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The German Yellowjacket (Vespula germanica) Problem in the United States
(Hymenoptera: Vespidae)t
J. F. MACDONALD!,
R. D. AKRP,
AND
R. E. KEYEU.
Associate Professor, Department of Entomology, Purdue University, W. Lafayette, IN 47907; I
Entomologist, Department of Entomology, Washington State University, Pullman, WA 99164;2
Research Assistant, Department of Entomology, Cornell University, Ithaca, NY 148533
Yellowjackets are the familiar social wasps which
typically build large paper nests consisting of multiple
combs enclosed in an envelope (see Greene and Caron
1980). Workers of all yellowjacket species aggressively
defend their colony if it is disturbed and such workers
are probably responsible for many of the stinging episodes attributed to "wasps and bees" (Fluno 1961, Barr
1974). Although difficult to accurately document, the
number of serious reactions and deaths resulting from
yellowjacket stings is high enough to have stimulated
an increasingly greater interest of medical researchers
apiaries in New Zealand, and is a potential threat to
apiculture in this country, although significant hive predation is still undocumented (Dejong 1979).
Because of the rate and extent of German yellowjacket dispersal, and its successful establishment in
many urban centers in this country, we believe an informative paper will be valuable to entomologists, public health officials and control specialists who may have
to deal with this spreading problem. We have experienced a marked increase in inquiries from professionals
and the public regarding yellowjackets the past few
into
years
the
problems
of human
hypersensitivity
to yel-
lowjacket venom and venom desensitization methodology. Akre et al. (1980) reviewed the medical importance
of yellowjackets, as well as the economic impact of
yellowjackets on resorts, campgrounds, lumber operations, fruit harvests, etc., in those areas of the USA
periodically experiencing years of yellowjacket abundance.
In addition to periodic problems with native yellowjacket species, we are now faced with a new yellowjacket problem. Since the late 1960's, the German
yellowjacket, Vespula germanica (Fab.), has become
firmly established over much of northeastern North
America from Montreal and Ottawa, Canada in the north
(Menke and Snelling 1975; Morse et al. 1977) and along
the eastern seaboard south into New Jersey, Delaware
and Washington D.C. (Lord 1978). Based on recent
reports and personal observations, during the past few
years the German yellowjacket has spread westward
across the northern midwest into Indiana, Illinois and
Michigan, and was reported from Winnipeg, Manitoba
in 1979 and Minneapolis, Minnesota in 1980.
The establishment of V. germanica in this country is
a cause for concern since the German yellowjacket,
more so than any native yellowjacket, appears to be
closely associated with human foods and food processing industries (Spradbery 1973a; Keyel 1980). Moreover, in this country the preferred nesting site of the
German yellowjacket is inside structures and not underground (Menke and Snelling 1975; Morse et al. 1977).
Both worker foraging behavior and structural nesting
bring the German yellowjacket into conflict with humans, and this species is emerging as the most important
urban/industrial pest yellowjacket (Akre et al. 1980). In
addition, the German yellowjacket is a serious pest of
t Journal paper number 768v, Purdue Agricultural Experimem
Station, and Scientific paper number 5381, Washington State University, College of Agriculture Research Center, Pullman. Work supported
by Purdue Agricuhural Experiment Station project number 58045 and
project number 4037 at Washington State University. Financed in part
by NSF Grant BNS-76-81400 to Washington State University. Received
for publication April 14, 1980.
"'19XO Entomological
Society of America
and have become
aware
that many entomologists,
and most public officials and the public, possess little
understanding of social wasp biology and, accordingly,
will likely not be able to deal effectively with this new
problem. Here, we summarize the natural history of the
German yellowjacket and provide practical information
on safe and effective control measures.
History of the German Yellowjacket in the
United States
Vespula germanica is native to Europe, northern Africa, and temperate Asia (Spradbery 1973a) but has been
introduced into numerous countries, including several
in the Southern Hemisphere: New Zealand (Thomas
1960); Tasmania (Spradbery 1973b); South Africa and
Chile (Edwards 1976); and Australia (Smithers and Holloway 1977). Once introduced and established, the German yellowjacket became a very serious pest in New
Zealand (Thomas 1960; Perrott 1975), and has always
been a serious pest over much of Europe, particularly
in urban, recreational,
and even commercial areas
(Spradbury 1973a).
