Bumblebee on:
[Wikipedia]
[Google]
[Amazon]
A bumblebee (or bumble bee, bumble-bee, or humble-bee) is any of over 250 species in the
The word "bumblebee" is a compound of "bumble" and "bee"—'bumble' meaning to hum, buzz, drone, or move ineptly or flounderingly. The generic name ''
On this hypothesis, the molecular data suggest that the Bombini are 25 to 40 million years old, while the Meliponini (and thus the clade that includes the Bombini and Meliponini) are 81 to 96 million years old, about the same age as the corbiculate group.
However, a more recent phylogeny using transcriptome data from 3,647 genes of ten corbiculate bee species supports the single origin of eusociality hypothesis in the corbiculate bees. They find that Bombini is in fact sister to Meliponini, corroborating that previous finding from Sophie Cardinal and Bryan Danforth (2011). However, Romiguier et al. (2015) shows that Bombini, Meliponini, and Apini form a monophyletic group, where Apini shares a most recent common ancestor with the Bombini and Meliponini clade, while Euglossini is most distantly related to all three, since it does not share the same most recent common ancestor as Bombini, Meliponini, and Apini. Thus, their analysis supports the single origin of eusociality hypothesis within the corbiculate bees, where eusociality evolved in the common ancestor of Bombini, Apini, and Meliponini.
The fossil record for bees is limited, with around 14 species that might possibly be Bombini having been described by 2019. The only ''Bombus'' relatives in Bombini are the late Eocene '' Calyptapis florissantensis'' from the Florissant Formation, USA, and '' Oligobombus cuspidatus'' from the Bembridge Marls of the
genus
Genus ( plural genera ) is a taxonomic rank used in the biological classification of living and fossil organisms as well as viruses. In the hierarchy of biological classification, genus comes above species and below family. In binomial n ...
''Bombus'', part of Apidae, one of the bee
Bees are winged insects closely related to wasps and ants, known for their roles in pollination and, in the case of the best-known bee species, the western honey bee, for producing honey. Bees are a monophyly, monophyletic lineage within the ...
families. This genus is the only extant group in the tribe Bombini, though a few extinct related genera (e.g., '' Calyptapis'') are known from fossil
A fossil (from Classical Latin , ) is any preserved remains, impression, or trace of any once-living thing from a past geological age. Examples include bones, shells, exoskeletons, stone imprints of animals or microbes, objects preserved ...
s. They are found primarily in higher altitudes or latitudes in the Northern Hemisphere, although they are also found in South America, where a few lowland tropical species have been identified. European bumblebees have also been introduced to New Zealand and Tasmania
)
, nickname =
, image_map = Tasmania in Australia.svg
, map_caption = Location of Tasmania in AustraliaCoordinates:
, subdivision_type = Country
, subdi ...
. Female bumblebees can sting repeatedly, but generally ignore humans and other animals.
Most bumblebees are social
Social organisms, including human(s), live collectively in interacting populations. This interaction is considered social whether they are aware of it or not, and whether the exchange is voluntary or not.
Etymology
The word "social" derives from ...
insects that form colonies
In modern parlance, a colony is a territory subject to a form of foreign rule. Though dominated by the foreign colonizers, colonies remain separate from the administration of the original country of the colonizers, the '' metropolitan state' ...
with a single queen. The colonies are smaller than those of honey bee
A honey bee (also spelled honeybee) is a eusocial flying insect within the genus ''Apis'' of the bee clade, all native to Afro-Eurasia. After bees spread naturally throughout Africa and Eurasia, humans became responsible for the current cosmop ...
s, growing to as few as 50 individuals in a nest. Cuckoo bumblebees are brood parasitic and do not make nests or form colonies; their queens aggressively invade the nests of other bumblebee species, kill the resident queens and then lay their own eggs, which are cared for by the resident workers. Cuckoo bumblebees were previously classified as a separate genus, but are now usually treated as members of ''Bombus''.
Bumblebees have round bodies covered in soft hair (long branched setae) called 'pile', making them appear and feel fuzzy. They have aposematic (warning) coloration, often consisting of contrasting bands of colour, and different species of bumblebee in a region often resemble each other in mutually protective Müllerian mimicry
Müllerian mimicry is a natural phenomenon in which two or more well-defended species, often foul-tasting and sharing common predators, have come to mimicry, mimic each other's honest signal, honest aposematism, warning signals, to their mutuali ...
. Harmless insects such as hoverflies often derive protection from resembling bumblebees, in Batesian mimicry, and may be confused with them. Nest-making bumblebees can be distinguished from similarly large, fuzzy cuckoo bees by the form of the female hind leg. In nesting bumblebees, it is modified to form a pollen basket, a bare shiny area surrounded by a fringe of hairs used to transport pollen
Pollen is a powdery substance produced by seed plants. It consists of pollen grains (highly reduced microgametophytes), which produce male gametes (sperm cells). Pollen grains have a hard coat made of sporopollenin that protects the gametop ...
, whereas in cuckoo bees, the hind leg is hairy all round, and they never carry pollen.
Like their relatives the honeybees, bumblebees feed on nectar, using their long hairy tongues to lap up the liquid; the proboscis
A proboscis () is an elongated appendage from the head of an animal, either a vertebrate or an invertebrate. In invertebrates, the term usually refers to tubular mouthparts used for feeding and sucking. In vertebrates, a proboscis is an elonga ...
is folded under the head during flight. Bumblebees gather nectar to add to the stores in the nest, and pollen to feed their young. They forage using colour and spatial relationships to identify flowers to feed from. Some bumblebees steal nectar, making a hole near the base of a flower to access the nectar while avoiding pollen transfer. Bumblebees are important agricultural pollinators, so their decline in Europe, North America, and Asia is a cause for concern. The decline has been caused by habitat loss
Habitat destruction (also termed habitat loss and habitat reduction) is the process by which a natural habitat becomes incapable of supporting its native species. The organisms that previously inhabited the site are displaced or dead, thereby ...
, the mechanisation of agriculture, and pesticides.
Etymology and common names
The word "bumblebee" is a compound of "bumble" and "bee"—'bumble' meaning to hum, buzz, drone, or move ineptly or flounderingly. The generic name ''Bombus
A bumblebee (or bumble bee, bumble-bee, or humble-bee) is any of over 250 species in the genus ''Bombus'', part of Apidae, one of the bee families. This genus is the only extant group in the tribe Bombini, though a few extinct related gene ...
'', assigned by Pierre André Latreille
Pierre André Latreille (; 29 November 1762 – 6 February 1833) was a French zoology, zoologist, specialising in arthropods. Having trained as a Roman Catholic priest before the French Revolution, Latreille was imprisoned, and only regained hi ...
in 1802, is derived from the Latin
Latin (, or , ) is a classical language belonging to the Italic branch of the Indo-European languages. Latin was originally a dialect spoken in the lower Tiber area (then known as Latium) around present-day Rome, but through the power ...
word for a buzzing or humming sound, borrowed from Ancient Greek
Ancient Greek includes the forms of the Greek language used in ancient Greece and the ancient world from around 1500 BC to 300 BC. It is often roughly divided into the following periods: Mycenaean Greek (), Dark Ages (), the Archaic p ...
().
According to the Oxford English Dictionary
The ''Oxford English Dictionary'' (''OED'') is the first and foundational historical dictionary of the English language, published by Oxford University Press (OUP). It traces the historical development of the English language, providing a com ...
(OED), the term "bumblebee" was first recorded as having been used in the English language in the 1530 work ''Lesclarcissement'' by John Palsgrave, "I bomme, as a bombyll bee dothe." However the OED also states that the term "humblebee" predates it, having first been used in 1450 in ''Fysshynge wyth Angle'', "In Juyll the greshop & the humbylbee in the medow." The latter term was used in ''A Midsummer Night's Dream
''A Midsummer Night's Dream'' is a comedy written by William Shakespeare 1595 or 1596. The play is set in Athens, and consists of several subplots that revolve around the marriage of Theseus and Hippolyta. One subplot involves a conflict ...
'' () by William Shakespeare
William Shakespeare ( 26 April 1564 – 23 April 1616) was an English playwright, poet and actor. He is widely regarded as the greatest writer in the English language and the world's pre-eminent dramatist. He is often called England's natio ...
, "The honie-bags steale from the humble Bees." Similar terms are used in other Germanic languages, such as the German (Old High German
Old High German (OHG; german: Althochdeutsch (Ahd.)) is the earliest stage of the German language, conventionally covering the period from around 750 to 1050.
There is no standardised or supra-regional form of German at this period, and Old High ...
), Dutch or Swedish
Swedish or ' may refer to:
Anything from or related to Sweden, a country in Northern Europe. Or, specifically:
* Swedish language, a North Germanic language spoken primarily in Sweden and Finland
** Swedish alphabet, the official alphabet used by ...
.
