Invasive Species Compendium

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Aethina tumida

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Datasheet

Aethina tumida

Summary

  • Last modified
  • 16 November 2018
  • Datasheet Type(s)
  • Invasive Species
  • Vector of Animal Disease
  • Natural Enemy
  • Preferred Scientific Name
  • Aethina tumida
  • Taxonomic Tree
  • Domain: Eukaryota
  •   Kingdom: Metazoa
  •     Phylum: Arthropoda
  •       Subphylum: Uniramia
  •         Class: Insecta
  • Summary of Invasiveness
  • The small hive beetle, Aethina tumida, is a scavenger and parasite of honeybee colonies. Adult beetles and larvae feed on honeybee larvae, pollen, honey and brood. There are some reports of infestation of ...

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Pictures

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PictureTitleCaptionCopyright
Small hive beetle (Aethina tumida); dorsal view of adult, museum set specimen. Florida, USA.
TitleAdult
CaptionSmall hive beetle (Aethina tumida); dorsal view of adult, museum set specimen. Florida, USA.
Copyright©Jeffrey W. Lotz/Florida Dept. of Agriculture & Consumer Services/Bugwood.org - CC BY 3.0 US
Small hive beetle (Aethina tumida); dorsal view of adult, museum set specimen. Florida, USA.
AdultSmall hive beetle (Aethina tumida); dorsal view of adult, museum set specimen. Florida, USA.©Jeffrey W. Lotz/Florida Dept. of Agriculture & Consumer Services/Bugwood.org - CC BY 3.0 US
Small hive beetle (Aethina tumida); adults on comb. USA.
TitleAdults on comb
CaptionSmall hive beetle (Aethina tumida); adults on comb. USA.
Copyright©Jessica Lawrence/Eurofins Agroscience Services/Bugwood.org - CC BY 3.0 US
Small hive beetle (Aethina tumida); adults on comb. USA.
Adults on combSmall hive beetle (Aethina tumida); adults on comb. USA.©Jessica Lawrence/Eurofins Agroscience Services/Bugwood.org - CC BY 3.0 US
Small hive beetle, (Aethina tumida) larvae on a comb of honey; Moultrie, Georgia, United State. 01 September, 2004
TitleLarvae on comb
CaptionSmall hive beetle, (Aethina tumida) larvae on a comb of honey; Moultrie, Georgia, United State. 01 September, 2004
Copyright©James D. Ellis/University of Florida/Bugwood.org - CC BY 3.0 US
Small hive beetle, (Aethina tumida) larvae on a comb of honey; Moultrie, Georgia, United State. 01 September, 2004
Larvae on combSmall hive beetle, (Aethina tumida) larvae on a comb of honey; Moultrie, Georgia, United State. 01 September, 2004©James D. Ellis/University of Florida/Bugwood.org - CC BY 3.0 US
Aethina tumida (small hive beetle); dissected larvae releasing entomopathogenic nematodes.
TitleDissected larvae releasing entomopathogenic nematodes
CaptionAethina tumida (small hive beetle); dissected larvae releasing entomopathogenic nematodes.
Copyright©Andrew GS Cuthbertson/Crown Copyright
Aethina tumida (small hive beetle); dissected larvae releasing entomopathogenic nematodes.
Dissected larvae releasing entomopathogenic nematodesAethina tumida (small hive beetle); dissected larvae releasing entomopathogenic nematodes.©Andrew GS Cuthbertson/Crown Copyright

Identity

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Preferred Scientific Name

  • Aethina tumida

International Common Names

  • English: Small hive beetle

Summary of Invasiveness

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The small hive beetle, Aethina tumida, is a scavenger and parasite of honeybee colonies. Adult beetles and larvae feed on honeybee larvae, pollen, honey and brood. There are some reports of infestation of Bombus species. A native of Africa, A. tumida has been introduced to the USA, Canada, Mexico, Jamaica, Australia and Italy, and reported but not substantiated in Egypt. Although it is considered to be a minor pest in Africa, it is a major problem in areas where it has been introduced. The adult beetles are able to fly several kilometres, aiding the rapid spread of infestation. Heavy infestations may result in desertion of the hive by bees. Insecticides are used to treat infestations, but residues in honey may result (OIE, undated).

Small hive beetle infestation is on the list of diseases notifiable to the World Organisation for Animal Health (OIE).

