to pro-
PN-6528
duce either xanthic (very yellow) or melanic (very FIGURE 9 — Nest of DoUchovespula maculata with half of
black) individuals. (2) Similarly, yellowjacket envelope removed to expose the multiple combs.
VERTEX
OCELLI
COMPOUND
EYE
OCULO-
MALAR
SPACE --
ANTENNA
MANDIBLE
FIGURE 10 —Worker yellowjacket illustrating structures used in the key, pages 12 t« 22.
eastern species of the V. vulgaris group as well likely arose as such), it is significant that V.
as a western member, V. pensylvanica (Saus- flavopilosa exists farther south than the ap-
sure) (Jacobson et al., 1978). V. flavopilosa main- parent southernmost distribution of V. vulgaris
tains its identity sympatrically (occurring in in the Eastern States.
same geographical area) with V. maculifrons Since the key was constructed, Wagner {1978)
throughout its range, and with V. vulgaris and presented evidence to show that Dolichovespula
V. germánica in its northern distribution. Since saxonica (Fab.), previously considered Palearc-
V. flavopilosa could conceivably be a hybrid be- tic, is widely distributed in the Nearctic. This
tween V. maculifrons and V. vulgaris (and most species (as defined) is closely related to, and eas-
YELLOWJACKETS OF AMERICA NORTH OF MEXICO 11
ily confused with, D. norvegicoides (Sladen). As nest at Eagle River, Alaska. No collection dates
part of the evidence, he reported on two colon- were given for either colony. The first nest was
ies, one with a nest in a wall void at King's reported to have a tiered envelope similar to
Mountain, Alaska; the other a supraterrestrial nests of D. saxonica in Europe, whereas the sec-
(on the surface of the ground in a hollow cavity) ond nest had an interconnecting envelope that
FIGURE 11 — a, Occipital carina extending to base of mandible (Vespula vulgaris group species); b, occipital carina ending
above base of mandible (Vespula rufa group species).
formed a spongy fibrous mass rather than Therefore, before deciding on the validity of
distinct sheets. The latter nest was 10.7 cm in the occurrence of D. saxonica in North America,
diameter and had one worker comb and one we would like to see more evidence comparing
reproductive comb. The number of combs was nest architecture, series of workers, males, and
not given for the first nest. Both nests were queens, and behavior of colony members of D.
small, the first contained the foundress queen, saxonica and D. norvegicoides.
55 workers, and a D. árctica (Rohwer) female; D. saxonica is not included in the key.
the second lacked a foundress queen and con-
tained 41 workers, 11 new queens, and 26 males.
Additional evidence used to support the conten- Identification of Members of Vespula
tion that D. saxonica occurs in North America Species Groups
was based on morphological analyses of work-
ers, queens, and males. One critical character The species of Vespula are divided into the V.
used in identifying the species was based on the vulgaris species group (scavengers, often pestifer-
ratio between head-width and the length of the ous) and the V. rufa (L.) species group (strictly
oculo-malar space. A second distinguishing fea- predators on live prey, usually nonpestiferous).
ture was the sequence of tyloides on male anten- These groups are discussed more fully in the ''bio-
nae, while the last concerned the color patterns logy" section. Although the separation between
on abdominal terga II to V. them is based on morphological features as well
An examination of European literature on D. as biological ones, these characteristics are diffi-
saxonica revealed a number of discrepencies cult to see and interpret. Since they are not essen-
with the above report. For example, the nest tial for species identification, they have not been
architecture reported by Kemper and Dohring included in the key, and are given separately in
(1967, fig* 31) does not show a tiered envelope, this section.
nor did Lóken (1965) report this as a character- Members of the V. vulgaris group have the
istic oiD, saxonica nests. Lóken (1964) also gave occipital carina (a ridge on the rear of the head)
a different tyloide sequence for males, In addi- well developed throughout its length and
tion, the gaster color pattern of D. saxonica il- extending to the base of the mandibles (fig. 11
lustrated by Kemper and Dohring (1967) is a). In addition, the first gastral tergum is not
vastly different from that mentioned by Wagner narrowed anteriorly and is not depressed. Its
(1978) or illustrated by Yamane (1975). hairs are usually pale gray or yellow.
Evidently, there are also problems or areas of Members of the V. rufa group have the occi-
confusion in Norway concerning the relationship pital carina prominent dorsally but ending well
between D. saxonica and the closely related D. above the base of the mandibles (fig. 11 b). The
norwegica (Fab.) as Lóken (1964) concluded her first gastral tergum is narrowed anteriorly with
discussion of D. saxonica with (we) ''need more a slight depression behind its anterior margin.
investigation to settle the status of this wasp." The hairs on this tergum are black.
We also measured a series of seven workers The occipital carina is almost completely con-
from a D. norvegicoides nest from Montana, cealed on intact specimens because of the dense
which showed two of these workers had head- hair on the head and the difficult viewing angle.
width to oculo-malar space ratios intermediate It can be best seen (under magnification) by
in value between D. saxonica and D. placing the severed head in clear fluid, such as
norvegicoides as given by Wagner (1978). alcohol or water.
Key to Vespinae in North America

Key to Genera
1. Vertex extending greatly above compound eyes; distance between lateral ocellus and occipital
carina much greater than distance between lateral occelli (fig. 12a); one very large (>20mm),
brown and yellow species.... genus Vespa L.
Vertex not or only slightly extending above compound eyes; distance between lateral ocellus
and occipital carina about the same as distance between lateral ocelli (fig. 126) 2
VERTEX
FIGURE 12.
2 Oculo-malar space narrow, compound eyes touching or nearly touching base of mandibles (fig.
13a) genus Vespula Thomson
Oculo-malar space broad, compound eyes remote from base of mandibles (fig. 136) .... genus
Dolichovespula Rohwer
OCULO-MALAR
SPACE
FIGURE 13.
Key to Species
Vespa
Gaster resembling figure 14 y. crabroL. (European hornet)
FIGURE 14.
DoUchovespula
1. Dorsal surface of first 3 terga entirely black (fig. 15a); large species (>15mm) ^^^^^^^^^^''^^^¡¡^^
First 3 terga with pale markings; smaller species «15mm)
2 Pale markings white; usually with reddish spot on sides of tergum II (fig. 1^6); rarely occur-
' ring south of Alaska and northern Canada D, albida (Sladen)
Pale markings yellow
FIGURE 15.
3. Yellow genal band continuous (fig. 16a), though sometimes deeply notched; yellow of terga I
and II sharply incised, usually interrupted medially (fig. 166); common D. arenana (Fab.)
(aerial yellowjacket)
Yellow genal band usually widely interrupted (fig. 16c); yellow of terga I and II not sharply in-
cised or interrupted medially (fig. 16d), but pattern sometimes closely approaches that of D,
arenaria; not common /). norvegicoides (Sladen)
QXP
FIGURE 16.
Vespula
Pale marking white 2

Pale markings yellow 3
Terga I and II with reddish markings (fig. 17a); rarely occurring outside Alaska and northern
Canada V, intermedia (Buysson)
Gaster without reddish markings; white band on posterior margin of tergum I usually inter-
rupted medially (fig. 176) V. consoftrina (Saussure) (blackjacket)
FIGURE 17.
3. Two yellow stripes on scutum (fig. 18a) 4

Scutum without stripes (fig. 186) 5
FIGURE 18.
4. Caster resembling figure 19a; not occurring west of the Rocky Mountains
V. squamosa (Drury) (southern yellowjacket)
Gaster resembling figure 196; not occurring east of the Rocky Mountains
V. sulphurea (Saussure) (CaUfornia yellowjacket)
FIGURE 19.
5. Usually with a continuous yellow ring dorsally around each compound eye (fig. 20a); medial
black mark on tergum I usually diamond shaped (fig. 206) V. pensylvanica (Saussure)
(western yellowjacket)
Head without a continuous yellow ring dorsally around each compound eye (fig. 20c) 6
FIGURE 20.
6. First antennal segment yellow ventrally (fig. 21a) 7

First antennal segment without yellow, usually entirely black (fig. 216), sometimes brownish
distally 9
7. Gaster resembling figure 21c; extensive black region on tergum II never with 2 yellow spots;
often with free, or nearly free, black spots on terga IV and/or V; not occurring west of the
Rocky Mountains V. vidua (Saussure)
Gaster not as in figure 21c 8
FIGURE 21.
8. Gaster resembling figure 22a, the 2 yellow spots on tergum II often faint or absent, some-
times expanded (fig. 226); medial margin of black region on tergum II broadly rounded; terga
III, IV, and V usually without free black spots; yellow genal band usually interrupted (fig
22c), rarely continuous y acadica (Sladen)
Gaster resembling figure 22e, or with black regions expanded, varying to fig. 22(1; when black
regions expanded, medial margin of black region on tergum II tapering sharply; terga III, IV
or V usually with some free, or nearly free, black spots; yellow genal band usually continuous
(fig. 22/), sometimes with central black spot, rarely with narrow interruption; occurring prin-
cipally west of the Rocky Mountains V. aíropi7osa(Sladen) (prairie yellowjacket)
FIGURE 22.
Yellow genal band interrupted with black (fig. 236), sometimes only slightly; gaster resemb-
ling figure 23a, tergum I varying as shown V. vulgaris (h,) (common yellowiacket)
Yellow genal band continuous (fig. 23c) 1^
FIGURE 23.
10. Gaster usually resembling figure 24a, black regions sometimes expanded, varying to figure
246; tergum II usually with free, or nearly free, black spots; medial black mark on tergum I
usually diamond shaped V. germánica (Fah,) (German yellowjacket)
Gaster resembling figures 24c or 24d; tergum II usually without 2 completely free black
spots; not occurring west of the Rocky Mountains
IL Medial black mark on tergum I usually anchor-shaped (fig. 24c), although varying in width;
common V. maculifrons (Buysson) (eastern yellowjacket)
Medial black mark on tergum I usually V-shaped (fig. 24d), although varying in width; not
common V. flavopilosa Jacobson (hybrid yellowjacket)
FIGURE 24.
Yellowjacket and Hornet Worker correspond exactly with the drawings, and
Gaster Patterns Drawn to Scale (fig. 25) extreme variant specimens may be encountered
that will not be readily identifiable. The practice
This section is intended as a quick reference of collecting a series of specimens whenever pos-
guide to the gaster patterns of worker yellow- sible will greatly facilitate identification of such
jackets and hornets, and not as a substitute for
individuals.
the preceding key. Very few specimens will
D. arenaria D. norvegicoides
Vespa crabro Dolichovespula
maculata
Vespula V. suiphurea V. intermedia V. vidua

Squamosa
V. acadica- V. acadica- V. atropiiosa- V. atropitosa- V. pensylvanica

melanic xanthic melanic xanthic
V vulgaris V. maculifrons V. flavopilosa V. germanicax- V. germánica-

anthic melanic
FIGURE 25 — Yellowjacket and hornet worker gaster patterns drawn to scale.

Vespa
tern of these 2 species highly variable; mel-
anic form of V. acadica without 2 yellow
V. crabro L. — Very large (>20 mm); dark brown
spots on second tergum resembles V. vidua,
and deep yellow; anterior third of the first
but V. acadica almost never has free black
tergum reddish brown.
spots on fourth and fifth terga; melanic form
DoUchovesp ula of V. acadica with 2 yellow spots resembles
melanic V. atropilosa, but medial margin of
D maculata (L.) — Large (>15 mm); gaster black region on second tergum is broadly
mostly black; white markings on last 3 rounded in V. acadica, sharply tapering in
terga. V. atropilosa; xanthic forms of V. acadica
and V. atropilosa resemble each other, but
D. albida (Sladen) — Black and white; usually
with lateral reddish spots on first 2 terga. V. atropilosa usually has some free or nearly
D, arenaria (Fab.) — Yellow of first terga free black spots on third, fourth, and fifth
terga.
sharply incised, usually interrupted medially.
D. norvegicoides (Sladen) — Yellow of first 2 V. pensylvanica (Saussure) — Diamond-shaped
terga continuous; yellow of most terga black mark on first tergum; similar to mel-
usually not as sharply incised as in D. aren- anic K germánica, but V. pensylvanica has
aria, continuous yellow ring dorsally around each
compound eye.
Vespula V. vulgaris (L.) — ResembHng V. maculifrons or
V. flavopilosa when yellow on first tergum is
V. squamosa (Drury) — Distinctive pattern as expanded, but V. vulgaris has yellow genal
illustrated; wide black ''hourglass" marking band interrupted with black.
on second tergum. V. maculifrons (Buysson) — Distinctive anchor
V. sulphurea (Saussure) — Distinctive pattern or arrow-shaped medial black mark on first
as illustrated; 2 yellow stripes on scutum tergum as illustrated.
distinguish V. sulphurea and V. squamosa V. flavopilosa Jacobson — ResembHng V. macu-
from all other species. lifrons, but medial black mark on first ter-
V. intermedia (Buysson) — Black and white; gum of most V. flavopilosa is V-shaped, with-
large red areas on first 2 terga. out the very narrow *'neck" of the mark on V.
V. consobrina (Saussure) — Black and white; maculifrons. Highly xanthic V. flavopilosa
white band on posterior margin of first may have 2 free black spots on second ter-
tergum usually interrupted medially. gum, and therefore closely resemble V, ger-
V. vidua (Saussure) - Extensive black region on mánica.
second tergum never with 2 yellow spots; V. germánica (Fab.) - Diamond-shaped black
usually with free or nearly free black spots mark on first tergum; usually with 2 free or
on fourth and fifth terga. nearly free black spots on second tergum.
V, acadica (Sladen)/V. atropilosa (Sladen) — Pat- Highly melanic form is not common.
BIOLOGY
General sis). Prey usually consists of live arthropods, al-

though several members will scavenge flesh
Vespidae (Stenogastrinae, Polistinae, Vespin- from dead animals.
ae) are extremely diverse (Akre and Davis, 1978). All colonies of Vespinae (Pro vespa, Vespa,
Behavior varies from presocial to eusocial. Most Vespula, Dolichovespula) are initiated by a sin-
yespids build nests of masticated vegetable fiber gle queen, and all build nests of paper carton.
initiated by a single queen (haplometrosis) or by The nests consist of a number of rounded combs
a number of queens that cooperate (pleometro- attached one below another. The combs are
usually covered with a many-layered envelope.

Some species primarily construct subterranean
nests, some aerial nests, and others utilize either
location. Colonies vary in size from less than a
hundred individuals to several thousand.
The Vespinae are the only wasps known to con-
struct special cells on the combs to rear queens.
Since there is only a single species of Vespa
(hornets) in North America, biological informa-
tion is included with the species discussion.
Newly produced yellowjacket [Vespula,
Dolichovespula) queens are the only members of
the colonies to survive the winter. These queens
overwinter in sheltered locations, such as under
loose tree bark or in decaying stumps, and
emerge from hibernation during the first warm
days of spring, usually in April or May (fig. 26).
Soon after emerging, they begin to feed on
flowers and other sources of nectar, and will also
catch and malaxate arthropod prey. Ovaries of
these queens are very small, but grow rapidly as
reproductive diapause is broken. During this
period, many queens are seen seeking nest sites,
alighting under the eaves of houses, or entering
cracks and crevices. After selecting a suitable
location, the queen gathers plant fibers to con-
struct the first cells of the nest. This fiber is
usually gathered from weathered wood but may
be collected from decayed wood or even Hving
plants.
The queen nest (fig. 27) ultimately consists of
IMÊÊ
PN-6530
FIGURE 27 — a, Queen nest; Vespula pensylvanica,- b, queen

PN-6529
nest; Dolichovespula arenaria; c, nest of V. atropilosa with
FIGURE 26 — Vespula pensylvanica queen in typical dia- queen and first workers; queen on edge of comb in bottom
pause position. of picture.
20 to 45 cells covered by a paper envelope. The

or near shrubs and other vegetation. Mating
queen lays eggs in the cells as they are construc-
may enhance successful overwintering, at least
ted and forages for nectar and arthropod prey to
in Vespula (MacDonald et al., 1974). The next
feed the developing larvae. In about 30 days, the
spring the cycle is repeated (fig. 29). More
first 5 to 7 workers emerge and assume such
detailed explanations of this cycle are found in
duties as cell and envelope building, foraging for
Duncan {1939), MacDonald et al. (1974) and
prey and nectar, and nest sanitatation (Akre et Spradbery (1973a).
al., 1976), After a short period of additional Yellowjackets do not store honey as do the
foraging, the queen no longer leaves the nest, bees and some of the other vespids. They feed
and her primary function is to lay eggs. Workers their larvae meat (arthropods, especially insects)
are much smaller than the queen, and under nor- and probably nectar and honeydew (a sugary
mal colony conditions they do not lay eggs. material excreted by plant lice and certain other
Until this time, colony growth is slow but be- plant-feeding insects). The adults feed on juices
comes exponential by midsummer as successive while malaxating prey in the nest. They also eat
broods of workers emerge. The general, multi- nectar and larval secretions. The exchange of ali-
tiered architecture of the vespine nest is appar- mentary liquids among colony members (trophal-
ent at this time with horizontally arranged laxis) is a prominent activity in the colony.
combs comprised of ventrally open cells, typical- Yellowjackets prey on a wide variety of in-
ly enclosed in a paper envelope. sects, spiders, other arthropods, and even mol-
Later in the season, the workers start building lusks to feed their larvae. All species of Dolicho-
the larger reproductive cells in which both males vespula will usually attack only Hve prey al-
and queens are produced; males are also reared though there are a few records of workers
in worker cells. The colony enters a declining scavenging from animal carcasses (Greene et al.,
phase shortly thereafter when workers pull lar- 1976). Vespula spp. are frequently divided into
vae from the comb and feed them to other larvae the V. vulgaris species group (genus Paraves-
pr discard them. During this period, in late sum- pula Blüthgen of European workers) and the V.
mer or autumn, workers of several species are rufa species group (genus Vespula of European
much more likely to sting, even when away from workers) (tables 1, 2; fig. 30; Bequaert, 1931;
the nest. When new queens and males emerge, MacDonald et al, 1974 and 1976). Members of the
they leave the nest, mate (fig. 28), and the fertil- V. rufa group have a shorter colony duration and
ized queens hibernate. Large numbers of males smaller colonies than members of the V. vulgaris
are often seen at this time swarming on hilltops group (fig. 31, table 3). More importantly, V. rufa
FIGURE 28 — Mating of Vespula pensylvanica; queen on left.