An old provincial name, "dumbledor", also denoted a buzzing insect such as a bumblebee or cockchafer, "dumble" probably imitating the sound of these insects, while "dor
DOR, Dor, or DoR may refer to:
Computer games and characters
* '' Advance Wars: Days of Ruin'', a turn-based tactics video game for the Nintendo DS
* Dor, a magician in the fictional Xanth universe; see Magicians of Xanth
* ''WWE Day of Reckoning ...
" meant "beetle".
In ''On the Origin of Species
''On the Origin of Species'' (or, more completely, ''On the Origin of Species by Means of Natural Selection, or the Preservation of Favoured Races in the Struggle for Life''),The book's full original title was ''On the Origin of Species by Me ...
'' (1859), Charles Darwin
Charles Robert Darwin ( ; 12 February 1809 – 19 April 1882) was an English natural history#Before 1900, naturalist, geologist, and biologist, widely known for his contributions to evolutionary biology. His proposition that all speci ...
speculated about "humble-bees" and their interactions with other species:
However, "bumblebee" remained in use, for example in ''The Tale of Mrs. Tittlemouse
''The Tale of Mrs. Tittlemouse'' is a children's book written and illustrated by Beatrix Potter, and published by Frederick Warne & Co. in 1910. The tale is about housekeeping and insect pests in the home, and reflects Potter's own sense of tid ...
'' (1910) by Beatrix Potter
Helen Beatrix Potter (, 28 July 186622 December 1943) was an English writer, illustrator, natural scientist, and conservationist. She is best known for her children's books featuring animals, such as '' The Tale of Peter Rabbit'', which was ...
, "Suddenly round a corner, she met Babbitty Bumble--"Zizz, Bizz, Bizzz!" said the bumble bee." Since World War II
World War II or the Second World War, often abbreviated as WWII or WW2, was a world war that lasted from 1939 to 1945. It involved the World War II by country, vast majority of the world's countries—including all of the great power ...
"humblebee" has fallen into near-total disuse.
Phylogeny
The bumblebee tribe Bombini is one of four groups of corbiculate bees (those with pollen baskets) in the Apidae, the others being the Apini (honey bees), Euglossini ( orchid bees), andMeliponini
Stingless bees, sometimes called stingless honey bees or simply meliponines, are a large group of bees (about 550 described species), comprising the tribe Meliponini (or subtribe Meliponina according to other authors). They belong in the family ...
(stingless bees). The corbiculate bees are a monophyletic
In cladistics for a group of organisms, monophyly is the condition of being a clade—that is, a group of taxa composed only of a common ancestor (or more precisely an ancestral population) and all of its lineal descendants. Monophyletic ...
group. Advanced eusocial behaviour appears to have evolved twice in the group, giving rise to controversy, now largely settled, as to the phylogenetic
In biology, phylogenetics (; from Greek φυλή/ φῦλον [] "tribe, clan, race", and wikt:γενετικός, γενετικός [] "origin, source, birth") is the study of the evolutionary history and relationships among or within groups o ...
origins of the four tribes; it had been supposed that eusocial behaviour had evolved only once, requiring the Apini to be close to the Meliponini, which they do not resemble. It is now thought that the Apini (with advanced societies) and Euglossini are closely related, while the primitively eusocial Bombini are close to the Meliponini, which have somewhat more advanced eusocial behaviour. Sophie Cardinal and Bryan Danforth comment that "While remarkable, a hypothesis of dual origins of advanced eusociality is congruent with early studies on corbiculate morphology and social behavior." Their analysis, combining molecular
A molecule is a group of two or more atoms held together by attractive forces known as chemical bonds; depending on context, the term may or may not include ions which satisfy this criterion. In quantum physics, organic chemistry, and bioch ...
, morphological and behavioural data, gives the following cladogram
A cladogram (from Greek ''clados'' "branch" and ''gramma'' "character") is a diagram used in cladistics to show relations among organisms. A cladogram is not, however, an evolutionary tree because it does not show how ancestors are related to ...
:
On this hypothesis, the molecular data suggest that the Bombini are 25 to 40 million years old, while the Meliponini (and thus the clade that includes the Bombini and Meliponini) are 81 to 96 million years old, about the same age as the corbiculate group.
However, a more recent phylogeny using transcriptome data from 3,647 genes of ten corbiculate bee species supports the single origin of eusociality hypothesis in the corbiculate bees. They find that Bombini is in fact sister to Meliponini, corroborating that previous finding from Sophie Cardinal and Bryan Danforth (2011). However, Romiguier et al. (2015) shows that Bombini, Meliponini, and Apini form a monophyletic group, where Apini shares a most recent common ancestor with the Bombini and Meliponini clade, while Euglossini is most distantly related to all three, since it does not share the same most recent common ancestor as Bombini, Meliponini, and Apini. Thus, their analysis supports the single origin of eusociality hypothesis within the corbiculate bees, where eusociality evolved in the common ancestor of Bombini, Apini, and Meliponini.
The fossil record for bees is limited, with around 14 species that might possibly be Bombini having been described by 2019. The only ''Bombus'' relatives in Bombini are the late Eocene '' Calyptapis florissantensis'' from the Florissant Formation, USA, and '' Oligobombus cuspidatus'' from the Bembridge Marls of the Isle of Wight
The Isle of Wight ( ) is a Counties of England, county in the English Channel, off the coast of Hampshire, from which it is separated by the Solent. It is the List of islands of England#Largest islands, largest and List of islands of England#Mo ...
. Two species of ''Bombus'' have been described from the Oligocene
The Oligocene ( ) is a geologic epoch of the Paleogene Period and extends from about 33.9 million to 23 million years before the present ( to ). As with other older geologic periods, the rock beds that define the epoch are well identified but ...
of Beşkonak, Bucak
Beşkonak is a village in the Bucak District
Bucak District is a Districts of Turkey, district of the Burdur Province of Turkey. Its seat is the town of Bucak, Burdur, Bucak.Bombus (Mendacibombus) beskonakensis'' and '' Bombus (Paraelectrobombus) patriciae''. Both species were originally placed in genera considered at the time of description as outside of ''Bombus'', being initially named ''Oligoapis beskonakensis'' and ''Paraelectrobombus patriciae'' respectively, however reexaminiation of the fore-wings lead to both being considered as ''Bombus'' species In 2012 a fossil bumblebee from the
The brightly coloured pile of the bumblebee is an aposematic (warning) signal, given that females can inflict a painful sting. Depending on the species and morph, the warning colours range from entirely black, to bright yellow, red, orange, white, and pink. Dipteran flies in the families Syrphidae (hoverflies), Asilidae (robber flies), Tabanidae (horseflies),
Nest size depends on species of bumblebee. Most form 
In fertilised queens, the ovaries only become active when the queen starts to lay. An egg passes along the 
After the emergence of the first or second group of offspring, workers take over the task of foraging and the queen spends most of her time laying eggs and caring for larvae. The colony grows progressively larger and eventually begins to produce males and new queens.Goulson, 2013. pp. 16–24
Bumblebee workers can lay unfertilised haploid eggs (with only a single set of
Bumblebees generally visit flowers that exhibit the bee pollination syndrome and these patches of flowers may be up to 1–2 km from their colony. They tend to visit the same patches of flowers every day, as long as they continue to find nectar and pollen there, a habit known as pollinator or flower constancy. While foraging, bumblebees can reach ground speeds of up to .
Bumblebees use a combination of colour and spatial relationships to learn which flowers to forage from. They can also detect both the presence and the pattern of electric fields on flowers, which occur due to atmospheric electricity, and take a while to leak away into the ground. They use this information to find out if a flower has been recently visited by another bee. Bumblebees can detect the temperature of flowers, as well as which parts of the flower are hotter or cooler and use this information to recognise flowers. After arriving at a flower, they extract nectar using their long tongues (" glossae") and store it in their
Bumblebees of the subgenus '' Psithyrus'' (known as 'cuckoo bumblebees', and formerly considered a separate genus) are brood parasites, sometimes called kleptoparasites, in the colonies of other bumblebees, and have lost the ability to collect pollen. Before finding and invading a host colony, a ''Psithyrus'' female, such as that of the ''Psithyrus'' species of '' B. sylvestris'', feeds directly from flowers. Once she has infiltrated a host colony, the ''Psithyrus'' female kills or subdues the queen of that colony, and uses pheromones and physical attacks to force the workers of that colony to feed her and her young. Usually, cuckoo bumblebees can be described as queen-intolerant inquilines, since the host queen is often killed to enable the parasite to produce more offspring, though some species, such as '' B. bohemicus'', actually enjoy increased success when they leave the host queen alive.
The female ''Psithyrus'' has a number of morphological adaptations for combat, such as larger mandibles, a tough cuticle and a larger venom sac that increase her chances of taking over a nest. Upon emerging from their cocoons, the ''Psithyrus'' males and females disperse and mate. The males do not survive the winter but, like nonparasitic bumblebee queens, ''Psithyrus'' females find suitable locations to spend the winter and enter diapause after mating. They usually emerge from hibernation later than their host species. Each species of cuckoo bee has a specific host species, which it may physically resemble. In the case of the parasitism of ''B. terrestris'' by '' B. (Psithyrus) vestalis'', genetic analysis of individuals captured in the wild showed that about 42% of the host species' nests at a single location had " osttheir fight against their parasite".
