Invasive Species Compendium

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Datasheet

Vespula germanica
(German wasp)

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Datasheet

Vespula germanica (German wasp)

Summary

  • Last modified
  • 08 November 2018
  • Datasheet Type(s)
  • Invasive Species
  • Natural Enemy
  • Host Animal
  • Preferred Scientific Name
  • Vespula germanica
  • Preferred Common Name
  • German wasp
  • Taxonomic Tree
  • Domain: Eukaryota
  •   Kingdom: Metazoa
  •     Phylum: Arthropoda
  •       Subphylum: Uniramia
  •         Class: Insecta
  • Summary of Invasiveness
  • Introduction and establishment of V. germanica has been recorded in New Zealand (1944), Australia (1959), South

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Pictures

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PictureTitleCaptionCopyright
Vespula germanica (German wasp); adult.
TitleAdult
CaptionVespula germanica (German wasp); adult.
Copyright©Richard Bartz, Munich (aka Makro Freak)/via wikipedia - CC BY 2.5
Vespula germanica (German wasp); adult.
AdultVespula germanica (German wasp); adult.©Richard Bartz, Munich (aka Makro Freak)/via wikipedia - CC BY 2.5
Vespula germanica (German wasp); worker, female, lateral view. ca.14mm in length. 10 km NW of Kitzbühel, Austria. September 2009.
TitleWorker
CaptionVespula germanica (German wasp); worker, female, lateral view. ca.14mm in length. 10 km NW of Kitzbühel, Austria. September 2009.
CopyrightPublic Domain - Released by Bernie Kohl/via wikipedia - CC0
Vespula germanica (German wasp); worker, female, lateral view. ca.14mm in length. 10 km NW of Kitzbühel, Austria. September 2009.
WorkerVespula germanica (German wasp); worker, female, lateral view. ca.14mm in length. 10 km NW of Kitzbühel, Austria. September 2009.Public Domain - Released by Bernie Kohl/via wikipedia - CC0
Vespula germanica (German wasp); worker, female, ventral view. ca.14mm in length. 10 km NW of Kitzbühel, Austria. September 2009.
TitleWorker
CaptionVespula germanica (German wasp); worker, female, ventral view. ca.14mm in length. 10 km NW of Kitzbühel, Austria. September 2009.
CopyrightPublic Domain - Released by Bernie Kohl/via wikipedia - CC0
Vespula germanica (German wasp); worker, female, ventral view. ca.14mm in length. 10 km NW of Kitzbühel, Austria. September 2009.
WorkerVespula germanica (German wasp); worker, female, ventral view. ca.14mm in length. 10 km NW of Kitzbühel, Austria. September 2009.Public Domain - Released by Bernie Kohl/via wikipedia - CC0
Vespula germanica (German wasp); worker, female, in habitat. ca.14mm in length. 10 km NW of Kitzbühel, Austria. September 2009.
TitleWorker
CaptionVespula germanica (German wasp); worker, female, in habitat. ca.14mm in length. 10 km NW of Kitzbühel, Austria. September 2009.
CopyrightPublic Domain - Released by Bernie Kohl/via wikipedia - CC0
Vespula germanica (German wasp); worker, female, in habitat. ca.14mm in length. 10 km NW of Kitzbühel, Austria. September 2009.
WorkerVespula germanica (German wasp); worker, female, in habitat. ca.14mm in length. 10 km NW of Kitzbühel, Austria. September 2009.Public Domain - Released by Bernie Kohl/via wikipedia - CC0
Vespula germanica (German wasp); rasping wood for nest building. July 2013.
TitleRasping wood
CaptionVespula germanica (German wasp); rasping wood for nest building. July 2013.
Copyright©Pjt56/Stuttgart, Germany/via wikipedia - CC BY-SA 3.0
Vespula germanica (German wasp); rasping wood for nest building. July 2013.
Rasping woodVespula germanica (German wasp); rasping wood for nest building. July 2013.©Pjt56/Stuttgart, Germany/via wikipedia - CC BY-SA 3.0
Vespula germanica (German wasp); workers landing on a fallen apple. 10 km NW of Kitzbühel, Austria. September 2009.
TitleWorkers
CaptionVespula germanica (German wasp); workers landing on a fallen apple. 10 km NW of Kitzbühel, Austria. September 2009.
CopyrightPublic Domain - Released by Bernie Kohl/via wikipedia - CC0
Vespula germanica (German wasp); workers landing on a fallen apple. 10 km NW of Kitzbühel, Austria. September 2009.
WorkersVespula germanica (German wasp); workers landing on a fallen apple. 10 km NW of Kitzbühel, Austria. September 2009.Public Domain - Released by Bernie Kohl/via wikipedia - CC0
Vespula germanica (German wasp); adult, extreme close-up of head capsule, showing characteristic facial pattern.
TitleHead capsule
CaptionVespula germanica (German wasp); adult, extreme close-up of head capsule, showing characteristic facial pattern.
Copyright©Richard Bartz, Munich (aka Makro Freak)/via wikipedia - CC BY 2.5
Vespula germanica (German wasp); adult, extreme close-up of head capsule, showing characteristic facial pattern.
Head capsuleVespula germanica (German wasp); adult, extreme close-up of head capsule, showing characteristic facial pattern.©Richard Bartz, Munich (aka Makro Freak)/via wikipedia - CC BY 2.5
Vespula germanica (German wasp); worker, female, ca.14mm in length. She is cleaning the tarsus of a front right leg between her mandibles. 10 km NW of Kitzbühel, Austria. September 2009.
TitleWorker
CaptionVespula germanica (German wasp); worker, female, ca.14mm in length. She is cleaning the tarsus of a front right leg between her mandibles. 10 km NW of Kitzbühel, Austria. September 2009.
CopyrightPublic Domain - Released by Bernie Kohl/via wikipedia - CC0
Vespula germanica (German wasp); worker, female, ca.14mm in length. She is cleaning the tarsus of a front right leg between her mandibles. 10 km NW of Kitzbühel, Austria. September 2009.
WorkerVespula germanica (German wasp); worker, female, ca.14mm in length. She is cleaning the tarsus of a front right leg between her mandibles. 10 km NW of Kitzbühel, Austria. September 2009.Public Domain - Released by Bernie Kohl/via wikipedia - CC0
Vespula germanica (German wasp); mated queen. nr. Primorsko, Bulgaria. October 2015.
TitleMated queen
CaptionVespula germanica (German wasp); mated queen. nr. Primorsko, Bulgaria. October 2015.
Copyright©Katya/Moscow, Russia/via wikipedia - CC BY-SA 2.0
Vespula germanica (German wasp); mated queen. nr. Primorsko, Bulgaria. October 2015.
Mated queenVespula germanica (German wasp); mated queen. nr. Primorsko, Bulgaria. October 2015.©Katya/Moscow, Russia/via wikipedia - CC BY-SA 2.0
Vespula germanica (German wasp); nest. Frankfurt Zoo, Germany. May2010.
TitleNest
CaptionVespula germanica (German wasp); nest. Frankfurt Zoo, Germany. May2010.
Copyright©JuTa/via wikipedia - CC BY-SA 3.0
Vespula germanica (German wasp); nest. Frankfurt Zoo, Germany. May2010.
NestVespula germanica (German wasp); nest. Frankfurt Zoo, Germany. May2010.©JuTa/via wikipedia - CC BY-SA 3.0