FIGURE 29 — Yellowjacket life cycle (Vespula pensylvanica): a. Mating; 6, fertilized queen in diapause during winter
months; c, queen nest beneath soil surface; d, nest at peak of colony development (J. Krispyn}.
group species forage only for live prey, whereas Zealand, where perennial colonies are not un-
members of the V. vulgaris group will also scav- common (Thomas, 1960). One nest was nearly 15
enge from animal carcasses, picnic tables, gar- ft tall, probably contained several million cells in
bage cans, and many other locations, which about 180 combs, and weighed an estimated
makes them pestiferous to man (MacDonald et 1,000 lb. Although the adult inhabitants of
al, 1976). these nests must have numbered into the tens of
Vespula nests are usually subterranean, al- thousands, counts were not made. For compari-
though some species will build their nests in hol- son, most nests of V. vulgaris group species
low logs, trees, in attics, between walls, or on contain 1,000 to 4,000 workers. Dolichovespula
eaves of houses (Duncan, 1939; Ebeling, 1975; nests are usually aerial, although all species
Green et al., 1970; Kemper and Dohring, 1967; probably nest underground on occasion and at
Spradbery, 1973a). Nest size varies from 300 to least two species (the Nearctic D. arenaria (Fab.)
120,000 cells, although most nests have 2,000 to and the Palearctic D. sylvestris (Scopoli)) may do
6,000 cells and are 8 to 15 cm in diameter. The this frequently (Greene et al., 1976; Archer,
largest nests are those of atypically perennial 1977a; Spradbery, 1973a). Most nests are small
colonies, which do not die out over winter but (300 to 1,500 cells), but those of vigorous D.
continue on for an additional season with arenaria colonies may contain up to 4,300 cells
multiple queens. For example, the nest of one and those of D. macúlala up to 3,500 cells.
such V. vulgaris colony in California was nearly A comparison of D. arenaria with V. pensyl-
4 ft long (Duncan, 1939); a V. squamosa (Drury) vanica (V. vulgaris species group) and V. atropi-
nest in Florida was 9 ft tall (Tissot and Robin- losa (V. rufa species group) is given in table 3.
son, 1954); however, the record for any vespine Although general comparisons cannot be made
species clearly belongs to V. germánica in New between Vespula and Dolichovespula, because
the necessary information is not available, some arthropods (table 1). Nearly all nests are subter-
of the characteristics compared in the table will ranean or in decaying logs or stumps
probably hold true for many species. D. arenaria (MacDonald et al., 1974, 1975a), The nest of
colonies (Greene et al., 1976) and at least some members of this group differs from that of the
European species of Dolichovespula (Spradbery,
V. vulgaris group by having broad supporting
1973a) often build their nests on the eaves of buttresses, a strong flexible envelope paper, and
houses or in other locations that bring them into a single comb of worker cells (table 2;
contact with man, and, therefore, may some- MacDonald et al., 1974; Spradbery, 1973a). The
times be regarded as general nuisances; how- nests of several species are also quite filthy and
ever, most Dolichovespula have small colonies, unkempt with remains of prey and dead workers
do not scavenge at picnic tables, and are not incorporated into the envelope. Yellowjackets of
troublesome unless their nest is disturbed. the V. rufa group are usually considered highly
Members of the K rufa group have small beneficial because of their prédation on a num-
colonies of 75 to 400 workers, nests of 500 to ber of economically important insects
2,500 cells, and are strictly predaceous on Uve (MacDonald et al., 1976); however, even these
TABLE 1,-Species groups, distnbution, and colony charactenstics of the subgenus Ves-
pula (subterranean nesting yellowjackets) in North Amenca. {Modified from
MacDonald et al., 1976)
Colonies
Species groups Distribution sampled Group colony parameters
Vespula vulgaris group
vulgaris Holarctic; ^59 Size: 500 to 5,000 workers at peak.
transcontinental
maculifrons Eastern ^78 Nest: 3,500 to 15,000 cells.
pensylvanica Western
Workers: Predators plus scavengers.
germánica Palearctic; now eastern •30 Decline: Late Sept. to Nov.-Dec.
North America
flavopilosa Eastern ^23
Vespula rufa group
acadica Transcontinental 54
Size: 75 to 400 workers at peak.
atropilosa Western •MO Nest: 500 to 2,500 cells.
consobrina Transcontinental Workers: Strictly predators.
vidua Eastern no Decline: Late Aug. to early Sept.
intermedia Transcontinental (Boreal) 0
aus triaca Holarctic, transcontinental 0 (Unknown, social parasite of V. acadica)
Uncertain affinities
(currently placed with
the V. rufa group)
squamosa Eastern, southeastern to no3 Size: 500 to 4,000 workers at peak.
Central America (social Nest: 2,500 to 10,000 cells.
parasite of V. maculifrons) Workers: Predators, reported to be
scavengers.
Decline: Late Sept. to late Nov.
sulphurea Western; southern Oregon, Unreported; based on Duncan {1939)
California into northern Mexico appears to exhibit a number of V.
vulgaris group characteristics.
'Spradbery, 1971. 'MacDonald et al., 1975a.
'MacDonald and Matthews, 198â.
^MacDonald and Matthews, 1976.
^MacDonald et al., 1974. 'MacDonald and Matthews, 1975.
'MacDonald, Matthews, and Jacobson, 1978. «MacDonald and Matthews, 198_b.
TABLE 2.—Nest architectural characteristics of sampling of social wasp colonies is the variety
genus Yespula—subterranean nest- and quantity of arthropods associated with the
ing yellowjackets nest. Subterranean yellowjacket colonies, with
large accumulations of wastes and bodies of
Nest architectural parameters
dead individuals in the soil beneath the nest,
Species group
Large nest—ultimately
support several species of associates, often in
Vespula vulgaris
group 3,500 to 15,000 cells. very large numbers. Expectedly, associated
Several worker-cell combs. fauna in aerial nests is less varied and occurs in
All suspensoria cordlike. much smaller numbers. Spradbery (1973a) pro-
Fragile envelope.' vided a general treatment of associates. This
Scalloped envelope paper.
discussion concentrates on the most common
V. rufa group Small nest—ultimately
500 to 2,500 cells.
and abundant associates found with Nearctic
One worker—cell comb. yellowjacket species. Colony associates can con,-
Top suspensoria buttresslike. veniently be separated into those actually invad-
Pliable envelope. ing the nest proper of active colonies and those
Laminar envelope paper.
developing in the soil beneath subterranean
V. squamosa Large nest—ultimately nests or invading abandoned (or nearly aban-
2,500 to 10,000 ceUs.
Several worker-cell combs.
doned) nests.
All suspensoria cordlike. Active yellowjacket and hornet colonies
Pliable envelope. harbor very few nest invaders. The most
Laminar envelope paper. important of these is an ichneumon wasp,
'Envelope either gray {V. pensylvanica, V. germánica) or Sphecophaga vesparum burra (Cresson). This
tan(K vulgaris, V. flavopilosa, V. maculifrons). pupal parasite (parasitoid) is a frequent occu-
pant in nests of the V. rufa group, less common
and less abundant in Dolichovespula nests, and
yellowjackets are sometimes pestiferous to man rarely found in nests of the V. vulgaris group
as they frequently nest in manmade structures (with the exception of V. vulgaris nests, where it
or in the yards of homes. In all cases, the wasps occurs frequently). Adverse effects on incipient
are a stinging hazard only after the colony is V. atropilosa colonies may occur, but heavy
disturbed. parasitism is undocumented in other species.
Members of the V. vulgaris group have larger The only additional common nest invaders are
colonies, larger nests, and a longer colony dura- cockroaches (Parcoblatta Hebard spp.), which
tion (table 1). These yellowjackets are frequently occur in nests of V. squamosa in the southeast.
pestiferous. For example, V. pensylvanica These scavengers roam about the nest proper in
commonly nests near well-watered yards in the over 70 percent of V. squamosa colonies but
dry areas of the Pacific Northwest, and numer- elicit no response from their hosts. Indeed, the
ous telephone calls were received from home- cockroaches may assist in nest sanitation; a
owners during 1974-77 in Pullman, Wash., for close relationship with the yellowjackets is sug-
assistance in removing or killing colonies. Per- gested by the deposition of cockroach oothecae
haps more importantly, workers of all species (egg cases) inside the layers of the nest envelope
belonging to this group will scavenge for flesh (MacDonald and Matthews, 198 b). At least in
and sweets, which brings them into frequent the southeast, viable aerial colonies of D. macul-
contact with humans. This is especially true ata occasionally harbor cockroaches, but they
with the tremendous increase in recreational are a different species of Parcoblatta,
developments such as parks, roadside rests, re- In contrast to the paucity of nest-invading as-
sorts, and in activities such as backyard sociates, numerous associates are found in the
cooking, hiking, and camping. Concentrated nest cavity of subterranean colonies and the de-
garbage and other meat sources in some of these composing carton of abandoned and nearly
areas have created a vast food supply for these abandoned nests. Particularly productive is the
scavenger species. waste-soaked soil beneath subterranean nests,
One of the notable aspects of late-season which supports huge numbers of scavenger fly
larvae. The most prevalent and abundant possibly dying) queens; however, such invasion
species is Dendrophaonia querceti (Bouche) occurs in colonies in an advanced state of de-
(Diptera: Muscidae). Two other muscids, Fannia cline, and this species is not considered a signifi-
canicularis (L.) and F. scalaris (Fab.), are very cant mortality factor.
common, although the rather cryptic larvae are Appearance of scavenger associates is a late-
more difficult to detect. Eggs of all three flies season phenomenon associated with large nest
are laid on the outside of the nest envelope, from size and an accumulation of wastes and other
which the fly larvae drop into the soil beneath debris either inside the nest or in the soil be-
the nest. Occurring less frequently, but often in neath subterranean nests. In contrast, invaders
extremely large numbers inside a decomposing of active colonies appear early in the colony
subterranean nest, are larvae of Triphleba lugu- cycle and may be present throughout the nest-
bris (Meigen) (Diptera: Phoridae). Most likely ing season.
scavengers on dead brood in the nest, T. Among the vespine wasps are two obligate
lugubris invade cells containing developing (or social parasites, Dolichovespula árctica and Ves-
PN-6531
FIGURE 30 — Comparison of paper and nest suspensoria of Vespula atropilosa (V. rufa group species) and V. pensylvanica
(V. vulgaris group species). V. atropilosa has laminar paper (a) and mostly buttresslike suspensoria (c) V. pen-
sylvanica has scalloped paper (6) and cordlike suspensoria (d).
TABLE S.—Companson of nesting biology and behavior of 3 yellowjacket species. Data obtained from
nests collected in several northwestern States and observations of captive colonies in Pull-
man, Wash. {Modified from Greene et aL, 1976)
DoUchovespula arenaria Vespula atropilosa Vespula pensylvanica

Approximate colony duration 3 to 4 months 3 to 4 months 4 to 5 months.
Nest sites usually aerial, sometimes always protected, usually always protected, usually
protected or subterranean subterranean subterranean.
Envelope composition usually laminar laminar scalloped.
Queen nest pedical often coated with glossy
oral secretion uncoated uncoated.
Comb suspensoria buttresslike both types usually present cordlike.
Number of worker combs 1 to 4; often multiple 1 1 to 6; usually multiple.
Reproductive cells on worker often extensive on all limited except for final 1 or 2
combs worker combs limited worker combs.
Number of reproductive combs Oto 4 Oto 3 Oto 3.
Reproductive production on 1 or 2 phases of largely mixed queen and male largely homogeneous queen
reproductive combs homogeneous queen or production, no phases production.
male production
Worker to reproductive cell
ratios 0.3 to 1.9 0.5 to 2.6 1.5 to 34.9.
Oophagy by foundress queen +^ - -
Prey malaxation by foundress continues for most of
queen season ceases early in season ceases early in season.
Prey malaxation by new queens
and males + — —
Cap trimming by new queens + - —
Mauling of new queens by
workers + — —
Gastral vibration by foundress
queen and workers + + (rare) —
Nest sanitation most waste removed heavy waste accumulation most waste removed.
Prey diversity extensive, carrion usually more limited, carrion not extensive, may utilize carrion
not utilized utilized heavily late in season.
Pronounced spraying of venom -f- - —
Specophaga parasitism usually light usually heavy rare.
' -I-, behavior or biology present; —, behavior or biology absent.
pula aus triaca, which usurp colonies of other frequently usurps colonies of V. maûlifrons
yellowjacket species. These yellowjackets lack a (MacDonald and Matthews, 1975). There is also
worker caste and are inquilines (permanent nest evidence of a V. pensylvanica usurping a nest of
parasites), relying on the workers of the host to V vulgaris (Akre et al., 1977), intraspecific com-
rear their brood. The hosts for D. árctica are two petition as evidenced by dead queens found in
closely related yellowjackets, D. arenaria (Greene entrance tunnels (MacDonald et al., 1974), and
et al., 1976) and D. norvegicoides (Sladen) (R.E. widespread interspecific and intraspecific compe-
Wagner, Univ. Calif., Riverside; cited by tition in V. maculifrons colonies (MacDonald
Yamane, 1975). The host of V. austríaca in and Matthews, 198_a). These data suggest that
North America was only recently discovered to be inter- and intraspecific usurpation in yellow-
Vespula acadica (Reed et al., 1979). Its Palearctic jackets, rather than being uncommon, is wide-
(European) host is V. rufa (Spradbery, 1973a). spread.
In addition to obligatory social parasitism, at Predators of yellowjacket colonies are few;
least one species of yellowjacket is a facultative however, skunks dig up the nests of subterran-
parasite. Vespula squamosa, a species that has ean yellowjackets and eat the combs. For
workers, can estabUsh colonies of its own, but example, from 1974 to 1976 in the Pullman,
On the basis of isolated reports, V. crabro now

occurs along the eastern seaboard from New
England south into northern Georgia and Ala-
bama, and westward to the Mississippi River
from Tennessee north into the Ohio Valley. Scat-
tered northern populations probably exist from
the Dakotas to southern Ontario and Quebec,
and the wasp has been reported as far south as
New Orleans.
Few biological studies have been made of this
species since the works of Janet (1895, 1903);
however, the distribution of V. crabro in Europe
was given by GuigUa (1972), and some behavior-
al aspects of one subspecies were recently
investigated in Japan by Matsuura (1969, 1970,
1971, 1973, 1974). Little information is available
on this species in America, and much of the in-
formation presented here is taken from Sprad-
bery (1973a).
FIGURE 31—Schematic comparison of colony duration and
In Europe, V. crabro typically builds its nests
size of Vespula vulgaris species group. V. rufa species
in hollow trees. Nests are also found in thatched
group, and DoUchouespula ssp. based on number of
roofs, barns, attics, hollow walls of houses, and
workers at time of colony collection of V. pensylvanica, V.
atropilosa, and D. arenaria in Pullman, Wash.
abandoned beehives. Similar nest sites are used
in North America (fig. 33), and Duncan (1939)
even reported some subterranean nests.
Wash., area, one to three nests (V. atropilosa/V. Nests built in unprotected locations are cov-
pensylvanica) were recorded as skunk victims ered with a thick, brown envelope composed
each year. During 1977, 9 of 51 recorded V. pen- chiefly of coarse, decayed wood fibers, and,
sylvanica colonies were lost to these predators. therefore, very fragile. Brown envelope and car-
Other predators include coyotes (one V. vulgaris ton distinguish the nests of V. crabro from the
colony in the Blue Mountains of Washington, more common gray nests of D. maculata. The
1977) and bears (one V. vulgaris colony in moun- envelope of V. crabro nests is usually composed
tains near La Grande, Oreg., 1975). In Georgia of elongated shells or tunnels, which are distinct
and Indiana, raccoons are one of the principal from the scalloped tan envelopes of Vespula vul-
predators on colonies of V. maculifrons, V. flavo- garis or V. maculifrons. In addition, cell size of
pilosay and V. squamosa. Other minor predators Vespa is several times larger. Nests in protected
include birds, especially robins, which stand near locations, such as hollow trees or between walls
the nest entrance where they catch and eat re- of houses, typically have only a rudimentary
turning foragers. The impact of these predators envelope at the top of the nest, with most of the
on total yellowjacket populations is probably combs exposed.
slight. V. crabro nests are often large because of the
In the following sections, distribution data for large individual cells, but contain relatively few
the yellowjackets relies heavily on Miller (1961), cells. A typical mature colony has a nest of 1,500
to 3,000 cells in six to nine combs; the lower two
Hornets — Vespa crahro to four combs contain queen cells. A huge nest
collected in northern Georgia had 33 combs (near-
Vespa crabro (European hornet) was introduc- ly one-third reproductive), containing 5,566 cells
ed into the New York area between 1840 and (fig. 33 j). V. crabro colonies have a long seasonal
1860 (Shaw and Weidhaas, 1956\ van der Vecht, cycle with eclosión of reproductives occurring
1957). Originally Palearctic, its estimated distri- from late August to November in eastern sea-
bution in North America is given in figure 32. board States. At its peak of development, a large
colony consists of about 1,000 workers. Typical court" of workers, which are highly attracted to
colonies, however, are more likely to have 200 to her (Matsuura, 1968, 1974), Workers fly at night,
400 workers at their peak. and are attracted to lighted windows in homes.
Workers of V. crabro are predacious on a ''Sometimes ten or more will beat themselves
variety of insects, capturing large species such against a window causing people inside to panic
as grasshoppers and other orthopterans, flies, thinking they are trying to break the glass and
honey bees, and yellowjackets. Since they are invade the house to attack them. The very loud
powerful, agile wasps, they probably forage far- buzzing and the force with which they âttack'
ther for prey than the smaller yellowjackets. The the window is impressive" (C.W. Rettenmeyer,
queen of V. crabro is attended by a "royal Univ. Conn., personal commun.). In the autumn,
Vespa crabro
FIGURE 32 — Distribution of Vespa crabro. Lighter pattern indicates scattered occurrence.

PN-6532
FIGURE 33 —'Vespa cmbro: a. Typical nest site in hollow tree; b, male; c, queen on cell caps; d, fall queen emerging from
pupal case; e, nest in playhouse; /, bottom of same nest; g, mature larvae; h, worker on comb; i, girdling of
lilac {a-i, C.W. Rettenmeyer in Connecticut); and 7", nest with 33 combs in sweet gum tree near Täte, Pickens
County, Ga. (R.W. Matthews).
YELLOW JACKETS OF AMERICA NORTH OF MEXICO 35
V. crabro males are often attracted to lights as noticeably weaker dichotomy between queen and
well; therefore, in addition to morphological dif- worker castes than in observed Vespula spp., as
ferences, a number of behavioral differences well as a different pattern of reproductive pro-
exist between this true hornet and yellow- duction. A comparison of several behavioral
jackets. aspects of D. arenaria and two Vespula spp. is
Workers of V. crabro have been reported to given in table 3. Preliminary observations of D,
girdle twigs and branches of numerous trees and maculata reveal this species also differs greatly
shrubs, including lilac, birch, ash, horsechestnut, from Vespula.
dogwood, dahlia, rhododendron, and boxwood
(Hitchcock, 1970; Shaw and Weidhaas, 1956), Dolichovespula nests are usually aerial, al-
Much of this girdling is probably done for sap though all species probably nest underground on
and not for fiber collection as the workers are occasion and subterranean nests of D. arenaria
highly attracted to the girdled area, imbibe sap are not uncommon in some areas. Typical nests
constantly, but usually do not carry fibers away. are relatively small (300 to 1,500 cells), although
The plants are sometimes killed. This species is exceptionally vigorous D. arenaria colonies may
a pest of honey bees in apiaries in Japan and construct over 4,300 cells, and the largest D.
Europe (Matsuura and Sakagami, 1973; Sprad- maculata nests have approximately 3,500 cells.
bery, 1973a), but there are no reports of similar Mature nests are usually comprised of two to
depredations to honey bees in America. V. six combs. Envelope paper covering the nest
crabro is primarily a forest species having few usually consists of distinct laminar sheets, al-
contacts with man and presenting a minimal though there may be a high degree of scalloping
stinging hazard. and cellular construction in mature D. maculata
nests. The paper itself is much stronger, more
flexible, and more resistant to water damage
Yellowjackets than the comparatively brittle paper constructed
by Vespula spp. Peak worker populations
Genus Dolichovespula usually range from 200 to 700 individuals.
The five species of Dolichovespula occurring Dolichovespula are not attracted to protein
in the Nearctic Region are D, a/ft/da (Sladen), D. baits and usually forage only for live prey, but
árctica, D. arenaria, D. maculata, and D, norveg- occasionally will scavenge flesh from animal
icoides. Although this group has not been well carcasses (Greene et al., 1976). In this respect,
studied, available data indicate there are suffi- their behavior seems "intermediate" between
cient morphological, physiological, and behavior- the Vespula rufa group species (strictly live
al differences between Dolichovespula spp. and prey) and the Vespula vulgaris group (frequent
other yellowjackets to warrant their generic scavengers).
separation. Some of the more prominent
morphological differences include adult and lar-
val head structure, male genitalia, structures on Dolichovespula albida
the male antennae, and abdominal ganglia
(Duncan, 1939; Greene et al., 1976), Although the Dolichovespula albida is restricted almost
strong attraction of certain Vespula spp. to the entirely to the Hudsonian Zone of the North
synthetic compounds 2,4-hexadienyl butyrate American Boreal Region (fig. 34). Miller (1958,
(Fluno, 1973) and heptyl butyrate makes 1961) considered this a valid species rather than
these chemicals a valuable sampling method for a subspecies of the closely related D. norvegi-
them, no attraction has been observed for any coides since it retains its identity over its range
Dolichovespula spp. The venom chemistry of at even in areas in which it is sympatric with D.
least D, arenaria differs from that of Vespula norvegicoides. Practically nothing is known of
spp. (O'Connor and Erickson, 1965; A. Benton, the biology and behavior of this species. Be-
Penn. State Univ., personal commun.). Most im- quaert (1931) discussed two reports of subterran-
portantly, observations of D. arenaria behavior ean and partially subterranean nests; one was 5
(Greene et al., 1976) indicate this species has a inches in diameter.
Dolichovespula albida
FIGURE 34 — Distribution of Dolichovespula albida.
Dolichovespula árctica D: árctica is an obligatory social parasite in

colonies of D, arenaria (Greene et al., 1976) and
Dolichovespula árctica is distributed through- D. norvegicoides (R.E. Wagner, Univ. Calif.,
out the Boreal Region of North America, occur- Riverside; cited by Yamane, 1975), relying on
ring from Alaska to Arizona and across to the host workers to raise its own offspring. A D.
eastern seaboard (fig. 35). Investigations of its árctica queen invades a host nest early in the
biology have been limited (Taylor, 1939; Wheeler season before workers have emerged (Evans,
and Taylor, 1921), although recent studies have 1975). Although it may be attacked vigorously
begun to reveal more about its complex behavior by the foundress queen at first (fig. S6d), the
(Evans, 1975; Greene et al., 1978; Jeanne, 1977). parasite becomes established as a nestmate
within a day (Greene et al., 1978\ Jeanne, 1977). unparasitized colonies. It is still unknown how
The D, árctica queen may be highly aggressive the D. árcticas occupation of a colony usually
and dominant in most interactions with its terminates. In one chronologued colony, antag-
hosts, yet it feeds host larvae as well as its own onism between parasite and host workers gradu-
and even participates in limited cell construc- ally increased after the queen's death, culmina-
tion. ting in the slaughter of many workers by the
The parasite queen kills the host queen before larger, better-armored parasite; however, it is
much of her worker production is completed, likely that this D. árctica was finally killed by
shortening the Ufespan of the colony and reduc- the workers (Greene et al., 1978), Although a
ing the size of the mature nest relative to that of host colony produces no new queens, the D,
Dolichovespula árctica
FIGURE 35 — Distribution oí Dolichovespula árctica.

