Bumblebee

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Bombus
Bumblebee October 2007-3a.jpg
Bombus terrestris
Scientific classification
Kingdom:Animalia
Phylum:Arthropoda
Class:Insecta
Order:Hymenoptera
Family:Apidae
Subfamily:Apinae
Tribe:Bombini
Genus:Bombus
Latreille, 1802
Diversity
> 250 species and subspecies
 
  (Redirected from Bumble bee)
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Bombus
Bumblebee October 2007-3a.jpg
Bombus terrestris
Scientific classification
Kingdom:Animalia
Phylum:Arthropoda
Class:Insecta
Order:Hymenoptera
Family:Apidae
Subfamily:Apinae
Tribe:Bombini
Genus:Bombus
Latreille, 1802
Diversity
> 250 species and subspecies

A bumblebee, also written bumble bee, is a member of the bee genus Bombus, in the family Apidae. The parasitic or "cuckoo" bumblebees have sometimes been classified as a subgenus or genus, Psithyrus, but are now usually treated as members of Bombus. Bombus is the only extant genus in the tribe Bombini, though a few extinct related genera (e.g., Calyptapis) are known from fossils. Over 250 species are known,[1] found primarily in higher latitudes or at higher altitude in the Northern Hemisphere, although they also occur in South America; however, a few lowland tropical species are known. European bumblebees have been introduced to New Zealand and Tasmania.

Bumblebees are social insects which form colonies with a single queen. Colonies are smaller than those of honey bees, consisting often of fewer than 50 individuals in a nest. Female bumblebees can sting, repeatedly, but generally ignore humans and other animals. "Cuckoo" bumblebees do not make nests; 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.

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. 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, which is modified to form a pollen basket: a bare shiny concave surface, surrounded by a fringe of hairs used to transport pollen, whereas in cuckoo bees, the hind leg is completely hairy with no basket, and pollen grains are wedged into the hairs for transport.

Like their relatives the honey bees, bumblebees feed on nectar, using their long hairy tongues to lap up the liquid; the proboscis is folded under the head for flight. Bumblebees gather pollen to feed their young. They forage using colour and spatial relationships to identify flowers to feed from. Some bumblebees exhibit "nectar robbing", 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.

Etymology and common names[edit]

Beatrix Potter called Babbity Bumble a "bumble bee" in The Tale of Mrs Tittlemouse, 1910.

The word "bumblebee" is a compound of "bumble" + "bee" — "bumble" meaning to hum, buzz, drone, or move ineptly or flounderingly.[2] The generic name Bombus, assigned by Pierre André Latreille in 1802, is derived from the Latin word for a buzzing or humming sound.[3]

According to the Oxford English Dictionary (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."[4] 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."[5] The latter term was used in A Midsummer Night's Dream (circa 1600) by William Shakespeare, "The honie-bags steale from the humble Bees."[6] 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" meant "beetle".[7] In On the Origin of Species (1859), Charles Darwin speculated about "humble-bees" and their interactions with other species:[8]

I have [...] reason to believe that humble-bees are indispensable to the fertilisation of the heartsease (Viola tricolor), for other bees do not visit this flower. From experiments which I have tried, I have found that the visits of bees, if not indispensable, are at least highly beneficial to the fertilisation of our clovers; but humble-bees alone visit the common red clover (Trifolium pratense), as other bees cannot reach the nectar.

However, "bumblebee" remained in use, for example in The Tale of Mrs. Tittlemouse (1910) by Beatrix Potter, "Suddenly round a corner, she met Babbitty Bumble--"Zizz, Bizz, Bizzz!" said the bumblebee." Since World War II "humblebee" has fallen into near-total disuse.[9]

Phylogeny[edit]

The bumblebee tribe Bombini is one of four groups of corbiculate bees (those with baskets) in the Apidae, the others being the Apini (honey bees), Euglossini (orchid bees), and Meliponini (stingless bees). The corbiculate bees are a monophyletic group. Advanced eusocial behaviour appears to have evolved twice in the group, giving rise to controversy, now largely settled, as to the phylogenetic origins of the four tribes. 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."[10] Their analysis, combining molecular, morphological and behavioural data, gives the following cladogram:[10]

Corbiculate bees


Apini (honeybees)



Euglossini (orchid bees)





Bombini (bumblebees)



Meliponini (stingless bees)




On this hypothesis, the molecular data suggest that the Bombini are 25 to 40 mya old, while the Meliponini (and thus the clade that includes the Bombini and Meliponini) are 81 to 96 mya old, about the same age as the corbiculate group.[10]

The fossil record for bees is very incomplete, but by 2012 some 11 specimens that might possibly be Bombini, some poorly documented, had been described; some (such as Calyptapis florrisantensis from Florissant, USA, and Oligoapis beskonakensis from Beskonak, Turkey) dated from the Oligocene.[11] In 2012 a fossil bumblebee, Bombus (Bombus) randeckensis was found in the Miocene rocks of the Randeck Maar in southwestern Germany and confidently placed in the subgenus Bombus.[11] In 2014, a new bumblebee, Bombus cerdanyensis, was described from Late Miocene lacustrine beds of La Cerdanya, Spain.[12]

Taxonomy[edit]

Further information: List of world bumblebee species

Over 250 species have been described. For an overview of the differences between bumblebees and other bees and wasps, see characteristics of common wasps and bees.

