Bombus terrestris (bumble bee)
- Summary of Invasiveness
- Taxonomic Tree
- Notes on Taxonomy and Nomenclature
- Distribution Table
- History of Introduction and Spread
- Risk of Introduction
- Habitat List
- Hosts/Species Affected
- Biology and Ecology
- Means of Movement and Dispersal
- Pathway Causes
- Pathway Vectors
- Impact Summary
- Economic Impact
- Environmental Impact
- Risk and Impact Factors
- Uses List
- Detection and Inspection
- Similarities to Other Species/Conditions
- Prevention and Control
- Links to Websites
- Distribution Maps
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PicturesTop of page
IdentityTop of page
Preferred Scientific Name
- Bombus terrestris (Linnaeus, 1758)
Preferred Common Name
- bumble bee
Other Scientific Names
- Apis terrestris Linnaeus, 1758
- Bombus audax Harris, 1776
- Bombus canariensis Pérez, 1895
- Bombus maderensis Erlandsson, 1979
- Bombus terreftris Linnaeus, 1758
International Common Names
- English: large earth bumble bee
Local Common Names
- France: le bourdon terrestre
- Germany: Dunkle Erdhummel
- UK: buff tailed bumble bee; buff tailed bumblebee; buff-tailed humble-bee; bumblebee
Summary of InvasivenessTop of page
This species is native to the western Palaearctic region (central and southern Europe, North Africa, Madeira and the Canary Islands, east to Afghanistan). It is found in all the countries around the Mediterranean Sea except Egypt (Rasmont et al., 2008) and extends to the north up to the latitude of Helsinki and east to Altai (Pekkarinen and Kaarnam, 1994). Bumble bees are highly valued pollinators worldwide and substantially add to the value of crop production (Goulson, 2003a). B. terrestris has been introduced into New Zealand (e.g. Gurr, 1957; MacFarlane and Gurr, 1995), Tasmania (Cardale, 1993; Stout and Goulson, 2000), Brazil (Thorp, 2003), Chile (Torretta et al., 2006), Mexico (Stout and Goulson, 2000; Winter et al., 2006), and Japan (Washitani, 1998; Inoue et al., 2008). It appears that it was also introduced into mainland Australia (New South Wales) without persisting (W. Froggatt in Franklin, 1913). Recently it has spread from Chile to Argentina (Torretta et al., 2006).
B. terrestris is not considered invasive in its ‘native’ range. Following recent establishment, B. terrestris is considered by some to be invasive in Japan and Tasmania, Australia (which has no native bumble bees) (Winter et al., 2006; Hingston, 2006; Ings et al., 2006). Australia, USA and Canada are prohibiting the import of B. terrestris. China, South Africa, the Canary Islands and Norway, do not allow B. terrestris to be imported either (Velthuis and van Doorn, 2006; Winter et al., 2006); the Japanese government has included B. terrestris in its Invasive Alien Species Act (Velthuis and van Doorn, 2006). In New South Wales (Australia), the alteration of natural pollination dynamics caused by the presence of B. terrestris in other countries has prompted its listing as a “Key Threatening Process”, and in Victoria, Australia, it is listed as a “Potentially Threatening Process”, and B. terrestris importation is also prohibited in all states.
Taxonomic TreeTop of page
- Domain: Eukaryota
- Kingdom: Metazoa
- Phylum: Arthropoda
- Subphylum: Uniramia
- Class: Insecta
- Order: Hymenoptera
- Family: Apidae
- Genus: Bombus
- Species: Bombus terrestris
Notes on Taxonomy and NomenclatureTop of page
B. terrestris is the most intensely studied non-Apis bee. It is one of the most abundant and widespread bumble bee species in the western Palaearctic. It is a geographically variable species with a wide distribution in Europe, the near east and northern Africa, the Mediterranean Islands, Canary Islands and Madeira. It is now widely domesticated and large numbers of colonies from different subspecies are transported from country to country (Velthuis, 2002). This leads to more and more mixing between populations (Ings et al., 2005b).