Identity

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Preferred Scientific Name

  • Vespula germanica Fabricius, 1793

Preferred Common Name

  • German wasp

Other Scientific Names

  • Dolichovespula germanica
  • Paravespula germanica Blüthgen, 1938
  • Pseudovespula germanica
  • Vespa germanica

International Common Names

  • English: European wasp; German yellow jacket
  • Spanish: avispa alemana
  • French: guepe germanique

Local Common Names

  • Bulgaria: vosa
  • Denmark: tysk gedehams
  • Germany: Wespe, Deutsche
  • Italy: vespe
  • Latvia: lapsene
  • Lithuania: vaspa
  • Norway: tysk veps
  • Portugal: abispa
  • Russian Federation: ovsa
  • Sweden: tysk geting

EPPO code

  • VESPGE (Vespa germanica)

Summary of Invasiveness

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Introduction and establishment of V. germanica has been recorded in New Zealand (1944), Australia (1959), South Africa (1972), United States (New York 1891, Maryland 1968, California 1991), Canada (1971), Chile (1974) and Argentina (1978). Introductions have also been recorded in Iceland and Ascension Island. V. germanica can have significant negative impacts on horticulture, apiculture, tourism and outdoor social activities, as well as animal health and biodiversity. 

Taxonomic Tree

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  • Domain: Eukaryota
  •     Kingdom: Metazoa
  •         Phylum: Arthropoda
  •             Subphylum: Uniramia
  •                 Class: Insecta
  •                     Order: Hymenoptera
  •                         Family: Vespidae
  •                             Genus: Vespula
  •                                 Species: Vespula germanica

Notes on Taxonomy and Nomenclature

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Vespula germanica is a wasp species of Palaearctic origin within the order Hymenoptera, family Vespidae, subfamily Vespinae. This species is commonly known as the European wasp in Australasia, South Africa and South America, and the German wasp or yellowjacket elsewhere.

Description

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V. germanica is marked with bold black and yellow bands. The black bands have arrow-shaped black markings down the centre of the abdomen, and there are pairs of small black spots on the yellow bands. The wings are long and transparent, the antennae are black and the legs are mostly yellow. Workers measure 12-15 mm in length; queens are similar but larger (up to about 20 mm) (OzAnimals, 2012).

Distribution

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Some details of the endemic distribution of V. germanica in Europe were provided by P Kment (National Museum of Czech Republic), J Diller (Zoological State Collection, Munich, Germany) and V Repasi (Hungarian Natural History Museum, Budapest, Hungary) (personal communications 2008).

The distribution of V. germanica in North America was described by Akre et al. (1980). Subsequently, V. germanica has been recorded in Maryland (Morse et al., 1977), Washington State (KM Pickett, Department of Biology, University of Vermont, personal communication, 2008), Missouri (Hunt and Sanders, 1998), Wisconsin (R Jeanne, Department of Entomology, University of Wisconsin; J Jandt , Department of Ecology and Evolutionary Biology, University of Arizona, personal communications, 2008), Illinois (B Taylor, Department of Zoology, University of Wisconsin-Madison, personal communication, 2008), and Alabama, Maryland, Tennessee and Kentucky (Triplehorn Insect Collection of the Ohio State University, courtesy of J Wenzel, Department of Entomology, The Ohio State University, personal communication, 2008). V. germanica specimens have been confirmed in Arizona, Utah, Oregon and Indiana by B Jacobson (Greer Laboratories, Lenoir, North Carolina, personal communication, 2008). The species also occurs widely in central and southern coastal California (K Vissher and N Nisson, Department of Entomology, University of California, Riverside, personal communications, 2008).

V. germanica has not yet been recorded in Georgia, Texas, Oklahoma or Florida (USA) (JM Carpenter, Division of Invertebrate Zoology, American Museum of Natural History; M Goodisman, School of Biology, Georgia Institute of Technology, Atlanta; B Jacobson, J Jandt and H Reed, Department of Biology, Oral Roberts University, Tulsa and D Wahl, American Entomological Institute, Gainesville, Florida, personal communications 2008).

The first records of V. germanica in Canada were from Winnipeg, Manitoba in 1976 (Galloway and Preston, 1982) and the species now occurs in Ontario, Quebec and British Columbia (M Buck, Department of Environmental Biology, University of Guelph, Ontario; S McCann, Department of Biological Sciences, Simon Fraser University; J Huber, Canadian National Collection of Insects and Arachnids, Ottawa, personal communications, 2008). It also occurs in Halifax (Nova Scotia), Sydney and Lake Uist (Cape Breton Island) (DB McCorquodale, Department of Biology, Cape Breton University, Nova Scotia; C Majka, Nova Scotia Museum of Natural History, Halifax, personal communications, 2008).

In Chile, V. germanica has been recorded from Punta Arenas in the southern tip of South America to Coquimbo in the north (R Ripa, Instituto de Investigaciones Agropecurias, La Cruz, Chile, personal communication, 2008). In Argentina, the species is present throughout Patagonia as far north as Buenos Aires Province and southwards to Parque Nacional Los Glaciares (P d’Adamo, Laboratorio de Ecología de Insectos Forestales, INTA EEA, Bariloche, Argentina, personal communication, 2008).

Records for the distribution of V. germanica in South Africa were provided by AH Kirk-Spriggs (Albany Museum, Grahamstown) and G Tribe (ARC Plant Protection Research Institute, Stellenbosch, personal communications, 2008).