PN-6533
FIGURE 36 — Doächovespula árctica: o, D. árctica queen on outside of D. arenaria nest (S.C. White); b, same nest, D.
arenaria queen entering (CF. Roush); c, D. árctica queen on comb of D. arenaria nest; and d, D. arenaria queen
attacking D. árctica, which assumes a submissive posture.
árctica apparently cannot prevent the host DoUchovespula arenaria

workers' ovaries from developing, and some may
start ovipositing during the final days of the Dolichovespula arenaria (the aerial yellow-
parasite's occupation. Additional large numbers jacket) is transcontinentally distributed in the
of unfertilized eggs may be oviposited by Boreal Region of North America, occurring from
workers following the parasite's death, and a north-central Alaska to as far south as Arizona
sufficient degree of colony cohesion may be and New Mexico (fig. 37). It is "one of the com-
maintained in some cases to rear a substantial monest wasps in North America" (Miller, 1961)
brood of host male offspring to adulthood and one of the most widely distributed. Al-
(Greene et al., 1976; 1978). though there are numerous papers on various
aspects of its behavior and nest architecture, the September. Nests are usually aerial and are con-
only comprehensive study of its social biology is structed from a few centimeters above ground in
that of Greene et al. (1976). grass, shrubs, and bushes to the very tops of
D. arenaria nest construction begins as early trees (fig. 38). They are also commonly found on
as March in California, with many colonies de- houses and in sheds, garages, and many similar
clining in mid-June and totally dying out by manmade structures. Colonies may sometimes
July (Duncan, 1939). In the mid-Atlantic States, be situated beneath rocks or even below ground,
colonies may be mature and producing reproduc- with workers excavating soil to allow for nest
tives by late June. In Washington, queens be- expansion as in Vespula spp. (Greene et al.,
come active in April and nests are initiated from 1976). During the summer of 1977, 14 subterranean
May to early June, with colonies entering de- nests with excavating workers were located in
cline in August. A few may endure throughout the mountains near La Grande, Oreg., and near
Dolichovespula arenaria
FIGURE 37 — Distribution of Dolichovespula arenaría.
40 AGRICULTURE HANDBOOK 552. U.S. DEPARTMENT OF AGRICULTURE
PN-6534
FIGURE 38 — DoUchovespula arenaria: a, Typical nest site on eave of house, Elberton, Wash; b, tjfpical nest site in ever-
greens about 38 cm above soil, Pullman, Wash.; c, nest taken from eave of house, glued to a board, and trans-
planted in another location; d, envelope removed from same nest showing combs and workers; e, subterranean
nest in mountains near La Grande, Oreg. (C.F. Roush); /, worker placing fiber (seen as dark band) on nest
envelope.
Harrison, Idaho, suggesting that such nests tion in Pullman, Wash., for 1975. Colonies in
may not be uncommon in similar habitats Eastern States may average considerably
throughout western North America. smaller.
Colony size of D. arenaria may vary greatly Commonly encountered nest associates of D.
from year to year. For example, most nests con- arenaria in southeastern Washington are the ich-
structed by colonies in southeastern Washing- neumonid, Sphecophaga vesparum burra, the
ton in 1974 were small, with diameters rarely social parasite, D. árctica, and the pyralid moth,
exceeding 10 cm and the largest nest containing Vitula edmandsae serratilineella Ragonot
only 2,662 cells (Greene et al., 1976). Conversely, (Greene et al., 1976). The rate of parasitism by
one nest in 1973 (25 cm in diameter and 30 cm the ichneumonid was low, and did not seem to
long) contained 4,277 cells, one in 1975 had be an adverse factor in colony development. A
4,290 cells, and one in 1976 had 4,359 cells. Al- diverse assortment of other arthropods, such as
though the largest nests collected by Greene et Parcoblatta sp. in nests in North Carolina and
al. (1976) had a maximum of six combs. Spencer earwigs in nests in Washington, are probably
(1960) mentioned two seven-combed nests from incidental.
British Columbia. Adult worker populations D. arenaria workers usually forage only for
during peak colony development vary from live prey, but may utilize carrion on occasion
about 200 to 700, whereas total adult production (Greene et al., 1976). Searching patterns and the
of some colonies over the season may reach wide range of prey attacked indicate their forag-
6,500 or more. Table 4 shows colony composi- ing habits are quite similar to those of V. pensyl-
TABLE 4.—D. arenaria colony composition in 1975, Pullman, Wash., and vicinity^
Spheco-
Cap- Emp- phaga Multiple
]Lar- ped ty occupied brood Total
Colony Date Workers Queens Males Eggs vae brood cells cells cells cells Combs
A-48 June 16 3 0* 0 74 11 0 0 0 85 2
A-49 16 4 0* 0 52 8 1 2 0 61 2
A-50 18 0 0* 0 34 5 0 0 0 39 1
A-51 21 5 0* 0 100 31 0 0 0 131 2
A-52 30 17 0* 0 214 34 0 0 1 248 2
A-56 July 17 316 0* 1 748 488 54 0 35 1,290 4
A-47 23 62 0* 0 416 153 2 0 0 571 3
A-57 Aug. 8 314 67 38 362 789 718 107 0 103 1,976 5
A-58 8 478 83* 0 309 792 854 176 0 11 2,131 5
A-59 12 295 0* 0 116 457 321 68 0 18 962 4
A-62 14 520 42 111 239 869 485 459 0 4 2,052 5
A-63 14 697 0* 188 296 1,346 668 546 16 89 2,872 5
A-47 Sept. 3 103 0 67 60 174 206 949 0 1 1,389 5
A-48 3 0 0 0 0 0 12 138 0 0 150 2
A-52 3 73 84 7 5 57 140 2,800 0 0 3,002 6
A-64 3 472 100 18 38 588 150 2,049 0 1 2,825 6
A-66 8 10 32 1 0 3 43 1,618 3 0 1,667 5
A-67 15 89 0 3 36 223 11 2,594 0 0 2,864 6
Asterisk (*) indicates presence of foundress queen in addition to queen total given. Only one brood per
cell tabulated; cells containing both larva and egg tabulated as larvae. Eggs and larvae not tabulated sep-
arately for first 7 colonies. Transplanted colonies A-47, A-48, and A-52 were analyzed twice (Greene et al.,
1976).
vanica. Prey includes grasshoppers, spittlebug D. maculata queens first become active in
adults, leafhoppers, Lygus Hahn bugs, tree April and May in southeastern Washington with
crickets, lacewings, caterpillars, flies, and nests usually established in late May or early
spiders. June. Reproductives are produced in late July
The behavior of D. arenaria within the nest and August, and colonies enter decline in
was discussed by Greene et al. (1976). In many August or September. In the mid-Atlantic
aspects, it is similar to that exhibited by other States, nest construction begins as early as late
yellowjackets (Akre et al., 1976); however, D. April. Nests are usually situated in vegetation,
arenaria displays some important differences, in- from shrubs or vines at ground level to 20 m or
cluding prey malaxation by the foundress queen higher in trees. They may also be built on rock
for most of her life, prey malaxation by new overhangs, electric power poles, houses, sheds,
queens and males, cap trimming by new queens, or other manmade structures (fig. 40 b,d).
mauling of new queens by workers, and gastral Nearly all are constructed in exposed locations,
vibration by the foundress queens and workers although Greene et al. {1976) mentioned litera-
(table 3). Some of these activities have been ob- ture reporting one subterranean nest and one
served in D. maculata colonies but not in other inside a hollow tree.
yellowjacket species. Since D. maculata is a large wasp, its nests
D. arenaria workers do not ordinarily scavenge can sometimes attain impressive dimensions,
for protein and so are not usually serious picnic with diameters of 35 cm and lengths of more
pests; however, like most yellowjackets, they than 60 cm (fig. 40Ä); however, much of the nest
may be attracted to sources of sugar in late bulk is made up of thick, multilayered envelope,
summer. Isolated D. arenaria workers, apparent- and the cells comprising the combs, although
ly more so than other yellowjacket species, have larger than those constructed by any other
an annoying and disconcerting tendency to buzz yellowjacket, are comparatively few in number.
around the heads of people, sometimes following The most populous colonies construct 3,500 cells
a person through the woods for a considerable in five combs, but the majority of nests contain
distance. They may be foraging for flies, which less than 2,000 cells in three or four combs. Peak
often are found with any large animal. Since D. worker population is typically 100 to 400, al-
arenaria often constructs nests on manmade though one Maryland nest collected in mid-Sep-
structures, the chances for encounters resulting tember contained 636 workers.
in stings to humans are greatest in this Nest associates of D. maculata have not been
situation. While small colonies may be quite studied in detail. The ichneumonid Sphecophaga
mild and not easily aroused, larger colonies are vesparum burra often occurs in low densities,
formidable when disturbed. Attacking workers and many of the incidental arthropods found
can even spray venom out of the sting (Greene with D. arenaria also have been collected in D.
et al., 1976). Control of some D. arenaria colonies maculata nests. In Georgia and North Carolina,
located in close proximity to human activity is cockroaches (Parcoblatta sp.) are found in some
therefore warranted. nests. Recent investigations in Maryland sug-
gest that a small chalcid wasp feeding on the
DoUchovespula maculata brood may be important in limiting size and
Dolichovespula maculata, the baldfaced **hor- number of colonies in spring and early summer.
net," is an atypically large, black and white yel- D, maculata workers occasionally scavenge for
lowjacket, which is probably the most widely protein (Greene et al, 1976; Payne and Mason,
distributed member of the Nearctic Vespinae. It 1971), but most forage only for live prey. Flies
occurs throughout the Canadian, Transition, are commonly taken, and, in some areas, other
Upper Sonoran, and Upper Austral Zones, and yellowjackets are an important prey item
extends into the Lower Austral Zone (fig. 39). It (Howell, 1973). One nest collected in Maryland
has been collected virtually in every State and contained the gasters of 44 V. maculifrons, 9 V.
Canadian Province, from north-central Alaska to squamosa, 1 V. flavopilosa, and 1 honey bee,
central Florida and the **Big Thicket" of south- Apis mellifera L. The particulate detritus in this
eastern Texas. nest had a yellowish tone due to the extreme
abundance of discarded yellowjacket parts and 1954; Gibo et al., 1974a,b, 1977\ Rau, 1929), a
bits of integument. D. maculata workers are comprehensive sociobiological study of this
powerful, agile wasps and occasionally attack re- species has only recently begun. Preliminary re-
latively large insects such as cicadas. There is sults indicate its behavior may be quite different
even a report of an attack on an adult humming- from that reported for D, arenaria.
bird (Grant, 1959), although this is undoubtedly D. maculata has adapted to nesting in human
an unusual occurrence. population centers throughout its range, and can
Although there are several papers on various sometimes be found well within the boundaries
aspects of the behavior of D. maculata (Balduf, of large cities. Its imposing size is often suffi-
Dolichovespula maculata
FIGURE 39 — Distribution of Dolichovespula maculata.

PN-6535
FIGURE 40 — Dolichovespula maculata: a, b. Queen nest with funnels 11 cm and 6.5 cm, respectively. La Grande, Greg. (C.F.
Roush); c, mature nest, Pullman, Wash.; d, mature nest on brick wall of house, Maryland; e, nest in weeds,
Connecticut; /, nest with envelope removed exposing combs, Connecticut; g, mature nest fastened to tree
branch, Connecticut (e-g, C.W. Rettenmeyer); h, nest 35 X 60 cm, Missouri (R. Getting); i, workers around
nest entrance, Connecticut (C.W. Rettenmeyer).
cient to alarm people, but its aggressiveness commonly recognized by laypersons. Its
fortunately does not match its appearance. Al- abandoned nests are displayed in almost every
though, as with any yellowjacket, accidental park nature center and sometimes hang as
jostling of a nest or pressing on an individual ornaments in homes. Unless a colony is located
will result in stings, our experience has been in close proximity to human activity, this
that workers are usually not as sensitive to dis- species should not be regarded as a pest but
turbance around the nest as some of the other, rather as a colorful and beneficial part of the
smaller species. Furthermore, it has a tendency American insect fauna.
to nest fairly high in trees (more so than any
other Nearctic wasp), reducing the likelihood of Dolichovespula norvegicoides
contact between humans and many colonies. The
baldfaced '^hornet" is often mentioned in Dolichovespula norvegicoides is restricted al-
introductory field biology guides, as much for most entirely to the Canadian and Hudsonian
its conspicuous architecture as its striking Zones of the Boreal Region (fig. 41). Most
appearance, and is one of the handful of insects aspects of the biology of this species are incom-
■^«2^
Dolichovespula norvegicoides
FIGURE 41 — Distribution oi Dolichovespula norvegicoides.

pletely known and data on its behavior are aspects of the biology of V. rufa group species
totally lacking. The few recorded nests have exist. The first and only detailed investigation
been aerial and quite small. One examined in of a V. rufa group species was of V. atropilosa in
Vermont on August 6 by Bequaert (1931) was in Washington. Other species have been studied
dense bushes about 10 cm off the ground. The somewhat, and only V. intermedia and North
colony consisted of 50 workers and the found- American V. austriaca remain unknown.
ress queen—reproductives had not yet emerged; Vespula acadica. — Vespula acadica (forest yel-
however, Bequaert previously collected males in lowjacket) is restricted almost entirely to the
the Adirondacks as early as July 19. A nest Canadian Zone. (fig. 42).
collected in late October at Granite Lake in the V. acadica was reported as an aerial nester by
Cabinet Wilderness, 10 to 15 miles south of Sladen (1918); however, two of three nests found
Libby, Mont., was located in a shrub 15 cm by MacDonald et al. (1975a) were subterranean,
above ground. The nest was 12.5 cm in diameter and one was in a decaying log on the forest floor.
and had two combs with a total of 234 cells. From 1975 to 1977, eight additional nests were
Twenty-three dead workers were in the nest. collected—six in forested areas near La Grande,
Worker populations of D. norvegicoides prob- Oreg., and two in northern Idaho near Harrison
ably seldom approach the size commonly (Roush and Akre, 1978), Four of these nests were
attained by its close relative, D. arenaria. in decaying logs, two were under logs, and the
R.E. Wagner (cited by Yamane, 1975) reported other two were subterranean, one in duff (partly
the social parasite, D, árctica, is sometimes decayed vegetable matter such as leaves and pine
found in colonies of D. norvegicoides. Since needles) and one in the soil (fig. 43). Entrance tun-
there have been few collections, the incidence of nels varied from 7 to 40 cm, usually 15 to 25 cm.
parasitism is unknown. The deepest subterranean nest was 15 cm under
the soil surface. Nests were comprised of one
Genus Vespula worker-producing comb and one to three repro-
ductive-producing combs. The largest mature col-
Vespula rufa Species Group ony consisted of 425 workers, 70 males, and 78
The Nearctic members of the V. rufa species queens. The nest had four combs with a total of
group include: V, acadica, V. atropilosa, V. 1,791 cells. The seasonal cycle and size of the col-
consobrina (Saussure), V, intermedia (Buysson), onies were comparable to V atropilosa colonies
V. vidua (Saussure), V, austriaca, V. squamosa, collected during the same year on comparable
and V. sulphurea (Saussure). The first five dates.
species are a morphologically and biologically V. acadica workers prey only on live arthro-
cohesive group; their taxonomic status has pods. They are predators on caterpillars, flies,
varied from treatment as varieties of the and hemipterans. Other prey preferences are un-
**parent'' species, V, rufa, to full species (Miller, known.
1961). The social parasite V. austriaca is closely No study of nest associates has been made,
related to its Palearctic host V. rufa, and there but five of the eight nests collected had 2 to 20
is no reason to treat it as other than a socially cells containing Sphecophaga cocoons, probably
parasitic member of the V, rufa group. V. S. vesparum burra, A few nests had eggs of Fan-
squamosa and V. sulphurea are of uncertain nia spp. on the outside envelope.
status (MacDonald et al., 1976), The former is a Being primarily a forest species, this yellow-
frequent facultative parasite of V. maculifrons, a jacket has httle contact with man; however,
member of the V. vulgaris group. Since social when colonies are disturbed, workers of this
parasites are usually closely related species, this yellowjacket may be quite aggressive and per-
casts doubt on the current placement of V. squa- sistent and sting repeatedly.
mosa within the V. rufa group. The status of V. Vespula atropilosa,—Vespula atropilosa (prai-
sulphurea is also uncertain because few data are rie yellowjacket) is found in western North Amer-
available on its nesting biology and colony ica where it is restricted to the Canadian and
duration. Transition Zones of the Boreal Region (fig. 44).
A number of superficial accounts of various Detailed biological and behavioral investigations
were reported by Akre et al. (1976) and MacDon- (1957), Buckell and Spencer (1950), and Miller
ald et al. (1974, 1975a. 1975b). (1961). MacDonald et al. (1974, 1975a) reported
V. atropilosa is abundant in prairie and open locations of 40 nests of V. atropilosa near
forest areas, but becomes increasingly less abun- Pullman, Wash. Thirty-six were subterranean,
dant in heavy forest where it is replaced by V. and the remaining four were in various locations
acadica. It also commonly nests in yards, —under sod pile, inside decayed 4- by 4-inch tim-
pastures, golf courses, and similar areas. ber (fig. 45/), in tree hollows, and beneath steps.
V. atropilosa was reported as a subterranean From 1974 to 1977, an additional 55 colonies were
nester by Bequaert (1931), Bohart and Bechtel located; 3 were nesting in between walls of hous-
Vespula acadica
FIGURE 42 — Distribution of Vespula acadica.

es, and the other nests were subterranean. Most TABLE 5.—Comparison of mature colonies (with
nests were built in rodent burrows with the nest reproductive cells) o/Vespula atropi-
located 10 to 30 cm into the tunnel and 10 to 15 losa, 1971-75
cm below the soil surface.
In the spring of 1977, an old (1976) nest of V. Average Most
CeUs in CeUs in number workers
atropilosa, containing 730 cells in 2 combs, was smallest largest of cells collected
found in a truly aerial position in Pullman. The Year Nests nest nest per nest per colony
nest was 150 cm above the ground—located
- Number -
underneath a concrete overhang, which was part
1971 6 942 2,676 1,890 368
of some steps—and was fastened to concrete on 1972 5 484 1,068 884 76
two sides and to plywood on the top. 1973 16 549 2,175 1,236 270
Table 5 compares mature (with reproductive 1974 14 603 2,497 1,313 504
cells) V. atropilosa colonies from 1971 to 1975. 1975 13 278 1,556 897 484
All colonies collected during 1976-77 were ex-
PN-6536
FIGURE 43 — Vespula acadica: a. Queen nest in hollow log. Laird Park, Idaho; 6, nest in pine needles and duff of forest floor,
Harrison, Idaho; c, nest in decayed tree root. La Grande, Oreg. (CF. Roush); d, workers in trophallaxis on
comb of nest.
Vespula atropilosa
FIGURE 44 — Distribution of Vespula atropilosa.
eluded as they were used in experiments. Nest live prey. They often attack spiders, phalangids,
size varied from year to year with the nest of the flies, caterpillars, hemipterans, and some homop-
smallest mature colony comprised of 278 cells; terans, but seldom attack other Hymenoptera or
the largest, 2,676. When collected, the greatest beetles.
number of workers in a colony was 504. This is Nest associates of V. atropilosa were studied
typical of colonies and nests of all members of by MacDonald et al. (1975b). Most associates are
the V. rufa group, which are only one-fifth to scavengers in debris below the nest or on fungi
one-fourth as large as those of members of the growing on the nest carton. Nests of V.
V. vulgaris group (table 1). atropilosa are quite filthy, with remnants of
V. atropilosa workers are predators only on bodies and other debris incorporated into the
50
PN-6537
FIGURE 45 — VespuUi atropilosa: a-c. Nest built in cardbotu-d box in shed; b, ventral view; c, envelopes removed; d, mature
larva chewing on insect flesh; e, egg, larvae, prepupa; f, nest in hollow 4X4 inch beam.
nest envelope and carton, probably providing dependent on host workers to rear its brood of
scavengers still another supply of food. The new males and queens. Various workers (Carpen-
pupal parasite, Sphecophaga vesparum burra, ter and Pack-Beresford, 1903\ Pack-Beresford,
appears to adversely affect development of 1904\ Sharp, 1903\ Weyrauch, 1937) described cer-
young colonies. This is an abundant species and tain aspects of the biology of K austríaca and its
some colonies invariably nest in yards, causing host V, rufa as summarized in Spradbery (1973a),
homeowners concern and often warranting re-
moval; however, workers are usually no problem V. austríaca is sympatric with a number of
unless the colony is disturbed. potential V. rufa group hosts (Miller, 1961);
Vespula austriaca.—Vespula austríaca, a rarely however, our studies indicated that V. acadica
collected species, is widespread throughout the was the most likely host. During the summer of
Boreal Region of North America (fig. 46). An ob- 1978, two colonies of V. acadica, each with a V.
ligatory social parasite of Palearctic V, rufa, V. austríaca queen, were collected in northern
austríaca does not possess a worker caste and is Idaho near the small town of Harvard. These are
Vespula austríaca
FIGURE 46 — Distribution of Vespula austríaca.

the first records of this social parasite being thews, 1976) or in the walls of houses (Gaul, 1948).
taken from nests of a Nearctic yellowjacket (Reed Five of nine nests located in Washington during
et al., 1979). 1974-77 were in rodent burrows, the remaining
Vespula consobrina.-Vespula consobrina (black- four were in the walls of buildings.
jacket) is a black and white yellowjacket found The colony cycle is short like that of other V.
in forested areas throughout the Canadian and rufa group species, with colonies decUning and
Transition Zones of the Boreal Region of North producing reproductives by mid-September in
America (fig. 47). Nests are typically in subter- Pullman and by September 1 in North CaroUna
ranean rodent burrows but may be aboveground (MacDonald and Matthews, 1976). V. consobrina
in logs or rock cavities (MacDonald and Mat- colonies are small; the average size of four
Vespula consobrina
FIGURE 47 - Distribution of Vespula consobrina.