DescriptionTop of page
Bumble bees are social insects (eusocial), and the vast majority have an annual life cycle. Mated queens emerge from hibernation in the spring, and attempt to found a nest in which they rear daughter workers. In optimum conditions, the number of workers may reach 300 to 500 in some species by mid-summer, when new queens and males are reared. These leave the and nest, mate and the new queens enter hibernation, while the rest of the population dies off. B. terrestris form annual colonies and new colonies are initiated each spring by mated solitary queens (Goulson, 2003b). In warmer areas, colonies may reach large sizes, with colonies of more than 1000 individuals being recorded in Tasmania (Buttermore, 1997).
DistributionTop of page
B. terrestris is widely distributed in the western Palaearctic region. Its distribution is typically Mediterranean, extending from the Canary Islands in the west, to the Altai in the East, and from the Anti-Atlas Mountains of Morocco in the south to southern Finland in the north (Rasmont, 1983; Estoup et al., 1996; Rasmont et al., 2008). B. terrestris is found to the latitude of Helsinki and east to the Altai, from ca. 30° to 60° N (Pekkarinen and Kaarnam, 1994). It is not found in Egypt and is absent from high alpine levels, the deserts and the arid, sub-desertic steppes (Rasmont et al., 2008). Within its wide distribution, there are important subspecific differences in morphological characters (e.g. coat colour and behaviour, learning performance), which underline the genetic differentiation among subspecies (Coppée et al., 2008).
Distribution TableTop of page
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/Region||Distribution||Last Reported||Origin||First Reported||Invasive||Reference||Notes|
|Armenia||Present||Native||Rasmont et al., 2008|
|Azerbaijan||Present||Native||Rasmont et al., 2008|
|Georgia (Republic of)||Present||Native||Rasmont et al., 2008|
|Iran||Present||Native||Rasmont et al., 2008||In the north|
|Japan||Present||Introduced||Invasive||Washitani and Matsumura, 1998; Inoue et al., 2008|
|Kazakhstan||Present||Native||Rasmont et al., 2008|
|Lebanon||Present||Native||Rasmont et al., 2008|
|Pakistan||Present||Native||Rasmont et al., 2008|
|Syria||Present||Native||Rasmont et al., 2008|
|Uzbekistan||Present||Native||Rasmont et al., 2008|
|Algeria||Present||Native||Rasmont et al., 2008|
|Libya||Present||Native||Rasmont et al., 2008|
|Morocco||Present||Native||Rasmont et al., 2008|
|-Canary Islands||Present||Introduced||Widmer et al., 1998|
|Mexico||Present||Introduced||Stout and Goulson, 2000; Flanders et al., 2003; Winter et al., 2006||Shipments into Jalisco in 1995 and 1996 for greenhouse tomato pollination|
|Argentina||Present||Introduced||Torretta et al., 2006|
|Brazil||Present||Introduced||Costa and Lordello, 1988; Thorp, 2003|
|Chile||Present||Introduced||Torretta et al., 2006||Introduced for pollination|
|Albania||Present||Native||Rasmont et al., 2008|
|Andorra||Present||Native||Rasmont et al., 2008|
|Austria||Present||Native||Rasmont et al., 2008|
|Belgium||Present||Native||Rasmont et al., 2008|
|Bulgaria||Present||Native||Estoup et al., 1996|
|Croatia||Present||Native||Rasmont et al., 2008|
|Cyprus||Present||Native||Rasmont et al., 2008|
|Czech Republic||Present||Native||Rasmont et al., 2008|
|Czechoslovakia (former)||Present||Native||Rasmont et al., 2008|
|Denmark||Present||Native||Rasmont et al., 2008|
|Estonia||Present||Native||Rasmont et al., 2008|
|Finland||Present||Pekkarinen and Kaarnam, 1994|
|France||Present||Native||Estoup et al., 1996|