Based upon a few collection records, V. germanica
was periodically introduced into northeastern North
America in the past century, but did not become established until about 1968, apparently in Maryland (Morse
et al. 1977). Records of the early establishment and
dispersal of V. germanica unfortunately are limited to
Menke and Snelling (1975) and Morse and co-workers
at Cornell University.
In 1972, V. germanica workers already made up
about half the yellowjackets in student collections at
Cornell, and by 1974 the German yellowjackel had become the major pest yellowjacket foraging at human
food sources and nesting in human structures in the Ithaca, New York vicinity (Morse et al. 1977). Although
the proportion of structural yellowjacket nests due to V.
germanica in Ithaca, NY has declined from 75% in 1972
(Morse et al. 1977) to 35% and 39% of such nests in
1977 and 1978, respectively, V. germanica has remained the most abundant yellowjacket at human food
0013-X754 80 0404-3607$00.75
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Vol. 26, no. 4, 1980
ESA BULLETIN
sources even when scantily represented on nearby natural carbohydrate sources such as honeydew (Keyel
1980).
Morse et aI. (1977) attempted to trace the spread of
V. germanica westward during the summer of 1976.
They collected it throughout northern Ohio, and in the
same year a few specimens appeared in student collections at Purdue University. However, V. germanica was
present in northeastern Ohio earlier since we have workers collected Aug 5, 1971 from Rock Creek, Ohio. Interestingly, the German yellowjacket has been perceived
as being different from native species. For example, by
the mid-1970 people in northern Ohio and around Temperance, Michigan noted yellowjackets they called
"newcomers."
Few yellowjacket problems occurred in
these areas over the previous 10 years or so, but now
these "newcomers"
were abundant enough to cause
concern as they scavenged on fallen fruit, other sweets,
in animal carcasses, and on meats eaten by humans. By
1978, V. germanica was a cause of problems in southern
Michigan, and 2 colonies were discovered on the Michigan State University campus that summer (P. Landolt,
pers. comm.).
Initially reaching west central Indiana in 1976, V.
germanica has become the most reported yellowjacket
in major urban areas throughout the state, and is the
most common yellowjacket species on the Purdue University campus. Indeed, in 1978, 11 colonies were found
in buildings within a 1/2 kn) radius of Entomology Hall.
During late summer and autumn 1978 and 1979, the
Department of Entomology, Purdue University, the Indiana State Board of Health, Indianapolis, and the State
Entomologist's Office, Indianapolis, each were receiving several calls weekly regarding yellowjacket problems. Most reports were from homeowners discovering
yellowjacket colonies inside their homes, but several
food processing industries called seeking a remedy for
yellowjackets disrupting plant operations. Such problems had not been reported in Indiana prior to 1978. In
addition, pest control operators reported a significant
increase over the past 2 to 3 seasons in calls to treat
structural colonies.
Based on the above accounts and numerous requests
for information and control recommendations from urban areas in Indiana, Illinois, and Ohio, the major dispersal route of V. germanica during the mid-1970s was
westward across the upper Midwest, apparently from
one urban area to another. The saltatorial spread of V.
germanica is probably related to the synanthropy exhibited by this species making dispersal of inseminated
queens via commerce very likely. In fact, entry into
New Zealand in 1944 was traced to overwintering
queens in airplane engine crates shipped into the country
from England (Thomas 1960). Once introduced into a
new area, dispersal is again probably facilitated by lines
of commerce. Study of the initial invasion in New Zealand showed founding queens ventured less than I000M
from a parent colony before establishing a new colony,
and the rate and pattern of dispersal was best explained
by the close relationship with rail and highway routes
(Thomas 1960).