Distribution Table

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The distribution in this summary table is based on all the information available. When several references are cited, they may give conflicting information on the status. Further details may be available for individual references in the Distribution Table Details section which can be selected by going to Generate Report.

Continent/Country/RegionDistributionLast ReportedOriginFirst ReportedInvasiveReferenceNotes

Asia

AfghanistanPresentNative Not invasive Giordani Soika, 1962; Gusenleitner, 1972; Archer, 1998
ArmeniaPresentNative Not invasive Kostylev, 1929; Archer, 1998
ChinaPresentPresent based on regional distribution.
-BeijingPresentNative Not invasive Yamane et al., 1980; Carpenter and Kojima, 1997
-GansuPresentNative Not invasive Carpenter and Kojima, 1997; Archer, 1998
-HebeiPresentNative Not invasive Carpenter and Kojima, 1997; Archer, 1998
-HeilongjiangPresentNative Not invasive Carpenter and Kojima, 1997; Archer, 1998
-JilinPresentNative Not invasive Yamane et al., 1980
-LiaoningPresentNative Not invasive Carpenter and Kojima, 1997; Archer, 1998
-Nei MengguPresentNative Not invasive Carpenter and Kojima, 1997
-QinghaiPresentNative Not invasive Carpenter and Kojima, 1997
-SichuanPresentNative Not invasive Yamane et al., 1980
-TibetPresentNative Not invasive Yamane et al., 1980
-XinjiangPresentNative Not invasive Archer, 1998
Georgia (Republic of)PresentNative Not invasive Kobakhidze, 1962; Archer, 1998
IndiaPresentPresent based on regional distribution.
-Jammu and KashmirPresentNative Not invasive Das and Gupta, 1989
IranPresentNative Not invasive Blüthgen and Gusenleitner, 1970; Archer, 1998
IraqPresentNative Not invasive Dvorák and Landolt, 2006
IsraelPresentNative Not invasive Ishay et al., 1986; Dubatolov, 2002
JordanPresentNative Not invasive Haddad et al., 2007Above 1000m altitude
KazakhstanPresentNative Not invasive Archer, 1998; Dubatolov, 2005
Korea, DPRPresentNative Not invasive Yamane et al., 1980
Korea, Republic ofPresentNative Not invasive Carpenter and Kojima, 1997
KyrgyzstanWidespreadNative Not invasive Dubatolov and Milko, 2004
LebanonPresentNative Not invasive Ebner, 1930
MongoliaPresentNative Not invasive Gusenleitner, 1991; Archer, 1998
PakistanPresentNative Not invasive Das and Gupta, 1989; Dvorak, 2007
SyriaPresentNative Not invasive Carpenter and Kojima, 1997; Archer, 1998
TajikistanPresentNative Not invasive Carpenter and Kojima, 1997; Archer, 1998
TurkeyPresentNative Not invasive Archer, 1998; Yildirim and Kojima, 1999
TurkmenistanPresentNative Not invasive Carpenter and Kojima, 1997; Dubatolov, 2002
UzbekistanPresentNative Not invasive Carpenter and Kojima, 1997; Archer, 1998; Dubatolov, 2002

Africa

AlgeriaPresentNative Not invasive Archer, 1998
MoroccoPresentNative Not invasive Gusenlaitner, 1977; Archer, 1998
Saint HelenaPresentPresent based on regional distribution.
-AscensionPresentIntroducedYarrow, 1967
South AfricaLocalisedIntroduced Invasive Whitehead and Prins, 1975First record at Kirstenbosch, Cape peninsula 1972
Spain
-Canary IslandsWidespreadIntroduced Invasive Erlandsson, 1978; Archer, 1998
TunisiaPresentNative Not invasive Archer, 1998

North America

CanadaPresentPresent based on regional distribution.
-AlbertaLocalisedIntroduced Invasive M Buck, Royal Alberta Museum, Canada, pers. comm., 2010Edmonton, first records in 2009
-British ColumbiaLocalisedIntroduced Invasive Archer, 1998In southern areas only
-ManitobaWidespreadIntroduced Invasive Buck et al., 2008First recorded 1976
-New BrunswickWidespreadIntroduced Invasive Akre et al., 1980
-Newfoundland and LabradorPresentIntroduced Invasive M Buck, Royal Alberta Museum, Canada, pers. comm., 2010Mt Pearl
-Nova ScotiaWidespreadIntroduced Invasive Akre et al., 1980
-OntarioWidespreadIntroduced Invasive Buck et al., 2008First recorded 1971
-QuebecWidespreadIntroduced Invasive Buck et al., 2008
-SaskatchewanLocalisedIntroduced Invasive Archer, 1998
USAPresentPresent based on regional distribution.
-CaliforniaWidespreadIntroduced Invasive Vetter et al., 1995First nests reported 1991
-ConnecticutWidespreadIntroduced Invasive Akre et al., 1980
-DelawareWidespreadIntroduced Invasive Menke and Snelling, 1975
-GeorgiaWidespreadIntroduced Invasive Akre et al., 1980
-IdahoPresentIntroduced Invasive Archer, 1998
-IllinoisWidespreadIntroduced Invasive Akre et al., 1980
-IndianaWidespreadIntroduced Invasive Macdonald et al., 1980First records 1976
-IowaWidespreadIntroduced Invasive Akre et al., 1980
-MaineWidespreadIntroduced Invasive Akre et al., 1980
-MarylandWidespreadIntroduced Invasive Morse et al., 1977Established 1968
-MassachusettsWidespreadIntroduced Invasive Akre et al., 1980
-MichiganWidespreadIntroduced Invasive Akre et al., 1980First record 1978
-MinnesotaWidespreadIntroduced Invasive Akre et al., 1980
-MissouriLocalisedIntroduced Invasive R Jeanne, University of Wisconsin, USA, personal communication, 2009; Hunt and Sanders, 1998
-New HampshireWidespreadIntroduced Invasive Akre et al., 1980
-New JerseyWidespreadIntroduced Invasive Wagner and Reierson, 1971First nest reported 1971
-New YorkWidespreadIntroduced Invasive Morse et al., 1977First record 1891
-North CarolinaWidespreadIntroduced Invasive Akre et al., 1980
-OhioWidespreadIntroduced Invasive Macdonald et al., 1980First records 1971
-OregonPresentIntroduced Invasive Archer, 1998
-PennsylvaniaWidespreadIntroduced Invasive Menke and Snelling, 1975
-South CarolinaWidespreadIntroduced Invasive Akre et al., 1980
-TennesseeWidespreadIntroduced Invasive Akre et al., 1980
-VermontWidespreadIntroduced Invasive Akre et al., 1980
-VirginiaWidespreadIntroduced Invasive Akre et al., 1980
-WashingtonPresentIntroduced Invasive Archer, 1998
-West VirginiaWidespreadIntroduced Invasive Akre et al., 1980
-WisconsinWidespreadIntroduced Invasive Akre et al., 1980Now dominates the 12 other spp. of Vespula