|-Corsica||Present||Native||Estoup et al., 1996|
|Germany||Present||Native||Rasmont et al., 2008|
|Greece||Present||Native||Estoup et al., 1996|
|Hungary||Present||Native||Rasmont et al., 2008|
|Ireland||Present||Native||Rasmont et al., 2008|
|Italy||Present||Native||Estoup et al., 1996|
|Latvia||Present||Native||Rasmont et al., 2008|
|Liechtenstein||Present||Native||Rasmont et al., 2008|
|Lithuania||Present||Native||Rasmont et al., 2008|
|Luxembourg||Present||Native||Rasmont et al., 2008|
|Macedonia||Present||Native||Rasmont et al., 2008|
|Malta||Present||Native||Rasmont et al., 2008|
|Moldova||Present||Native||Rasmont et al., 2008|
|Monaco||Present||Native||Rasmont et al., 2008|
|Montenegro||Present||Native||Rasmont et al., 2008|
|Netherlands||Present||Native||Rasmont et al., 2008|
|Norway||Present||Native||Rasmont et al., 2008|
|Poland||Present||Native||Estoup et al., 1996|
|Portugal||Present||Native||Rasmont et al., 2008|
|-Madeira||Present||Introduced||Widmer et al., 1998||Could be a subspecies|
|Romania||Present||Native||Rasmont et al., 2008|
|Russian Federation||Present||Present based on regional distribution.|
|-Central Russia||Present||Native||Rasmont et al., 2008|
|-Southern Russia||Present||Native||Rasmont et al., 2008|
|San Marino||Present||Native||Rasmont et al., 2008|
|Serbia||Present||Native||Rasmont et al., 2008|
|Slovakia||Present||Native||Rasmont et al., 2008|
|Slovenia||Present||Native||Rasmont et al., 2008|
|Spain||Present||Native||Estoup et al., 1996|
|Sweden||Present||Native||Rasmont et al., 2008|
|Switzerland||Present||Native||Estoup et al., 1996|
|UK||Present||Native||Rasmont et al., 2008|
|Ukraine||Present||Native||Rasmont et al., 2008|
|Yugoslavia (former)||Present||Native||Rasmont et al., 2008|
|Yugoslavia (Serbia and Montenegro)||Present||Native||Rasmont et al., 2008|
|Australia||Present||Present based on regional distribution.|
|-Tasmania||Present||Cardale, 1993; Semmens et al., 1993; Stout and Goulson, 2000|
|New Zealand||Present||Introduced||Invasive||Hopkins, 1914; Gurr, 1957; MacFarlane and Gurr, 1995|
History of Introduction and SpreadTop of page
(Several subspecies of) B. terrestris has been introduced in many countries as a pollinator of greenhouses crops such as tomatoes and peppers. It has escaped from the greenhouses and formed feral populations in several countries where it has been exported to. In Israel, Dafni and Shmida (1996) reported that the numbers of honeybees and solitary bees declined during the last two decades on Mt. Carmel where the range of B.terrestris had expanded. In New Zealand [?], where it has been introduced and where conditions are similar to its native environment, B. terrestris has demonstrated a rapid rate of range expansion up to 90 kilometers per year (Hopkins, 1914). Everywhere it has been introduced, it is competing with native pollinators for nest sites and food resources, leading to a decrease of many native species (Hingston and McQuillan, 1998; 1999; Dafni and Schmida, 1996; Dafni, 1998; Hergstrom et al., 2002; Matsumura et al., 2004; Hingston, 2005; 2006; Inoue et al., 2007). In Britain, where the south-eastern European subspecies Bombus terrestris dalmitinus is imported each year in large quantities (Ings et al., 2006), there is a great risk of competition with the endemic Bombus terrestris audax. There is a high risk of introduction and transmission of parasites (Ings et al., 2005a) and out-competition, particularly because the introduced subspecies has superior foraging efficiency and reproductive rate (Ings et al., 2006).