mature nests collected in Pullman in 1974 was interactions with other Vespula spp. workers. For
only 555 cells, ranging from 490 to 690 cells (fig. example, a single worker successfully fended off
48). MacDonald and Matthews (1976) reported many V. pensylvanica workers at a dish of honey
on two North Carolina colonies with 1,437 and and usually chased them away.
731 cells with less than 100 workers per colony. As with other Vespula spp., aggressiveness
Other reports also indicate the small size of V. varies considerably relative to colony size and
consobrina colonies (Dow, 1930; Gaul, 1948; history. Some are distinctly more easily aroused
Taylor, 1939). Nests consisted of one worker- to sting than others although Gaul (1948)
producing comb and from one to three reproduc- regarded V. consobrina as not aggressive. Be-
tive-producing combs. cause of its infrequent contacts with man, this
The little information available indicates a les- species poses little problem for humans in most
ser incidence of nest associates with V. conso- areas of North America; however, in forested
brina colonies than with V. atropilosa; however, areas of western North Carolina, the blackjacket
one colony collected in Pullman in 1974 is responsible for numerous stinging episodes
contained 33 adult phorid flies (Triphleba among loggers.
lugubris); 28 were gravid females. Both North Vespula intermedia.—Vespula intermedia, a
Carolina colonies harbored eggs and larvae of black, ivory, and red species, is restricted almost
Dendrophaonia querceti and Fannia spp. entirely to the Hudsonian Zone of the Nearctic
(Diptera: Muscidae). Boreal Region (fig. 49). It is rarely collected, and
V. consobrina colony development and social no nest has been found (Miller, 1961). Since other
behavior are not well known, but studies indi- members of the V. rufa group are attracted to
cate they are similar in most respects to that synthetic organic compounds such as heptyl bu-
reported for V. atropilosa by Akre et al. (1976). tyrate, use of attractant traps might be of value
Foraging is for live prey only, with spiders and in surveying for this species.
phalangids readily accepted. Other arthropods Vespula vidua.—Vespula vidua is restricted al-
utilized as prey are similar to those attacked by most entirely to the Transition and Upper Aus-
V. atropilosa. tral Zones of eastern North America (fig. 50). The
Regarding interspecific interactions, one in- nesting biology of this species was studied by
stance of colony "drift" was recorded when MacDonald and Matthews (1976), but detailed be-
several V. consobrina workers joined a V. havioral information is lacking.
atropilosa colony where they oviposited as Seemingly less restricted to forested habitats
evidenced by the production of numerous V. than V. acadica and V. consobrina, V. vidua
consobrina males later in the season (Akre et al., commonly nests in disturbed areas (yards,
1976). V. consobrina workers appear aggressive in pastures) as well as forests from Minnesota and
PN-6538
FIGURE 48 — Vespula consobrina: a. Mature nest; 6, nest, envelope removed.
Vespula intermedia
FIGURE 49 — Distribution of Vespula intermedia.
Iowa to the New England States, then south Although most nests of V. vidua are subter-
into northern Georgia. The relative abundance ranean, it has also been reported as nesting in
of V. vidua in many of these areas is unknown. decaying logs (MacDonald and Matthews, 1976),
In many respects—nest habitats and locations, and in manmade structures (Ebeling, 1975, and
size and architecture (fig. 51), seasonal duration Shew2). The seasonal cycle of V. vidua is short
and size of colonies, adult size—K vidua is the as is common for V. rufa group members. The
eastern counterpart of V. atropilosa. Indeed,
without the presence of adults or developed
pupae, a nest of V. vidua cannot be 2Shew, R.L. The biology and control of vespine wasps. M.S.
distinguished from a nest of V. atropilosa. thesis, Ohio State University. 59 pp. 1966. [Mimeographed.]
largest colony (six analyzed) constructed a nest odors. This species is not a stinging hazard to
of 2,447 cells and had an estimated adult worker humans unless the nest is located in structures
population of 385 at its peak of development or yards where the colony can be disturbed by
(MacDonald and Matthews, 1976). Nests con- human activities.
sisted of one worker-producing comb and from
one to three reproductive-producing combs. Uncertain Status of Vespula squamosa and V. sulphurea
Although Wagner and Reierson (1971a)
reported V. vidua workers as pests around These two species are unique among yellow-
picnic tables, they were probably capturing in- jackets in possessing a characteristic mesono-
sects in the area or momentarily attracted to tum with two long, broad, longitudinal yellow
Vespula vidua
FIGURE 50 — Distribution of Vespula vidua.
PN-6539
FIGURE 61 — Vespula vidua: a. Mature nest taken from honey bee nest box, New York (R. Keyel); b, mature nest. North
Carolina.
stripes. Accordingly, they have been considered studied by MacDonald and Matthews {1975,
closely related (Duncan, 1939), and united with 198_b).
the V. rufa species group on the basis of adult Easily recognized by its color pattern and ex-
morphology (Bequaert, 1931). Studies have now treme female caste dimorphism, V. squamosa is
shown that both species have seven ovarioles an intriguing species. Unassociated until the
whereas other members of the rufa group, mem- early 1900's when a colony was excavated, the
bers of the vulgaris group, and four Dolichoves- very large and predominately orange queen
pula spp. have only six ovarioles (Kugler et al., differs markedly from the worker and male,
1976). Biological study of V. sulphurea is totally which are more typical of yellowjackets in size
lacking, and V. squamosa has only recently re- and color. Also curious is the success of V. squa-
ceived attention (MacDonald and Matthews, mosa in the southern distribution of Nearctic
1975, 198 b). These studies revealed V. squa- Vespula. Since the discovery of a V. squamosa
mosa differs from other V. rufa group members queen in the nest of V. vidua (Taylor, 1939), V.
in a number of ways, including seasonal dura- squamosa has been considered a facultative social
tion and size of colonies, nest size and architec- parasite of Vespula.
ture, and, possibly, foraging behavior. Indeed, Working in Georgia, MacDonald and Matthews
V. squamosa appeeirs more similar biologically (1975) found V. squamosa to be a facultative so-
to the V. vulgaris group. cial parasite of V. maculifrons, a member of the V.
Unfortunately, little data are available on V. vulgaris group. The parasite queen usurps the
sulphurea, but it may share biological as well as nest from the host queen and assumes complete
morphological attributes with V. squamosa. In- control of the colony. Host workers rear the first
depth behavioral studies are needed, and until brood of V. squamosa workers and eventually a
then we prefer to consider these two species as of colony consisting solely of V. squamosa results.
uncertain status. The relationship appears to be facultative since
Vespula squamosa.—Vespula squamosa (south- about 20 percent of mature V. squamosa colonies
ern yellowjacket) is widely distributed from Iowa exhibit no evidence of a V. maculifrons heritage.
to Texas and across to the eastern seaboard, and The remaining 80 percent either contain host
occurs in the southeastern part of the Transition, workers (early in the season) or the nests have
Upper Austral, and Lower Austral Zones of the clearly visible remnants of V. maculifrons nest
Nearctic Region (fig. 52), and also in southern construction. The incipient host nest with much
Mexico and Guatemala. Nesting biology was smaller, tan cells is in striking contrast to the sur-
rounding larger, gray cells constructed by para- siderably. Accordingly, discussion of the nesting
site workers; thus, although host workers may biology of V. squamosa must take into account
not be present, the early history of a V. squamosa the requirements of host species.
colony can be determined by analyzing nest
architecture. Most V. squamosa colonies (usurped or pure)
The above account depicts the host-parasite are located in disturbed habitats, particularly in
relationship in Georgia. In other areas, a differ- yards, parks and roadsides, whereas a few are in
ent host species may be involved, and the per- "natural" habitats. Among the latter, nearly all
centage of nests parasitized probably varies con- colonies are in pine forests, a few are in mixed
^«?î^
Vespula squamosa
FIGURE 52 — Distribution of Vespula squamosa.

forests but rarely in hardwoods despite the tive cells, although a complete count could not be
abundance of V. maculifrons colonies. made because of the nest's poor condition.
Coincident with its host, V. squamosa nests Although the behavior of V. squamosa has not
typically are subterranean (MacDonald and Mat- been studied, some of the workers definitely
thews, 1975, 198_b; Miller, 1961)\ however, aerial scavenge for protein (R. Matthews, Univ. of Ga.,
nests have been reported (Tissot and Robinson, personal commun.) and may be a nuisance at pic-
nics. This behavior is not typical for V. rufa
1954), and in the Washington, D.C., area where
group species.
V. maculifrons commonly nests in house walls,
V. squamosa is an important pest species be-
V. squamosa colonies are also frequently ob-
cause of the great number of nests found in
served in these locations. Colonies typically per-
urban and recreational areas. Colonies are typi-
sist into autunm with most producing reproduc-
cally large and disturbances of the nests usually
tives from late August into November; this
result in multiple stings.
seems intermediate between the longer cycle of
the V. vulgaris group and the shorter cycle of Vespulasulphurea.—^Vespula sulphurea (Cali-
the V. rufa group. Colony size is variable but fornia yellowjacket) is restricted almost entirely
most contain 500 to 4,000 workers at their peak to the Upper Sonoran fauna of California, but it
and ultimately construct a nest of 2,500 to also occurs in southern Oregon, western Nevada,
10,000 cells (table 6). southern Arizona, and northern Baja, Mexico
(fig. 54; Bohart and Bechtel, 1957), Duncan {1939)
Sometimes, perennial colonies of this species mentioned three nests that he collected, and in-
persist in subtropical areas of its distribution. cluded several photographs of one.
For example, Tissot and Robinson {1954) Based on the Umited data available, V. sul-
recorded a nest of V, squamosa of 6 ft by 30 phurea is a subterranean nester. One of the col-
inches by 12 inches with 17 combs, and another onies collected by Duncan {1939) in early October
was 9 ft tall with a circumference of 9 ft 10 inches 1930 was comprised of 134 workers, 6 new
and contained 39 combs. A perennial, subter- queens, and the foundress queen. The nest con-
ranean colony located 13 miles west northwest of tained a few male pupae. Seven additional sub-
Gainesville, Fla., was killed in early February terranean colonies have been collected in Califor-
1977 (J. Sharp, USDA, SEA, Gainesville). The nia by R. E. Wagner (Univ. of Calif., Riverside)
nest had 14 comb levels and 120,130 cells (fig. 53 (table 7). Three nests had three worker combs,
0, b); at least 1,350 of these cells were reproduc- two of these also had a single primary reproduc-
TABLE 6.—Summary of mature Vespula squamosa colonies collected after

October 15, 1974-75, in and around Athens, Ga, {MacDonald and
Matthews, 198__a)
Parameter 1974 1975

Colonies located '75 '46
Largest Average Largest Average
Total cells per nest 9.906 5.703 (N = 26) 9,802 5,034 (N = 9)
Average Percent of total Average Percent of total
ceUs cells
Queen cells per nest 1,001 (N = 26) 17 percent 842 (N = 9) 19 percent
Maximum workers per colony 3,697 3,193
Combs per nest Average Average
Worker 4.5 3.9
Mixed 0.6 0.6
Queen 23 2.0
Total 7.1 6.3
'Of 75 V. squamosa colonies excavated. 62 were initially founded by V. maculifrons.
'Of 39 V. squamosa colonies excavated. 37 were initially founded by V. maculifrons.
.^Mm
r f" .y^l 7^
r ,-. " »1
i)-^
..^Î^^^^Bj
Í -^^- .
^^J^HEr'UBrK- " .-." - î r■ "
um '----... N. ..^.,....^. ^^^[^K
PN-6540
FIGURE 53 — Vespula squamosa: a, Excavation of perennial nest 13 miles west-northwest of Gainesville, Fla., Feb. 2,1977, by
J. Ryan, left, and J. James, right (J.F. Sharp); 6, same nest, envelope removed exposing combs; c, mature
nest, Athens, Ga.; d, atypical nest envelope with highly colored stripes; most nests are uniformly gray.
tive-cell comb. Partial reproductive combs are Vesputa vulgaris Species Group
often placed wherever space is available in the
nest, even adjacent to the first worker comb; The Nearctic members of the V. vulgaris
however, no reproductive cells have been found group include V. vulgaris, V. flavopilosa, V.
on worker combs. maculifrons, and V. pensylvanica. A fifth spe-
"This species has a slightly shorter active cies, V. germánica, was introduced into the
season than V. atropilosa, the first workers ap- United States (Menke and Snelling, 1975), has
pearing a week or so later and the last workers recently become abundant in the East, and
disappearing a couple of weeks before those of V. appears to be gradually spreading westward
atropilosa" (R. E. Wagner, personal conmiun.). across the country (Morse et al., 1977).
Workers of V. sulphurea are attracted to heptyl Workers of the V. vulgaris group species are
butyrate, but other aspects of their behavior are notorious scavengers and frequently are pesti-
unreported. Members of this species are not very ferous to man, especially in recreation areas and
abundant, and workers have not been reported as at outdoor picnics. Accordingly, these important
picnic pests. pests have been investigated much more than
Vespula sulphurea
FIGURE 54 — Distribution of Vespula sulphurea.
other Vespula spp. European species have been MacDonald and Matthews (198_a), and Preiss.^
studied in detail by a number of investigators
(Spradbery, 1973a), as have some of the Oriental ^Preiss, F.J. Nest site selection, microenvironment and pré-
species (Iwata, 1976). Biological investigations dation of yellowjacket wasps, Vespula maculifrons (Buys-
in America were reported by Akre et al. (1975, son), (Hymenoptera: Vespidae) in a deciduous Delaware
1976), Duncan {1939), Jacobson et al. (1978), woodlot. M.S. thesis. University of Delaware. Newark.
MacDonald et al. (1974, 1975a, 6, 1976, 1980), 81 pp. 1967. [Mimeographed.]
TABLE 7.—Summary o/Vespula sulphurea colonies collected in California

(After RE. Wagner)
Adiilts
Worker Reproductive
Colonv Date FaU combs combs
No. Locality collected Workers Queens queens Males and No. cells and No. cells
1 Clear Lake 7-27-72 580 1 0 0 (•) (')

2 Clear Lake 9-11-72 804 0 23 287 (') (')
9-12-72 1,086 1 31 363 3 - 3,258 1+4 partial
3 Clear Lake
- 769
4 Clear Lake 9-13-73 392 1 7 44 n n
5 Clear Lake 9-24-72 556 0 103 258 (') (')
Malibu Can- 7-03-75 462 1 0 0 3 - 1,641 0-0
6
yon, Santa
Monica
Mountains
8-27-75 105 1 0 0 3 - 1,080 1 - 33
7 Mill Creek,
San Bernar-
dino Mtns.
'Not available.
Vespula flavopilosa.—Vespula flavopilosa (hy- as a species, no studies of behavior or nest associ-

brid yellowjacket) occurs from Minnesota to the ates are in the literature.
east coast and south to northern Georgia (fig. 55). For the same reason, the pest status of V. flavo-
Future collection records will undoubtedly extend pilosa is unevaluated; however, since workers are
scavengers, V, flavopilosa probably presents a
this known distribution.
Similar to V. maculifrons and V. vulgaris in pest problem similar to that of other V. vulgaris
most aspects of its biology, V. flavopilosa con- group species. Queens of this species establish
structs subterranean nests (fig. 56) of tan, fragile nests along roadsides and in yards. Three V. fla-
carton (not gray carton as found in nests of V. ger- vopilosa nests in structures have been found in
mánica and V. pensylvanica) (MacDonald et al, New York (R. Key el, Cornell Univ., personal com-
1980); however, V. flavopilosa carton is noticeably mun.). Based on a small collection from Minne-
sturdier, and cell size is significantly larger than sota, V. flavopilosa workers are attracted to hep-
that found in V. maculifrons and V. vulgaris tyl butyrate to some degree; response to protein
nests. Accordingly, workers and queens of the baits is unevaluated.
Vespula germánica.—Native to Europe, Vespu-
new species are larger and similar to V. germánica
la germánica (German yellowjacket) has been in-
and V. pensylvanica in this respect. At least in
troduced into New Zealand (Thomas, 1960), Tas-
the southeast, colony seasonal cycle is shorter,
mania (Spradbery, 1973b), South Africa (White-
ending a month or so earUer than that of V. macu-
lifrons; thus, colony size was noticeably smaller. head and Prins, 1975), Chile (Pena et al, 1975),
Sydney, AustraUa (R. Edwards, Rentokil Ltd.,
Typical V. flavopilosa colonies in western North
Sussex, England, personal commun.) and the
Carolina completed their cycle by early October,
United States (Menke and SnelUng, 1975). The
during which time a peak worker population of
known distribution of this species in North Amer-
500 to 1,000 constructed from 2,000 to 5,000 cells
consisting of nearly 20 percent queen cells; there- ica, as of 1977, is given in figure 57.
In Europe, V. germánica nests are usually sub-
fore, nests of V. flavopilosa collected in late sum-
mer contained 2,500 to 4,000 fewer worker cells terranean, but they may be aerial or in roofs,
than those of V. maculifrons collected at the same attics, and between the walls of houses
(Spradbery, 1973a). In North America, nearly all
time and locality (table 8).
Since V. flavopilosa was only recently described reported nests have been in structures. Eighteen
of 22 colonies studied in Tompkins County, New mánica in New Zealand (Thomas, 1960) and Tas-
York, during 1977, built nests between the walls mania. In New Zealand, one aerial nest was 14 ft
of buildings (R. Keyel, Cornell Univ., personal 11 inches by 5 ft by 2 ft, and a subterranean
commun.). Colonies are usually annual, but nest (47 by 40 by 38 inches) with 27 combs
perennial colonies are not infrequent in New weighed about 100 lb. A subterranean nest in
Zealand (Thomas, 1960) and Tasmania (Sprad- Tasmania measured 6 by 2.5 by 4 ft, had 30
bery, 1973b). combs, and was comprised of 1.5 million cells.
From these data, it was estimated that the larg-
Spradbery (1973a) discussed the huge nests est New Zealand nest had 3 to 4 milHon cells
constructed by perennial colonies of V. ger- and weighed 1,000 lb.
Vespula flavopilosa
FIGURE 55 — Distribution of Vespula flavopilosa.