437
Natural History of the German Yellowjacket
Vespula is divided into the V. rufa (L.) and V. vulgaris (L.) species groups both on the basis of morphology and several aspects of biology. Most significantly,
workers of the V. vulgaris group, in addition to capturing live insect prey, turn increasingly to scavenging in
late summer and autumn. Particularly attractive meat
sources include animal carcasses, dead insects, meat in
garbage, and numerous meats eaten by humans. Foraging for the latter 2 sources plus for sweets found in
garbage, at picnics, and in association with some food
processing industries frequently brings workers into
conflict with humans. Furthermore, V. vulgaris group
species often attain very high population densities in
recreational and residential areas where their large colonies may remain active into early winter. Thus, colony
duration may be I to 3 months longer and colony size
much larger than that of other temperate social wasps.
Because of worker scavenging behavior, large colony
size, long seasonal cycle, and frequent high population
density, species of the V. vulgaris group constitute our
most serious social wasp problem (MacDonald et a!.
1976).
The V. vulgaris group in North America consists of
4 native species: the eastern yellowjacket, V. maculifrons (Buysson); the western yellowjacket, V. pensylvanica (Saussure); a Holarctic species, V. vulgaris; and
a recently described eastern yellowjacket species, V.flavopilosa Jacobson (Jacobson et a!. 1978). Vespula germanica is also in the V. vulgaris group, adding a fifth
member to the group in North America.
Comprehensive
treatment of yellowjacket biology
may be found in Spradbery (l973a) and Akre et a!.
(1980) which provide the basis for the general biology
presented below. Later, we concentrate on the biology
of V. germanica in North America, pointing out interesting deviations in biology compared to European populations.
All northern temperate yellowjacket species possess
an annual seasonal cycle. Each species is represented
over the winter solely by inseminated queens which independently establish a colony in early May. Colony
growth is very slow at first, but the worker population
gradually increases and colonies reach their developmental peak by mid to late summer. Associated with
this developmental peak of the annual cycle, workers
shift from building small cells to constructing special
large cells (queen cells) in which reproductives (queens
and some males) are reared.
Queens are produced until the end of the seasonal
cycle, typically late autumn to early winter, and large
colonies may produce several hundred to a few thousand
new queens. Mating typically occurs away from the parent colonies. Males die soon thereafter while inseminated queens quickly enter protected sites to overwinter.
In the parent colonies, the worker force dwindles rapidly
and the foundress dies, marking the termination of the
seasonal cycle since temperate yellowjacket colonies do
not possess secondary reproductives.
Throughout much of the summer, yellowjacket work-
438
ESA BULLETIN
ers are beneficial predators on a variety of insects including adult flies, moths, caterpillars, beetle larvae,
true bugs and leafhoppers, and Orthoptera. In Europe
and in the USA (Schmidtrnann 1977), German yellowjacket workers display a propensity for preying on
adult muscoid flies, including the stable fly, Stomoxys
calcitrans (L.), and the face fly,. Musca autumnalis
DeGeer. Predation on adult flies is probably associated
with fly abundance in habitats supporting the German
yellowjacket. By late summer, however, foraging for
sweets increases as does scavenging for meat. This foraging behavior is related to the increasing nutritional
demands of the colonies associated with queen rearing,
and compounded by a decline in natural prey as the
season progresses.
Nearctic V. germanica colonies appear to be behaviorally similar to their European counterparts with the
notable exception of the nesting site. According to
Vol. 26, no. 4,1980
onies may reach larger size more often in the USA than
in their native land. Based on estimates of cell utilization
established by Spradbery (1973a), the largest V. germanica colony (Frankfort, IN) produced an estimated
23,000 workers prior to switching to queen production
in late summer compared to an estimated 17,000 workers in the largest colony reported by Spradbery.
As alluded by Morse et aI. (1977), success of V.
germanica in North America in part must be related to
the structural nest site. Although rigorous study is lacking, the structural nest site appears to offer a number of
advantages in comparison with the subterranean site.
Incipient yellowjacket colonies are quite vulnerable and
the protected structural nest site probably affords a more
favorable environment during this critical period. For
example, numerous incipient colonies are destroyed by
inundation during spring rains, while mature terrestrial
colonies are subject to heavy vertebrate predation. The
$pradbery (1971), 27 of 30 V. germanica nests in Eng-
structural site provides protection against such adversi-
land were subterranean, and only 1 was in an outbuilding; no colony was found in an occupied human structure. To date, we know of very few subterranean V.
germanica colonies in North America (Menke and
Snelling 1975; Keyel 1980). Nearly all other reported
German yellowjacket colonies have been from occupied
structures or from outbuildings such as barns and sheds.