IntroductionsTop of page
|Introduced to||Introduced from||Year||Reason||Introduced by||Established in wild through||References||Notes|
|Natural reproduction||Continuous restocking|
|Argentina||Chile||Intentional release (pathway cause)||Yes||Torretta et al. (2006)|
|Chile||Europe||1998||Intentional release (pathway cause)||Yes||Ruz (2002); Ruz and Herrera (2001)||Ssp. audax|
|Israel||Europe||1960s||Intentional release (pathway cause)||Yes||Dafni (1998)||Escaped from glasshouses|
|Japan||Europe||1991||Intentional release (pathway cause)||Yes||Inari et al. (2005); Inoue et al. (2007); Matsumura et al. (2004); Nagamitsu et al. (2007)||Ssp. terrestris and ssp. dalmatinus|
|Mexico||Europe||1995 & 1996||Intentional release (pathway cause)||Yes||Flanders et al. (2003)||Escaped from glasshouses|
|New Zealand||England and Wales||1875||Intentional release (pathway cause)||Yes||Dumbleton (1949); Goulson and Hanley (2004); Gurr (1964); Hopkins (1914); MacFarlane and Gurr (1995)||Subspecies Bombus audax|
|Tasmania||1992||Intentional release (pathway cause)||Yes||Buttermore (1997); Hingston et al. (2001); Semmens et al. (1993)||Feral population of ssp. audax discovered near Hobart|
Risk of IntroductionTop of page
There are other dangers associated with commercial trafficking of bumble bees. B. terrestris is now naturalised in several countries where it was first introduced for pollination of greenhouse crops. There are substantiated concerns regarding possible competitive effects of this species on native bumble bees and other native pollinators, which may in turn reduce the reproductive capacity of native plants (Hingston et al., 2002). Studies have shown that B. terrestris has four times the reproductive output of native species in Japan (Matsumara et al., 2004) and that there are considerable overlaps in forage use and timing of foraging (Inari et al., 2005). Some greenhouses are not secure and a recent study in Canada found that 73% of pollen carried by workers returning to commercial colonies originated from plants outside the greenhouse (Morandin et al., 2001; Whittington and Winston, 2004). B. terrestris was allowed into Mexico in 1995 and 1996 without the knowledge of the USA or Canadian regulatory agencies (Flanders et al., 2003). These bumble bees were infected with the microsporidian Nosema bombi, an internal parasite of bumble bees, prompting the destruction of the colonies intended for Jalisco, Mexico, and a retraction of import permits (Winter et al., 2006). The use of B. terrestris for greenhouse production in Mexico has since been replaced by Bombus impatiens, with up to 55,000 colonies sold per year (Velthuis and van Doorn, 2006), since importation by Koppert de Mexico in 2001 (Martinez Guzman, 2005).
HabitatTop of page
Bumble bees are among the most important pollinators of temperate zone plants because of their diverse body and proboscis sizes, ability to sonicate, dense pile, long activity periods, and adaptability to a wide variety of temperatures and climate types (Winter et al., 2006). Bumble bees are adapted to a diversity of climates and habitats, and are active even when light intensity is low. Due to their relatively large body sizes and dense pile, they are able to continue foraging even at temperatures as low as 10°C and as high as 32°C, with observations of Bombus terrestris dalmatinus at temperatures as low as 2°C (Winter et al., 2006). Their increased motility allows them to continue flower visits for most of the year, unlike honeybees, which are mostly inactive at temperatures below 16°C (Heinrich, 1979). Bumble bees can forage during adverse climatic conditions, even flying during light rain, visiting from 20-50 flowers per minute with high pollination efficiency.
Habitat ListTop of page
|Terrestrial – Managed||Cultivated / agricultural land||Principal habitat|
|Protected agriculture (e.g. glasshouse production)||Secondary/tolerated habitat|
|Managed forests, plantations and orchards||Principal habitat|
|Managed grasslands (grazing systems)||Secondary/tolerated habitat|
|Rail / roadsides||Principal habitat|
|Urban / peri-urban areas||Secondary/tolerated habitat|
|Terrestrial ‑ Natural / Semi-natural||Natural forests||Principal habitat|
|Natural grasslands||Principal habitat|
|Scrub / shrublands||Secondary/tolerated habitat|
|Coastal areas||Principal habitat|
Hosts/Species AffectedTop of page
Bumble bees are second only to honeybees as commercial pollinator insects. B. terrestris is a specialist pollinator of a number of European plant species, either because they require a bee of a certain size (e.g. foxglove, Digitalis spp.), weight (e.g. Scotch broom, Cytisus scoparius), or require buzz pollination to release pollen from poricidal anthers (e.g. many Solanaceae). This may facilitate an increase in the abundance and distribution of weed species in countries where European plants have been introduced. The presence of B. terrestris may also disrupt pollination of native plant species (Hingston and McQuillan, 1998).