Bumblebees, despite their ability to sting, are eaten by certain predators. Nests may be dug up by 
The great grey shrike is able to detect flying bumblebees up to away; once captured, the sting is removed by repeatedly squeezing the insect with the mandibles and wiping the abdomen on a branch. The European honey buzzard follows flying bees back to their nest, digs out the nest with its feet, and eats larvae, pupae and adults as it finds them.
Bumblebees are parasitised by tracheal mites, ''
In 2006 the bumblebee researcher
According to 20th-century 
The calculations that purported to show that bumblebees cannot fly are based upon a simplified linear treatment of
The orchestral interlude '' Flight of the Bumblebee'' was composed (c. 1900) by 
Emily Dickinson made a bumblebee the subject of her parody of Isaac Watts's well-known poem about honeybees, "How Doth the Little Busy Bee" (1715). Where Watts wrote "How skilfully she builds her cell! How neat she spreads the wax!", Dickinson's poem, "The Bumble-Bee's Religion" (1881), begins "His little Hearse-like Figure / Unto itself a Dirge / To a delusive Lilac / The vanity divulge / Of Industry and Morals / And every righteous thing / For the divine Perdition / of Idleness and Spring." The letter was said to have enclosed a dead bee.
In 1847, 
The scientist and illustrator Moses Harris (1731–1785) painted accurate watercolour drawings of bumblebees in his ''An Exposition of English Insects Including the Several Classes of Neuroptera, Hymenoptera, & Diptera, or Bees, Flies, & Libellulae (1776–80)''.
Bumblebees appear as characters, often eponymously, in children's books. The surname Dumbledore in the
Bumblebees
'. Scottish Natural Heritage, 2003. * Macdonald, Murdo & Nisbet, G.
'
HBRG
2006. .
Supplement 2
(2007). * Michener, C.D. ''The Bees of the World''. Johns Hopkins University Press, 2000. * Schweitzer, Dale F. et al.
Conservation and Management of North American Bumble Bees
'. Washington D.C.: U.S. Forest Service, 2012.
Bumblebees of the world – find species by region, species groups, colour pattern
nhm.ac.uk
Bumblebee Conservation Trust
IUCN's Bumblebee Specialist Group
''Bombus'' Identification GuideDiscover Life: List of SpeciesWorldwide Species Map
Deciphering the Mystery of Bee Flight
{{good article Aposematic animals Articles containing video clips Extant Eocene first appearances Insects in culture
Miocene
The Miocene ( ) is the first geological epoch of the Neogene Period and extends from about (Ma). The Miocene was named by Scottish geologist Charles Lyell; the name comes from the Greek words (', "less") and (', "new") and means "less recent" ...
was found in Germany's Randeck Maar and classified as '' Bombus (Bombus) randeckensis''. In 2014, another species, '' Bombus cerdanyensis'', was described from Late Miocene lacustrine beds of La Cerdanya, Spain, but not initially placed into any subgenus, The species ''Bombus trophonius
''Bombus trophonius'' is an extinct species of bumble bee known from a Miocene fossil found in Europe. It belongs to the ''Bombus'' subgenus ''Cullumanobombus'' and is considered most similar to the living species '' Bombus rufocinctus'' of Nort ...
'' was described in October 2017 and placed in ''Bombus'' subgenus ''Cullumanobombus''. A redescription of the Bombini fossil record by Dehon ''et al'' (2019
File:2019 collage v1.png, From top left, clockwise: Hong Kong protests turn to widespread riots and civil disobedience; House of Representatives votes to adopt articles of impeachment against Donald Trump; CRISPR gene editing first used to experim ...
) resulted in the synonymization of the genus ''Oligoapis'' with ''Bombus'' subgenus ''Mendacibombus'', and the placement of genus ''Paraelectrobombus'' as ''Bombus'' subgenus ''Paraelectrobombus'', rather than as a genus in Electrobombini. The subgenus ''Cullumanobombus'' was expanded to include not only ''Bombus trophonius'' but also ''Bombus randeckensis'' which was moved from subgenus ''Bombus'' and ''Bombus pristinus
A bumblebee (or bumble bee, bumble-bee, or humble-bee) is any of over 250 species in the genus ''Bombus'', part of Apidae, one of the bee families. This genus is the only extant group in the tribe Bombini, though a few extinct related gener ...
'', first described by Unger (1867
Events
January–March
* January 1 – The Covington–Cincinnati Suspension Bridge opens between Cincinnati, Ohio, and Covington, Kentucky, in the United States, becoming the longest single-span bridge in the world. It was renamed a ...
). Within the subgenus ''Melanobombus'' only ''Bombus cerdanyensis'' is present from the fossil record. An additional three species, '' "Bombus" luianus'', '' "Bombus" anacolus'' and '' "Bombus" dilectus'' have been attributed to ''Bombus'' from the Middle Miocene Shanwang formation of China by Zhang, ( 1990) and Zhang ''et al'' (1994
File:1994 Events Collage.png, From left, clockwise: The 1994 Winter Olympics are held in Lillehammer, Norway; The Kaiser Permanente building after the 1994 Northridge earthquake; A model of the MS Estonia, which sank in the Baltic Sea; Nelson Ma ...
). Due to not being able to study Zhang's type specimen
In biology, a type is a particular specimen (or in some cases a group of specimens) of an organism to which the scientific name of that organism is formally attached. In other words, a type is an example that serves to anchor or centralizes the ...
s, but only illustrations of the fossils, Dehon ''et al'' did not place the three species within any specific subgenera, and considered all three as "species inquirenda", needing fuller re-examination. Two other species were not examined at all by Dehon ''et al'', '' Bombus? crassipes'' of the Late Miocene Krottensee deposits in the Czech Republic, and '' Bombus proavus'' from the Middle Miocene Latah Formation, USA.
Taxonomy
The genus ''Bombus'', the only one extant genus in the tribe Bombini, comprises over 250 species; for an overview of the differences between bumblebees and other bees and wasps, see characteristics of common wasps and bees. The genus has been divided variously into up to 49 subgenera, a degree of complexity criticised by Williams (2008). The cuckoo bumblebees ''Psithyrus'' have sometimes been treated as a separate genus but are now considered to be part of ''Bombus'', in one or more subgenera. Examples of ''Bombus'' species include ''Bombus pauloensis
''Bombus pauloensis'' is a neotropical bumblebee, formerly known as ''Bombus atratus'', that is found throughout regions of South America, including Colombia, Ecuador, Brazil, and Argentina. It lives in social colonies that include a founder que ...
'', '' Bombus dahlbomii'', '' Bombus fervidus'', ''Bombus lapidarius
''Bombus lapidarius'' is a species of bumblebee in the subgenus ''Melanobombus''. Commonly known as the red-tailed bumblebee, ''B. lapidarius'' can be found throughout much of Central Europe. Known for its distinctive black and red body, this so ...
'', '' Bombus ruderatus'', and ''Bombus rupestris
''Bombus rupestris'' is a species of cuckoo bumblebee present in most of Europe except Iceland. In the Balkans it is found in montane and alpine habitats northwards from Central Greece. It is also found in Turkey.
Description
The female is much ...
.''
:::Subgenera of the genus ''Bombus''
General description
Bumblebees vary in appearance, but are generally plump and densely furry. They are larger, broader and stouter-bodied than honeybees, and their abdomen tip is more rounded. Many species have broad bands of colour, the patterns helping to distinguish different species. Whereas honeybees have short tongues and therefore mainly pollinate open flowers, some bumblebee species have long tongues and collect nectar from flowers that are closed into a tube. Bumblebees have fewer stripes (or none), and usually have part of the body covered in black fur, while honeybees have many stripes including several grey stripes on the abdomen. Sizes are very variable even within species; the largest British species, ''B. terrestris'', has queens up to long, males up to long, and workers between long. The largest bumblebee species in the world is '' B. dahlbomii'' of Chile, up to about long, and described as "flying mice" and "a monstrous fluffy ginger beast".Distribution and habitat
Bumblebees are typically found intemperate climate
In geography, the temperate climates of Earth occur in the middle latitudes (23.5° to 66.5° N/S of Equator), which span between the tropics and the polar regions of Earth. These zones generally have wider temperature ranges throughout ...
s, and are often found at higher latitude
In geography, latitude is a coordinate that specifies the north– south position of a point on the surface of the Earth or another celestial body. Latitude is given as an angle that ranges from –90° at the south pole to 90° at the north po ...
s and altitudes than other bees, although a few lowland tropical species exist. A few species ('' B. polaris'' and '' B. alpinus'') range into very cold climates where other bees might not be found; ''B. polaris'' occurs in northern Ellesmere Island in the high Arctic, along with another bumblebee '' B. hyperboreus'', which parasitises its nest. This is the northernmost occurrence of any eusocial insect. One reason for their presence in cold places is that bumblebees can regulate their body temperature
Thermoregulation is the ability of an organism to keep its body temperature within certain boundaries, even when the surrounding temperature is very different. A thermoconforming organism, by contrast, simply adopts the surrounding temperatur ...