For example, in the Lafayette, IN, area, 42/43 V. germanica colonies found in 1977-80 were inside structures; 1 colony was subterranean. Many more structural
yellowjacket colonies were reported but not confirmed
as to species (e.g. a pest control operator with a background in social insect biology destroyed nearly 50
structural yellowjacket colonies in Lafayette homes in
1977-78). In comparison, during 1977-78,47
subterranean but only 7 structural eastern yellowjacket (V.
maculifrons) colonies were located around Lafayette,
IN. The same pattern existed in Ithaca, NY, where during 1977-78, 24 of 26 V. germanica colonies were
structural; the other 2 colonies were initiated in railroad
ties and later expanded into the soil below. In comparison, of 53 eastern yellowjacket colonies in the Ithaca,
NY area, 46 were subterranean, 5 structural and 2 in
railroad ties.
Propensity for structural nesting by Nearctic V. germanica is probably a result of a behavioral difference
among founding queens. Instead of searching for a nest
site in soil cavities, Nearctic V. germanica queens apparently search vertical surfaces of structures for potential nest sites. Commonly employed access holes into
structures include chinks in mortar, gaps where walls
and eaves or where window frames and walls meet, and
holes left from electrical conduits. Such access holes
lead into common nest sites including wall voids, attics
(often in insulation materials), and basements (space
between floor and basement drop ceiling).
Due to the difficulty of sampling nests from structural
sites, few Nearctic V. germanica colonies have been
excavated and analyzed. However, of the few colonies
sampled in the USA, half of them are larger than the
largest colony found by Spradbery (1971) in England
(Table I). This suggests that German yellowjacket col-
ties. Extended colony duration also is facilitated by the
protected structural site. Lastly, the propensity of V.
germanica workers to exploit garbage late in the seasonal cycle may facilitate a longer seasonal cycle.
Possibility of Perennial Colonies
One of the most frequent questions asked about a
yellowjacket colony is if it will persist in the same site
the following season. Perennial V. germanica colonies
are not without precedent as they have occurred in Tasmania (Spradbery 1973b), New Zealand (Perrott 1975),
Algeria and Morocco (Vuillaume et al. 1969), and Chile
(Jeanne 1980). Authors report second-season colonies
of enormous size with a thousand or more functional
queens, several hundred thousand workers and nests of
nearly a million cells. Very few second-season colonies
of other species have been reported in the USA, however, and they are the exception and are largely restricted to areas with mild winters such as California
(Duncan 1939) and Florida (Tissot and Robinson 1954;
Akre et aI. 1980). However, Spencer (1960) reported
a "perennial" V. pensylvanica colony in the attic of a
Vancouver, British Columbia house. Otherwise, temperate yellowjacket colonies are inherently limited to a
single season.
Among temperate yellowjackets, a foundress queen
typically dies prior to cessation of colony activity. Since
colonies do not possess secondary queens, continuation
of a colony would require re-queening. While re-queening of V. germanica colonies in the USA is possible, a
major obstacle would have to be overcome; namely, the
circumvention of ovarian diapause that is characteristic
of newly emerged northern temperate social wasp
queens in autumn, and which persists throughout the
overwintering period. Spradbery (1973b) postulates that
the ability of V. germanica queens produced near the
end of the seasonal cycle in Tasmanian colonies to circumvent ovarian diapause (and thus join the parent colony and assume egg-laying) is related to photoperiod.
While day length is decreasing during the period of new
queen production in both the southern and northern hem-
Vol. 26, no. 4, 1980
ESA BULLETIN
ispheres, the seasonal cycle is shifted in the southern
hemisphere so that in Tasmania duration of the dark
period is a few hours shorter. Thus, Spradbery speculates, newly inseminated queens are somehow able to
"join" the present colony (or possibly another nearby
colony) and then initiate ovarian development. If, indeed, photoperiod differences are as significant to requeening as Spradbery postulates, then re-queening of
V. germanica colonies in the USA may be precluded.
Unfortunately, we have no explanation for the few reported perennial Vespula colonies in the USA.