South America

ArgentinaLocalisedIntroduced Invasive Willink, 1980; Edwards, 1984; D'Adamo et al., 2002First established in Depto. Mina, northern Patagonia in 1978, now widespread throughout Patagonia
ChileLocalisedIntroduced Invasive Peña et al., 1976First recorded in Santiago in 1974

Europe

AlbaniaWidespreadNative Not invasive Archer, 1998; Cetkovic, 2002
AustriaWidespreadNative Not invasive Gusenleitner, 1981; Archer, 1998
BelarusPresentNative Not invasive Archer, 1998; Agunovich, 2007; Shlyakhtenok, 2007
BelgiumWidespreadNative Not invasive Archer, 1998
BulgariaWidespreadNative Not invasive Archer, 1998; Cetkovic, 2002
CroatiaWidespreadNative Not invasive Archer, 1998; Cetkovic, 2002
CyprusWidespreadNative Not invasive Georghiou, 1977
Czech RepublicWidespreadNative Not invasive Archer, 1998; Dvorak and Straka, 2007
DenmarkWidespreadNative Not invasive Archer, 1998
EstoniaPresentNative Not invasive Remm, 1983
FinlandLocalisedNative Not invasive Pekkarinen and Hulden, 1995Irregular occurrence
FranceWidespreadNative Not invasive Archer, 1998; Gereys, 2006
-CorsicaPresentNative Not invasive Dvorák and Landolt, 2006
GermanyWidespreadNative Not invasive Archer, 1998; Oehlke, 2001
GreeceWidespreadNative Not invasive Archer, 1998; Cetkovic, 2002
HungaryWidespreadNative Not invasive Móczár, 1995
IcelandLocalisedIntroduced Not invasive Olafsson, 1991Rarely in Reykyavik area
IrelandPresentNative Not invasive Archer, 1998; Ings and Roberts, 2002
ItalyWidespreadNative Not invasive Giordani-Soika and Borsato, 1995Also Sicily and Sardinia
-SardiniaPresentGiordani-Soika and Borsato, 1995
-SicilyPresentGiordani-Soika and Borsato, 1995
LatviaPresentTumss, 1968
LithuaniaPresentNative Not invasive Dubatolov, 2002
LuxembourgWidespreadNative Not invasive Sauber and Hoffmann, 1974
MacedoniaPresentNative Not invasive Cetkovic, 2002
MaltaPresentIntroduced Invasive Schembri, 1985
MontenegroPresentNative Not invasive Cetkovic, 2002
NetherlandsWidespreadNative Not invasive Hensen, 1985
NorwayLocalisedNative Not invasive Løken, 1978Southern Norway
PolandWidespreadNative Not invasive Archer, 1998; Skibinska, 2004
PortugalWidespreadNative Not invasive Diniz, 1978Also present Madeira Is.
-MadeiraWidespreadIntroduced Invasive Saunders, 1903; Blüthgen, 1940; Erlandsson, 1978; Smit, 2000
RomaniaWidespreadNative Not invasive Archer, 1998
Russian FederationPresentPresent based on regional distribution.
-Central RussiaWidespreadNative Not invasive Archer, 1998; Dubatolov, 2005
-Eastern SiberiaPresentNative Not invasive Dubatolov, 2005
-Russian Far EastPresent, few occurrencesNative Not invasive Yamane et al., 1980; Dubatolov, 2005Lake Khanka, Sakhalin Island (Kurzenko, 2004)
-Southern RussiaWidespreadNative Not invasive Archer, 1998
-Western SiberiaPresentNative Not invasive Dubatolov, 2005
SerbiaWidespreadNative Not invasive Cetkovic, 2002
SlovakiaWidespreadNative Not invasive Dvorak and Straka, 2007
SloveniaWidespreadNative Not invasive Archer, 1998; Cetkovic, 2002
SpainWidespreadNative Not invasive Madero Montero, 1988
SwedenWidespreadNative Not invasive Erlandsson, 1971; Archer, 1998Southern Sweden
SwitzerlandWidespreadNative Not invasive Archer, 1998; Neumeyer, 2000
UKWidespreadNative Not invasive Edwards, 1984Recorded on Ascension Is in South Atlantic (yarrow, 1967)
-Channel IslandsWidespreadNative Not invasive Ings and Roberts, 2002
UkraineWidespreadNative Not invasive Archer, 1998; Dubatolov, 2002

Oceania

AustraliaPresentPresent based on regional distribution.
-Australian Northern TerritoryAbsent, formerly presentIntroduced Invasive Australian Broadcasting Corporation, 2005One nest found in Alice Springs, January 2005
-New South WalesWidespreadIntroduced Invasive Spradbery and Maywald, 1992Introduced 1975/78, widespread SE half of State
-QueenslandLocalisedIntroduced Invasive Spradbery and Maywald, 1992Restricted to SE of state, nests reported northwards to Maryborough
-South AustraliaLocalisedIntroduced Invasive Spradbery and Maywald, 1992Port Lincoln, Port Augusta and Adelaide region since 1978
-TasmaniaWidespreadIntroduced Invasive Spradbery and Maywald, 1992First nest reported in Hobart in 1959
-VictoriaWidespreadIntroduced Invasive Spradbery and Maywald, 1992First reported in Melbourne 1977
-Western AustraliaLocalisedIntroduced Invasive Spradbery and Maywald, 1992First nests reported in Perth 1977; nests in Albany 1984-85 but since absent; single nests in Kalgoorlie 1988 and 1991
New ZealandWidespreadIntroduced Invasive Clapperton et al., 1989Established Hamilton, North Island in 1945 and South Island in 1955
Norfolk IslandEradicatedIntroduced Not invasive Naumann, 1990Single colony destroyed in 1982

History of Introduction and Spread

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The global endemic distribution of V. germanica includes Europe north to 62°N, the Mediterranean region including coastal North Africa (but excluding Egypt) and eastwards to Palaearctic Asia, northern India, southern Russia, China and Korea south to 23°N. The family includes many Vespula species although only two other species are invasive, Vespula vulgaris in Australia and New Zealand (Clapperton et al., 1989; Matthews et al., 2000; Bashford, 2001) and Vespula pensylvanica in Hawaii (Akre et al., 1980).