Taxonomic Tree

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  • Domain: Eukaryota
  •     Kingdom: Metazoa
  •         Phylum: Arthropoda
  •             Subphylum: Uniramia
  •                 Class: Insecta
  •                     Order: Coleoptera
  •                         Family: Nitidulidae
  •                             Genus: Aethina
  •                                 Species: Aethina tumida

Distribution

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A. tumida is native to sub-Saharan Africa; it has been introduced to the USA, Canada, Mexico, Jamaica, Australia and Italy, and reported but not substantiated in Egypt (Thomas, 1998; OIE, 2013; FERA, 2010; Delaplane, 1998; Cuthbertson et al., 2013b; Mutinelli et al., 2014). The Distribution table contains records for all countries where it has been introduced, but in the native range only for those countries where records are readily available.

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

Africa

EgyptAbsent, unreliable recordIntroducedFERA, 2010Reported in 2000 but not substantiated.
KenyaPresentNativeTorto et al., 2010
NigeriaPresentNativeAkinwande et al., 2013
South AfricaPresentNativeSpiewok et al., 2007
SudanPresentNativeEl-Niweiri et al., 2008
UgandaPresentNativeRoberts, 1971
ZimbabwePresentNativePapadopoulo, 1964

North America

CanadaPresentPresent based on regional distribution.
-AlbertaAbsent, formerly presentIntroducedKozak, 2010Reported in 2006; control measures taken and species did not become established.
-ManitobaAbsent, formerly presentIntroducedFERA, 2010; Kozak, 2010Reported in 2002 and 2006; control measures taken and species did not become established.
-OntarioLocalisedIntroducedKozak, 2010; Kozak, 2012Detected in 2010. Restricted to a quarantine area in southern Ontario.
-QuebecPresentIntroducedFERA, 2010Detected in 2008. As of 2010, not yet well established.
MexicoPresentIntroducedFERA, 2010Confirmed in 2007.
USAWidespreadIntroduced Invasive FERA, 2010First reported in Florida in 1988; now very widespread.
-FloridaPresentIntroduced Invasive FERA, 2010First reported 1998.
-HawaiiPresentIntroduced Invasive FERA, 2010First reported 2010.

Central America and Caribbean

JamaicaPresentIntroducedFERA, 2010Confirmed in 2005.

Europe

ItalyLocalisedIntroduced2014Mutinelli et al., 2014Calabria (plus one migratory apiary in Sicily)
PortugalAbsent, intercepted onlyIntroducedMurilhas, 2005; FERA, 2010Intercepted and eradicated in a consignment of bees from Texas in 2004.

Oceania

AustraliaPresentPresent based on regional distribution.
-New South WalesPresentIntroduced Invasive FERA, 2010First found in 2002.
-QueenslandPresentIntroduced Invasive FERA, 2010First found in 2002.
-VictoriaPresent, few occurrencesIntroducedAnnand, 2008
-Western AustraliaLocalisedIntroducedFERA, 2010

History of Introduction and Spread

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A. tumida is native to southern Africa. The first record of this beetle in the western hemisphere was in May 1998, when it was found in a commercial apiary in St. Lucie County, Florida, USA. It was soon detected in Indian River and 4 other counties, where it caused significant damage to honey bee colonies (Thomas, 1998). Beetle specimens were found in beehives near Atlanta, Georgia and confirmed as A. tumida in July 1998 (Delaplane, 1998). The intra- or interstate movement of honey bee colonies was prohibited until they could be inspected, to prevent further spread (Thomas, 1998).

Nevertheless, the species spread rapidly and is now very widespread throughout the USA, including Hawaii where it was first recorded in 2010. It has also reached parts of Canada, as well as Mexico and Jamaica (FERA, 2010).

It was first found in Queensland and New South Wales, Australia, in October 2002, and is now considered endemic in those states, with some beetles recorded in Victoria and Western Australia (FERA, 2010).

It was first recorded in southern Italy in September 2014 (Mutinelli et al., 2014).

Introductions

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Introduced toIntroduced fromYearReasonIntroduced byEstablished in wild throughReferencesNotes
Natural reproductionContinuous restocking
Florida Africa 1998 Yes Thomas (1998)
Georgia 1998 Yes Delaplane (1998)
Manitoba USA   No FERA (2010); Kozak (2010) With beeswax

Risk of Introduction

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Adults and larvae of the small hive beetle are found in active hives and on stored bee equipment. Routine tasks carried out by apiarists can aid the spread of beetles or provide room for them to become established away from the cluster of protective bees (Delaplane, 1998).