, via solar radiation, internal mechanisms of "shivering" and radiative cooling from the abdomen (called heterothermy). Other bees have similar physiology
Physiology (; ) is the scientific study of functions and mechanisms in a living system. As a sub-discipline of biology, physiology focuses on how organisms, organ systems, individual organs, cells, and biomolecules carry out the chemic ...
, but the mechanisms seem best developed and have been most studied in bumblebees. They adapt to higher elevations by extending their wing stroke amplitude. Bumblebees have a largely cosmopolitan distribution but are absent from Australia (apart from Tasmania where they have been introduced) and are found in Africa only north of the Sahara. More than a hundred years ago they were also introduced to New Zealand, where they play an important role as efficient pollinators.
Biology
Feeding
The bumblebee tongue (theproboscis
A proboscis () is an elongated appendage from the head of an animal, either a vertebrate or an invertebrate. In invertebrates, the term usually refers to tubular mouthparts used for feeding and sucking. In vertebrates, a proboscis is an elonga ...
) is a long, hairy structure that extends from a sheath-like modified maxilla. The primary action of the tongue is lapping, that is, repeated dipping of the tongue into liquid.
The tip of the tongue probably acts as a suction cup and during lapping, nectar may be drawn up the proboscis by capillary action. When at rest or flying, the proboscis is kept folded under the head. The longer the tongue, the deeper the bumblebee can probe into a flower and bees probably learn from experience which flower source is best-suited to their tongue length. Bees with shorter proboscides, like ''Bombus bifarius
''Bombus bifarius'', the two-form bumblebee, is a species of eusocial bumblebee of the subgenus ''Pyrobombus''. ''B. bifarius'' inhabits mountainous regions of western North America, primarily the states of Colorado and Utah. Its common name refe ...
'', have a more difficult time foraging nectar relative to other bumblebees with longer proboscides; to overcome this disadvantage, ''B. bifarius'' workers were observed to lick the back of spurs on the nectar duct, which resulted in a small reward.
Wax production
The exoskeleton of theabdomen
The abdomen (colloquially called the belly, tummy, midriff, tucky or stomach) is the part of the body between the thorax (chest) and pelvis, in humans and in other vertebrates. The abdomen is the front part of the abdominal segment of the tors ...
is divided into plates called dorsal tergites and ventral sternites. Wax is secreted from gland
In animals, a gland is a group of cells in an animal's body that synthesizes substances (such as hormones) for release into the bloodstream ( endocrine gland) or into cavities inside the body or its outer surface ( exocrine gland).
Structure
...
s on the abdomen and extruded between the sternites where it resembles flakes of dandruff. It is secreted by the queen when she starts a nest and by young workers. It is scraped from the abdomen by the legs, moulded until malleable and used in the construction of honeypots, to cover the eggs, to line empty cocoons for use as storage containers and sometimes to cover the exterior of the nest.
Coloration
The brightly coloured pile of the bumblebee is an aposematic (warning) signal, given that females can inflict a painful sting. Depending on the species and morph, the warning colours range from entirely black, to bright yellow, red, orange, white, and pink. Dipteran flies in the families Syrphidae (hoverflies), Asilidae (robber flies), Tabanidae (horseflies), Oestridae
Botflies, also known as warble flies, heel flies, and gadflies, are a family of flies known as the Oestridae. Their larvae are internal parasites of mammals, some species growing in the host's flesh and others within the gut. ''Dermatobia homin ...
(bot or warble flies) and Bombyliidae
The Bombyliidae are a family of flies, commonly known as bee flies. Adults generally feed on nectar and pollen, some being important pollinators. Larvae are mostly parasitoids of other insects.
Overview
The Bombyliidae are a large family of ...
(bee flies, such as '' Bombylius major'') all include Batesian mimics of bumblebees, resembling them closely enough to deceive at least some predators.
Many species of ''Bombus'', including the group sometimes called ''Psithyrus'' (cuckoo bumblebees), have evolved Müllerian mimicry
Müllerian mimicry is a natural phenomenon in which two or more well-defended species, often foul-tasting and sharing common predators, have come to mimicry, mimic each other's honest signal, honest aposematism, warning signals, to their mutuali ...
, where the different bumblebees in a region resemble each other, so that a young predator need only learn to avoid any of them once. For example, in California a group of bumblebees consists of largely black species including '' B. californicus'', '' B. caliginosus'', '' B. vandykei'', '' B. vosnesenskii'', '' B. insularis'' and '' B. fernaldae''. Other bees in California include a group of species all banded black and yellow. In each case, Müllerian mimicry provides the bees in the group with a selective advantage. In addition, parasitic (cuckoo) bumblebees resemble their hosts more closely than would be expected by chance, at least in areas like Europe where parasite-host co-speciation is common; but this too may be explained as Müllerian mimicry, rather than requiring the parasite's coloration to deceive the host ( aggressive mimicry).
Temperature control
Bumblebees are active under conditions during which honeybees stay at home, and can readily absorb heat from even weak sunshine.Macdonald, 2003. p. 6 The thick pile created by long setae (bristles) acts as insulation to keep bumblebees warm in cold weather; species from cold climates have longer setae (and thus thicker insulation) than those from the tropics. The temperature of the flight muscles, which occupy much of the thorax, needs to be at least before flight can take place. The muscle temperature can be raised by shivering. It takes about five minutes for the muscles to reach this temperature at an air temperature of .Chill-coma temperature
The chill-coma temperature in relation to flying insects is the temperature at which flight muscles cannot be activated. Compared to honey bees and carpenter bees, bumblebees have the lowest chill-coma temperature. Of the bumblebees '' Bombus bimaculatus'' has the lowest at . However, bumblebees have been seen to fly in colder ambient temperatures. This discrepancy is likely because the chill-coma temperature was determined by tests done in a laboratory setting. However, bumblebees live in insulated shelters and can shiver to warm up before venturing into the cold.Communication and social learning
Bumblebees do not have ears, and it is not known whether or how well they can hear. However, they are sensitive to the vibrations made by sound travelling through wood or other materials. Bumblebees do not exhibit the " bee dances" used by honeybees to tell other workers the locations of food sources. Instead, when they return from a successful foraging expedition, they run excitedly around in the nest for several minutes before going out to forage once more. These bees may be offering some form of communication based on the buzzing sounds made by their wings, which may stimulate other bees to start foraging. Another stimulant to foraging activity is the level of food reserves in the colony. Bees monitor the amount of honey in the honeypots, and when little is left or when high-quality food is added, they are more likely to go out to forage. Bumblebees have been observed to partake in social learning. In a 2017 study involving ''Bombus terrestris
''Bombus terrestris'', the buff-tailed bumblebee or large earth bumblebee, is one of the most numerous bumblebee species in Europe. It is one of the main species used in greenhouse pollination, and so can be found in many countries and areas wh ...
'', bees were taught to complete an unnatural task of moving large objects to obtain a reward. Bees who first observed another bee complete the task were significantly more successful in learning the task than bees who observed the same action performed by a magnet, indicating the importance of social information. The bees did not copy one another exactly: in fact, the study suggested that the bees were instead attempting to emulate one another's goals.
Reproduction and nesting
Nest size depends on species of bumblebee. Most form colonies
In modern parlance, a colony is a territory subject to a form of foreign rule. Though dominated by the foreign colonizers, colonies remain separate from the administration of the original country of the colonizers, the '' metropolitan state' ...
of between 50 and 400 individuals, but colonies have been documented as small as ~20 individuals and as large as 1700. These nests are small compared to honeybee hives, which hold about 50,000 bees. Many species nest underground, choosing old rodent
Rodents (from Latin , 'to gnaw') are mammals of the Order (biology), order Rodentia (), which are characterized by a single pair of continuously growing incisors in each of the upper and lower jaws. About 40% of all mammal species are roden ...
burrows or sheltered places, and avoiding places that receive direct sunlight that could result in overheating. Other species make nests above ground, whether in thick grass or in holes in trees. A bumblebee nest is not organised into hexagonal combs like that of a honeybee; the cells are instead clustered together untidily. The workers remove dead bees or larvae from the nest and deposit them outside the nest entrance, helping to prevent disease. Nests in temperate regions last only for a single season and do not survive the winter.