The impact of V. germanica as an invasive species has been greatest in Australasia after its introduction and subsequent establishment near a New Zealand Air Force depot in Hamilton, New Zealand in 1944-45. The species had been recorded once before in 1922 but did not become established at that time (Thomas, 1960). The Hamilton introduction, with seven nests reported in 1945, was probably due to the accidental importation during the previous year of queen wasps hibernating on wooden crates containing aircraft spare parts from the United Kingdom. Remains of dead queens were subsequently discovered on the crates. The pest species quickly spread throughout the North Island of New Zealand with 6,000 nests recorded in 1951 (Thomas, 1960) and reached the South Island by the mid 1950s. In 1959 two nests were discovered in Hobart, Tasmania, Australia, and the species spread throughout the island State in the ensuing 15 years (Spradbery and Maywald, 1992). The establishment of V. germanica on the mainland of Australia appears to have occurred as a result of a ship offloading cargo (probably timber from New Zealand) during a single voyage to the ports of Sydney, Melbourne, Adelaide and Perth, with the first nests recorded in the New South Wales, Victoria, South Australia and Western Australia port capitals during 1977 and 1978. Indeed, the first interceptions of V. germanica in Australia were queens discovered in consignments of New Zealand timber shipped to Sydney in 1954 (Chadwick and Nikitin, 1969). The outbreaks in Perth and Albany (Western Australia) have largely been contained or eradicated through vigorous search and destroy activities, but in the other Australian states, the European wasp has spread and now occupies the southeastern half of New South Wales as far north as Queensland, throughout Victoria, and in suburban and rural Adelaide in South Australia. Individual nests have been reported in Kalgoorlie (Western Australia), Maryborough (SE Queensland) and Alice Springs (Northern Territory). 

In the United States, the first record of V. germanica was from a specimen in the Los Angeles Museum of Natural History, collected in Ithaca, New York, in 1891 (Menke and Snelling, 1975). The species remained uncommon in New York from that time until the mid 1960s and was then recorded in Maryland in 1968 (Morse et al., 1977) and later in Delaware, New Jersey, Pennsylvania and Washington in the northeastern United States; it subsequently spread southwards to the Carolinas, Tennessee and Georgia, with its establishment on the western seaboard of California in 1991 (Vetter et al., 1995). In Canada, V. germanica was first recorded in Ontario in 1971 and Manitoba in 1976 (Buck et al., 2008).

The first record of V. germanica in South Africa was a specimen dated 1972, apparently from a nest that had been destroyed in the Kirstenbosch area of the western Cape Peninsula (Whitehead and Prins, 1975).

V. germanica became established in Santiago, Chile, in 1974 and was then discovered approximately 1300 km south at Ponto Aisen after five years (Edwards, 1984). The first record of V. germanica for Argentina was in 1978 at Neuquen and the species has since dispersed rapidly throughout Patagonia (Willink, 1980).

Introductions

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Introduced toIntroduced fromYearReasonIntroduced byEstablished in wild throughReferencesNotes
Natural reproductionContinuous restocking
Argentina Chile 1978 Hitchhiker (pathway cause) Yes Accidental as hibernating queens
Australian Northern Territory South Australia 2005 Hitchhiker (pathway cause) No Accidental as hibernating queens. ABC News Online
California 1991 Hitchhiker (pathway cause) Yes Vetter et al. (1995) Accidental as hibernating queens
Chile 1974 Hitchhiker (pathway cause) Yes Peña et al. (1976) Accidental as hibernating queens
Indiana 1976 Hitchhiker (pathway cause) Yes Macdonald et al. (1980) Accidental as hibernating queens
Manitoba 1976 Hitchhiker (pathway cause) Yes Buck et al. (2008) Accidental as hibernating queens
Maryland 1968 Hitchhiker (pathway cause) Yes Menke and Snelling (1975); Morse et al. (1977) Accidental as hibernating queens
Michigan 1978 Hitchhiker (pathway cause) Yes Akre et al. (1980) Accidental as hibernating queens
New Jersey 1971 Yes Accidental as hibernating queens
New South Wales New Zealand 1977 Hitchhiker (pathway cause) Yes Spradbery and Maywald (1992) Accidental as hibernating queens
New York 1891 Hitchhiker (pathway cause) Yes Menke and Snelling (1975) Accidental as hibernating queens
New Zealand UK 1944 Hitchhiker (pathway cause) Yes Accidental as hibernating queens
Ohio 1971 Hitchhiker (pathway cause) Yes Macdonald et al. (1980) Accidental as hibernating queens
Ontario 1971 Hitchhiker (pathway cause) Yes Buck et al. (2008) Accidental as hibernating queens
Queensland New South Wales 1988 Hitchhiker (pathway cause) Yes Spradbery and Maywald (1992) Accidental as hibernating queens
South Africa 1972 Hitchhiker (pathway cause) Yes Whitehead and Prins (1975) Accidental as hibernating queens
Tasmania New Zealand 1959 Hitchhiker (pathway cause) Yes Spradbery and Maywald (1992) Accidental as hibernating queens

Risk of Introduction

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An insect pest with a life history stage that includes a hibernating, independent, inseminated adult female which is capable of establishing a social insect colony without further interaction or inputs with other members of the species, has a high potential for geographical dispersal and for successful colonization of new territories. The modern age of travel by ships, planes and road vehicles provides immense potential for dispersal of such organisms. If that insect impacts adversely on humans and is transported to a place where the climate and other resources are suitable for its establishment, and where natural enemies probably do not occur, the stage is set for major pest outbreaks.

The social wasp, V. germanica, occupies a very broad climatic variation in its endemic northern hemisphere range: from dry, hot North African coastal areas to the cool, moist latitudes of Scandinavia and Asia, from the Middle East to Korea. Climatic homologues occur throughout the southern hemisphere and North America and it is here on four different continents that this species in recent times has been accidently introduced and successfully established, and has become a notorious invasive species.