PN-6541
FIGURE 56 — Vespula flavopilosa mature nest. New York: o, Bottom view; b, top view (R. Keyel).
TABLE 8.—Vespula flavopilosa and V. maculifrons colo-

nies collected during 1974 in northwestern
Georgia and southwestern North Carolina
(MacDonald et al, 1980) ^
Maxi-
Worker cells in nest Worker mum
Species Colonies Largest X combs workers
Number - - -
V. flavopilosa 23 3,844 1,871 (N = 19) 3.2 1,332
V. maculifrons 32 9,700 4,422 (N = 14) 4.8 4,839
^6,211 (N= 7) 5.3
'Since most colonies were sampled early in the period of queen-cell con-
struction, only comparisons of worker cells and worker cell combs were
made.
' V. maculifrons colonies in which queen cell construction had begun.
Typical nests are considerably smaller and are in the Washington, D.C., area. One located with-
similar in size to nests of V. pensyluanica. in the framework of a house porch and examined
Spradbery {1973a) described a colony collected in in late August contained 11,540 cells in seven
early September in Great Britain, which was combs (fig. 58). Another, situated in an attic,
comprised of 1,820 workers, 447 males, 147 new measured nearly 80 cm in diameter by 80 cm long
queens, and the foundress. The nest had 8 (J. Nixon, Silver Spring, Md., personal commun.)
combs with a total of 6,341 cells, including 1,214 (fig. 59). The nest was comprised of only six
queen cells. Nests built by mature colonies in combs. The large size was mostly due to
Israel have 1 to 15 combs (Ishay and Brown, multiple envelope layers, probably a response to
1975). high temperatures in the attic. As reported from
Very little data are available on V. germánica New York and Ohio (Morse et al, 1977), colonies
colonies in North America, although pest control in the mid-Atlantic States may be active well into
operators report large nests are not uncommon December.
Vespula germánica
FIGURE 57 — Distribution of Vespula germánica.
The most significant work on the behavior of sounds produced by larvae (Ishay and Brown,
V, germánica is that of Montagner (1966) who 1975), and nest usurpation (Nixon, 1935).
reported a dominance hierarchy among workers. V. germánica workers accept a wide variety of
More dominant workers solicit and receive food arthropods as prey and are opportunistic in the
from foragers, but dominance is not correlated exploitation of food sources. Most workers
to ovarian development. Other works on V. ger- forage within several hundred meters of the
mánica behavior include studies on foraging nest, but some will travel up to 1,200 m (Ferro,
(Free, 1970), worker behavior (Archer, 1972b), 1976). They are also notorious scavengers for
Vespula maculifrons.— Vespula maculifrons

(eastern yellowjacket) is widespread in the Aus-
tral Region of North America from southern
Manitoba to Montana and New Mexico and from
there across to the eastern seaboard (fig. 60). Bio-
logical investigations of this species were con-
ducted by Balduf (1968a, 1968b), Haviland [1962),
MacDonald and Matthews {198 a), Preiss,^ and
Shew. 5
^Preiss, F.J. Nest site selection, microenvironment and

prédation of yellowjacket wasps, Vespula maculifrons
(Buysson), (Hymenoptera: Vespidae) in a deciduous Delaware
woodlot. M.S. thesis. University of Delaware. Newark. 81 pp.
1967. [Mimeographed.]
PN-6542 ^Shew, R.L. The biology and control of vespine wasps. M.S.
FIGURE 58 — Vespula germánica nest, Washington, D.C., thesis, Ohio State University. 59 pp. 1966. [Mimeographed.]
area (J. Nixon).
protein and are attracted in great numbers to

sweets.
V. germánica is a problem to beekeepers
(Walton and Reid, 1976), a general nuisance in
bakeries, markets, parks, and butcher shops
(Kemper and Dohring, 1967), and, in New
Zealand, has been responsible for the closing of
several schools because of the great number of
nesting colonies in surrounding playgrounds.
However, they are also beneficial as a great
number of adult flies of wool maggots (blow
flies, Calliphoridae; Thomas, 1960) and muscid
flies (Schmidtmann, 1977) are taken as prey.
This species has become more abundant and
seems to be replacing V. maculifrons as the
most abundant pest yellowjacket in some areas
of the East (Morse et al., 1977). As it becomes
more firmly established, it has spread slowly
westwards and reached Indiana as of 1976.
Indeed, five colonies were located in houses in
Lafayette, Ind., during the late summer and fall
of 1977. None could be removed for sampling as
they were deep in hollow walls or flooring.
During the summer of 1978, several colonies of
V. germánica were located on the campus of
Michigan State University. Subsequently, this
species was found distributed throughout much
of the State and was responsible for many of the
reported yellowjacket problems. Obviously, the
PN-6543
known distribution of this species is rapidly
FIGURE 59 — Vespula germánica nest in attic of house,
expanding. Washington, D.C., area (J. Nixon).
Vespula maculifrons
FIGURE 60 — Distribution of Vespula maculifrons. Lighter pattern indicates scattered occurrence.
V, maculifrons typically builds its nests (fig. Haviland (1962), Preiss,^ and Shew.^ Other nest
61) in subterranean locations in yards, along locations have also been described such as in de-
roadsides and creek banks, in hardwood forests, caying stumps (Green et al., 1970 and Shew'),
and in urban environments. MacDonald and
Matthews (198_a) reported 143 of 145 nests in
the southeast were subterranean and commonly ^See footnote 4.
found in creek banks in hardwood forests. Sub- See footnote 5.
terranean nest locations were also reported by See footnote 5.
attics (Balduf, 1968a, 1968b; MacDonald and of August or September. The seasonal cycle is
Matthews, 198-a; Simon and Benton, 1968), in somewhat shorter than in the South, and colo-
the walls of structures (Balduf, 1968b, Green et nies typically are smaller.
al., 1970), and even in abandoned cars and inside Haviland {1962) collected 10 nests that ranged
the hull of a boat. In the Washington, D.C., from 9.5 to 30 cm in diameter (table 9). The en-
area, nests in walls now constitute nearly half of velope of these nests was tan-brown and very
homeowner complaints regarding V. maculifrons fragile. The largest had eight comb levels and
(J. Nixon, Silver Spring, Md., personal contained 2,800 adults. Simon and Benton (1968)
commun.). This species also frequently nests in observed, and later collected on December 5 in
structures in Indiana. Pennsylvania, a nest from the loft of a barn that
Colonies in Georgia have a very long seasonal contained 10,960 cells, including 880 queen cells.
cycle. Reproductive production does not begin The adult population of the colony was 5,062
until late September but continues well into De- (598 queens, 3,255 males, and 1,209 workers). A
cember. In the North, colonies are initiated in large colony of V. maculifrons in the southeast
May or June and usually peak in the latter part typically consists of 3,000 to 5,000 workers at
\
<áS^mBP*^^-'^^^ÜK? 'HP5
i
" ^^^^^^^^^^^Ê^ÎS^^-a^
^111 1 ^
PN-6544
FIGURE 61 — Vespula maculifrons: a. Workers attacking a grub; 6, workers feeding on a fallen peach; c, worker in nest
entrance with a load of soil, subterranean colony in a Delaware woodlot {a-c, W.D. Lord); d, combs of a nest
indicating size.
TABLE 9.—Analysis of 10 nests of Vespula maculifrons from

Washington, D.C. (Modified from Haviland, 1962)
Adults
Nest Date Fall Work- Total Queen
No. examined Combs queens Males ers cells cells
M,.r^hr.y-
1 6-27-53 4 0 0 127 1,101 0

2 7-07-56 4 0 0 126 1,495 0
3 7-30-50 5 0 0 390 2,344 0
4 8-08-56 2 0 0 590 2,836 0
5 8-12-56 5 0 0 635 2,665 0
6 9-15-50 5 1 30 1,026 2,967 424 (14 percent)
7 9-16-60 6 120 1,023 1,281 9,762 866 (9 percent)
8 9-27-61 5 54 414 764 4,538 158 (3 percent)
9 9-28-60 8 43 654 2,192 7,755 681 (9 percent)
10 10-26-54 4 12 5 7 1,683 76 (5 percent)
its peak and constructs a nest of 10,000 to infested with Sphecophaga vesparum burra, and
15,000 cells of which nearly 30 percent are queen then only a few cells were parasitized.
cells (MacDonald and Matthews, 198_a\ table In Georgia and western North Carolina, K
10 of this handbook). maculifrons is the primary host of the facul-
Virtually every mature V. maculifrons colony tative social parasite, V. squamosa (MacDonald
in Georgia and North Carolina harbored large and Matthews, 1975). An estimated 40 percent
numbers of scavenger fly larvae in the soil be- of the V. maculifrons colonies in this area are
neath the nest. Larvae of Dendrophaonia quer- usurped by queens of the parasite (MacDonald
ceti were by far the predominant associate, and and Matthews, 198_b),
Fannia larvae were present in lower numbers in There are isolated behavioral observations on
nearly every nest cavity. Larvae of Triphleba V. maculifrons (Balduf, 1968a\ Beamer, 1925\
lugubris were rarely encountered, but when pre- Simon and Benton, 1968), but no studies on
sent in nearly abandoned nests they totally in- foraging behavior or interactions within the
vaded the nest proper. Only 3 of 145 nests were nest.
TABLE 10.—Summary of mature Vespula maculifrons colonies collected after October 15,
1974-75 in and around Athens, Go. (MacDonald and Matthews, 198_ a)
Parameter 1974 1975

Colonies located 30 55
Largest Average Largest Average
Total cells per nest 11.407 6,461 (N = 12) 14,105 9,196 (N = 10)
Average Percent of total Average Percent of total
cells cells
Queen cells per nest 1,622 (N = 12) 25 percent 2,780 (N = 10) 30 percent
Maximum workers per colony 4,875
1
Combs per nest Average Average
Worker 3.9 4.8
Mixed 0.9 0.5
Queen 2.8 3.6
Total 7.7 8.9
^Due to late-season collecting, a large colony was not sampled at its peak
YELLOW JACKETS OF AMERICA NORTH OF MEXICO 69
Although V. maculifrons workers prey upon fewer associates than V, atropilosa. In addition,
economically important insects, such as earwigs the ichneumonid parasite, Sphecophaga ves-
(Kurczewski, 1968) and fall webworm larvae parum burra, was found in a V. pensylvanica
(Morris, 1972), this aspect of their biology nest for the first time in southeastern Washing-
usually has been ignored. In most areas in which ton in 1977. The nest contained one adult para-
it occurs, V. maculifrons is the primary pest site, one empty cocoon, and four intact cocoons;
yellowjacket because of its scavenging habits thus, only one of the 186 analyzed nests from
and the great number of colonies that build 1971-77 contained this parasite. This may be
nests in yards, golf courses, recreational areas, due to the behavior of V. pensylvanica workers,
and buildings. In some areas of the northeast, it which immediately attack and kill introduced
appears that V. maculifrons is being replaced by Sphecophaga, whereas V. atropilosa workers fre-
V. germánica as the primary pest species, and in quently ignore adult Sphecophaga when they are
the southeast V. squamosa competes with V. introduced.
maculifrons for primary pest status. V. pensylvanica workers will accept a wide
Vespula pensylvanica.—Vespula pensylvanica variety of prey (Akre et al., 1976; MacDonald et
(western yellowjacket) occurs in the Canadian al., 1974). Included are slugs, phalangids, spi-
and Transition Zones in western North America. ders, grasshoppers, flies, cercopids, and bugs.
This species also occurs in Hawaii on the islands The preponderance of prey collected were mem-
of Kauai (WilUams, 1927), Oahu (WilUams, 1937), bers of Hemiptera-Homoptera. Members of this
Maui, and Hawaii ( J. W. Beardsley, Univ. Hawaii species also scavenge extensively for protein,
at Manoa, personal commun.), and in Mexico especially later in the year.
(Ebeling, 1975; Snelling, 1970), Distribution is The behavior of V. pensylvanica colony mem-
given in figure 62. bers in the nest was observed by Akre et al.
Nests of V, pensylvanica are usually subterra- (1976). The division of labor among workers is
nean (Bohart and Bechtel, 1957; Duncan, 1939; not as sharply defined as in honey bees, and
Smith, 1956), and most are constructed in rodent workers performed a number of activities during
burrows (MacDonald et al., 1974), A few nests the same day. Social interactions similar to the
are built in other dark cavities such as in attics dominance hierarchy described by Montagner
or between the walls of houses (Buckell and (1966) for V. germánica and V. vulgaris were
Spencer, 1950; Ebeling, 1975; Spencer, 1960). In exhibited by the workers of V. pensylvanica. In
Pullman, Wash., subterranean nests were typi- another study (Akre et al., 1975), 80 percent of
cally 10 to 15 cm below the soil surface (fig. 63 the workers foraged within 1,100 ft of the nest.
d, e) with entrance tunnels of 10 to 30 cm (Mac- V. pensylvanica is the primary pest yellow-
Donald et al., 1974). jacket in the West from Washington to Califor-
Duncan {1939) tabulated one colony of V. pen- nia. Periodic population outbreaks, associated
sylvanica comprised of 4,768 workers and the with warm, dry springs, occur every 3 to 5 years
foundress queen. The colony probably would or slightly longer, which create severe problems
have grown for another month if it had not been for people engaged in logging, raising fruit, or in
collected, but it had already constructed a nest recreation-associated activities. A number of
of 10,468 cells. He mentioned another nest of colonies establish their nest in yards or recre-
17.25 by 16 by 10 inches but gave no other data. ation areas, and the chance of colony distur-
Table 11 lists the size of mature (producing bance resulting in stings is great. Since this
queens) colonies collected 1971-77 in Pullman, species is a scavenger, it is a constant com-
Wash., and La Grande, Oreg. The largest nest panion at picnics or at food dispensing facilities
collected contained 12,316 cells, and the maxi- and frequently must be controlled.
mum number of workers in a colony was 3,933. Vespula vulgaris,—Vespula vulgaris (common
Most colonies had nests of 4,000 to 10,000 cells. yellowjacket) is a Holarctic species transconti-
MacDonald et al. (1975b) discussed nest nentally distributed in the Nearctic Region (fig.
associates occurring with V. pensylvanica and 64). In western North America, this species is
V. atropilosa. As a rule, at least until colony de- prevalent in heavily forested areas. In Hawaii, V.
cline, V. pensylvanica has cleaner nests and vulgaris occurs only on the island of Maui (Ho-
TABLE 11,—Size of mature (producing queens) Vespula pensylvanica

colonies 1971-77^
Few- Average Average Total

Most est number Fewest Most number Maximum analyzed
Year Nests combs combs of combs cells cells of cells workers nests
Ar..»v,A^-
1971 3 8 4 6.0 8,595 12,316 10,751 2,266 6

1972 6 8 5 6.3 1,422 8,013 4,116 1,125 9
1973 28 7 4 5.9 2.547 6,689 4,623 1,151 47
1974 16 8 6 7.0 2,926 10,438 6,051 1,883 31
1975 13 7 4 6.1 1,342 6,760 3,859 1,777 55
1976^ 0 7 3 5.0 703 4,919 2,494 924 11
1977 18 9 4 6.3 1,574 11,838 6,842 3,933 30
1971-
77» 84 9 4 6.3 1,342 12,316 5,435 3,933 178
'Colonies from 1971-74 are only from Pullman, Wash.; colonies were also collected
from La Grande, Oreg., during 1975-77,
^Due to late season, no mature colonies were collected. Totals include aU nests
»Excluding 1976.
warth, 1975). It also occurs in Mexico (Ebeling, personal commun.). The nest envelope and
1975), Detailed biological studies of this species, combs are usually constructed of decayed wood
such as those of Archer (1972a, b, c, 1973, 1977b), fibers and are red- to tan-brown and very brittle.
Guiglia {1972), Kemper (1961), Kemper and Dbh- In England, the nest is initiated in May and
ring (1967), and Potter^ were done in Europe. ultimately contains an average of nine combs
These data and much of his own have been sum- when the colony peaks development in Septem-
marized by Spradbery (1973a), No in-depth bio- ber (Spradbery, 1973a). In Washington, northern
logical study of this species in America is avail- Idaho, and northeastern Oregon, the nest is also
able, but Balduf (1968b) made some observations initiated in May or early June, and colonies peak
on nests and seasonal development of colonies in in September. Some colonies are still active in
Minnesota. October, but rarely in November; however, nests
In Europe, most V. vulgaris nests are subter- are comprised of only three to five combs.
ranean, although some nests are constructed in The maximum number of cells in a nest ana-
the walls of houses or even in aerial locations lyzed by Spradbery (1973a) was 16,832 including
(Spradbery, 1973a; Weyrauch, 1935), In the 3,888 queen cells. In Germany, Kemper (1961)
Western United States, this species usually reported a nest with 21,692 cells, including
builds its nests in rotten logs or stumps, in for- 4,028 queen cells. Duncan (1939) illustrated one
est duff, or in the soil (fig. 65). Entrance tunnels large nest of V, vulgaris in California con-
to nests vary from 4 to 100 cm, usually 10 to 15 structed by a perennial colony that had 22 func-
cm, with most nests found about 10 cm below tional queens at the time of collection. The nest
the soil surface. Some colonies, especially in had 21 comb levels, and was 46 by 40 by 30
western Washington, construct nests between inches. No count was made of the adult popula-
the walls of houses. Similarly, in New York, 11 tion but they filled four 1-gallon jars. Four gal-
of 21 nests located during 1977 were between lons of V. pensylvanica, a larger species, would
the walls of a building (R. Keyel, Cornell Univ.. be over 60,000 workers. Another California col-
ony contained 2,951 workers when killed. Analy-
^Potter, N. B. A study of the biology of the common wasp, ses of 19 mature nests collected during 1975-77
Vespula vulgaris L., with special reference to the foraging in Washington, Idaho, and Oregon showed them
behavior, Ph.D. thesis, Univ. Bristol, England. 75 pp. 1964. to be much smaller, averaging 2,100 cells, with
[Mimeographed.] 10 percent reproductive cells. The largest nest
contained 4,972 cells. No study has been made Behavioral interactions inside the nest were
of the nest associates of V. vulgaris in North studied by Montagner (1966) and Potter.^« Pot-
America; however, 7 of the 19 nests mentioned ter's studies included foraging behavior and daily
above contained cocoons of the pupal parasite,
Sphecophaga vesparum burra. ^°See footnote 9.
FIGURE 62 — Distribution of Vespula pensylvanica. Lighter pattern indicates scattered occurrence *

PN-6545
FIGURE 63 — Vespula pensylvanica: a. Mud turret surrounding nest entrance late in the season; b, c, workers entering and
leaving, guard workers; d, e, subterranean nest near Washington State University golf course; /, workers
scavenging flesh from fish carcass.
activities of the members of the colony, whereas variety of prey, similar to that attacked by V.
Montagner studied social dominance (hierarchy) pensylvanicUy including a variety of caterpillars,
among the workers of the colony with certain in- small beetle larvae, hemipterans, homopterans,
dividuals being dominant and demanding food and flies. In addition, workers of this species are
from foragers. notorious scavengers attracted to nearly any
V. vulgaris workers are predators on a wide protein or sugar source.
FIGURE 64 — Distribution of Vespula vulgaris.