In the early spring, the queen comes out of diapause and finds a suitable place to create her colony. Then she builds wax cells in which to lay her eggs which were fertilised the previous year. The eggs that hatch develop into female workers, and in time, the queen populates the colony, with workers feeding the young and performing other duties similar to honeybee workers. In temperate zones, young queens ( gynes) leave the nest in the autumn and mate, often more than once, with males (drone
Drone most commonly refers to:
* Drone (bee), a male bee, from an unfertilized egg
* Unmanned aerial vehicle
* Unmanned surface vehicle, watercraft
* Unmanned underwater vehicle or underwater drone
Drone, drones or The Drones may also refer to:
...
s) that are forcibly driven out of the colony. The drones and workers die as the weather turns colder; the young queens feed intensively to build up stores of fat for the winter. They survive in a resting state (diapause), generally below ground, until the weather warms up in the spring with the early bumblebee
The early bumblebee or early-nesting bumblebee (''Bombus pratorum'') is a small bumblebee with a wide distribution in most of Europe and parts of Asia. It is very commonly found in the UK and emerges to begin its colony cycle as soon as Februar ...
being the species that is among the first to emerge. Many species of bumblebee follow this general trend within the year. '' Bombus pensylvanicus'' is a species that follows this type of colony cycle. For this species the cycle begins in February, reproduction starts in July or August, and ends in the winter months. The queen remains in hibernation until spring of the following year in order to optimize conditions to search for a nest.
In fertilised queens, the ovaries only become active when the queen starts to lay. An egg passes along the oviduct
The oviduct in mammals, is the passageway from an ovary. In human females this is more usually known as the Fallopian tube or uterine tube. The eggs travel along the oviduct. These eggs will either be fertilized by spermatozoa to become a zygote, ...
to the vagina where there is a chamber called the spermatheca
The spermatheca (pronounced plural: spermathecae ), also called receptaculum seminis (plural: receptacula seminis), is an organ of the female reproductive tract in insects, e.g. ants, bees, some molluscs, oligochaeta worms and certain othe ...
, in which the sperm from the mating is stored. Depending on need, she may allow her egg to be fertilised. Unfertilised eggs become haploid males; fertilised eggs grow into diploid
Ploidy () is the number of complete sets of chromosomes in a cell, and hence the number of possible alleles for autosomal and pseudoautosomal genes. Sets of chromosomes refer to the number of maternal and paternal chromosome copies, respecti ...
females and queens. The hormones that stimulate the development of the ovaries are suppressed in female worker bees, while the queen remains dominant.
To develop, the larva
A larva (; plural larvae ) is a distinct juvenile form many animals undergo before metamorphosis into adults. Animals with indirect development such as insects, amphibians, or cnidarians typically have a larval phase of their life cycle.
Th ...
e must be fed both nectar for carbohydrate
In organic chemistry, a carbohydrate () is a biomolecule consisting of carbon (C), hydrogen (H) and oxygen (O) atoms, usually with a hydrogen–oxygen atom ratio of 2:1 (as in water) and thus with the empirical formula (where ''m'' may or ...
s and pollen for protein
Proteins are large biomolecules and macromolecules that comprise one or more long chains of amino acid residues. Proteins perform a vast array of functions within organisms, including catalysing metabolic reactions, DNA replication, respon ...
. Bumblebees feed nectar to the larvae by chewing a small hole in the brood cell into which they regurgitate nectar. Larvae are fed pollen in one of two ways, depending on the bumblebee species. Pocket-making bumblebees create pockets of pollen at the base of the brood-cell clump from which the larvae feed themselves. Pollen-storing bumblebees keep pollen in separate wax pots and feed it to the larvae.
After the emergence of the first or second group of offspring, workers take over the task of foraging and the queen spends most of her time laying eggs and caring for larvae. The colony grows progressively larger and eventually begins to produce males and new queens.Goulson, 2013. pp. 16–24
Bumblebee workers can lay unfertilised haploid eggs (with only a single set of chromosome
A chromosome is a long DNA molecule with part or all of the genetic material of an organism. In most chromosomes the very long thin DNA fibers are coated with packaging proteins; in eukaryotic cells the most important of these proteins ar ...
s) that develop into viable male bumblebees. Only fertilised queens can lay diploid eggs (one set of chromosomes from a drone, one from the queen) that mature into workers and new queens.
In a young colony, the queen minimises reproductive competition from workers by suppressing their egg-laying through physical aggression and pheromones. Worker policing leads to nearly all eggs laid by workers being eaten. Thus, the queen is usually the mother of all of the first males laid. Workers eventually begin to lay male eggs later in the season when the queen's ability to suppress their reproduction diminishes. Because of the reproductive competition
Competition is a rivalry where two or more parties strive for a common goal which cannot be shared: where one's gain is the other's loss (an example of which is a zero-sum game). Competition can arise between entities such as organisms, ind ...
between workers and the queen, bumblebees are considered "primitively eusocial".
Although a large majority of bumblebees follow such monogynous colony cycles that only involve one queen, some select ''Bombus'' species (such as ''Bombus pauloensis
''Bombus pauloensis'' is a neotropical bumblebee, formerly known as ''Bombus atratus'', that is found throughout regions of South America, including Colombia, Ecuador, Brazil, and Argentina. It lives in social colonies that include a founder que ...
'') will spend part of their life cycle in a polygynous phase (have multiple queens in one nest during these periods of polygyny).
Foraging behaviour
Bumblebees generally visit flowers that exhibit the bee pollination syndrome and these patches of flowers may be up to 1–2 km from their colony. They tend to visit the same patches of flowers every day, as long as they continue to find nectar and pollen there, a habit known as pollinator or flower constancy. While foraging, bumblebees can reach ground speeds of up to .
Bumblebees use a combination of colour and spatial relationships to learn which flowers to forage from. They can also detect both the presence and the pattern of electric fields on flowers, which occur due to atmospheric electricity, and take a while to leak away into the ground. They use this information to find out if a flower has been recently visited by another bee. Bumblebees can detect the temperature of flowers, as well as which parts of the flower are hotter or cooler and use this information to recognise flowers. After arriving at a flower, they extract nectar using their long tongues (" glossae") and store it in their crop
A crop is a plant that can be grown and harvested extensively for profit or subsistence. When the plants of the same kind are cultivated at one place on a large scale, it is called a crop. Most crops are cultivated in agriculture or hydropo ...
s. Many species of bumblebees also exhibit "nectar robbing": instead of inserting the mouthparts into the flower in the normal way, these bees bite directly through the base of the corolla
Corolla may refer to:
*Corolla (botany), the petals of a flower, considered as a unit
*Toyota Corolla, an automobile model name
* Corolla (headgear), an ancient headdress in the form of a circlet or crown
* ''Corolla'' (gastropod), a genus of moll ...
to extract nectar, avoiding pollen transfer.
Pollen is removed from flowers deliberately or incidentally by bumblebees. Incidental removal occurs when bumblebees come in contact with the anthers of a flower while collecting nectar. When it enters a flower, the bumblebee's body hairs receive a dusting of pollen from the anthers. In queens and workers this is then groomed into the corbiculae (pollen baskets) on the hind legs where it can be seen as bulging masses that may contain as many as a million pollen grains. Male bumblebees do not have corbiculae and do not purposively collect pollen. Bumblebees are also capable of buzz pollination, in which they dislodge pollen from the anthers by creating a resonant vibration with their flight muscles.
In at least some species, once a bumblebee has visited a flower, it leaves a scent mark on it. This scent mark deters bumblebees from visiting that flower until the scent degrades. This scent mark is a general chemical bouquet that bumblebees leave behind in different locations (e.g. nest, neutral, and food sites), and they learn to use this bouquet to identify both rewarding and unrewarding flowers, and may be able to identify who else has visited a flower. Bumblebees rely on this chemical bouquet more when the flower has a high handling time, that is, where it takes a longer time for the bee to find the nectar once inside the flower.
Once they have collected nectar and pollen, female workers return to the nest and deposit the harvest into brood cells, or into wax cells for storage. Unlike honeybees, bumblebees only store a few days' worth of food, so are much more vulnerable to food shortages. Male bumblebees collect only nectar and do so to feed themselves. They may visit quite different flowers from the workers because of their different nutritional needs.
Asynchronous flight muscles
Bees beat their wings about 200 times a second. Their thorax muscles do not contract on each nerve firing, but rather vibrate like a plucked rubber band. This is efficient, since it lets the system consisting of muscle and wing operate at its resonant frequency, leading to low energy consumption. Further, it is necessary, since insect motor nerves generally cannot fire 200 times per second. These types of muscles are calledasynchronous muscles
Asynchronous muscles are muscles in which there is no one-to-one relationship between electrical stimulation and mechanical contraction. These muscles are found in 75% of flying insects and have convergently evolved 7-10 times. Unlike their synchro ...
and are found in the insect wing systems in families such as Hymenoptera, Diptera, Coleoptera
Beetles are insects that form the order Coleoptera (), in the superorder Endopterygota. Their front pair of wings are hardened into wing-cases, elytra, distinguishing them from most other insects. The Coleoptera, with about 400,000 describ ...
, and Hemiptera. Bumblebees must warm up their bodies considerably to get airborne at low ambient temperatures. Bumblebees can reach an internal thoracic temperature of 30 °C (86 °F) using this method.