Habitat List

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CategoryHabitatPresenceStatus
Other
Soil Principal habitat
Terrestrial-managed
Buildings Principal habitat Harmful (pest or invasive)
Cultivated / agricultural land Principal habitat Harmful (pest or invasive)
Disturbed areas Principal habitat Harmful (pest or invasive)
Rail / roadsides Principal habitat Harmful (pest or invasive)
Urban / peri-urban areas Principal habitat Harmful (pest or invasive)
Terrestrial-natural/semi-natural
Natural forests Secondary/tolerated habitat Harmful (pest or invasive)
Riverbanks Principal habitat Harmful (pest or invasive)

Host Animals

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Animal nameContextLife stageSystem
Apis mellifera
Bos indicus (zebu)
Bos taurus (cattle)

List of Symptoms/Signs

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SignLife StagesType
Pain / Discomfort Signs / Skin pain Sign
Reproductive Signs / Mastitis, abnormal milk Sign
Reproductive Signs / Teat injury, cut, tear Sign
Skin / Integumentary Signs / Parasite visible, skin, hair, feathers Sign
Skin / Integumentary Signs / Skin ulcer, erosion, excoriation Sign

Biology and Ecology

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V. germanica is a social wasp with a typically annual life cycle. Queens produced during late summer and autumn mate with one or more males, storing the sperm within a sperm-storage organ, the spermatheca. They then pass the winter months in hibernation in well-insulated places such as leaf litter, crevices in tree bark and also buildings and outhouses [see pictures]. In spring, the queens emerge from hibernation, feed at nectar sources and begin searching for suitable nesting sites. Typically such sites are hidden away: underground, behind retaining walls and rockeries in gardens, and also in cavity walls or roof spaces in buildings. The abandoned burrows of rodents and rabbits are also utilized. Only very rarely does V. germanica construct an exposed nest [see pictures]. There is sometimes competition for nest sites and spring queens may die during fights while invading or defending a nest site (Spradbery, 1991). The founding queen builds her ‘embryo’ nest after collecting wood from old, seasoned (grey-coloured) dead trees, wooden poles and fences by scraping off the fibres, mixing with saliva and working with its serrated mandibles to produce thin strips of wet ‘paper’ that resemble papier-mâché or carton. The embryo nest consists of a comb of about 30 hexagonal cells with their openings facing downwards attached to the substrate with a stout pillar of wasp carton and surrounded by several round layers of umbrella-like envelopes for insulation, the whole structure being about the size and shape of a large golf ball [see pictures].

Once adult worker wasps (daughters of the founding queen) are reared to maturity, they assume all foraging and nest-building duties and the queen remains within the nest. Worker activities include excavating the nest site by removing soil, often after softening it with collected water, to allow nest enlargement. Foraging for food entails seeking protein to feed the developing larvae such as arthropod prey and carrion, and carbohydrates for adults such as nectar, tree sap, fruit and honey dew.

As the nest develops, more combs are added below the original queen-built comb, suspended by stout carton pillars and the outer, insulating envelope is re-cycled as the nest grows larger. By late summer the nest is about the size of a football with several thousand adult workers in the colony. At this time, queen-rearing cells, which are twice the volume of the worker cells, are constructed and the new queens and also males (which may be reared in both worker and queen cells) are produced.

After feeding and laying down quantities of body fat, the young queens leave the nest, mate and then finally seek hibernation quarters where they will remain until spring. V. germanica colonies have the capacity to overwinter by re-queening whereby the new queens, instead of leaving the nest to hibernate, remain in the nest, develop their ovaries and begin laying eggs. Such behaviour has been observed occasionally in North Africa, on the Greek island of Skopolos and in the South of France, but is a very common phenomenon in Australasia where about 10 per cent of colonies of V. germanica continue for 12 months or more, some perennial nests growing to a prodigious size (Spradbery, 1988) [see pictures].

Climate

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ClimateStatusDescriptionRemark
Cf - Warm temperate climate, wet all year Preferred Warm average temp. > 10°C, Cold average temp. > 0°C, wet all year
Cs - Warm temperate climate with dry summer Preferred Warm average temp. > 10°C, Cold average temp. > 0°C, dry summers
Cw - Warm temperate climate with dry winter Preferred Warm temperate climate with dry winter (Warm average temp. > 10°C, Cold average temp. > 0°C, dry winters)

Latitude/Altitude Ranges

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Latitude North (°N)Latitude South (°S)Altitude Lower (m)Altitude Upper (m)
65 47 0 0

Air Temperature

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Parameter Lower limit Upper limit
Absolute minimum temperature (ºC) <0
Mean annual temperature (ºC) 26
Mean maximum temperature of hottest month (ºC) 33
Mean minimum temperature of coldest month (ºC) 18

Natural enemies

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Natural enemyTypeLife stagesSpecificityReferencesBiological control inBiological control on
Aspergillus flavus
Linepithema humile Predator
Melittobia australica Parasite
Sphecophaga vesparum Parasite Larvae/Pupae to genus Donovan and Read, 1987; Field and Darby, 1991; Read et al., 1990 New Zealand
Sphecophaga vesparum burra Parasite New Zealand
Steinernema feltiae Parasite

Notes on Natural Enemies

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The wasp colony with its paper structure, large quantities of protein in the form of immature and adult wasps plus the detritus that accumulates underground below the nest, provides a rich habitat for predators, parasites and commensals, despite the defensive stings and jaws possessed by adult wasps. Numerous fly and beetle species, parasitoids such as ichneumonid wasps, mites, nematodes, and predators such as birds and badgers have all been recorded living at the expense of wasp colonies (Spradbery, 1973). However, with the exception of the ichneumonid parasitoid, Sphecophaga vesparum, none have been considered appropriate as biological control agents. In the case of S. vesparum, this species was introduced from the United States and Europe to New Zealand where it was reared on field-collected V. germanica brood, host specificity tests were carried out and it was subsequently released into the field (Donovan and Read, 1987). Parasitoid establishment was confirmed although wasp nest densities appear unaffected (Beggs et al., 1996). A similar exercise using S.vesparum from New Zealand was carried out in Victoria, Australia (Field and Darby, 1991).

Means of Movement and Dispersal

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Foraging worker wasps of the genus Vespula can travel up to 3000 m but the majority forage within 400 m of their nest site (Akre et al., 1975). Queens of the related social wasp, Vespula rufa, have been observed in mass movements during the spring exodus from hibernation quarters near the coast of Sweden and it was estimated that some had flown at least 50km (Rudebeck, 1965). Undoubtedly, when searching for suitable nest sites in the spring, queens of V. germanica can probably disperse several km but no confirmed distances have been determined.  