A further obstacle to re-queening may be associated
with mating behavior. Although little is known about
Vespine mating behavior, mating probably does not occur or is very rare inside the parent colony between
siblings. If new queens leave the parent colony to mate,
an orientation flight would be necessary to enable mated
queens to return to the parent colony. Alternatively,
mated queens would have to discover and somehow join
an alien colony. Two points of evidence appear to preclude re-queening in the parent colony. First, dissections
of queens collected inside nests excavated in autumn
have shown all such queens uninseminated. For example, all 25 queens sampled from a Frankfort, IN, V.
germanica colony collected Nov. 2, 1978 had empty
spermathecae, including several that appeared to be in
the typical overwintering posture inside the nest envelope. Second, Vespula queens leaving parent colonies
in autumn typically do not make an orientation flight
about the nest entrance, although such behavior has been
reported (Spradbery 1973a).
To summarize, the possibility of re-queening leading
to second season colonies inside structures seems unlikely. More likely is the frequent occurrence of annual
colonies remaining active well into winter with persistence being facilitated by favorable environmental conditions inside certain structures. For example, Morse et
al. (1977) mentioned late-season active colonies, and
we have reports of active structural colonies in midDecember in Washington D.C. (1. Nixon pers. comm.)
and in late November-early December in northern Ohio
(P. Landolt pees. comm.). Workers of a structural Lafayette, IN, colony were still foraging when the colony
was destroyed the day before Thanksgiving. Such colonies, however, will not persist over winter, and (unless
re-queened) will not become active the following spring.
Abatement of the German YeUowjacket
MacDonald et al. (1976) evaluated the status of yellowjacket abatement in the USA and reviewed the salient research; this report is the basis for the discussion
below. In summary, there is no effective area-wide
abatement technique for the German yellowjacket. Short
of the avoidance of outdoor eating in heavily infested
areas, we can offer no other ameliorative measure than
the practice of strict sanitation to remove attractive, colony supporting foods. This requires tight fitting receptacle lids and regular garbage pickup, and policing of
recreational areas. Industries producing quantities of
sweet effluents may be forced to alter production tech-
439
niques and/or production schedules to eliminate attractive materials or to produce them only after the annual
cycle is completed.
The 2 area-wide abatement techniques that have enjoyed varying, but very limited success against I or 2
west coast species have not worked against any eastern
yellowjacket species. For example, use of traps baited
with artificial attractants has thus far only abated the
western yellowjacket, V. pensylvanica, and then on only
one occasion under specific conditions in Oregon. In
other areas of the west, depletion trapping has not
worked and it is totally ineffective against eastern species. In fact, attractant traps do not even serve as a
population monitoring method since eastern species are
rarely collected, and Perrott (1975) reported attractant
trapping was ineffective against V. germanica in New
Zealand. Future use of attractants will likely be limited
to enhancing meat bait removal by foraging workers as
reported by Wagner and Reierson (1969). Surprisingly,
however, the basis for the attractancy, or lack thereof,
has still not been investigated.
Exploitation of the scavenging behavior of V. vulgaris
group workers has been accomplished by mixing a slow
acting, non-repellent insecticide into an attractive meat
such as tuna, hamburger, horsemeat or commercial cat
foods. This is an environmentally sound abatement strategy and has effectively abated populations of the western yellowjacket and V. vulgaris in some areas of California. Unfortunately this method has not worked
against eastern species and the failure is apparently due
to subtleties in foraging behavior. Still, this is a potentially useful strategy, but research is sorely lacking into
suitable meat baits. Encouraging is the report by Perrott
(1975) that several meat baits are attractive to the German yellowjacket in New Zealand, including freezedried meats such as barracuda. Research related to the
foraging behavior of V. germanica is needed in this
country, particularly since the biotype established may
differ in aspects of behavior other than nest location.