PN-6546
FIGURE 66 — a, 6, SttbtaTanean Vespula vulgaria nests, Harrison, Idaho.
This species is a nuisance in food-dispensing programs nearly ever year (Grant et al., 1968).
facilities in Germany (Kemper and Döhring, The 1973 outbreak of yellowjackets in the Paci-
1967), is described as being the most annoying fic Northwest included high populations of V.
to humans of any yellowjacket in Norway vulgaris in forested areas such as western Wash-
(Lóken, 1964), and is second only to V. pensyl- ington and northern Idaho. In 1974, severe prob-
vanica as a pest species in western North Amer- lems were caused by high populations of this
ica (Ebeling, 1975). In San Mateo County, CaUf., species in Alaska (Davis, 1978).
these yellowjackets are the subject of control
ECONOMIC IMPORTANCE
It is impossible to accurately assess the true Beneficial Aspect
economic importance of yellowjackets in North
America because information pertaining to the Reports mentioning the beneficial aspect of
beneficial aspects of these wasps remains largely yellowjackets include those by Gaul {1952),
unreported, and even the pest status of several Greene et al. {1976), Kuhlhorn {1961), MacDonald
species is not fully documented. For example, al- et al. {1974), and Schmidtmann {1977). These
though researchers have frequently observed studies showed that yellowjackets readily at-
yellowjackets preying on defoliators in forests, tacked house flies and blow flies. From these
little has been published about this prédation data, and from observations of yellowjacket
and its possible values. Similarly, no data are workers preying on flies in large concentrations
available on the beneficial nature of yellowjac- in places like mink and cattle ranches, it can be
kets in other situations, such as in agriculture or inferred that yellowjackets help to control fly
urban environments. Conversely, the pestiferous populations. Other economically important in-
nature of scavenger species has been only spor- sects used by yellowjackets as prey are Lygus
adically and incompletely reported with no com- bugs, several species of Homoptera including
prehensive evaluation of possible economic im- spittle bugs, various caterpillars, and grass-
pact. hoppers (MacDonald et al, 1974). Fall webworm
larvae, Hyphantria cunea Drury, are taken as mation was gathered from accumulated news-
prey by Vespula maculifrons, V. vulgaris, and paper and magazine articles, journals, technical
Dolichovespula arenaria (Morris, 1972). White et bulletins, personal communications from other
al. (1969) and Howell and Davis (1972) showed V. researchers, and letters from numerous citizens
pensylvanica, and perhaps workers of other throughout North America who have had yel-
species of yellowjackets, attacked codling low jacket problems.
moths, Laspeyresia pomonella (L.), within their
experimental plots and interfered with their test Losses in Agricultural Areas
results. In addition, R. W. Zwick (Oreg. State
Univ., personal commun.) showed that workers Yellowjackets often become serious pests in
of D. arenaria could completely control pear fruit-growing regions, and during years of abun-
psylla on pear under caged conditions. The dance they may stop harvesting operations. This
above reports imply that yellowjackets are bene- occurred at a peach orchard in Oregon in 1969
ficial and help control populations of a number when thousands of Vespula pensylvanica workers
of insect pests. Still needed are studies on yel- were attracted to the fruit. Pickers were frequent-
lowjacket prédation that show where it is of ly stung as they moved their ladders over yellow-
possible benefit to man and his crops. We be- jackets feeding on the soft, windfallen fruit on the
lieve that many species will be found to be highly ground. Losses due to harvesting delays affect or-
beneficial once data are collected. chard workers in lost wages and in medical ex-
penses that may be incurred. The grower must
pay higher production costs, and additional loss-
Economic Losses es result from lowered income from reductions in
fruit quality.
Although most yellowjackets can be regarded
Yellowjackets have also been troublesome to
as beneficial, a few pestiferous species are re-
growers in pear and apple orchards (Buckell and
sponsible for economic losses (which can be con-
Spencer, 1950; R. W. Zwick, Oreg. State Univ.,
siderable) in agricultural, recreational, and ur-
personal commun.). The problem occurs during
ban-suburban environments during years of
years of peak populations of the aerial-nesting
peak populations or outbreaks. Again, there is a
species Dolichovespula arenaria (aerial yellow-
lack of information as to the economic losses
which can be attributed directly to the activities jacket) and D. maculata (baldfaced "hornet")
since many colonies build their nests in the fruit
of yellowjackets. Most information on this sub-
trees. Laborers are commonly stung as they per-
ject is found in vague, fragmentary reports,
form routine chores in the orchard that disturb
which, at best, are only attempts to estimate
losses in a particular locality. For example, and redisturb the colonies.
Poinar and Ennik (1972) cited a report by Similar problems also occurred from thous-
Hawthorne, ^^ stating that 1968 losses to agri- ands of foraging Vespula and Dolichovespula
cultural operations in California were estimated spp. workers attracted to a fruit processing
at $200,000. These losses were attributed to dis- plant in Toppenish, Wash., during August and
ruptions due to Vespula spp. bothering and September 1975. Most yellowjacket colonies had
stinging pickers, feedlot workers, laborers at entered decline, and the workers were attracted
meat processing plants, and other persons in- by the many sweet juices and fruit remnants
volved in agriculturally oriented occupations. scattered throughout the plant. Some laborers
A brief description of some of the many prob- were stung, but more importantly, many were
lems attributed to yellowjackets follows. Infor- terrified of the yellowjackets and production de-
clined greatly. Changes were made in the plant
i^Hawthorne, R. M. Estimated damage and crop loss caused operations so that less food debris and juices
by insect/mite pests. 1968. Calif. Dept. Agrie. Bur. Ent., 11 were available to attract yellowjackets, and
pp. 1969. [Mimeographed.] other control procedures were employed before
the problem was alleviated.
Duncan (1939) and Howell (1973) reported yel- tions of V. germánica on their bee colonies since
lowjackets feeding on grapes and removing the this yellowjacket was introduced into the coun-
juices. This appears to be especially true in re- try in 1945. During years of high yellowjacket
cent years in newly developed vineyards of wine populations, the wasps cause havoc and consid-
grapes in the Pacific Northwest. For example, erable financial losses to the industry. A survey
several growers in Oregon and Washington by Walton and Reid (1976) revealed that 88.6
reported losses of nearly half their crop of red percent of the beekeepers they surveyed con-
grape varieties (mostly pinot noir) during 1975 sidered V. germánica a nuisance and 73.6 per-
due to yellowjackets. Specimens examined from cent a cause of financial loss. During the 1975-
one vineyard were V. pensylvanica, but other 76 season, V. germánica workers were estimated
species might also be involved. According to the to have destroyed 3,900 colonies and partly
growers, the yellowjackets land on the larger affected more than 10,000 others.
clusters, sever the skin of one or more grapes and In North America, beekeepers also experience
feed on the juice. They reported that many clus- financial losses due to colony destruction by yel-
ters were damaged and, perhaps equally impor- lowjackets, especially V. pensylvanica and V.
tant, the remaining grapes are susceptible to the germánica. The yellowjackets may also delay
brown or bunch rot fungus. In addition, pickers operations during honey extraction because of
were frequently stung (yellowjackets feeding on the apprehension or actual stinging of personnel
the back side of the clusters were not seen), result- at the apiary. Another, indirect, problem for bee-
ing in delays in harvesting. keepers in many fruit-producing regions occurs
Nearly identical problems are experienced when the yellowjackets pierce ripe pears,
with the grape crop and yellowjackets in Penn- peaches, and other fruits. If there is a lack of
sylvania and New York. During 1976, many nectar, large numbers of honey bees (which only
grapes in Pennsylvania were damaged, and occasionally pierce sound fruit) are attracted by
many delays occurred in harvesting because the juices of the pierced fruit. This sometimes
pickers were stung. Unfortunately, no concrete causes severe problems for fruit growers and
data were collected on these incidents, but the pickers, and the apiarist may be asked to re-
problems were described as "considerable." The move or control the hives.
species involved were probably V. maculifrons Yellowjackets are also a problem for commer-
and perhaps K germánica. In New York, yel- cial Christmas tree growers, especially during
lowjackets also cause some problems to grape pruning operations in midsummer. Laborers
growers (W. B. Robinson, Cornell Univ., personal often do not see the nests on the opposite side of
commun.). The damage seems to be most ex- the tree until it is too late, and they are fre-
tensive in small personal lots of grapes, and quently stung repeatedly when the tree is vi-
sweet, early ripening, tender-skinned varieties brated.
are the most susceptible. The yellowjackets Yellowjackets may become equally trouble-
sever the skin of the fruit to get at the juices. In some in logging and sawmill operations. During
some of these small plots, as much as 95 percent years of high yellowjacket populations in for-
of early season grapes are lost to yellowjacket ests, operators of heavy equipment, such as
damage. Later varieties of grapes that ripen in graders, tractors, skidders, trucks, and loaders,
cold weather are much less susceptible. Again, are prone to attack from yellowjackets whose
the species of yellowjackets involved has not nests are disturbed by vibrations or displace-
been determined, but V. maculifrons and V. ment, and much time is lost while the wasps are
germánica workers are probably the principal controlled. Occasionally, workers at sawmills are
pests, with some damage by workers of Vespa troubled by yellowjackets. During the summer
crabro. of 1973, during a "population outbreak" of
In agricultural areas, it is probably the bee- yellowjackets in the Pacific Northwest, one saw-
keeper who consistently experiences the most mill near Wenatchee, Wash., closed after person-
problems with yellowjackets. These problems nel at the mill refused to tolerate these insects
are shared by beekeepers throughout the world. any longer. The mill opened only after the work-
In New Zealand, they have combated depreda- ers were convinced that a control program was
well underway. Similarly, a building contractor areas in southern California because of yellow-
constructing a house near Moscow, Idaho, stopped jackets. Howell et al. (1974) stated that yellow-
all building until the future homeowner instituted jackets are considered one of the most serious
control procedures for the extremely abundant pests in outdoor recreational areas in Georgia.
yellowjackets. The delay was costly to both the Although yellowjackets undoubtedly create sim-
contractor and the homeowner in lost wages, pen- ilar problems in other areas of North America,
alties, and time. no additional information is available.
"In a recent 5-year period, insect stings Officials of private organizations and govern-
accounted for over 5 percent of all Forest Ser- mental agencies providing recreational facilities
vice medical treatment and lost-time accidents" are becoming increasingly concerned about
(Putnam, 1977). These stings are especially finding better ways to protect their patrons
numerous and frequent when Forest Service from yellowjackets. They are especially con-
personnel are fighting forest fires. The fire- cerned about persons hypersensitive to the ve-
fighters are commonly stung and must be re- noms. Most are planning to increase their bud-
moved from the firelines as they encounter gets to contain contingency plans for yellow-
hordes of yellowjackets flying above their jacket control programs.
smoke-filled subterranean nests. The firefighters
are also disturbed by yellowjackets during meals Losses in Urban and Suburban Areas
when the scavenger species are attracted to the
food and juices being served. To partially alle- During years of high yellowjacket popula-
viate these types of problems, the Forest Service tions, these wasps are responsible for economic
has published a bulletin concerning the control losses in urban and suburban areas, just as they
of stinging and biting insects at campsites are in agricultural and recreational areas. These
(Putnam, 1977). losses occur at city parks, playgrounds, zoos,
amusement parks, and athletic facilities. The
Losses in Recreational Areas presence of large numbers of yellowjackets at
these places causes considerable alarm among pa-
Everyone who has camped or picnicked at trons and results in severely reduced atten-
recreational facilities, especially in forested dance. Revenues from admissions drop pro-
areas, is aware of the nuisance of scavenging portionally. Operators of concession stands also
yellowjackets. These pests are sometimes re- suffer financially. Customers refuse to buy food
sponsible for considerable financial losses to gov- and beverages since this material is highly at-
ernment agencies and private organizations that tractive to yellowjackets. During 1973, the Port-
provide the facilities since attendance revenues
land, Greg., zoo was experiencing all the above
are often drastically reduced during years of yel- difficulties, but once an effort was made to cover
lowjacket outbreaks. For example, during the all garbage cans in the area, to remove all food
1973 outbreak in the Pacific Northwest, private, remnants, and to introduce a limited control
county. State, and Federal parks and recreation program by baiting, the yellowjacket density
areas were nearly deserted. Revenue dropped quickly dropped to acceptable levels.
considerably, and many bitter complaints about
yellowjackets were received by the Forest Ser- In some communities, yellowjacket problems
vice, National Park Service, Bureau of the Inter- are severe enough nearly every year to neces-
ior, and the Army Corps of Engineers, all main- sitate organized control programs. Often local,
taining recreational facilities of one type or county, or State agencies provide important bio-
another in this area. In addition, resorts in Incrical and ecological studies and attempt to
northern Washington and Idaho lost consider- control these pests. For example, one large
able revenue until vigorous yellowjacket control county mosquito abatement district in Californ-
programs were instituted. Poinar and Ennik ia spent nearly 6 percent of its operational bud-
(1972) estimated losses at certain resorts in Cal- get from 1969 to 1972 for evaluation and control
ifornia reached $5,000 annually per resort. of pestiferous yellowjacket species in their dis-
Wagner and Reierson (1969) reported similar re- trict.
ductions in attendance at many recreational Yellowjackets can also be a serious and costly
occupational hazard to certain types of employees lems with yellowjackets, especially at mess fa-
in cities and suburbs. Garbage collectors and cilities and garbage collection sites.
operators of heavy equipment at municipal land- The greatest number of yellowjacket problems
fills and refuse dumps are sometimes repeatedly occur with homeowners on their own property.
stung, requiring significant numbers of them to With the increased interest in barbecues and
leave their jobs to seek medical attention. In outdoor cooking, the number of encounters with
1973, during the peak populations of V. pensyl- yellowjackets has increased. Many people claim
vanica and V. vulgaris in the Pacific Northwest, they are unable to enjoy their own yards during
personnel of one industrial accident insurance outbreak years, and are greatly annoyed during
group became alarmed at the number of compen- typical years. Although some effective control
sation requests they processed from employees procedures are available to homeowners, many
stung by yellowjackets. employ pest control specialists to achieve con-
Other employees in urban-suburban areas af- trol, especially when the yellowjackets build
fected by yellowjackets include workers at food their nests in or close to their homes.
canneries (several companies have reported cost-
ly delays); meat processing plants; jam, jelly, Although, during certain years, yellowjackets
and pickle factories; and wineries. The most im- are responsible for large economic losses in agri-
portant single factor in many of these cases is culture, in recreation industry, and in the urban-
not the stinging, but the annoyance factor and suburban environment, it should be stressed
the threat of being stung. The employees are that most people are terrified of Hymenoptera
frightened of yellowjackets, and it affects their and yellowjackets in particular. There is no way
work greatly. to determine the economic value of this stress
Personnel at military bases, often located in or upon people, but its importance must be recog-
adjacent to suburban areas, also encounter prob- nized.
MEDICAL IMPORTANCE
Yellowjacket Sting Apparatus form egg pods or glue the eggs to the substrate.
In aculeate Hymenoptera, the ovipositor has
Morphology become a stinging device that no longer func-
tions in egg laying. The genital opening (gono-
Although cartoonists delight in portraying the pore) from which the eggs pass is anterior to the
sting or "stinger" of wasps and bees as a con- sting apparatus, which is flexed up out of the
stantly protruding spike, the real sting is part of way during oviposition. The accessory glands
a complex apparatus hidden in a cavity at the have also been modified: One has become a poi-
end of the abdomen. It is actually a modified son gland ("acid" gland); the function of the
ovipositor, or egg-laying device, which com- other (Dufour's or "alkaline" gland) is unknown,
prises the external genitalia of many female in- although it may be associated with the produc-
sects. The visible portion of a typical ovipositor tion of pheromones.
consists of three pairs of more or less elongate Detailed discussions of the origin of insect
structures, or valves, which are used to insert genitalia (including ovipositors) and their rela-
the eggs into a substrate such as plant tissue or tionships to other appendages were presented by
soil. One pair of valves often serves as a sheath Smith (1969, 1970). The morphology and func-
and is not a piercing structure; the other two tion of the venom apparatus of Hymenoptera was
pairs form a hollow shaft, which pierces the sub- discussed by Maschwitz and Kloft (1971), and a
strate by a back and forth sawing motion of one specific discussion of the yellowjacket sting ap-
pair held in position by the other. The eggs then paratus was presented by Spradbery (l97Sa).
pass down through the shaft. Accessory glands As shown in figure 66, the sting apparatus in
(usually two) within the body of the female often its normal, retracted position is located between
inject secretions, through the ovipositor, which abdominal tergum 8 (a small sclerite concealed
beneath tergum 7) and sternum 7. In figures 67 picts one-half of the apparatus from the side.
and 68, the apparatus is illustrated with the From each small triangular plate (gonocoxite 8
hoodlike tergum 8 removed. Each side consists or valvifer I), arises a long, slender lancet (go-
primarily of three plates. Figure 67 illustrates napophysis 8 or válvula I). The triangular plate
these paired plates ventrally, and figure 68 de- articulates ventrally with a large oblong plate
ABDOMINAL
TERGA
FIGURE 66 — Yellowjacket worker abdominal terga and sterna pulled apart to show location of sting apparatus (relaxed
position—only one-half visible) and associated glands. The sting, that is, shaft of stylet plus lancets, is partly
visible behind the semitransparent sting sheath.
FIGURE 67 — Ventral view of relaxed yellowjacket worker sting apparatus, partially flattened out. Compare with lateral view
of one-half of apparatus, figs. 66 and 68a). Note that the lancet on the right (unshaded) overlaps the one on the
left (shaded) shortly after they converge.
Abbreviations for figures 67 and 68
OP oblong plate SS sting sheath

QP quadrate plate L lancet
TP triangular plate s shaft of stylet
ap anterior process of oblong plate b bulb of stylet
PD poison duct a basal arm of stylet
AL anal lobe
FIGURE 68 — Semidiagrammatic lateral views of one-half of yellowjacket worker sting apparatus, eighth tergum removed.
Nonagitated and stinging yellowjackets with enlarged apical gastral segments also illustrated, a, Sting fully
retracted and concealed behind oblong plate and sting sheath in normal, rekxed position within body of wasp.
6, c. Sting at maximum thrust. Both quadrate and oblong plates are displaced upward. As the sting slides
against the inner surface of the curved oblong plate, it pulls on the plate's lower edge, flattening the plate
somewhat and thus broadening its lateral configuration. At the apex of the oblong plate, the flexible sting
sheath is bent downward and parallels the sting. 6, Quadrate plate shifted posteriorly, triangular plate
(shaded) rocked upward, lancet retracted, c. Quadrate plate shifted anteriorly, triangular plate rocked down-
ward, lancet extruded. (See fig. 67 legend for explanation of abbreviations.)
(gonocoxite 9 or val vifer II). From an anterior angular plates are thus displaced forward in
process on the oblong plate arises a basal arm alternate strokes, each sliding on its track
of the stylet (gonapophyses 9 or valvulae II): against the shaft. The tips of the lancets are
these arms fuse into a swollen, hollow bulb, equipped with tiny barbs to facilitate penetra-
which then narrows into a tapering shaft. The tion, and as shown in figure 69, they literally
stylet interlocks along its entire length with the saw through the victim's flesh as each in turn is
two lancets, which slide freely along its ventral thrust forward and anchored in place by the
edges. Where they converge at the bulb, they barbs. Thus, the action of the sting closely
form a channel with the shaft through which the resembles that of many ovipositors during egg
poison flows. As seen in figure 68, the oblong laying. (The barbs on a yellowjacket or hornet
plates overlap the sting bulb and part of the sting are much smaller than those of the honey
shaft. The distal ends of the plates are pro- bee. Consequently, the sting does not normally
longed into sting sheaths (gonostyli 9 or valvu- become fixed in a person's flesh, and the wasp
lae III), which laterally cover the remainder of may quickly withdraw with a upward pull of its
the sting shaft when it is retracted. abdomen and sting again.)
The remainder of each half of the apparatus
consists of a large quadrate plate, modified from Venoms
abdominal tergum 9, which overlaps the oblong
plate and, like the latter, articulates with the tri- Most aculeate wasps are solitary and use their
angular plate. A hair-covered anal lobe derived stings primarily for subduing prey. Their ven-
from the tenth abdominal segment is connected oms are also specialized for this purpose and
by a membrane to the posterior of the appa- most cause slight and temporary pain to hu-
mans. The social Vespidae (including yellowjack-
ratus. The poison gland consists of two slender
filaments which arise from one end of a prom- ets), however, use the sting primarily as a defen-
inent, heavily muscled reservoir, or poison sac sive weapon, and the venom contains materials
(fig. 66). This in turn empties, via a narrow neck, that cause intense pain to vertebrates.
into the sting bulb, as does Dufour's gland (fig. Yellowjacket venoms were investigated by
67). Geller et al. (1976), Habermann (1972), and Yo-
shida et al. (1976). Feingold (1973) discussed the
materials present in the venom and their action
Sting Function
on the human body. Table 12 lists the active
constituents of wasp (yellowjacket), bee, and hor-
An attacking yellowjacket grips firmly with
net (Vespa crabro) venoms. A more complete
its legs (the hornet, Vespa crabro, often holds on
listing, by species, was compiled by Geller et al.
with its mandibles as well), elevates the oblong
(1976).
and quadrate plates to expose the sting, then
Yellowjacket venoms are highly complicated
plunges the tip of the interlocked lancets and
mixtures of pharmacologically or biologically
stylet into the skin with a downward thrust of
active agents or both (Habermann, 1972). These
the abdomen. Simultaneously, contraction of the
agents cause the contraction of smooth muscle,
poison sac muscles injects the venom into the
stimulate glands of external secretion, increase
sting bulb and through the channel formed by
capillary permeability, produce vasodilation
lancets and shaft, much like a hypodermic nee-
with a resulting fall in blood pressure, destroy
dle; however, penetration is not a matter of a
normal tissue barriers, and cause intense pain.
single stroke. The curious person who inspects a
The effects of individual constituents were given
bee or wasp for the first time is usually sur-
by Feingold (1973).
prised at how flexible the sting is and the diffi-
culty in piercing human skin with it once the in-
sect is dead. As shown in figure 68, on each side Yellowjacket Stings
of the apparatus the quadrate plate rapidly
moves back and forth during stinging, pivoting Prior to the 1960's, literature on Hymenoptera
the triangular plate on its articulation with the as medically important insects was meager, and
oblong plate. The lancets attached to the tri- their capabilities to cause death of humans was
STYLET
LANCET
FIGURE 69 — Ventral views of sting tip showing: sting penetration by alternate forward strokes of the lancets, sliding along
the shaft of the stylet, and lancets pulled out of their track on the shaft. Although the lancets lie side-by-side
for a short distance after converging at the base of the shaft, their expanded, bladelike ends overlap at the
tip. The facing blade surfaces are each concave, creating a tubular poison channel between them.
TABLE 12,—Pharmacologically and biochemi- sects may transmit pathogenic micro-organisms

cally active constituents of wasp, bee, and hor- by way of the sting.
net venoms (modified from Habermann, 1972) Although Parrish (1963) used death certifi-
[H-, present; —, absent; ?, questionable] cates for some of his information, Fluno (1961)
suggested such documents do not include all
Substance Wasp Bee Hornet deaths due to wasp stings. He pointed out that
Biogenic amines reactions to stings are not even categorized in
Histamine + -f- + morbidity reports at some hospitals; a code is
Serotonin + — + used which combines all types of venoms in-
Dopamine + + - cluding those of snakes, scorpions, wasps, and
Noradrenalin + + - spiders. Parrish (1959) was of the opinion that
Acetylcholine - - +
many coroners are not aware of anaphylactic
Protein polypeptide t oxins
(nonenzymatic)
shock as a cause of death; therefore, many
Melitten - + — deaths reported as heart attacks or heat strokes
Apamin - + - may have been caused by hymenopteran stings.
MOD peptide - + - For these reasons, the exact number of fatalities
Minimie - + - caused by yellowjackets is unknown; however, it
Kinins (') n is probably much higher than suspected.
Enzymes In recent years, there appears to be an in-
Phospholipase A + + + creasing amount of interest and concern about
Phospholipase B + 7 -f
+ + yellowjackets by the general public. Much of
Hyaluronidase —
this concern probably stems from increased en-
'Vespula kinin.
^Hornet kinin. counters with these wasps due to the present
emphasis on outdoor activities such as camping,
picnicking, and hiking. Fluno (1961) stated that
about half of the 10,000 requests for information
about wasps and their control received yearly by
largely undocumented. More recent reports, US DA were accompanied by statements that
such as those by Parrish (1959, 1963), Leclercq one or more members of the family reacted se-
and Lecomte (1975), and Lecomte and Leclercq verely to wasp stings. Although there are no re-
(1973), have drawn attention to the medical im- cent data on the number of requests received
portance of Hymenoptera. Parrish (1963) ana- yearly, the number has apparently increased
lyzed fatalities due to venomous animals in the greatly. In addition, thousands of requests for
United States from 1950 to 1959. The study re- information on wasps are received by univer-
vealed that 229 of 460 recorded deaths were due sities and other State and Federal agencies.
to the stings of hymenopterans such as yellow- There has also been a corresponding aware-
jackets, other wasps, ants, and bees. Barnard ness on the part of physicians and other medical
(1973), in a publication of reports of the Insect researchers of problems related to hymenop-
Sting Subcommittee of the American Academy of teran stings. This has resulted in a number of
Allergy, reported more than 400 fatal reactions in papers on various aspects of these subjects,
a 10-year period due to one or more stings. The which were reviewed and listed for 1953-70 by
reports indicated four main types of pathology Barr (1971) and for January 1970 to November
were responsible for the fatalities. One hundred 1972 by Nowak (1972).
autopsies revealed 69 deaths were due to respira-
tory problems, 12 due to anaphylactic shock, 12 Fatal Reactions to Stings
to vascular involvements, and 7 to neurological
problems. One to 2 percent of the 400 fatalities Great variability exists among individuals in
were attributed to bacterial septicemia. Benton their reactions to insect stings; however, a con-
and Heckman (1969) suggested that most infec- siderable amount of information has been pub-
tions probably result from scratching the wound lished on the subject, which provides a better
site, although there is a possibility stinging in- understanding of the medical problems involved.
Zeleznick et al. {1977) were of the opinion that rel- "A shock or anaphylactic reaction would ex-
atively few deaths occur from anaphylactic shock hibit any of these symptoms in addition to two
due to stings by wasps, hornets, yellowjackets, or more of the following: (1) lowered blood pres-
and bees; however, they noted estimates of 0.4 sure, (2) cyanosis, (3) collapse, and (4) inconti-
to 0.8 percent of the population that experienced nence and unconsciousness."
a systemic reaction to stings. Therefore, the po- While immediate reactions are an instant
tential for these types of reactions is great cause for concern, delayed reactions resulting in
(Frazier, 1976). Perlman {1962) found severe re- death may occur several hours after the sting.
actions to arthropod bites and stings statisti- Bukantz {1975) cited the results of 100 autopsies
cally no more frequent in patients with general that showed 59 of the deaths occurred within 60
allergic disorders than in the general population, min, 22 between 1 and 6 h, 7 between 6 and 96
nor were they correlated with heredity. h, and 9 after more than 96 h. The interval of
time before death was unknown for three of the
Symptoms of Allergic Reaction victims.
Feingold (1973) reported the following symp-

toms concerning local reactions to a sting (fig.
70). He stated, "Immediately following the
sting, intense burning is experienced at the site
of the sting to be followed after several minutes
by swelling and intense itching. The swelling
may be localized to a few centimeters immedi-
ately surrounding the sting or the edema may
involve an entire extremity or other segment of
the body."
Symptoms of a generalized systemic reaction
may range from mild to severe. Frazier {1976)
stated, "Such reactions can be delayed, pre-
senting serum sicknesslike symptoms of fever,
headache, malaise, urticaria, lymphadenopathy,
and polyarthritis. It is an immediate reaction,
however, that presents the physician with a
medical emergency. Even a sligh't systemic re-
action with symptoms of generalized urticaria,
itching, malaise, and anxiety should be assessed
and treated on a long-term basis in the realiza-
tion that the next time the patient is stung the
results may be far more serious, even life-threat-
ening.
"A moderate systemic reaction may be
marked by any of the symptoms mentioned and
two or more of the following: (1) constriction of
throat or chest, (2) abdominal pain, nausea, vom-
iting, (3) dizziness, (4) wheezing, and (5) general-
ized edema.
"A severe systemic reaction may include any of
PN-6547
these symptoms and two or more of the fol-
lowing: (1) labored breathing, (2) difficulty in FIGURE 70 — Localized sting reactions to Vespula pensyl-
vanica: a. Swollen foot about 6 hours after 2 stings by a single
swallowing, hoarseness, or thickened speech, (3)
worker; b, sting blister on ankle, 4.2 X 2 X 1.5 cm high, 36
weakness, (4) confusion, and (5) a feeHng of im- hours after being stung twice by a single worker. This is a pri-
pending disaster. mary reaction, not a secondary infection.
Ancillary Factors and Sting Fatalities Long-Term Treatment