A study by Spiewok at al. (2008), to investigate the dispersal of small hive beetles, revealed that there was a 95% re-infestation rate of beetle-free colonies, among 10 infested apiaries in South Africa, Australia and the USA. Apiaries in Maryland remained uninfested, but those in Australia were shown to have high infestation numbers. Factors affecting infestation of new apiaries seem to be apiary density, beetle population levels, and ongoing mass beetle reproduction. Apiaries in forested habitats showed higher infestation levels, which was thought to be due to the presence of wild/feral colonies.

The movement of bees, equipment and supplies worldwide has assisted in spreading bee diseases to all areas where bees are raised (OIE, undated).

FERA (2010), assessing the risk to the UK, judge that there is a significant risk that the species could be introduced to Europe (including the UK).

Pathogen Characteristics

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Adult A. tumida are dark brown to almost black, broad, flattened, measuring approximately 5.7 mm long and 3.2 mm wide. Just after pupation, the adults are red, but quickly become blackish. They move quickly and are difficult to pick up (Delaplane, 1998).

The larvae are elongate, whitish and with small spines in rows along the back (Cuthbertson et al., 2013a). They are similar in appearance to wax moth larvae; however, the legs of small hive beetles are larger, more obvious and positioned near to the head. The larvae pupate in the soil outside the hive. The pupae are whitish brown (Delaplane, 1998).

Under its native conditions of southern Africa, A. tumida requires 38-81 days to develop from egg to adult, and can produce up to 5 generations per year under suitable conditions (Delaplane, 1998; Cuthbertson et al., 2008). The adults are attracted to bee colonies to reproduce; however, they can survive and reproduce in other natural environments, feeding on other resources such as fruit, which makes the species very difficult to eradicate (OIE, 2012).

Habitat List

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CategorySub-CategoryHabitatPresenceStatus
Terrestrial

Host Animals

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Animal nameContextLife stageSystem
Apis melliferaDomesticated host, Wild host
Bombus impatiensDomesticated host
Bombus terrestris (bumble bee)Experimental settings

Notes on Natural Enemies

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An article published in Bee World in 2005 suggested that pseudoscorpions (Ellingsenius fulleri and E. indicus) may protect bees from pests such as the small hive beetle (Donovan and Paul, 2005).

While investigating the use of bottom boards to monitor hive beetles in bee colonies, Torto et al. (2010) reported that the ant Pheidole megacephala was a key predator of larvae at a site in Kenya.

Under laboratory conditions entomopathogenic nematodes have shown excellent control potential against pupating larvae in the ground (Cuthbertson et al., 2012).

Pathway Causes

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CauseNotesLong DistanceLocalReferences
Animal productionOn bees Yes Yes OIE, 2012
HitchhikerOn apiculture products or equipment Yes Yes OIE, 2012
Self-propelled Yes OIE, 2012

Pathway Vectors

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VectorNotesLong DistanceLocalReferences
Host and vector organisms Yes Yes OIE, 2012
Machinery and equipment Yes OIE, 2012

Economic Impact

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Infestations of A. tumida can cause considerable financial loss to beekeepers (Delaplane, 1998). Time and labour to detect and control the beetles, and losses in honey production and pollination, are the main economic losses suffered by the beekeeping industry (Calderón et al., 2006).

Adult A. tumida eat bee eggs and the larvae consume brood, pollen and honey and heavily damage wax comb (Calderón et al., 2006). Beekeepers in Florida have suffered considerable damage and colony loss, where beetle larvae have tunnelled through combs, killed bee brood and ruined combs; abandoning of combs and entire colonies by bees once they have become infested has been reported in Florida (Delaplane, 1998).

The beetles defecate in the honey and cause it to ferment; this produces a frothy mess in supers and honey houses. Honey that is contaminated is no longer saleable and is also unpalatable to bees so it cannot be used as bee feed (Delaplane, 1998).

Within two years of the discovery of A. tumida in the USA, at least 20,000 bee colonies had been destroyed by it, costing many millions of dollars. It has had a serious detrimental effect on the beekeeping industry in Australia as well (FERA, 2010).