Accidental introduction of V. germanica queens during the hibernation phase of their life cycle is inevitable in a world with modern transport systems. The movement of materials in which queens are hibernating such as boxes and other merchandise by ship, rail or plane to a geographical assortment of destinations has occurred many times over the past seventy years or so. Introductions to countries in the southern hemisphere and North America with a similar climate to their endemic range in the northern hemisphere, such as Australia, New Zealand, South Africa, United States, Canada, Chile and Argentina all bear witness to the success of this passive transport. Once established within a new country or region, the accidental spread of hibernating queens via road and rail transport enables further dispersal and potential colonization of all suitable habitats. This has certainly been the case in Australia where V. germanica has now spread from the original seaports of introduction to all parts of the island continent that are suitable, with only Western Australia authorities succeeding in limiting its local spread (Spradbery and Maywald, 1992).

Pathway Causes

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CauseNotesLong DistanceLocalReferences
HitchhikerEurope to New Zealand, New Zealand to Australia, within Australia Yes Yes Spradbery and Maywald, 1992; Thomas, 1960

Pathway Vectors

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VectorNotesLong DistanceLocalReferences
Bulk freight or cargo Yes Yes
Containers and packaging - non-wood Yes Yes
Containers and packaging - wood Yes Yes

Impact Summary

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CategoryImpact
Cultural/amenity Negative
Environment (generally) Negative
Human health Negative

Impact

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V. germanica can have significant negative impacts on horticulture, apiculture, tourism and outdoor social activities, as well as animal health and biodiversity.

Such impacts occur primarily in countries where the wasp has been accidentally introduced although there are examples of adverse V. germanica impacts on the quality of life and also human fatalities in countries where this species is endemic. In Australia, wasps kill livestock when animals such as goats eat fallen fruit containing foraging wasps. Several dog deaths have been reported, ferrets have been killed when sent down rabbit burrows containing wasp nests, and a horse died after trampling a nest in the Snowy Mountains in Victoria, Australia.

Although the wasps’ impact on horticulture can be costed relatively easily, the major impact of this pest species is in urban and tourist areas where economic costs are more difficult to gauge. Fowler (1983) noted long ago that “unlike agricultural systems, it is often not possible to approach urban pests on a cost-benefit basis. Human attitudes have yet to be assessed fully to produce an urban counterpart to the agricultural economic threshold, due in part to the psycho-sociological research needed to establish a suitable tolerance level”.

Economic Impact

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Beekeepers in New Zealand and Tasmania, Australia, suffer significant losses to V. germanica predation of adult bees and invasion of hives. Some 2% of New Zealand apiaries are destroyed annually and 10,000 hives severely affected (Walton and Reid, 1976). Vineyards in Australia are adversely affected by foraging wasps which spoil fruit and intimidate pickers, and grape growers in Oregon and Washington State can lose half their crops to Vespula wasps. A similar species in California caused annual losses to agriculture of US $200,000 in 1970s values (Poinar and Ennik, 1972).

Vespula can have a marked impact on tourism when they are present in large numbers and foraging aggressively for food. For example, on the Greek island of Skopolos where V. germanica occasionally overwinters, “populations of social wasps in some years reach such numbers that they pose a definite risk to the future of tourism and agriculture on the island” (Cainadas, 1987). Social wasps have also been reported as major problems at holiday resorts in the United States where swimming, boating and fishing (especially cleaning fish outdoors) proved impossible to endure and guests departed resorts prematurely (Akre, 1983). At Miami University, Florida, US, the authorities were forced to re-schedule most of their outdoor activities in 1975 due to plagues of V. germanica (Akre, 1983). In Tasmania, tourists visiting the Gordon River on cruise boats have refused to leave the safety of the toilets for fear of wasps in the area and in southern Tasmania, several popular camping sites are uninhabitable at times because of V. germanica numbers (Gullan, 1999).

Environmental Impact

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V. germanica requires large quantities of protein to rear its brood. Sources of protein are typically insects and carrion but they are also predatory on nestling birds (Moller, 1990). The quantity of arthropods collected by foraging wasps is remarkable, with a recorded maximum annual collection of 99kg of prey by a single, overwintered nest in New Zealand (Harris, 1996). Prey was mainly flies together with honey bees, moths and spiders and bird remains. This weight of prey is approximately equivalent to 3.5 million blowflies. Where the density of nests is high, wasps can destroy virtually all insect life in a local area with the subsequent disappearance of insectivorous birds (Barr et al., 1996). In parts of Tasmania where the wasps had collected all other insect life, they were observed preying cannibalistically on each other (Spradbery, 1988).

In the fragile ecosystems of southwestern Tasmania and other parts of the world where V. germanica is prevalent in wilderness areas, the loss of insect pollinators of rare or endangered flora cannot be over-emphasized.

In New Zealand, V. germanica wasps (and the sympatric V. vulgaris) collect large quantities of the honeydew produced by scale insects in the Nothofagus beech forests. This carbohydrate resource is also the main food source for many native arthropods and birds and, where foraging by wasps leads to a depletion of this resource, it adversely affects the breeding success of birds such as the endangered kaka (Beggs and Wilson, 1991).

Social Impact

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Large numbers of V. germanica workers attracted to food sources can affect the quality of life and render outdoor socializing an unpleasant experience (Spradbery, 1988). Schools in parts of Australia have been closed or eating outdoors banned because of wasp activity, while popular picnic and barbeque spots have been closed to the public due to high wasp densities. Sharing food outdoors in suburbia with family and friends when wasps are abundant can be a health hazard for many Australians and New Zealanders. While the nuisance value of the ‘wasp season’ in Europe may last a few weeks at most, in Australasia it continues for months and, with the propensity for this species to successfully overwinter, even all year round in some coastal regions.