Anyone experiencing any of the symptoms of
Ancillary factors, such as age of victim, type
sensitization to stings should seriously consider
of insect involved, and the number of stings,
desensitization procedures. Frazier (1976) stated
were presented by Barnard (1973). From 100
"the physician's responsibility. . does not end
postmortem cases, it was determined that 64
with the successful treatment of the acute epi-
were under 50 years of age. '*Bees" were respon-
sode. The patient is totally vulnerable and may
sible for 44 of the deaths, *'wasps and hornets"
die quickly if stung again. Desensitization
for 26, **yellowjackets" for 18, and stings of un-
should be begun at once."
known origin for 12. Since identifications of the
Desensitization involves injection of extracts
stinging insects are usually unreliable, these made from entire insects (Frazier, 1976) or injec-
figures should only be taken as rough estimates. tions prepared from pure venom (Lichtenstein,
These data showed 78 of the victims had 1 to 4
1975\ Sobotka et al., 1976\ Zeleznick et al, 1977).
stings, 16 had more than 4; 43 percent of the However, there was some controversy over which
stings were on the head and trunk, 27 percent on method was better (Reisman, 1975). Frazier
the extremities, and 9 percent on the trunk. (1976), who uses the whole-body extract, recom-
mended that a severely allergic patient be kept on
Emergency Treatment maintenance dosage indefinitely rather than for a
set period since there is no good knowledge on the
Frazier (1976) stressed that the most impor- permanence of this desensitization procedure.
tant step in treating systemic reactions is the Hunt et al. {1978) have shown that pure venom is
immediate subcutaneous injection of 0.2 to 0.5 superior to whole-body extracts for desensitiza-
ml of epinephrine (1:1000) for an adult and not tion.
more than 0.3 ml for a child. Feingold (1973)
advised epinephrine injections for all patients Emergency Medical Kits
with a history of previous stings since a pre- Most allergists recommend epinephrine
vious mild reaction is no index of severity of syringe kits for immediate treatment when pro-
subsequent reactions. He stated, 'Tor epineph- fessional medical help is not available. For
rine to be effective, it must be administered im- example, Frazier (1976) prescribes an insect
mediately without waiting for the development sting kit for his patients, who are instructed to
of symptoms. Once the reaction is established, keep it with them at all times. The kit contains a
reversal is very difficult. If signs and symptoms preloaded syringe of epinephrine, antihistamine
develop following the initial dose, repeat the epi- tablets, and several tablets of phénobarbital.
nephrine within fifteen to twenty minutes." Similar kits are now available commercially if
Barnard (1973) reported that 67 percent of the prescribed by a physician. In addition to a
patients whose sting was severe but nonfatal syringe of epinephrine, these kits contain anti-
had received early treatment with aqueous epi- histamine tablets and/or an inhaler that contains
nephrine. He also mentioned an appreciable an effective bronchodilator. Directions for use
number of fatal reactions where isoproterenol are present in each kit.
had been the sole treatment. This type of kit should be a standard item in
Cold compresses at the site of the sting pre- first aid supplies at public facilities such as
vent rapid dissemination of toxin and relieve, at campgrounds and parks. Persons entrusted with
least to some extent, the immediate burning the public safety, such as scout leaders, forest
sensation and pain. Other immediate treatments rangers, park service personnel, lifeguards, and
include applying a proteolytic enzyme (such as golf and tennis professionals, should be in-
household meat tenderizer) to the sting site. structed how to recognize severe allergic reac-
This offers some reUef as do commercial prepa- tions and how to administer the medications
rations. Antihistamines may be administered provided in the kit (Frazier, 1976). Persons with
orally or parenterally; however, they should not severe allergic reactions to stings should also
be used as a substitute for epinephrine (Fein- wear a medical warning bracelet and carry a
gold, 1973), similar card in their wallet.
Avoidance of Stings should try to remain calm in the presence of yel-
lowjackets. If wasps are gently brushed off the
Reisman (1975) suggested several simple meas- body when they alight instead of being swatted,
ures that help avoid stings. They include: (1) chances of being stung are reduced considerably.
Not wearing perfumes, hair sprays, suntan lo- When yellowjackets are present at picnic tables,
tion, and cosmetics, as yellowjackets are often the same principle holds: Move slowly and delib-
attracted to these compounds; (2) wearing light- erately instead of rapidly, and the wasps will
colored clothing such as white and tans; (3) not rarely sting. When a yellowjacket is discovered
walking barefoot outside; (4) exercising care in a moving automobile, the car should be driven
when gardening, mowing a lawn, and cutting off the main road quickly and the wasp removed
shrubbery; and (5) avoiding outdoor cooking and or killed. It usually poses no threat to the driver
eating during the yellowjacket and bee season. as long as all movements are careful and unhur-
Even more important than the above, people ried.
CONTROL
Even though most yellowjackets are benefi- such as propoxur (1.5 lb per gallon emulsifiable
cial, when aerial nesters build their nests in in- concentrate at the rate of 8 oz per gallon of wa-
convenient locations, or when populations of ter) or carbaryl (5 percent dust) into the entrance
scavenger species reach intolerable levels, con- hole, which is then plugged with cotton or simi-
trol measures must be employed. It is often dif- lar material. If the plug and immediate area are
ficult to determine when population levels are saturated with insecticide, foragers that have
high enough to warrant control since some peo- spent the night elsewhere will return in the
ple tolerate large numbers of yellowjackets morning and contact this material. Research re-
whereas others are alarmed at the sight of a sults also show that aerosols containing pyre-
single worker. thrins, rotenone, and a cooHng agent (to lower
MacDonald et al. (1976) noted that when body temperature) are also effective in control-
abatement becomes necessary, there are two ling subterranean colonies if the nest is fairly
general approaches of control—destruction of close to the entrance hole.
individual yellowjacket colonies and area-wide
abatement of worker populations. Akre and
Davis (1978) listed nest destruction, use of poi- Nests in Houses
son baits, proper management of garbage, and
trapping with synthetic lures as being the most Yellowjackets nesting in between walls of
commonly used methods of abatement. Other houses or other structures make control diffi-
useful methods employ the use of baits over cult. Morse et al. (1977) described the tendency
water containing a wetting agent and traps of V. germánica to nest within the walls of
using fish or other meat as bait. houses in New York and cautioned homeowners
not to block the entrance hole as the yellowjac-
kets may chew through the interior walls to es-
Nest Destruction cape. Vespa crabro, Vespula maculifrons, V. squa-
Destruction of colonies within the nest re- mosoy V. vidua (J. Nixon, Silver Spring, Md.,
mains one of the principal methods of control- personal commun.), V. pensylvanica, and V. vul-
ling yellowjackets, especially by homeowners. garis also frequently build nests in houses; other
The best time for control is after dark when for- species, to a much lesser extent. Because control
aging activity has ceased, and the maximum of these colonies is difficult, the homeowner may
number of workers are in the nest. wish to seek professional assistance.
One effective method of treatment, indicated
Subterranean Nests by research results, involves the use of a synthe-
Research data indicate that subterranean colo- tic pyrethroid and carbaryl dust (J. Nixon,
nies are easily killed by pouring insecticides Silver Spring, Md., personal commun.). The plas-
tic wand of the aerosol generator containing the or beef bait, was very effective against V. pen-
pyrethroid is quickly placed into the entrance sylvanica (C. A. Johansen, Wash. State Univ.,
hole, and 3.5 to 10.5 g of the material are re- personal commun.). The first program of yel-
leased during a 10- to 30-sec burst. The entrance lowjacket control, utilizing a nonrepellent, per-
hole is then plugged with steel wool, and the sistent insecticide in conjunction with meat
wool and surrounding area are dusted with baits, was proposed by Grant (1963). Later,
about one ounce (28 g) of 5 percent carbaryl. Grant et al. (1968) pubhshed the results of a 5-
This method has the advantage that treatment year testing program that utiHzed cooked horse
can take place during the day; returning for- meat and chlordane. This material effectively
agers chew at the steel wool coated with insecti- controlled populations of V. pensylvanica and V.
cide dust and succumb quickly. Protective cloth- vulgaris in parks and suburban areas in San
ing (bee suit, veil, gloves) should be worn during Mateo County, Cahf. Wagner and Reierson
the entire operation. Although Morse et al. (1969, 1971b) also reported excellent control of V.
(1977) recommended the entrance hole not be pensylvanica in parts of southern California
plugged, this pertained only to healthy colonies, using a mixture of fish-flavored cat food and
not to those being treated with insecticides. mirex, a slow-acting insecticide, used in con-
With the control method outlined above, there is junction with a chemical attractant. Similar pro-
little danger of workers chewing a new exit and grams have been tried by Keh et al. (1968) and
emerging inside the house. Workers from only 1 Rohe and Madon^^ in CaHfornia.
of 50 nests treated in this manner were able to To be acceptable, the insecticide must not be
find or create a new exit, and, even then, less repellent to yellowjackets, and it must be rela-
than 10 workers were found dead in a closet tively slow acting so that it can be distributed
below the nest. throughout the colony by trophallaxis.
The most promising new material, tested and
Aerial Nests evaluated by Ennik (1973), is an encapsulated
formulation of diazinon mixed with tunafish cat
Colonies of yellowjackets in aerial nests may food. Using this bait at eight test sites in north-
be killed by using one of several recently devel- ern California, foraging worker populations of V.
oped aerosol products that contain a quick pensylvanica and V. vulgaris were reduced an
knockdown insecticide (Davis, 1978). The toxi- estimated 75 to 90 percent within 2 days of ex-
cants are propelled distances of 3 m (10 ft) or posure to the bait. The effectiveness of this
more by the aerosol. Nests of this type may also material is due in part to the microencapsulation
be sprayed with propoxur (1.5 lb per gallon of the diazinon in a plastic material that masks
emulsifiable concentrate at the rate of 8 oz per its repellent odors. According to F. Ennik (Calif.
gallon of water). Heath Dept., personal commun.), the technique
of microencapsulation of insecticides will make
Use of Poison Baits available many compounds that are extremely
repellent to yellowjackets but are relatively safe
When a mixture of insecticide and bait is used to use and are biodegradable. He also noted that
for control, yellowjacket workers collect the bait preliminary tests with microencapsulated insect
at stations placed in the field, carry it back to the growth regulators appears promising as an al-
nest, and the material is then distributed to other ternate to the use of insecticides.
members of the colony by trophallaxis Encapsulated diazinon is registered for use in
(MacDonald et al., 1976). Death of the colony a bait formulation in California and Washing-
usually occurs within 2 weeks.
The use of a bait combined with an insecticide i2Rohe, D.L., and M.B. Madon. A field evaluation of
is an old technique, having been in use at least 40 mirex and chlordane poisoned baits for the control of
years ago in Washington when lead arsenate was the ground-nesting yellowjacket Vespula pensylvanica
used as the toxicant. About 1950, a wettable (Saussure), on Santa Catalina Island, Calif. Progress
powder formulation of chlordane became Report. 1968. Calif. Dept. Pub. Health, Bureau Vector
available and this material, together with a fish Control, lip. 1969. [Mimeographed]
ton. Only the insecticide is available commer- tages of using poison baits for yellowjacket
cially, bait is provided by the user. One teaspoon control, which included the following: (l)Speci-
of the insecticide is mixed with 6 oz of a suitable ficity for the target species (pestiferous scav-
meat bait (cooked meats, tunafish, and various engers) without disturbance to the beneficial,
cat foods). The poisoned bait is then placed in a strictly predaceous species; (2) minimal contami-
container constructed to permit foraging yellow- nation of the environment; (3) elimination of the
jackets access, but to exclude birds and small need to locate individual yellowjacket colonies
animals (fig. 71). The container(s) is then placed for destruction; (4) possible control of the yellow-
(perhaps suspended from a tree branch or placed jackets early in the season before populations
on a platform) in the area in which control is reach a nuisance level; and (5) destruction of col-
desired. Every 2 or 3 days the bait must be onies before the development of new reproduc-
replaced as yellowjackets will not scavenge tives late in the season.
spoiled meat. More explicit directions are in- Although several poison bait programs have
cluded with the insecticide. apparently been succesful, nearly all have been
MacDonald et al. (1976) listed several advan- restricted to the abatement of a single species,
V. pensylvanica, and only in California. The
pestiferous species in the Eastern States, such
as V. maculifrons and V. squamosa, are un-
affected by this technique as they are apparent-
ly not attracted to the baits currently in use.
Effectiveness of this method of control on V.
vulgaris outside California is unknown, but the
workers of this species are attracted to protein
baits. Similarly, little is known about abatement
possibilities for V. germánica using the poison
bait technique since this species has only re-
cently become a serious problem and no testing
has been done. Perrott (1976) found a mixture of
mirex and canned fish substantially reduced
worker populations of this species in certain
areas of New Zealand; however, other research-
ers report that the overall abatement program in
New Zealand is not as successful.
MacDonald et al. (1976) suggested the failure
of this technique can be attributed mainly to
pest species being differentially attracted to the
baits. This necessitates the development of
attractive, long-lasting baits specifically com-
pounded for each pest species. Even after attrac-
tive baits have been found and a control pro-
gram is initially successful, commercial proc-
essing may alter the original attractiveness
(Rogers, 1972). Another factor greatly influ-
encing the success of these programs is the dif-
ferent biotypes of species occurring in different
FIGURE 71 — Dispensing station for insecticide-treated
areas. For example, colonies of V. pensylvanica
meat bait (S. Peck). The 1- X 1-inch stake can be made any
height, but 36 to 40 inches places the station at a conven- in southern California appear heavily committed
ient height. The top is removable to replace the meat. Yel- to scavenging, whereas colonies of this species
lowjackets can pass through the hardware cloth with ease, in Washington and Idaho, while still pestiferous,
whereas children, pets, and wild animals are prevented are not so totally committed to this method of
from coming into contact with the bait.
obtaining protein. This greatly influences the
acceptability of baits and, therefore, abatement enough to alleviate problems, but may create an
by the poison bait method. added hazard for residents by exposing them to
additional yellowjackets attracted into the area.
If this technique is ever to become an important
Chemical Lures consideration in abatement programs, addition-
al, highly specific lures will have to be developed
Davis et al. {1967) discovered that the synthet-
or discovered for all major pest species. In
ic material 2,4-hexadienyl butyrate was a high-
addition, these attractants will have to be ef-
ly specific attractant for the western yellow- fective over a wider range of environmental con-
jacket, V. pensylvanica. Another material, hep- ditions so that they will be useful in most areas
tyl butyrate, was discovered to be even more of North America.
attractive and was used in small (87 oz) carton Additional uses of synthetic yellowjacket
traps positioned on the periphery of a 22-acre attractants include the use of this technique as a
peach orchard in Oregon to effectively depress monitor for colony growth, to estimate the
yellowjacket worker populations so that fruit- number of workers in an area, or to increase the
pickers could resume harvesting (Davis et al., number of workers visiting poison baits. Figure
1973). Abatement of the population to this level 72 shows a good correlation between numbers of
occurred within 4 days. In similar experiments workers caught in attractant traps and colony
at highway rest stops and overnight campsites growth in the field. Similar data, collected from
along the Columbia River in Oregon and Wash- specific localities, could be used in intelligent
ington, use of attractant traps baited with this control programs to create the least disturbance
synthetic lure depressed V. pensylvanica worker to beneficial species and to time selected control
populations below troublesome levels. procedures for maximum effectiveness against
Difficulties with the attractant trapping tech- pest species. Ennik (1973) used traps baited with
nique are similar to those experienced with poi- heptyl butyrate to monitor V. pensylvanica pop-
soned baits. Attractants presently available are ulations that had been subjected to poison bait
effective only for the western yellowjacket, V. formulations, and Wagner and Reierson (1969)
pensylvanica. Other pest species, especially found the addition of heptyl crotonate to poi-
those in the Eastern States, are only slightly soned bait tripled the amount removed by yel-
attracted (Grothaus et al, 1973; Howell et al, lowjacket workers.
1974). In addition, perimeter trapping with
attractant traps has been effective only in areas
limited in size (20 to 30 acres) and only in speci-
fic localities such as the orchards described by
Davis et al. (1973). Heptyl butyrate seems to be
most attractive to yellowjackets when used in
dry areas with low humidity. These synthetic
lures are highly attractive to workers and
queens of beneficial, strictly predacious species,
such as V. atropilosa, and trapping should be
timed to minimize the number of individuals of
these species that are caught (MacDonald et al.,
1973).
Although a commercial trap is available,