Biology and EcologyTop of page
ClimateTop of page
|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)|
Means of Movement and DispersalTop of page
Evidence points to the fact that B. terrestris is able to naturalize easily, even with limited numbers of founding queens (Buttermore et al., 1998) and can spread quickly once introduced. In New Zealand [?], where B. terrestris has been introduced and conditions are similar to its native environment, the bumble bee has demonstrated a rapid rate of range expansion up to 90 kilometers per year (Hopkins, 1914). It maintains higher population densities than semi-social and solitary bees across a broad range of habitats and geographical regions, and is a generalist forager (Goulson, 2003a,b), allowing B. terrestris populations to potentially occupy a wide diversity of niches also used by different species of pollinator (Winter et al., 2006).
Impact SummaryTop of page
|Environment (generally)||Positive and negative|
Economic ImpactTop of page
The commercial production of bumble bees has developed into a thriving branch of agribusiness, generating an estimated yearly economic value of US$1.25 billion in pollination services in the USA alone (Ghazoul, 2005). The benefits to growers include reduced costs from not having to pollinate mechanically using shaker tables or by hand with electronic vibrating wands, ease in monitoring bumble bee activity, increased fruit yields, little or no need for pesticides, and improved fruit quality leading to higher sales prices (Velthuis and van Doorn, 2006). Bumble bees, notably several subspecies of Bombus terrestris, have been shipped throughout the world in vast numbers since the late 1980s, and around countries in Europe, where they are now competing with local native populations, leading some to extinction (Goka et al., 2001; Goulson, 2003a). Five species of bumble bees are used commercially, Bombus impatiens and Bombus occidentalis in Northern America, Bombus ignitus and Bombus lucorum in East Asia, and B. terrestris in Europe, South America, Asia, and New Zealand. Commercially-reared bumble bees can escape from greenhouses in relatively large proportions (Morandin et al., 2001) if growers are not cautious in preventing their accidental release. The use of non-native bumble bee species is a major ecological concern as they have been shown to escape and naturalize quickly (Ruz and Herrera, 2001; Hingston et al., 2002; Matsumura et al., 2004; Velthuis and van Doorn, 2006). Introduced pollinators could change the local flora by increasing the pollination rate of exotic weeds, causing increased costs in weed control programmes and threaten native pollinators by increasing competition (Goulson, 2003a,b). Furthermore, imported bumble bees carry parasites that potentially threaten native bumble bee and honeybee populations (Goka et al., 2001), potentially causing even more stress and losses to endemic species and populations.
Environmental ImpactTop of page
Worldwide, there are over 250 species of bumble bees (Michener, 2000), which form the monophyletic genus Bombus within the family Apidae (Hymenoptera) (Kawakita et al., 2004). Native bumble bees face threats from introduced parasites and diseases, including Nosema bombi, the microorganism Crithidia bombi, the tracheal mite Locustacarusbuchneri, and hymenopteran brood parasitoids such as Melittobia acasta and Melittobia chalybii, which can be difficult to detect when inspecting commercial colonies and may be spread from commercial to wild colonies by greenhouse production facilities (Winter et al., 2006).
In new environments, B. terrestris may threaten populstions of native pollinators not only by introducing new diseases, but also by competing for resources, or by disrupting genetic adaptations by hybridizing with native species.