Environmental Impact

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Impact on Habitats

A decline in bee numbers has been attributed to various bee pests and diseases, such as A. tumida. Bee decline will have a significantly negative affect on pollination in habitats where plants rely on bees. The value of pollination is estimated to exceed the value of products from beehives many-fold (Delaplane and Mayer, 2000).

Impact on Biodiversity

A decline in native bees, such as A. mellifera, due to infestation by small hive beetles will have a negative impact on bee biodiversity (Cuthbertson and Brown, 2009).

Risk and Impact Factors

Top of page Invasiveness
  • Proved invasive outside its native range
  • Highly mobile locally
  • Has high reproductive potential
Impact outcomes
  • Host damage
  • Negatively impacts agriculture
  • Negatively impacts livelihoods
  • Reduced native biodiversity
  • Damages animal/plant products
Impact mechanisms
  • Pest and disease transmission
  • Predation
Likelihood of entry/control
  • Highly likely to be transported internationally accidentally

References

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Akinwande KL; Badejo MA; Ogbogu SS, 2013. Challenges associated with the honey bee (Apis mellifera adansonii) colonies establishment in south western Nigeria. African Journal of Food, Agriculture, Nutrition and Development, 13(2):7467-7484. http://www.ajfand.net/Volume13/No2/Akinwande12175.pdf

Annand N, 2008. Small hive beetle management options. Orange, New South Wales, Australia: New South Wales Department of Primary Industries, 7 pp. http://www.dpi.nsw.gov.au/__data/assets/pdf_file/0010/220240/small-hive-beetle-management-options.pdf

Buchholz S; Merkel K; Spiewok S; Pettis JS; Duncan M; Spooner-Hart R; Ulrichs C; Ritter W; Neumann P, 2009. Alternative control of Aethina tumida Murray (Coleoptera: Nitidulidae) with lime and diatomaceous earth. Apidologie, 40(5):535-548. http://www.edpsciences.org/journal/index.cfm?edpsname=apido

Cabanillas HE; Elzen PJ, 2006. Infectivity of entomopathogenic nematodes (Steinernematidae and Heterorhabditidae) against the small hive beetle Aethina tumida (Coleoptera: Nitidulidae). Journal of Apicultural Research, 45(1):49-50.

Calderón RA; Arce H; Ramírez JF, 2006. The small hive beetle Aethina tumida Murray, an important problem affecting honey bees. (El pequeño escarabajo de la colmena Aethina tumida Murray, un problema importante que afecta las abejas melíferas.) Ciencias Veterinarias (Heredia), 24(1):49-55.

Cuthbertson AGS; Brown MA, 2009. Issues affecting British honey bee biodiversity and the need for conservation of this important ecological component. International Journal of Environmental Science and Technology, 6(4):695-699. http://www.ceers.org/ijest

Cuthbertson AGS; Mathers JJ; Blackburn LF; Marris G, 2013. Lifecycle of the Small hive beetle, Aethina tumida. Bee Craft, 95(5):32-33.

Cuthbertson AGS; Mathers JJ; Blackburn LF; Powell ME; Marris G; Pietravalle S; Brown MA; Budge GE, 2012. Screening commercially available entomopathogenic biocontrol agents for the control of Aethina tumida (Coleoptera: Nitidulidae) in the UK. Insects, 3(3):719-726. http://www.mdpi.com/2075-4450/3/3/719

Cuthbertson AGS; Mathers JJ; Blackburn LF; Wakefield ME; Collins LE; Luo WeiQi; Brown MA, 2008. Maintaining Aethina tumida (Coleoptera: Nitidulidae) under quarantine laboratory conditions in the UK and preliminary observations on its behaviour. Journal of Apicultural Research, 47(3):192-193. http://www.ibra.org.uk

Cuthbertson AGS; Wakefield ME; Powell ME; Marris G; Anderson H; Budge GE; Mathers JJ; Blackburn LF; Brown MA, 2013. The small hive beetle Aethina tumida: a review of its biology and control measures. Current Zoology, 59(5):644-653. http://www.actazool.org/temp/%7B02A6F36B-3FC8-4D97-AFFA-33FBBA383F27%7D.pdf

Delaplane KS, 1998. The small hive beetle, Aethina tumida. A new beekeeping pest. Tifton, Georgia, USA: University of Georgia, 2 pp. http://www.bugwood.org/factsheets/small_hive_beetle.html

Delaplane KS; Mayer DF, 2000. Crop pollination by bees. Wallingford, UK: CABI Publishing, xv + 344 pp. http://www.cabi.org/CABeBooks/default.aspx?site=107&page=45&LoadModule=PDFHier&BookID=26

Donovan BJ; Paul F, 2005. Pseudoscorpions: the forgotten beneficials inside beehives and their potential for management for control of varroa and other arthropod pests. Bee World, 86(4):83-87.