Localized yellowjacket abatement is achieved via the
destruction of colonies, but even this is complicated
when the German yellowjacket is the offender due to
the structural nest site. For example, one Lafayette, IN,
colony was suspended (totally exposed) at head height
from the ceiling of a dark basement and constituted a
serious stinging hazard. One particularly insidious aspect of the structural nest site was noted with 3 Lafayette, IN, colonies in 1979. In each case, a homeowner
called to report "masses of yellowjackets"
chewing
through an interior wall and emerging into the home.
Apparently restricted in normal downward nest expansion in the structural wall void, worker yellowjackets
initiated lateral nest expansion leading them eventually
to chew through the wall. In these 3 cases the homeowner had no knowledge of an existing colony. An additional manifestation of undiscovered structural colonies occurred in several homes this autumn in Lafayette,
IN. Homeowners called to report "huge yellowjackets
suddenly appearing one morning at windows and on
curtains". In these cases, numerous newly produced
queens and males were found. They apparently repre-
440
ESA BULLETIN
sented reproductives emerging from the parent colony
in a wall void toward an interior light source instead of
to the exterior via the access hole used by workers.
Since the above problems can arise during the season,
we recommend destroying yellowjacket colonies in
structures as soon as located, although destruction of a
viable colony will result in eventual decomposition and
possibly some structural damage and offensive odor; if
possible, removal of such colonies is advisable.
Destruction of individual German yellowjacket colonies poses difficulties not typically encountered with
aerial and subterranean nesters. Unfortunately for control purposes, the actual nest site in relation to the access
hole is quite variable and not predictable. Thus, the nest
may be a few centimeters or many meters distant from
the access hole. Furthermore, the nest may be deep into
an interior wall, along an outside wall, up into an attic,
down into a basement wall or situated between a floor
and a basement ceiling. For example, one Lafayette,
IN, colony used an access hole above a first floor window frame yet the nest was located over 30 meters away
in the attic insulation.
Choice of an appropriate control method is very important because ineffective control of a structural colony
can result in surviving workers finding or making another exit from the wall void or other structural locations. Exact location of a structural nest should be determined prior to deciding on a control measure but this
is not always possible because of the cryptic nest site.
The access hole into the structure should never be
plugged without first correctly applying the proper insecticide. Furthermore, most household insecticides dispensed from spray cans will not penetrate deeply enough
into the structural void and will not penetrate the thick
protective nest envelope. Several ineffective control efforts by homeowners in the Lafayette, IN, area led to
homes filled with displaced yellowjackets that had
chewed through interior wallboard walls.
The following procedures and materials are recommended. If the colony is located near the access hole,
1% resmethrin applied as an aerosol is effective. The
plastic wand of the aerosol generator is inserted into the
access hole and the material released for 30-45 seconds,
after which the hole should be plugged with steel wool
sprinkled with 5% carbaryl (Apicidel) dust (Akre et al.
1980). Resmethrin provides relatively fast knockdown
of adult yellowjackets and the dusted steel wool is
chewed by workers attempting to escape the wall void
and by workers returning to the nest from the outside.
However, resmethrin may not always penetrate the nest
envelope to kill the queen and developing brood; thus,
adults could emerge for several days following treatment
(J. Nixon pers. comm.).
When the nest is located away from the access hole,
around a corner, or when one is unsure of nest location
but wishes to insure success, an additional step is advised. Carbaryl dust (5%) should be forcibly blown deep
in all possible directions into the wall void (amount varies with the situation) and followed with the resmethrin
application mentioned above. Agitated workers will repeatedly track the carbaryl dust into the nest interior
Vol. 26, no. 4, 1980
where it will contact the queen and brood, and effect
colony control within a few days. This step is essential
when the nest is deep into the structure in situations
precluding adequate penetration of resmethrin. Application of carbaryl dust only around the entrance probably will not effect control and may result in workers
finding or making another exit hole.
We trust entomologists will assist in providing information about the necessity for thoughtful and effective
control applications (outlined above) when dealing with
structural yellowjacket colonies. Experience in rectifying misguided and ineffective control of the German
yellowjacket the past 2 years has impressed upon us the
need to prevent such failures from occurring in the future.
Future Considerations
The German yellowjacket will likely be introduced
into many more areas around the world, and probably
continue its westward march across the USA. Based on
the propensity for movement of overwintering queens
via commerce, V. germanica may spread saltatorially
via urban centers to the western states. Somewhat surprising is the absence of (or undetected) establishment
of V. germanica in the southern states.