The use of non-native bumble bee species is a major ecological concern (Velthuis and van Doorn, 2006). Introduced pollinators could change the local flora, e.g., by increasing the pollination rate of exotic weeds, and threaten native pollinators by increasing competition (Goulson, 2003a,b). These concerns are reinforced by results showing that even the introduction of non-native subspecies, such as Bombus terrestris sassaricus from Southern Europe to Western Europe, could lead to the displacement of native bees (Ings et al., 2005a).
Risk and Impact FactorsTop of page Invasiveness
- Invasive in its native range
- Has a broad native range
- Abundant in its native range
- Highly adaptable to different environments
- Is a habitat generalist
- Capable of securing and ingesting a wide range of food
- Highly mobile locally
- Fast growing
- Has propagules that can remain viable for more than one year
Uses ListTop of page
- Commercial pollinator
DiagnosisTop of page
Detection and InspectionTop of page
B. terrestris is a relatively large, primitively eusocial bee native to Europe. B. terrestris is generally more heavily built and hairier than the honeybee, Apis mellifera. B. terrestris queens are 30-35 mm long; workers are more variable in size, ranging from 8-22 mm long; and males are similar in size and appearance to large workers. B. terrestris are black with one yellow or ochre band across the front of the thorax and a second yellow or ochre band across the abdomen. The tip of the abdomen is either buff or white.
Similarities to Other Species/ConditionsTop of page
The world bumble bee (Bombus) fauna consists of approximately 250 known species; most of them are found in temperate parts of the northern hemisphere. Several distinct populations occur in the natural range of B. terrestris that show distinct colour and size variations (Chittka et al., 2004; Rasmont et al., 2008). Genetic studies have shown that several distinct subspecies exist, some of which can be considered as distinct species altogether. Examples include populations from Sardinia hosting Bombus terrestris sassaricus, the Canary Islands with Bombus terrestris canariensis, and the British Isles with Bombus terrestris audax (Estoup et al., 1996; Widmer et al., 1998). Rasmont et al. (2008) recognise nine morphological subspecies in B. terrestris: ssp. terrestris; ssp. africanus (North Africa), ssp. audax (British Isles), ssp. calabricus (Italy and Sicily), ssp. canariensis (Canary Islands), ssp. dalmatinus (Balkans, Urals and Asia), ssp. lusitanicus, ssp. sassaricus (Sardinia) and ssp. xanthopus (Corsica). For taxonomic differences and distribution within subspecies see the papers from Widmer et al. (1998); Rasmont et al. (2008); Coppée et al. (2008); the website of the Natural History Museum, London devoted to bumble bees (http://www.nhm.ac.uk/research-curation/research/projects/bombus/bo.html) and Professor David Goulson’s (Stirling University) webpage (http://www.sbes.stir.ac.uk/people/goulson/index.html).
Prevention and ControlTop of page
Invasive Species Management
There is no known control of B. terrestris once it has been introduced and/or has escaped in the natural environment. The only management strategy is through exclusion and restriction of importation into new areas or countries where it is not endemic and can impact on the local environment. Currently the best option is to promote the commercial use of native bumble bees within their countries of origin, rather than import the exotic species (B. terrestris) with the potential for damaging consequences.
ReferencesTop of page
Buttermore RE; Pomeroy N; Hobson W; Semmens T; Hart R, 1998. Assessment of the genetic base of Tasmanian bumble bees (Bombus terrestris) for development as pollination agents. Journal of Apicultural Research, 37(1):23-25.