attractant trapping with synthetic lures is not
an effective control or abatement technique for FIGURE 72 —Weekly heptyl butyrate trap collection and
the homeowner. One or two traps placed in a weekly averages from excavated colonies. Each station on
yard will not lower the worker population the abscissa represents 1 week (MacDonald et al., 1974).
Management of Garbage colonies, this agent is of Hmited use. Therefore,
no biological control agent for yellowjackets
The proper management of garbage denies exists, and this method of control is considered
scavenging yellowjacket workers a readily avail- totally infeasible at the present time.
able source of protein that could have been used
to feed larvae during the period of exponential
growth of the colony. When workers are forced Control of Yellowjacket Queens
to forage mostly for live prey, the colony proba-
bly does not grow as rapidly since more time is Spradbery (1973a) cited two attempts to regu-
spent to feed each larva. late yellowjacket populations by the destruction
Morse et al. (1977) determined over 80 percent of hibernating queens. In New Zealand, a boun-
of the yellowjackets collected on food and gar- ty was paid for each V. germánica queen sent to
bage in and near restaurants and parks in Ith- the Department of Agriculture (Thomas, 1960),
aca, N.Y., were V. germánica, a species notorious A similar project was carried out in Cyprus.
for its scavenging habits. Similarly, most of the Results of both projects showed that mass des-
hordes of V. pensylvanica workers pestering pa- truction of queens had virtually no effect on yel-
trons of the Portland, Greg., zoo in 1973 were at- lowjacket populations the following summer.
tracted into the area by garbage in numerous Spradbery thus remarked, "It is evident from
cans with no, or ill-fitting, lids. A general policing the study of wasp populations that, with a mor-
of the area and a limited control program allevi- tality of 99.9 percent of potential queens and
ated much of the problem. In southern California, incipient colonies being necessary to maintain
Wagner (1961) found that treating trash contain- the same annual number of colonies, the destruc-
ers once a week with an aqueous spray of 0.75 per- tion of winter queens is likely to have little or no
cent dichlorvos at picnic sites reduced V. pensyl- effect on wasp populations. Indeed, a culling of
vanica worker populations to about 1 percent of winter queens may even cause an increase in the
those in untreated test areas. Akre and Retan number of wasp nests by reducing the competi-
(1973) also reported the effectiveness of dichlor- tion for suitable nest sites in the spring."
vos strips attached to the inside of garbage can
lids.
Fish or Protein Bait
(No Insecticide) Traps
Biological Control
Various traps using fish or other meat baits
Biological control of yellowjackets shows Httle have been constructed to collect workers of
promise as an abatement method (MacDonald et scavenging species of yellowjackets. While
al., 1976). Spradbery (1973a) described parasites, shape and size of these traps vary, nearly all use
predators, and pathogens as having Httle effect a reverse screen cone so workers can readily
on yellowjacket populations. Biotic agents only enter the trap but cannot find their way back out.
have an effect in weakened colonies, levels of Disadvantages of using the trap include (1)
parasitism are low, and only small numbers of keeping the bait fresh (yellowjackets will not
colonies are affected. He suggested that regula- scavenge spoiled flesh) by replacing it every other
tion of yellowjacket populations on an annual day, and (2) killing and removing all collected
basis would require that colonies be weakened or workers.
destroyed before queen production was initiated.
A neoaplectanid nematode was investigated Fish/Wetting Agent/Water Traps
by Poinar and Ennik (1972) as a possible bio-
logical control agent, but the propagation of this One of the oldest and still effective control
parasite requires 100 percent relative humidity. methods for yellowjacket workers involves sus-
Since this condition is not met in yellowjacket pending a raw fish several centimeters above a
pan filled with water to which a wetting agent Loon Lake, Wash., causing many guests to can-
has been added (fig. 73). The skin on the sides of cel reservations or curtail their activities. Nine
the fish should be broken or cut to give ready fish traps were placed at various locations
access to the flesh. around the resort, and within one week nearly
Typical worker behavior is to cut a piece of 1,000 yellowjacket workers were captured. Con-
flesh from the carcass, then to fly a short dis- tinued use of these traps in combination with
tance where it aUghts to trim the piece to a man- several other control procedures reduced, within
ageable size. The initial piece is frequently so 2 weeks, the number of workers flying in the
heavy the worker has difficulty flying and drops area to a level that guests would tolerate.
toward the substrate. Thus, when flying from Since these traps are easily constructed, use
the fish bait, many workers fall into the water, no toxic materials, and are specific for scavenger
and since the surface film has been reduced by species, this method of reducing yellowjacket
the wetting agent, they sink and drown. populations in specific areas remains one of the
During the outbreak of yellowjackets in the best. Two serious disadvantages of this method
Pacific Northwest in 1973, V. pensylvanica are the unavailability of rough fish to use as
workers were extremely abundant at a resort at bait, and removal of fish from traps by dogs or
cats; however, if the latter is a serious problem, a
chicken wire or hardware cloth cage can be placed
around the trap.
Repellent Jackets
Several commercial manufacturers produce
jackets impregnated with chemicals repellent to
biting fhes. These jackets are of no value in re-
pelling yellowjackets.
Control Summary
1. Yellowjackets, including scavenger species,
should not be controlled or eliminated unless
they are pestiferous because all yellowjackets
are beneficial.
2. All colonies should be exterminated at night
when the workers are least active and the max-
imum number are within the nest.
3. Subterranean colonies can be eliminated
with the use of insecticides such as carbaryl or
propoxur. These insecticides are poured into the
entrance tunnel which is then plugged with cot-
ton. The plug and surrounding soil should also
be treated to kill any foragers returning the next
day. Certain aerosols will also effectively control
colonies if the nest is near the entrance hole.
4. Colonies in wall voids are effectively con-
trolled by use of a synthetic pyrethroid gener-
ator and application of carbaryl dust to the
plugged entrance hole.
FIGURE 73 — Raw fish, with sides cut to expose flesh, sus-
pended above a pan of water to which a wetting agent has
5. Aerial nests can be easily treated with aero-
been added. Yellowjacket workers attempt to fly away with sol products containing a quick knockdown in-
a large piece of flesh, fall into the water, and drown. secticide. Several types of commercial products
will propel a thin stream of insecticide and sol- 8. Synthetic lure traps containing heptyl buty-
vent up to 3 m or more. Aerial nests may also be rate or other attractants have been used effec-
sprayed with conventional insecticides such as tively to abate populations of V. pensylvanica in
propoxur. some arid areas of Washington and Oregon. In
6. Simple traps using a fish as bait with a pan most areas of North America, these lures have
beneath containing water and a wetting agent no value as most species are not attracted.
can be quite effective in lowering populations of Hopefully, future research will uncover addition-
yellowjackets in small areas. These traps have al compounds that are differentially attractive
the advantage of being specific for scavenging to some of the other species. This type of trap-
species. Other traps using fish as bait and a re- ping is not recommended for homeowners.
verse cone to trap entering workers are effective 9. One of the best methods of reducing contact
but not as convenient as other methods. with yellowjackets (and perhaps even reducing
7. At the present time, poison bait formula- populations) in urban areas is to control garbage
tions are effective only against V. pensylvanica by making it less available to scavenging work-
and, to a lesser extent, against V. vulgaris. The ers. Treatment of trash containers routinely
only effective programs utilizing this technique with dichlorvos sprays or placing an insecticide
were in California. More research is needed to impregnated strip in the garbage can lid also
discover acceptable baits for pestiferous species helps.
in other areas.
ACKNOWLEDGMENTS
It is impossible to write a handbook of this necticut, Storrs; CF. Roush, Department of En-
scope without the help and cooperation of a tomology, Washington State University, Pull-
large number of scientists who gave freely of man; R. Key el. Department of Entomology, Cor-
their data, their photographs, and their time. nell University, Ithaca, N.Y.; J. Nixon, Allied
Without their help, many sections of this publi- Exterminating Services, Silver Spring, Md.; R.E.
cation would be much less informative. While Wagner, Department of Entomology, University
this small note cannot begin to express our real of California, Riverside; R.W. Matthews, De-
appreciation, we extend it as a grateful thank partment of Entomology, University of Georgia,
you. Athens; J. Sharp, Insect Attractants and Basic
We thank the following for the loan of yellow- Biology Laboratory, USDA, Gainesville, Fla.;
jacket specimens: J.D. Lattin and G. Ferguson, CA. Johansen, Department of Entomology,
Department of Entomology, Oregon State Uni- Washington State University, Pullman; F.
versity, Corvallis; R.H. Washburn, USDA, Ennik, California Department of Health, Berk-
Palmer, Alaska; T.J. Zavortink, CaHfornia Acad- eley; R.W. Zwick, Oregon State University Ex-
emy of Science, San Francisco; P.J. Hurd, periment Station, Hood River; and W.B. Robin-
Smithsonian Institution, Washington, D.C.; son, Department of Food Science and Tech-
A.S. Menke, Systematic Entomology Labora- nology, Cornell University, Ithaca, N.Y.
tory, U.S. National Museum, Washington, D.C.; J. Schuh, Klamath Falls, Oreg., A.A. Anto-
M. Favreau, American Museum of Natural His- nelli, Western Washington Experiment and Ex-
tory, New York City; H.V. Daly, Division of En- tension Center, Puyallup, and W. Baker, Friday
tomology and Parasitology, University of Cali- Harbor, Wash., are acknowledged for the many
fornia, Berkeley; and W.J. Turner, M.T. James yellowjacket specimens and nests they pre-
Entomological Collection, Washington State Uni- sented to us during the course of conducting the
versity, Pullman. research that led to this handbook.
Appreciation is extended to the following re- Photographs used in illustrating some of the
searchers who permitted us to cite their unpub- text were obtained from: C.W. Rettenmeyer; R.
lished data or observations: C.W. Rettenmeyer, Keyel; J. Nixon; CF. Roush; R. Getting, De-
Biological Sciences Group, University of Con- partment of Entomology, Georgia Agricultural
Experiment Station, Experiment, Ga.; and W.D. mology, University of Georgia, Athens, drew che
Lord, Department of Entomology, University of illustrations of the Polistes nest and the yellow-
Delaware, Newark. jacket Hfe cycle. All other illustrations are by Al
Special thanks are due to J. Nixon for his crit- Greene. Photographs not credited to other per-
ical reading of sections of the manuscript and in sonnel were taken by the authors.
advice with chemical control, to C.F. Roush for Much of the information contained in the
his criticism of parts of the manuscript and for handbook was obtained from research projects
his many helpful assistances, and to D.M. Jack- at Washington State University, funded by
son, Department of Entomology, Washington National Science Foundation (grants GB-30922
State University, Pullman, for his untiring help and BNS 76-81400), the USDA, Science and
with data, illustrations, collating materials, and Education Administration (cooperative agree-
checking literature. ment), and the USDA, Douglas-fir Tussock
Moth Expanded Research and Development
Several illustrations used in this publication Program (GSRS grant 704-15-16); from projects
were prepared by G. Shinn, University of Wash- at the University of Georgia, Department of En-
ington, Seattle, and S. Peck, Washington State tomology; and from Purdue University Exper-
University. Joan Krispyn, Department of Ento- iment Station project 58045.
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APPENDIX
Glossary of Selected Terms Cyanosis. — A bluish discoloration of the skin

Used in the Text due to inadequate oxygénation of the blood.
Discoidal cell. — A closed (surrounded by veins)
Aculeate. — Pertaining to the stinging members cell or cells in the middle of the wing.
of the order Hymenoptera; that is, the bees, Dominance hierarchy. — A behavioral domina-
ants, and wasps. tion of certain members of a colony over cer-
Alimentary liquid. — Any material in the food tain other members; a "pecking order.''
canal of adults and larvae. Duff.— Decayed leaves and branches of the sur-
Anaphylactic shock. — A sometimes violent face of the forest floor.
physiological response to the introduction of
Eclosión. — Emergence of an adult from a pupal
a foreign substance such as venom into the
case or cocoon.
body of a person hypersensitive to that sub-
stance. Electrophoresis. — Migration of different sub-
Apiary. — Any place where honey bees ( Hymen- stances at different rates within an electrical
optera: Apidae) are kept. field, used to separate small amounts of chem-
Associate. — Arthropods (spiders, mites, sow- ical components of a mixture.
bugs, and other insects) found in or around Encapsulated formulation. — Insecticide coated
yellowjacket colonies and nests. Many are with a plastic material (microcapsules) causing
scavengers, a few are parasites. the toxic component to release slowly.
Biotype. — A group of organisms having the Envelope. — Sheaths of paper or masticated
same genetic makeup but possibly different plant fiber surrounding the combs of a nest.
physical characteristics.
Epinephrine. — An adrenal hormone that stimu-
Brood. — All the immature members of a colony lates organs or tissues (heart rate, smooth
including eggs, larvae, and pupae.
muscle of lung, and blood vessels) controlled
Cap. — A covering of silk secreted by a larva over by the autonomie nervous system.
the opening of a cell shortly before it forms a
Eusocial. — A condition in which a group of in-
pupa.
dividuals exhibits division of labor, with sterile
Carton. — Material comprising the combs and members of the group working on behalf of ac-
suspensoria of yellowjacket nests. tively reproducing members and with two or
Caste. — A group of morphologically distinct in- more generations (that is, foundress and her
dividuals within a colony often having distinc- daughter workers) contributing to the group
tive behavior (that is, workers, queens, and welfare.
males). Facultative parasite. — An organism capable of
Chronologued colony. — A colony the activities supporting itself (free living) but which may oc-
of which have been recorded on a daily basis casionally obtain its livelihood at the expense
from beginning through decline to death. of another organism called the host.
Cocoon. — A silken enclosure secreted by a larva Female. — In reference to poHstine wasps, an
just before pupation; exposed end called a overwintering, fertilized individual that coop-
**cap." erates with other such individuals to construct
Colony. — Individuals, other than a single mated a nest; one usually becomes dominant.
pair, which cooperate to construct a nest or Forager. — An individual colony member, usual-
rear offspring. ly a member of the worker caste, that searches
Comb. — A single layer of regularly arranged for and obtains food, water, or building mater-
cells or cocoons. ials.
Cryptic. — ConceaUng or camouflaging, as in Foundress. — An individual, usually a fertilized
cryptically colored moths often difficult to see female, that founds a colony, all the members
against their substrates. of which would be her daughters or sons.
Gaster. — The prominent part of the abdomen, Opportunistic. — Ability to accept whatever re-
separated from the other parts of the body by sources (food, nest, or sites) are available at
a thin connecting segment called the petiole or the time.
pedicel. Paper wasp. — Members of the genus Polis tes.
Girdle. — A band made around a tree by removal Parasitism. — An association in which one or-
of a ring of bark, usually kiUing the tree. ganism, called the parasite, lives at the ex-
Gravid. — Full of ripe eggs. pense of another organism, called the host,
Haplometrosis. — The founding of a colony by a without killing it.
single fertilized female or foundress. Parasitoid. — A type of parasitic association in
Hornet. — Large wasps belonging to the genus which the host is eventually killed by the para-
Vespa, although erroneously applied to the site.
bald-faced "hornet," D. maculata. Pleometrosis. — The founding of a colony by
Incipient. — An initial or early stage of a colony more than one fertilized female or foundress.
or its nest. Polyarthritis. — Simultaneous inflamation with
Incised. — Cut into or notched. associated pain of several joints.
Presocial. — A condition in which a group of in-
Inquiline. — In the broadest terms, any individ-
dividuals exhibits only one or two of the follow-
ual occurring in the nest of another individual.
ing traits: Cooperation in care of immatures,
Interspecific. — Involving individuals of two or sterile members working in behalf of actively
more species. reproducing members, or with two or more
Intraspecific. — Involving individuals of the generations contributing to group welfare.
same species. Queen. — A member or an egg-producing caste
Lymphadenopathy. — An abnormal state of the as opposed to a worker caste or a male caste.
lymph glands often associated with swelling of Reproductive diapause. — In the adult, a temp-
the glands. orary cessation in ability to develop mature
Malaise. — A feehng of being ill or depressed. oocytes (eggs).
Malaxating. — Chewing and squeezing of prey Reproductives. — Any individuals of a colony
by an adult yellowjacket, which softens prey (that is, males and females capable of being
and reduces it in size before it is given to lar- fertilized) able to reproduce.
vae. Royal court. — Vespa workers surrounding and
Mastication. — Chewing and mechanically break- licking the queen, which is highly attractive.
ing up food or building material. Scutum.— The dorsocentral part of an insect
Mauling. — Stereotyped interaction in which one thoracic segment.
worker vigorously bites another but apparent- Septicemia. — A disease state caused by patho-
ly produces no physical damage to the usually genic organisms in the blood; often called
motionless recipient. "blood poisoning."
Medial. — Toward the midline of the body. Social parasite. — A type of parasitism involving
Melanie. — Exhibiting a predominance of dark a social insect as host.
brown pigmentation. Species group. — A group composed of several
Mesonotum. — The upper surface of the meso- to many closely related species.
thorax. Subcutaneous. — Beneath the skin.
Nest sanitation. — A behavior pattern resulting Submarginal cell. — Closed cell in the insect
in rapid removal from the nest of dead or dy- wing, occurring just posterior to the leading
ing brood, unused prey, and other debris. edge of the wing.
Nest usurpation. — The adoption of a colony by Sympatric. — A condition in which several non-
an unrelated individual, usually by forcible interbreeding populations occupy the same
eviction or death of the foundress. geographic area.
Obligate parasite. — A parasitic association in Urticaria. — An allergic reaction of the skin to
which the parasite cannot complete its life cy- foreign substances such as venom, indicated
cle without the host. by itchiness and welts.
Ventral. — Closest to the lower surface or "bel- Worker. — A member of a nonreproductive or

ly'' area of the body. sterile caste, which contributes to a colony wel-
Vespine wasp. — Members of the Vespinae, in- fare by rearing offspring of reproductives.
cluding Provespa, Vespa, Vespula, and Doli-
chovespula. Xanthic. — Exhibiting a predominance of yellow
Wall void. — space between the outer and inner pigmentation.
walls of a building.
Wasp (that is, true Wasp). — Hymenoptera that Yellowjacket. — Any member of the ground or
have the ovipositor modified into a sting for aerial nesting wasps belonging to the genera
injecting venom into the host. Egg no longer Vespula and Dolichovespula.
passes down the modified ovipositor.
List of Chemicals Mentioned in This Publication

Pesticide
designation Chemical name
Carbaryl 1-naphthyl methylcarbamate
Chlordane 1,2,4,5,6,7,8,8-octachloro-3a,4,7,7,a-tetrahydro-4,
7-methanoindan (60% minimum and not over
40% of related compounds)
Diazinon 0,0-diethylO-(2-isopropyl-6-methyl-4-pyrimi-
dinyl) phosphorothioate
Dichlorvos 2,2- dichloro vinyl dime thy Iphospha te
Heptyl butyrate heptyl butyrate
Heptyl crotonate heptyl crotonate
Hexadienyl butyrate 2,4-hexadienyl butyrate
Mirex dodecachlorooctahy dro-1,3,4- metheno-1H- cy clo-

buta[c(i]pentalene
Propoxur o-isopropoxyphenyl methylcarbamate
Pyrethrins (An insecticidal constitutent of pyrethrum)
Resmethrin (5-benzyl-3-furyl) methyl trans-(-\-)-2,2-

dimethyl-3-(2-methylpropenyl) cyclopro-
panecar boxylate
Rotenone 1,2,12,12a-tetrahydro-2-isopropenyl-8,9-dimeth-
oxy[l]benzopyran-6(6a//)-one

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1573^9^

For sale by the Superintendent of Documents, U.S. Government Printing Office

This pubhcation reports research involving pesticides. It does

CAUTION: Pesticides can be injurious to humans, domestic

Trade names are used in this publication solely to provide spe-

Issued June 1981

Introduction human contact is with the social Vespidae. Ves-

FIGURE 2 — Distribution of Stenogastrinae (drawn by G. Shinn).

FIGURE 3 - Distribution of Ropalidüni (G. Shinn).

FIGURE 4 - Distribution of Polybüni (G. Shinn).

FIGURE 5 - Distribution of Polistini (G. Shinn).

FIGURE 6 — Distribution of Vespinae (G. Shinn).

identification in eastern North America is compli-

FIGURE 8 — a, Polistes queen on her nest; b, Polistes nest

separate genera, the taxonomic treatment of

Key to Vespinae in North America

Gaster resembling figure 14 y. crabroL. (European hornet)

First 3 terga with pale markings; smaller species «15mm)

Pale marking white 2

3. Two yellow stripes on scutum (fig. 18a) 4

6. First antennal segment yellow ventrally (fig. 21a) 7

Vespula V. suiphurea V. intermedia V. vidua

V. acadica- V. acadica- V. atropiiosa- V. atropitosa- V. pensylvanica

V vulgaris V. maculifrons V. flavopilosa V. germanicax- V. germánica-

FIGURE 25 — Yellowjacket and hornet worker gaster patterns drawn to scale.

General sis). Prey usually consists of live arthropods, al-

usually covered with a many-layered envelope.

FIGURE 27 — a, Queen nest; Vespula pensylvanica,- b, queen

20 to 45 cells covered by a paper envelope. The

FIGURE 28 — Mating of Vespula pensylvanica; queen on left.

DoUchovespula arenaria Vespula atropilosa Vespula pensylvanica

On the basis of isolated reports, V. crabro now

FIGURE 32 — Distribution of Vespa crabro. Lighter pattern indicates scattered occurrence.

FIGURE 34 — Distribution of Dolichovespula albida.

Dolichovespula árctica D: árctica is an obligatory social parasite in

FIGURE 35 — Distribution oí Dolichovespula árctica.

árctica apparently cannot prevent the host DoUchovespula arenaria

FIGURE 39 — Distribution of Dolichovespula maculata.

FIGURE 41 — Distribution oi Dolichovespula norvegicoides.

FIGURE 42 — Distribution of Vespula acadica.

FIGURE 44 — Distribution of Vespula atropilosa.

FIGURE 46 — Distribution of Vespula austríaca.

FIGURE 47 - Distribution of Vespula consobrina.

FIGURE 49 — Distribution of Vespula intermedia.

FIGURE 52 — Distribution of Vespula squamosa.

TABLE 6.—Summary of mature Vespula squamosa colonies collected after

Parameter 1974 1975

FIGURE 54 — Distribution of Vespula sulphurea.

TABLE 7.—Summary o/Vespula sulphurea colonies collected in California

1 Clear Lake 7-27-72 580 1 0 0 (•) (')

Vespula flavopilosa.—Vespula flavopilosa (hy- as a species, no studies of behavior or nest associ-

FIGURE 55 — Distribution of Vespula flavopilosa.

TABLE 8.—Vespula flavopilosa and V. maculifrons colo-

FIGURE 57 — Distribution of Vespula germánica.

Vespula maculifrons.— Vespula maculifrons

^Preiss, F.J. Nest site selection, microenvironment and

protein and are attracted in great numbers to

FIGURE 60 — Distribution of Vespula maculifrons. Lighter pattern indicates scattered occurrence.

TABLE 9.—Analysis of 10 nests of Vespula maculifrons from