Cuckoo bumblebees
Bumblebees of the subgenus '' Psithyrus'' (known as 'cuckoo bumblebees', and formerly considered a separate genus) are brood parasites, sometimes called kleptoparasites, in the colonies of other bumblebees, and have lost the ability to collect pollen. Before finding and invading a host colony, a ''Psithyrus'' female, such as that of the ''Psithyrus'' species of '' B. sylvestris'', feeds directly from flowers. Once she has infiltrated a host colony, the ''Psithyrus'' female kills or subdues the queen of that colony, and uses pheromones and physical attacks to force the workers of that colony to feed her and her young. Usually, cuckoo bumblebees can be described as queen-intolerant inquilines, since the host queen is often killed to enable the parasite to produce more offspring, though some species, such as '' B. bohemicus'', actually enjoy increased success when they leave the host queen alive.
The female ''Psithyrus'' has a number of morphological adaptations for combat, such as larger mandibles, a tough cuticle and a larger venom sac that increase her chances of taking over a nest. Upon emerging from their cocoons, the ''Psithyrus'' males and females disperse and mate. The males do not survive the winter but, like nonparasitic bumblebee queens, ''Psithyrus'' females find suitable locations to spend the winter and enter diapause after mating. They usually emerge from hibernation later than their host species. Each species of cuckoo bee has a specific host species, which it may physically resemble. In the case of the parasitism of ''B. terrestris'' by '' B. (Psithyrus) vestalis'', genetic analysis of individuals captured in the wild showed that about 42% of the host species' nests at a single location had " osttheir fight against their parasite".
Sting
Queen and worker bumblebees can sting. Unlike in honeybees, a bumblebee's stinger lacks barbs, so the bee can sting repeatedly without leaving the stinger in the wound and thereby injuring itself. Bumblebee species are not normally aggressive, but may sting in defence of their nest, or if harmed. Female cuckoo bumblebees aggressively attack host colony members, and sting the host queen, but ignore other animals unless disturbed. The sting is painful to humans, and not medically significant in most cases, although it may trigger an allergic reaction in susceptible individuals.Predators, parasites, and pathogens
Bumblebees, despite their ability to sting, are eaten by certain predators. Nests may be dug up by badger
Badgers are short-legged omnivores in the family Mustelidae (which also includes the otters, wolverines, martens, minks, polecats, weasels, and ferrets). Badgers are a polyphyletic rather than a natural taxonomic grouping, being united ...
s and eaten whole, including any adults present. Adults are preyed upon by robber flies and beewolves
Beewolves (genus ''Philanthus''), also known as bee-hunters or bee-killer wasps, are solitary, predatory wasps, most of which prey on bees, hence their common name. The adult females dig tunnels in the ground for nesting, while the territorial m ...
in North America. In Europe, birds including bee-eaters and shrike
Shrikes () are passerine birds of the family Laniidae. The family is composed of 34 species in four genera.
The family name, and that of the largest genus, '' Lanius'', is derived from the Latin word for "butcher", and some shrikes are also kno ...
s capture adult bumblebees on the wing; smaller birds such as great tits also occasionally learn to take bumblebees, while camouflage
Camouflage is the use of any combination of materials, coloration, or illumination for concealment, either by making animals or objects hard to see, or by disguising them as something else. Examples include the leopard's spotted coat, the b ...
d crab spiders catch them as they visit flowers.
The great grey shrike is able to detect flying bumblebees up to away; once captured, the sting is removed by repeatedly squeezing the insect with the mandibles and wiping the abdomen on a branch. The European honey buzzard follows flying bees back to their nest, digs out the nest with its feet, and eats larvae, pupae and adults as it finds them.
Bumblebees are parasitised by tracheal mites, ''Locustacarus buchneri
''Locustacarus buchneri'' is a parasitic mite that lives in the respiratory air sacs of bumblebee
A bumblebee (or bumble bee, bumble-bee, or humble-bee) is any of over 250 species in the genus ''Bombus'', part of Apidae, one of the bee f ...
''; protozoans including '' Crithidia bombi'' and '' Apicystis bombi''; and microsporidians including '' Nosema bombi'' and ''Nosema ceranae
''Nosema ceranae'' is a microsporidian, a small, unicellular parasite that mainly affects '' Apis cerana'', the Asiatic honey bee. Along with ''Nosema apis'', it causes the disease nosemosis, the most widespread of the diseases of adult honey ...
''. The tree bumblebee '' B. hypnorum'' has spread into the United Kingdom despite hosting high levels of a nematode that normally interferes with queen bees' attempts to establish colonies. Deformed wing virus
''Deformed wing virus'' (DWV) is an RNA virus, one of 22 known viruses affecting honey bees. While most commonly infecting the honey bee, ''Apis mellifera'', it has also been documented in other bee species, like '' Bombus terrestris'', thus, i ...
has been found to affect 11% of bumblebees in Great Britain.
Female bee moths (''Aphomia sociella
''Aphomia sociella'', also known as the bee moth and the bumble bee wax moth, is a small moth of the family Pyralidae (snout moths) and subfamily Galleriinae. Its body and forewings are typically reddish brown, tan, or dark green in color and ...
'') prefer to lay their eggs in bumblebee nests. The ''A. sociella'' larvae will then feed on the eggs, larvae, and pupae left unprotected by the bumblebees, sometimes destroying large parts of the nest.
Relationship to humans
Agricultural use
Bumblebees are important pollinators of both crops and wildflowers. Because bumblebees do not overwinter the entire colony, they do not stockpile honey, and therefore are not useful as honey producers. Bumblebees are increasingly cultured for agricultural use as pollinators, among other reasons because they can pollinate plants such as tomato ingreenhouse
A greenhouse (also called a glasshouse, or, if with sufficient heating, a hothouse) is a structure with walls and roof made chiefly of transparent material, such as glass, in which plants requiring regulated climatic conditions are grown.These ...
s by buzz pollination whereas other pollinators cannot. Commercial production began in 1987, when Roland De Jonghe founded the Biobest company; in 1988 they produced enough nests to pollinate 40 hectares of tomatoes. The industry grew quickly, starting with other companies in the Netherlands. Bumblebee nests, mainly of buff-tailed bumblebees, are produced in at least 30 factories around the world; over a million nests are grown annually in Europe; Turkey is a major producer.
Bumblebees are Northern Hemisphere animals. When red clover was introduced as a crop to New Zealand in the nineteenth century, it was found to have no local pollinators, and clover seed had accordingly to be imported each year. Four species of bumblebee from the United Kingdom were therefore imported as pollinators. In 1885 and 1886 the Canterbury Acclimatization Society brought in 442 queens, of which 93 survived and quickly multiplied. As planned, red clover was soon being produced from locally-grown seed. Bumblebees are also reared commercially to pollinate tomatoes grown in greenhouse
A greenhouse (also called a glasshouse, or, if with sufficient heating, a hothouse) is a structure with walls and roof made chiefly of transparent material, such as glass, in which plants requiring regulated climatic conditions are grown.These ...
s. The New Zealand population of buff-tailed bumblebees began colonising Tasmania
)
, nickname =
, image_map = Tasmania in Australia.svg
, map_caption = Location of Tasmania in AustraliaCoordinates:
, subdivision_type = Country
, subdi ...
, away, after being introduced there in 1992 under unclear circumstances.
Some concerns exist about the impact of the international trade in mass-produced bumblebee colonies. Evidence from Japan and South America indicates bumblebees can escape and naturalise in new environments, causing damage to native pollinators. Greater use of native pollinators, such as '' Bombus ignitus'' in China and Japan, has occurred as a result. In addition, mounting evidence indicates mass-produced bumblebees may also carry diseases, harmful to wild bumblebees
* and honeybees.
In Canada and Sweden it has been shown that growing a mosaic of different crops encourages bumblebees and provides higher yields than does a monoculture of oilseed rape, despite the fact that the bees were attracted to the crop.Goulson, 2013. pp. 169–172
Population decline
Bumblebee species are declining in Europe, North America, and Asia due to a number of factors, including land-use change that reduces their food plants. In North America, pathogens are possibly having a stronger negative effect especially for the subgenus ''Bombus''. A major impact on bumblebees was caused by the mechanisation of agriculture, accelerated by the urgent need to increase food production during the Second World War. Small farms depended on horses to pull implements and carts. The horses were fed on clover and hay, both of which were permanently grown on a typical farm. Little artificial fertiliser was used. Farms thus provided flowering clover and flower-rich meadows, favouring bumblebees. Mechanisation removed the need for horses and most of the clover; artificial fertilisers encouraged the growth of taller grasses, outcompeting the meadow flowers. Most of the flowers, and the bumblebees that fed on them, disappeared from Britain by the early 1980s. The last native British short-haired bumblebee was captured near Dungeness in 1988. This significant increase in pesticide and fertilizer use associated with the industrialization of agriculture has had adverse effects on the genus ''Bombus''. The bees are directly exposed to the chemicals in two ways: by consuming nectar that has been directly treated with pesticide, or through physical contact with treated plants and flowers. The species '' Bombus hortorum'' in particular has been found to be affected by the pesticides; their brood development has been reduced and their memory has been negatively affected. Additionally, pesticide use negatively affects colony development and size. Bumblebees are in danger in many developed countries due tohabitat destruction
Habitat destruction (also termed habitat loss and habitat reduction) is the process by which a natural habitat becomes incapable of supporting its native species. The organisms that previously inhabited the site are displaced or dead, thereby ...
and collateral pesticide damage. The European Food Safety Authority ruled that three neonicotinoid pesticides ( clothianidin, imidacloprid, and thiamethoxam
Thiamethoxam is the ISO common name for a mixture of ''cis-trans'' isomers used as a systemic insecticide of the neonicotinoid class. It has a broad spectrum of activity against many types of insects and can be used as a seed dressing.