CABI CEFAS CEH CSL IC UoG, 2005. UK non-native organism risk assessment scheme version 3. Bombus terrestris - subspecies not native to the UK e.g. B. terrestris terrestris, B. terrestris dalmatinus. York, UK: Fera. http://www.fera.defra.gov.uk/
Chittka L; Ings TC; Raine NE, 2004. Chance and adaptation in the evolution of island bumblebee behaviour. Population Ecology, 46(3):243-251. http://springerlink.metapress.com/(xtcmjrv3jppf10idpmhtr2yk)/app/home/contribution.asp?referrer=parent&backto=issue,4,11;journal,4,21;linkingpublicationresults,1:103139,1
Colla SR; Otterstatter MC; Gegear RJ; Thomson JD, 2006. Plight of the bumble bee: pathogen spillover from commercial to wild populations. Biological Conservation, 129(4):461-467. http://www.sciencedirect.com/science/journal/00063207
Coppée A; Terzo M; Valterova I; Rasmont P, 2008. Intraspecific variation of the cephalic labial gland secretions in Bombus terrestris (L.) (Hymenoptera: Apidae). Chemistry & Biodiversity, 5(12):2654-2661. http://www.chembiodiv.ch/
Costa JL da S; Lordello S, 1988. Role of insects in the dissemination of Fusarium disease of pineapple. (Papel da entomofauna na disseminação da fusariose do abacaxizeiro.) Fitopatologia Brasileira, 13(1):63-65.
Dafni A; Shmida A, 1996. The possible ecological implications of the invasion of Bombus terrestris (L.) (Apidae) at Mt. Carmel, Israel. In: The conservation of bees [ed. by Matheson, A.\Buchmann, S. L.\O'Toole, C.\Westrich, P.\Williams, I. H.]. London, UK: Academic Press for the Linnean Society of London and the International Bee Research Association, 183-200.
Dag A; Kammer Y, 2001. Comparison between the effectiveness of honey bee (Apis mellifera) and bumblebee (Bombus terrestris) as pollinators of greenhouse sweet pepper (Capsicum annuum). American Bee Journal, 141:447-448.
Estoup A; Solignac M; Cornuet J-M; Goudet J; Scholl A, 1996. Genetic differentiation of continental and island populations of Bombus terrestris (Hymenoptera: Apidae) in Europe. Molecular Ecology, 5(1):19-31.
Flanders RV; Wehling WF; Craghead AL, 2003. Laws and regulations on the import, movement, and release of bees in the United States. In: For Nonnative Crops, Whence Pollinators of the Future? [ed. by Strickler, K. \Cane, J. H.]. 99-111.
Hanley-Nicholls JR, 2008. A short study into the presence on foraging behaviour of bumblebees (Bombus spp.) on Skálanes nature reserve. A short study into the presence on foraging behaviour of bumblebees (Bombus spp.) on Skálanes nature reserve. Glasgow, UK: University of Glasgow, unpaginated. [University of Glasgow Iceland Expedition 2008.] http://www.slideshare.net/Skalanes/university-of-glasgow-iceland-expedition-2008-presentation
Harris M, 1776. An exposition of English insects, with curious observations and remarks, wherein each insect is particularly described; its parts and properties considered; the different sexes distinguished, and the natural history faithfully related. London, UK: Robson, viii+166 pp.
Hergstrom K; Buttermore R; Seeman O; McCorkell B, 2002. Environmental research on the impact of bumblebees in Australia and facilitation of national communication for/against further introduction. Horticulture Australia Project No: VG99033. Hobart, Australia: Horticulture Australia Ltd., unpaginated.
Hingston AB, 2005. Does the introduced bumblebee, Bombus terrestris (Apidae), prefer flowers of introduced or native plants in Australia? Australian Journal of Zoology, 53(1):29-34. http://www.publish.csiro.au/journals/ajz
Hingston AB, 2006. Is the exotic bumblebee Bombus terrestris really invading Tasmanian native vegetation? Journal of Insect Conservation, 10(3):289-293. http://www.springerlink.com/content/n0020114717x26n6/fulltext.pdf
Hingston AB; Marsden-Smedley J; Driscoll DA; Corbett S; Fenton J; Anderson R; Plowman C; Mowling F; Jenkin M; Matsui K; Bonham KJ; Ilowski M; McQuillan PB; Yaxley B; Reid T; Storey D; Poole L; Mallick SA; Fitzgerald N; Kirkpatrick JB; Febey J; Harwood AG; Michaels KF; Russell MJ; Black PG; Emmerson L(et al), 2002. Extent of invasion of Tasmanian native vegetation by the exotic bumblebee Bombus terrestris (Apoidea: Apidae). Austral Ecology, 27(2):162-172.
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