Ellis JD; Delaplane KS, 2007. The effects of three acaricides on the developmental biology of small hive beetles (Aethina tumida). Journal of Apicultural Research, 46(4):256-259.

Ellis JD; Spiewok S; Delaplane KS; Buchholz S; Neumann P; Tedders WL, 2010. Susceptibility of Aethina tumida (Coleoptera: Nitidulidae) larvae and pupae to entomopathogenic nematodes. Journal of Economic Entomology, 103(1):1-9. http://esa.publisher.ingentaconnect.com/content/esa/jee/2010/00000103/00000001/art00001

El-Niweiri MAA; El-Sarrag MS; Neumann P, 2008. Filling the Sudan gap: the northernmost natural distribution limit of small hive beetles. Journal of Apicultural Research, 47(3):184-185. http://www.ibra.org.uk

Eyer M; Chen YP; Schäfer MO; Pettis J; Neumann P, 2009. Small hive beetle, Aethina tumida, as a potential biological vector of honeybee viruses. Apidologie, 40(4):419-428. http://www.edpsciences.org/journal/index.cfm?edpsname=apido

FERA (Food and Environment Research Agency), 2010. The Small Hive Beetle: a serious threat to European apiculture. Sand Hutton, UK: Food and Environment Research Agency, 23 pp. https://secure.fera.defra.gov.uk/beebase/downloadDocument.cfm?id=17

Kokkinis MI, 2005. Small hive beetle [SHB], Aethina tumida. Deltion tes Ellenikes Kteniatrikes Etaireias = Journal of the Hellenic Veterinary Medical Society, 56(3):228-238.

Kozak P, 2010. Small Hive Beetle. Guelph, Ontario, Canada: Ontario Ministry of Agriculture, Food and Rural Affairs, 4 pp. http://www.omafra.gov.on.ca/english/food/inspection/bees/info-shb.pdf

Kozak P, 2012. Biosecurity practices for preventing the spread of Small Hive Beetle. Guelph, Ontario, Canada: Ontario Ministry of Agriculture, Food and Rural Affairs, 4 pp. http://www.omafra.gov.on.ca/english/food/inspection/bees/biosecurity.pdf

Muerrle TM; Neumann P; Dames JF; Hepburn HR; Hill MP, 2006. Susceptibility of adult Aethina tumida (Coleoptera: Nitidulidae) to entomopathogenic fungi. Journal of Economic Entomology, 99(1):1-6. http://miranda.esa.catchword.org/vl=5478027/cl=18/nw=1/rpsv/cw/esa/00220493/v99n1/s1/p1

Murilhas AM, 2005. Aethina tumida arrives in Portugal. Will it be eradicated? EurBee Newsletter, 2:7-9.

Mutinelli F; Montarsi F; Federico G; Granato A; Ponti MA; Grandinetti G; Ferrè N; Franco S; Duquesne V; Rivière MP; Thiéry R; Henrkix P; Ribière-Chabert M; Chauzat MP, 2014. Detection of Aethina tumida Murray (Coleoptera: Nitidulidae.) in Italy: outbreaks and early reaction measures. Journal of Apicultural Research, 53(5):569-575. http://dx.doi.org/10.3896/IBRA.1.53.5.13

Neumann P; Hoffmann D, 2008. Small hive beetle diagnosis and control in naturally infested honeybee colonies using bottom board traps and CheckMite+strips. Journal of Pest Science, 81(1):43-48. http://www.springerlink.com/content/4830215644411151/fulltext.pdf

OIE (Office International des Epizooties), undated. Diseases of bees. Paris, France: Office International des Epizooties, 6 pp. http://www.oie.int/fileadmin/Home/eng/Media_Center/docs/pdf/Disease_cards/BEES-EN.pdf

OIE (World Organisation for Animal Health), 2012. Terrestrial Animal Health Code, edition 21. Paris, France: Office International des Epizooties. http://www.oie.int/international-standard-setting/terrestrial-code/access-online/

OIE (World Organisation for Animal Health), 2013. Manual of Diagnostic Tests and Vaccines for Terrestrial Animals. Paris, France: World Organisation for Animal Health. http://www.oie.int/en/international-standard-setting/terrestrial-manual/access-online/

Papadopoulo P, 1964. Enemies of bees (1). Rhodesia Agricultural Journal, 61(6):114-115 pp.