Histo ...
) presented a high risk for bees. While most work on neonicotinoid toxicity has looked at honeybees, a study on ''B. terrestris'' showed that "field-realistic" levels of imidacloprid significantly reduced growth rate and cut production of new queens by 85%, implying a "considerable negative effect" on wild bumblebee populations throughout the developed world.
However, in another study, following chronic exposure to field-realistic levels of the neonicotinoid pesticide thiamethoxam, colony weight gain was not affected, nor were the number or mass of sexuals produced. Low levels of neonicotinoids can reduce the number of bumblebees in a colony by as much as 55%, and cause dysfunction in the bumblebees' brains. The Bumblebee Conservation Trust considers this evidence of reduced brain function "particularly alarming given that bumblebees rely upon their intelligence to go about their daily tasks." A study on ''B. terrestris'' had results that suggests that use of neonicotinoid pesticides can affect how well bumblebees are able to forage and pollinate. Bee colonies that had been affected by the pesticide released more foragers and collected more pollen than bees who had not been dosed with neonicotinoid. Although the bees affected by the pesticide were able to collect more pollen, they took a longer amount of time doing so.
Of 19 species of native nestmaking bumblebees and six species of cuckoo bumblebees formerly widespread in Britain, three have been extirpated, eight are in serious decline, and only six remain widespread. Similar declines have been reported in Ireland, with four species designated endangered
An endangered species is a species that is very likely to become extinct in the near future, either worldwide or in a particular political jurisdiction. Endangered species may be at risk due to factors such as habitat loss, poaching and inv ...
, and another two considered vulnerable
Vulnerable may refer to:
General
* Vulnerability
* Vulnerability (computing)
* Vulnerable adult
* Vulnerable species
Music
Albums
* ''Vulnerable'' (Marvin Gaye album), 1997
* ''Vulnerable'' (Tricky album), 2003
* ''Vulnerable'' (The Used album) ...
to extinction. A decline in bumblebee numbers could cause large-scale changes to the countryside, resulting from inadequate pollination of certain plants.
Some bumblebees native to North America are also vanishing, such as '' Bombus balteatus'', '' Bombus terricola'', '' Bombus affinis'', and '' Bombus occidentalis'', and one, ''Bombus franklini
Franklin's bumblebee (''Bombus franklini'') is known to be one of the most narrowly distributed bumblebee species, making it a critically endangered bee of the western United States. It is known only from a area in southern Oregon and northern C ...
'', may be extinct. In South America, ''Bombus bellicosus'' was extirpated in the northern limit of its distribution range, probably due to intense land use and climate change effects.
Conservation efforts
In 2006 the bumblebee researcher Dave Goulson
Dave Goulson (born 30 July 1965) is Professor of Biology (Evolution, Behaviour and Environment) at the University of Sussex. Specializing in the ecology and conservation of insects, particularly bumblebees, Goulson is the author of several boo ...
founded a registered charity, the Bumblebee Conservation Trust, to prevent the extinction "of any of the UK's bumblebees." In 2009 and 2010, the Trust attempted to reintroduce the short-haired bumblebee, ''Bombus subterraneus
The short-haired bumblebee (''Bombus subterraneus''), or short-haired humble-bee, is a species of bumblebee found in Eurasia, as well as in New Zealand, where it is an introduced species.
It lived in the British Isles among other parts of Europe ...
'', which had become extinct in Britain, from the British-derived populations surviving in New Zealand from their introduction there a century earlier. From 2011 the Trust, in partnership with Natural England
Natural England is a non-departmental public body in the United Kingdom sponsored by the Department for Environment, Food and Rural Affairs. It is responsible for ensuring that England's natural environment, including its land, flora and fauna ...
, Hymettus and the RSPB, has reintroduced short-haired bumblebee queens from Skåne
Scania, also known by its native name of Skåne (, ), is the southernmost of the historical provinces (''landskap'') of Sweden. Located in the south tip of the geographical region of Götaland, the province is roughly conterminous with Skåne ...
in southern Sweden to restored flower-rich meadows at Dungeness in Kent. The queens were checked for mites and American foulbrood disease. Agri-environment schemes spread across the neighbouring area of Romney Marsh have been set up to provide over 800 hectares of additional flower-rich habitat for the bees. By the summer of 2013, workers of the species were found near the release zone, proving that nests had been established. The restored habitat has produced a revival in at least five "Schedule 41 priority" species: the ruderal bumblebee, '' Bombus ruderatus''; the red-shanked carder bee, '' Bombus ruderarius''; the shrill carder bee, ''Bombus sylvarum
''Bombus sylvarum'', the shrill carder bee or knapweed carder-bee, is a species of bumblebee with a wide distribution across Europe, east to the Ural Mountains, and north to Great Britain, Ireland, and southern Scandinavia.
Description
It is a ...
''; the brown-banded carder bee, ''Bombus humilis
The brown-banded carder bee (''Bombus humilis'') is a bumblebee found in most of Europe west of Russia, with the exception of Ireland and Iceland. It is also found in Turkey, on the Tibetan plateau, northern China, eastern and southern Mongolia, ...
'' and the moss carder bee, ''Bombus muscorum
''Bombus muscorum'', commonly known as the large carder bee or moss carder bee, is a species of bumblebee in the family Apidae. The species is found throughout Eurasia in fragmented populations, but is most commonly found in the British Isles. '' ...
''.
The world's first bumblebee sanctuary was established at Vane Farm in the Loch Leven National Nature Reserve in Scotland in 2008. In 2011, London's Natural History Museum
A natural history museum or museum of natural history is a scientific institution with natural history collections that include current and historical records of animals, plants, fungi, ecosystems, geology, paleontology, climatology, and more ...
led the establishment of an International Union for Conservation of Nature
The International Union for Conservation of Nature (IUCN; officially International Union for Conservation of Nature and Natural Resources) is an international organization working in the field of nature conservation and sustainable use of natur ...
Bumblebee Specialist Group, chaired by Dr. Paul H. Williams, to assess the threat status of bumblebee species worldwide using Red List criteria.
Bumblebee conservation is in its infancy in many parts of the world, but with the realization of the important part they play in pollination of crops, efforts are being made to manage farmland better. Enhancing the wild bee population can be done by the planting of wildflower strips, and in New Zealand, bee nesting boxes have achieved some success, perhaps because there are few burrowing mammals to provide potential nesting sites in that country.
Misconception about flight
According to 20th-century folklore
Folklore is shared by a particular group of people; it encompasses the traditions common to that culture, subculture or group. This includes oral traditions such as Narrative, tales, legends, proverbs and jokes. They include material culture, r ...
, the laws of aerodynamics prove the bumblebee should be incapable of flight
Flight or flying is the process by which an object moves through a space without contacting any planetary surface, either within an atmosphere (i.e. air flight or aviation) or through the vacuum of outer space (i.e. spaceflight). This can b ...
, as it does not have the capacity (in terms of wing size or beats per second) to achieve flight with the degree of wing loading necessary.
The origin of this claim has been difficult to pin down with any certainty. John H. McMasters
John H. McMasters (April 26, 1939 – February 13, 2008) was an aeronautical engineer notable for his contributions to aerodynamics and engineering education.
Early life and education
McMasters was born in Taft, California and fell in love wit ...
recounted an anecdote about an unnamed Swiss aerodynamicist at a dinner party who performed some rough calculations and concluded, presumably in jest, that according to the equations, bumblebees cannot fly. In later years, McMasters backed away from this origin, suggesting there could be multiple sources, and the earliest he has found was a reference in the 1934 book ' by French entomologist Antoine Magnan (1881–1938); they had applied the equations of air resistance to insects and found their flight was impossible, but "One shouldn't be surprised that the results of the calculations don't square with reality".
The following passage appears in the introduction to ''Le Vol des Insectes'':
Magnan refers to his assistant André Sainte-Laguë. Some credit physicist Ludwig Prandtl (1875–1953) of the University of Göttingen
The University of Göttingen, officially the Georg August University of Göttingen, (german: Georg-August-Universität Göttingen, known informally as Georgia Augusta) is a public research university in the city of Göttingen, Germany. Founded i ...
in Germany with popularizing the idea. Others say Swiss gas dynamicist Jakob Ackeret (1898–1981) did the calculations.