Risk and Impact Factors

Top of page Invasiveness
  • Proved invasive outside its native range
  • Has a broad native range
  • Abundant in its native range
  • Highly adaptable to different environments
  • Is a habitat generalist
  • Tolerates, or benefits from, cultivation, browsing pressure, mutilation, fire etc
  • Pioneering in disturbed areas
  • Tolerant of shade
  • Capable of securing and ingesting a wide range of food
  • Highly mobile locally
  • Benefits from human association (i.e. it is a human commensal)
  • Long lived
  • Fast growing
  • Has high reproductive potential
Impact outcomes
  • Altered trophic level
  • Negatively impacts agriculture
  • Negatively impacts human health
  • Negatively impacts tourism
  • Reduced amenity values
  • Reduced native biodiversity
  • Threat to/ loss of native species
Impact mechanisms
  • Antagonistic (micro-organisms)
  • Induces hypersensitivity
  • Predation
Likelihood of entry/control
  • Highly likely to be transported internationally accidentally
  • Difficult/costly to control

Uses

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Species of Vespula and Vespa are used for human consumption in parts of southern Honshu, Japan, where the wasp pupae are an especially sought-after and tasty food which is rich in protein. Young Vespula nests are collected in early summer by enterprising wasp farmers and re-located to favourable sites such as greenhouses where they receive supplementary feeding and protection. When mature, the brood-filled combs are sold at local food markets. It is unlikely that such utilization would reduce the impact of V. germanica but the concept has some pecuniary potential for an enterprising entomocentric entrepreneur.

Similarities to Other Species/Conditions

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V. germanica is similar in appearance to the related Vespula vulgaris, but with some differences in markings. V. germanica has black dots in the yellow bands on the abdomen, and three black dots on the face, but V. vulgaris does not (OzAnimals, 2012).

Prevention and Control

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Prevention

To prevent the movement of hibernating queens of V. germanica would require an impossibly efficient quarantine operation and cannot be considered a feasible option. Killing hibernating or post-hibernation queens to prevent them from establishing nests in the spring is unlikely to have much impact. Indeed, in Cyprus many years ago the government offered a bounty for queen wasps in the spring and after a programme which cost dearly, that year was the worst ‘wasp year’ on record (Spradbery, 1973). Similarly in New Zealand, a small bounty was offered for each hibernating queen, but with a negligible impact on subsequent wasp densities (Thomas, 1960). Culling such queens may even improve the rate of successful establishment of nests by reducing competition for limited nest sites in the spring.

Public awareness programmes can offer the most cost effective approach to wasp control. Such programmes educate the public about the pest, raise awareness of its importance through local media sources, provide information on identification and offer advice on detection and control options for wasps and their nests through a dedicated hotline telephone number. Such activities can result in the widespread and early detection and treatment of nests, hopefully before the production of new queens in late summer/autumn.

Eradication

Once V. germanica has colonized an area and begun to consolidate, it is impossible to eradicate, especially if hibernating queens are constantly being re-imported from other infested centres through the movement of cargo. Eradication of V. germanica would entail discovering and treating every nest, including the very last one. Just one nest remaining undiscovered could produce several thousands of new queens to re-infest the area. The resources required for such an eradication programme would be intimidating. Eradication by genetic methods such as sterile male release does not appear to be a viable option for V. germanica.

Control

Biological control

Although biological control does not appear to be a potentially very useful option for V. germanica control, attempts have been made in Australasia by utilizing the ichneumonid parasitoid, Sphecophaga vesparum, which parasitizes Vespula brood in the nest. The initial releases were made in New Zealand with S. vesparum from the United States, later S. vesparumvesparum from Europe was used and also released in Victoria and Tasmania, Australia, and finally, S. vesparum burra from North America has more recently been introduced into New Zealand (Gullan, 1999). However, either the impact of the biotic agent has not been determined or the results have not been encouraging, despite its establishment. The other parasitic wasp species and the parasitic beetle, Metoecus paradoxus, which occur in colonies of Vespula species (Spradbery, 1973), do not seem to offer much encouragement for biological control programmes aimed at reducing the impact of V. germanica.

Nematode parasites have received scant attention as potential biological control agents. A fungus is being studied in New Zealand as a possible pathogen to be incorporated into baits (Beggs, 1999).

Chemical control

Chemical treatment of nests is the most widely used method for V. germanica control. Adult wasps that gain access to people’s homes can be treated with rapid-knockdown insecticide sprays.

Wasp nests are generally discovered by accident although attempts to locate nests by following returning foragers can be successful. When a nest has been located it is treated by the householder or professional pest controller using appropriate, registered chemicals, preferably insecticide powders. The operation should be conducted while wearing safety equipment such as a bee veil and gloves or after dark when flight activity has ceased. Nests built in cavity walls and roof spaces in houses are best left to professional pest management operators.

Wasp nests discovered on public land or in nature reserves would normally be the responsibility of the local council to treat.

Numerous commercial and homemade traps, including a number based on plastic bottles, with a variety of baits, from fruit juices to stale beer, are available for wasp control. While they may capture some wasps, they are unlikely to have any real impact on local wasp populations. In a study undertaken in the United States more than 500,000 wasps were captured in traps but there was no apparent reduction in the intensity of foraging in the trapping area (Reierson and Wagner, 1975). Because of the wide range of materials collected by wasps to feed themselves and their larvae, providing a lure that is competitively attractive to foraging workers limits the usefulness of such traps. In Central Europe diluted syrup was effective in attracting V. germanica and other Vespula species to traps (Dvorak and Landolt, 2006). In the United States there is ongoing research into chemically defined lures for social wasps, often based on the break-down products of fermentation such as isobutanol, mixed with acetic acid (Landolt et al., 2000). Poison baiting has considerable potential in reducing wasp populations, especially in areas where locating wasp nests is particularly difficult. The bait must be attractive to wasps but not to beneficial insects such a honey bees. Most wasp baits that have been evaluated in New Zealand, Australia and South America have been of animal protein. Among products that have been shown to be effective are tinned fish such as sardines (Spurr, 1993), chicken mince (Conolly et al., 2004) and kangaroo meat mince (R Bashford, Forestry Tasmania, Australia, personal communication, 2008).

A bait station requires that foraging wasps collect the bait which incorporates an insecticide, fly back to the nest and distribute this to the occupants before succumbing to the poison. Thus, the insecticide must be either slow acting or encapsulated. The poisons used successfully in baiting programmes to control V. germanica include sodium monofluoroacetate or compound 1080 (Spurr, 1991), sulfluramid (Spurr, 1993) and granular fipronil (Beggs, 1999; Conolly et al., 2004).  

Although the results of baiting programmes have demonstrated their efficacy (Conolly et al., 2004), when colonies are killed off and populations decline, wasps from adjacent areas can quickly re-invade the cleared areas (Beggs et al., 1998).

References

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10/06/08 Original text by:

Philip Spradbery, XCS Consulting, Canberra, Australian Capital Territory, Australia

Libor Dvorak, Municipal Museum Marianske Lazne, Goethovo namesti 11, 35301 Marianske Lazne, Czech Republic

 

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