Potential for new establishment throughout this country makes recognition of V. germanica imperative.
However, identification of Vespula species, which is
based largely on color patterns, is difficult without experience. Difficulties center around separation ofV. germanica from native Nearctic species such as V. maculifrons, V. vulgaris and V. jlavopilosa in the east, and
J
•
FIG. I.-Nest of a Vespula germanica colony in Frankfort,
Indiana home attic. The colony was killed with resmethrin Nov.
I, 1978, and collected the following day. Most of the visible
structure was envelope paper surrounding the 8 brood combs.
Vol. 26, no. 4, 1980
Table I.-Location
Location
Ithaca, NY
Ithaca, NY
Ithaca, NY
Frankfort, IN
Lafayette, IN
Lafayette, IN
England
ESA BULLETIN
and nest size of V. germanica
441
colonies from the USA and England."
Date
Aug. 25, 1977
Oct. 10, 1978
Dec. 1978
Nov. 2, 1978
Sep. 14, 1979
Oct. 29, 1979
Ju]y-Oct. ]96]
Estimated
Nest Size
(Cells)
Nest Site
Wall void
Wall void
Attic
Attic
Basement
Basement
Subterranean
12,850
7,850
18,370
17,810
11,112"
6,963
11,961
(largest)
•• Data based on 30 V. gumanica colonies studied by Spradbery (1911); 27 were subterranean and none were inside homes.
h Viable colony that had not shifted to quec:n·cell construction; all 11.112 cells were small, and most contained either worker or male pupae,
this colony probably would have remained active into December and would have constructed nearly 14,000 cells.
V. vulgaris and V. pensylvanica in the west. Well illustrated identification keys for Nearctic Vespula are provided by Alae et al. (1980) and for the V. vulgaris group
by Jacobson et al. (1978). Among eastern North America Vespula. the gray nest of V. germanica is easily
distinguished from the tan/orange, fragile nest of V.
maculifrons. V. vulgaris and V. j1avopilosai however,
a few other Vespula species build gray nests, including
V. pensylvanica and several less common species.
Establishment of the German yellowjacket in urban
areas of the USA will eventually bring about a necessity
for area-wide abatement. Presently, the most promising
potential tool is the development of a poison meat bait
program. Any possible success rests with discovering
an attractive bait, but this will require research into
worker foraging behavior. Success of a New Zealand
program based on freeze-dried meat (Perrott 1975) provides encouragement and, hopefully, incentive for similar research in this country.
A more novel approach (at least for yellowjackets)
might involve the exploitation of the natural chemical
compounds that mediate the social biology of colonies.
For example, rigorous study is needed into the basis of
attractancy of the synthetic compounds (e.g. heptyl butyrate) discovered to be specific for Vespula. We postulate that these attractants simulate natural compounds
that are in some way "biologically meaningful" to individuals in the genus Vespula. Enhancement could lead
to an effective means of iuring foraging workers away
from attractive food sources. Additionally, Vespula
queens appear to influence control of colony mates by
means of a queen pheromone (Landolt et al. 1977), and
it may be possible to develop an "anti-queen pheromone" and somehow disrupt colony cohesion. Research
into the chemical ecology of Vespula is in its infancy,
but hopefully this fertile area will attract future researchers.
Acknowledgment
We thank Dr. George Eickwort, Department of Entomology, Cornell University, and Drs. Gary Bennett
and Donald Paschke, Department of Entomology, Purdue University for their constructive criticism and suggestions for improving the manuscript.
REFERENCES
larvae Or eggs. Left alone,
CITED
Akre, R. E., A. Greene, J. F. MacDonald, P. Landolt, and
H. G. Davis. 1980. Yellowjackets of America North of
Mexico. USDA Agric. Handbook 552 (In press).
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Dejong, D. 1979. Social wasps, enemies of honey bees. Am.
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Edwards, R. 1976. The world distribution pattern of the German wasp Paravespula germanica (Hymenoptera: Vespidae). Entomol. Germ. 3:269-71.
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