The calculations that purported to show that bumblebees cannot fly are based upon a simplified linear treatment of oscillating
Oscillation is the repetitive or periodic variation, typically in time, of some measure about a central value (often a point of equilibrium) or between two or more different states. Familiar examples of oscillation include a swinging pendulum ...
aerofoils. The method assumes small amplitude oscillations without flow separation. This ignores the effect of dynamic stall (an airflow separation inducing a large vortex
In fluid dynamics, a vortex ( : vortices or vortexes) is a region in a fluid in which the flow revolves around an axis line, which may be straight or curved. Vortices form in stirred fluids, and may be observed in smoke rings, whirlpools in t ...
above the wing), which briefly produces several times the lift of the aerofoil in regular flight. More sophisticated aerodynamic analysis shows the bumblebee can fly because its wings encounter dynamic stall in every oscillation
Oscillation is the repetitive or Periodic function, periodic variation, typically in time, of some measure about a central value (often a point of Mechanical equilibrium, equilibrium) or between two or more different states. Familiar examples o ...
cycle.
Additionally, John Maynard Smith
John Maynard Smith (6 January 1920 – 19 April 2004) was a British theoretical and mathematical evolutionary biologist and geneticist. Originally an aeronautical engineer during the Second World War, he took a second degree in genetics ...
, a noted biologist with a strong background in aeronautics, has pointed out that bumblebees would not be expected to sustain flight, as they would need to generate too much power given their tiny wing area. However, in aerodynamics experiments with other insects, he found that viscosity
The viscosity of a fluid is a measure of its resistance to deformation at a given rate. For liquids, it corresponds to the informal concept of "thickness": for example, syrup has a higher viscosity than water.
Viscosity quantifies the inte ...
at the scale of small insects meant even their small wings can move a very large volume of air relative to their size, and this reduces the power required to sustain flight by an order of magnitude.
In music and literature
The orchestral interlude '' Flight of the Bumblebee'' was composed (c. 1900) by Nikolai Rimsky-Korsakov
Nikolai Andreyevich Rimsky-Korsakov . At the time, his name was spelled Николай Андреевичъ Римскій-Корсаковъ. la, Nicolaus Andreae filius Rimskij-Korsakov. The composer romanized his name as ''Nicolas Rimsk ...
. It represents the turning of Prince Guidon into a bumblebee so he can fly away to visit his father, Tsar Saltan, in the opera ''The Tale of Tsar Saltan
The Tale of Tsar Saltan, of His Son the Renowned and Mighty Bogatyr Prince Gvidon Saltanovich, and of the Beautiful Princess-Swan ( rus, «Сказка о царе Салтане, о сыне его славном и могучем богаты ...
'', although the music may reflect the flight of a bluebottle rather than a bumblebee. The music inspired Walt Disney
Walter Elias Disney (; December 5, 1901December 15, 1966) was an American animator, film producer and entrepreneur. A pioneer of the American animation industry, he introduced several developments in the production of cartoons. As a film p ...
to feature a bumblebee in his 1940 animated musical '' Fantasia'' and have it sound as if it were flying in all parts of the theater. This early attempt at " surround sound" was excluded from the film in later showings.
In 1599, during the reign of Queen Elizabeth I
Elizabeth I (7 September 153324 March 1603) was Queen of England and Ireland from 17 November 1558 until her death in 1603. Elizabeth was the last of the five House of Tudor monarchs and is sometimes referred to as the "Virgin Queen".
Eli ...
, someone, possibly Tailboys Dymoke, published ''Caltha Poetarum: Or The Bumble Bee'', under the pseudonym "T. Cutwode". This was one of nine books censored under the Bishop's Ban issued by the Archbishop of Canterbury John Whitgift and the Bishop of London Richard Bancroft.
Emily Dickinson made a bumblebee the subject of her parody of Isaac Watts's well-known poem about honeybees, "How Doth the Little Busy Bee" (1715). Where Watts wrote "How skilfully she builds her cell! How neat she spreads the wax!", Dickinson's poem, "The Bumble-Bee's Religion" (1881), begins "His little Hearse-like Figure / Unto itself a Dirge / To a delusive Lilac / The vanity divulge / Of Industry and Morals / And every righteous thing / For the divine Perdition / of Idleness and Spring." The letter was said to have enclosed a dead bee.
In 1847, Ralph Waldo Emerson
Ralph Waldo Emerson (May 25, 1803April 27, 1882), who went by his middle name Waldo, was an American essayist, lecturer, philosopher, abolitionist, and poet who led the transcendentalist movement of the mid-19th century. He was seen as a cham ...
published his poem " The Humble-Bee".
The entomologist Otto Plath wrote ''Bumblebees and Their Ways'' in 1934. His daughter, the poet Sylvia Plath, wrote a group of poems about bees late in 1962, within four months of her suicide, transforming her father's interest into her poetry.
The scientist and illustrator Moses Harris (1731–1785) painted accurate watercolour drawings of bumblebees in his ''An Exposition of English Insects Including the Several Classes of Neuroptera, Hymenoptera, & Diptera, or Bees, Flies, & Libellulae (1776–80)''.
Bumblebees appear as characters, often eponymously, in children's books. The surname Dumbledore in the Harry Potter
''Harry Potter'' is a series of seven fantasy novels written by British author J. K. Rowling. The novels chronicle the lives of a young wizard, Harry Potter, and his friends Hermione Granger and Ron Weasley, all of whom are students a ...
series (1997–2007) is an old name for bumblebee. J. K. Rowling said the name "seemed to suit the headmaster, because one of his passions is music and I imagined him walking around humming to himself". J. R. R. Tolkien, in his poem '' Errantry'', also used the name Dumbledor, but for a large bee-like creature.
Among the many books for younger children are ''Bumble the Bee'' by Yvon Douran and Tony Neal (2014); ''Bertie Bumble Bee'' by K. I. Al-Ghani (2012); ''Ben the Bumble Bee: How do bees make honey?'' by Romessa Awadalla (2015); ''Bumble Bee Bob Has a Big Butt'' by Papa Campbell (2012); ''Buzz, Buzz, Buzz! Went Bumble-bee'' by Colin West (1997); ''Bumble Bee'' by Margaret Wise Brown (2000); ''How the Bumble Came to Bee'' by Paul and Ella Quarry (2012); ''The Adventures of Professor Bumble and the Bumble Bees'' by Stephen Brailovsky (2010). Among Beatrix Potter
Helen Beatrix Potter (, 28 July 186622 December 1943) was an English writer, illustrator, natural scientist, and conservationist. She is best known for her children's books featuring animals, such as '' The Tale of Peter Rabbit'', which was ...
's "little books", Babbity Bumble and other members of her nest appear in ''The Tale of Mrs. Tittlemouse
''The Tale of Mrs. Tittlemouse'' is a children's book written and illustrated by Beatrix Potter, and published by Frederick Warne & Co. in 1910. The tale is about housekeeping and insect pests in the home, and reflects Potter's own sense of tid ...
'' (1910).
Military
TheUnited States Naval Construction Battalions
United States Naval Construction Battalions, better known as the Navy Seabees, form the U.S. Naval Construction Force (NCF). The Seabee nickname is a heterograph of the initial letters "CB" from the words "Construction Battalion". Depending upon ...
adopted the bumblebee as their insignia in 1942.
See also
* ''Ophrys bombyliflora
''Ophrys bombyliflora'', the bumblebee orchid, is a species of ''Ophrys'' (bee orchid), native from the Mediterranean region from Portugal and the Canary Islands to Turkey and Lebanon. The genus name ' is from the Greek in reference to the hairy ...
'', the bumblebee orchid
Notes
References
Sources
* Abbott, Carl, and Bartlett, John. "Bumble Bees". Encarta Encyclopedia. 2004 ed. * Anon. "Bees". World Book Encyclopedia, 1998 ed. * Benton, Ted. ''Bumblebees''. New Naturalist Series (#98). Collins, 2006. * Freeman, Scott. ''Biological Science''. Upper Saddle River, 2002. * Goulson, Dave. ''Bumblebees: Their Behaviour and Ecology'', 2003.Oxford University Press
Oxford University Press (OUP) is the university press of the University of Oxford. It is the largest university press in the world, and its printing history dates back to the 1480s. Having been officially granted the legal right to print book ...
. .
* Goulson, Dave. ''A Sting in the Tale''. Jonathan Cape, 2013.
* Hasley, William D. "Bees". Collier's Encyclopedia, 1990 ed.
* Macdonald, Murdo. Bumblebees
'. Scottish Natural Heritage, 2003. * Macdonald, Murdo & Nisbet, G.
'
HBRG
2006. .
Supplement 2
(2007). * Michener, C.D. ''The Bees of the World''. Johns Hopkins University Press, 2000. * Schweitzer, Dale F. et al.
Conservation and Management of North American Bumble Bees
'. Washington D.C.: U.S. Forest Service, 2012.
External links
Bumblebees of the world – find species by region, species groups, colour pattern
nhm.ac.uk
Bumblebee Conservation Trust
IUCN's Bumblebee Specialist Group
''Bombus'' Identification Guide
Deciphering the Mystery of Bee Flight
{{good article Aposematic animals Articles containing video clips Extant Eocene first appearances Insects in culture