Roberts E, 1971. A survey of beekeeping in Uganda. Bee World, 52(2):57-67.

Schäfer MO; Pettis JS; Ritter W; Neumann P, 2008. A scientific note on quantitative diagnosis of small hive beetles, Aethina tumida, in the field. Apidologie, 39(5):564-565. http://www.edpsciences.org/journal/index.cfm?edpsname=apido

Schäfer MO; Ritter W; Pettis J; Neumann P, 2010. Small hive beetles, Aethina tumida, are vectors of Paenibacillus larvae. Apidologie, 41(1):14-20. http://www.edpsciences.org/journal/index.cfm?edpsname=apido

Schäfer MO; Ritter W; Pettis JS; Teal PEA; Neumann P, 2009. Effects of organic acid treatments on small hive beetles, Aethina tumida, and the associated yeast Kodamaea ohmeri. Journal of Pest Science, 82(3):283-287. http://www.springerlink.com/content/b757803l567h7618/?p=44ff5f5bd86346bf8086df5eb7625e26&pi=10

Spiewok S; Duncan M; Spooner-Hart R; Pettis JS; Neumann P, 2008. Small hive beetle, Aethina tumida, populations II: dispersal of small hive beetles. Apidologie, 39(6):683-693. http://www.edpsciences.org/journal/index.cfm?edpsname=apido

Spiewok S; Pettis JS; Duncan M; Spooner-Hart R; Westervelt D; Neumann P, 2007. Small hive beetle, Aethina tumida, populations I: Infestation levels of honeybee colonies, apiaries and regions. Apidologie, 38(6):595-605. http://www.edpsciences.org/journal/index.cfm?edpsname=apido

Stedman M, 2006. Small Hive Beetle (SHB): Aethina tumida Murray (Coleoptera: Nititulidae). Glenside, South Australia, Australia: Primary Industries and Resources South Australia, 15 pp. http://www.pir.sa.gov.au/__data/assets/pdf_file/0015/41262/apiary_shb_fact_sheet_2006.pdf

Thomas MC, 1998. Florida pest alert - the small hive beetle. American Bee Journal, 138(8):565.

Torto B; Fombong AT; Arbogast RT; Teal PEA, 2010. Monitoring Aethina tumida (Coleoptera: Nitidulidae) with baited bottom board traps: occurrence and seasonal abundance in honey bee colonies in Kenya. Environmental Entomology, 39(6):1731-1736. http://esa.publisher.ingentaconnect.com/content/esa/envent/2010/00000039/00000006/art00005

Ward L; Brown M; Neumann P; Wilkins S; Pettis J; Boonham N, 2007. A DNA method for screening hive debris for the presence of small hive beetle (Aethina tumida). Apidologie, 38(3):272-280. http://www.edpsciences.org/journal/index.cfm?edpsname=apido

Links to Websites

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WebsiteURLComment
GISD/IASPMR: Invasive Alien Species Pathway Management Resource and DAISIE European Invasive Alien Species Gatewayhttps://doi.org/10.5061/dryad.m93f6Data source for updated system data added to species habitat list.
Global register of Introduced and Invasive species (GRIIS)http://griis.org/Data source for updated system data added to species habitat list.

Organizations

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World: IBRA, International Bee Research Association, Unit 6, Centre Court, Main Avenue, Treforest, RCT, CF37 5YR, UK, www.ibra.org.uk

World: OIE (World Organisation for Animal Health), 12, rue de Prony, 75017 Paris, France, http://www.oie.int/

UK: British Beekeepers’ Association, National Beekeeping Centre, Stoneleigh Park, Stoneleigh, Warwickshire, CV8 2LG, UK, www.britishbeekeepers.com

Contributors

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23/03/2012: Original text by:

Dr Claire Beverley, CABI, Nosworthy Way, Wallingford. OX10 8DE.

Distribution Maps

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