The genus Bombus has been divided variously into up to 49 subgenera, a degree of complexity criticised by Williams (2008).[13] 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.[13] Williams identifies 15 subgenera that are seen to be monophyletic, fewer in number, diagnosable from morphology, and at least roughly correspond to behavioural and ecological groupings:[13]

Distribution and habitat[edit]

Bumblebees are typically found in higher latitudes and/or high altitudes, though exceptions exist (a few lowland tropical species exist).[14] A few species (B. polais and B. alpinus) range into very cold climates where other bees might not be found; B. polaris can be found 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.[15] One reason for their presence in cold places is that bumblebees can regulate their body temperature, via solar radiation, internal mechanisms of "shivering" and radiative cooling from the abdomen (called heterothermy). Other bees have similar physiology, but the mechanisms seem best developed and have been most studied in bumblebees.[16] They adapt to higher elevations by extending their wing stroke amplitude.[17] They are absent from Australia, although they have been introduced to Tasmania, and are found in Africa only north of the Sahara.[18]

Biology[edit]

A common carder bumblebee Bombus pascuorum extending its tongue towards a Heuchera inflorescence

Feeding[edit]

Like all bee tongues, the bumblebee tongue (the proboscis) is a long, hairy structure that extends from a sheath-like modified maxilla. The primary action of the tongue is lapping, i.e. repeated dipping of the tongue into liquid.[19] During lapping, nectar is drawn up the proboscis by capillary action. When at rest or flying, the proboscis is kept folded under the head.

Wax production[edit]

The exoskeleton of the abdomen is divided into plates called dorsal tergites and ventral sternites. Wax is secreted from glands 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 maleable 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.[20]

Coloration: aposematism and mimicry[edit]

Cuckoo bumblebees, like this Bombus barbutellus, have similar aposematic (warning) coloration to nest-making bumblebees, and may also mimic their host species.

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.[21] Dipteran flies in the families Syrphidae (hoverflies), Asilidae (robber flies), Tabanidae (horseflies), Oestridae (bot or warble flies) and Bombyliidae (bee flies) all include Batesian mimics of bumblebees, resembling them closely enough to deceive at least some predators.[22]

Many species of Bombus, including the group sometimes called Psithyrus (cuckoo bumblebees), have evolved Müllerian mimicry, 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, P. insularis and P. fernaldae. Other bees in California include a group of species all banded black and yellow. In each case, Mullerian mimicry provides the bees in the group with a selective advantage.[22] 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.[23]

Temperature control[edit]

Bumblebees are active in conditions when honey bees stay at home, and can readily absorb heat from even weak sunshine.[24] 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.[25] The temperature of the flight muscles, which occupy much of the thorax, needs to be at least 30 °C (86 °F) 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 13 °C (55 °F).[26]

Communication[edit]

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.[20]

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. Rhese 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.[27] Another stimulant to foraging activity is the levels of food reserve in the colony. Bees monitor the level of honey in the honeypots, and when the level is low, or when high quality food is added, they are more likely to go out to forage.[28]

Reproduction and nesting[edit]

Nest of red-tailed bumblebee. Bombus lapidarius, showing wax pots full of honey
Further information: haplodiploidy and worker policing

Bumblebees form colonies of up to about 400 individuals; this is small compared to honeybee hives which hold about 50,000 bees. Many species nest underground, choosing old rodent burrows or sheltered places, and avoiding places that receive direct sunlight which 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 a honeybee hive; 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.[29]

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 fertilised eggs from the previous winter. 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 honey bee workers. In temperate zone species, in the autumn, young queens or gynes mate with males (drones) and die as the weather turns colder. The new, young queens leave the nest, mate, and feed intensively to build up stores of fat for the winter. These new queens survive in a resting state (diapause), generally below ground, during the winter.[30]

In fertilised queens, the ovaries are activated when the queen lays her egg. It passes along the oviduct to the vagina. In the vagina is a chamber called the spermatheca, where the queen stores sperm from her mating. The queen, depending on need, may allow her egg to be fertilised. Unfertilised eggs become haploid males; fertilised eggs grow into diploid females and queens.[31] The hormones that stimulate the development of the ovaries are suppressed in female worker bees, while the queen remains dominant. Before hibernation, queens eat as much as they can to increase their store of body fat; it is used up during hibernation.[32]

A queen buff-tailed bumblebee, Bombus terrestris, digging a nest hole

To develop, the larvae must be fed both nectar for carbohydrates and pollen for protein. 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.[33]

An above-ground nest, hidden in grass and moss, of the common carder bee, Bombus pascuorum. The wax canopy or involucrum has been removed to show winged workers and pupae in irregularly placed wax cells.

After the emergence of the first or second group of workers, 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.[32] Unlike the workers of more advanced social insects such as honeybees, bumblebee workers are fertile, and can lay unfertilised haploid eggs (with only a single set of chromosomes) 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.[34]

In a young colony, the queen minimises reproductive competition from workers by suppressing their egg-laying through physical aggression and pheromones.[35] Worker policing leads to nearly all eggs laid by workers being eaten.[36] 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.[37] Because of the reproductive competition between workers and the queen, bumblebees are considered "primitively eusocial".[10][36]

New queens and males leave the colony after maturation. Males in particular are forcibly driven out by the workers. The queens mate, often more than once, and find suitable locations for diapause, digging themselves a shallow burrow in the ground.[32]

Foraging behaviour[edit]

Further information: Bumblebee communication
A bumblebee loaded with pollen in its pollen baskets

Bumblebees generally visit flowers that exhibit the bee pollination syndrome. They can visit patches of flowers up to 1–2 km from their colony.[38] They tend to visit the same patches of flowers every day, as long as they continue to find nectar and pollen,[39] a habit known as pollinator or flower constancy. While foraging, bumblebees can reach ground speeds of up to 15 metres per second (54 km/h).[40]

Biting open the stem of a flower...
...and using its tongue to drink the nectar.
A bumblebee "nectar robbing" a flower

Bumblebees use a combination of colour and spatial relationships to learn from which flowers to forage.[41] They can also detect both the presence and the pattern of electric fields on flowers, which occur due to the positive static charges generated when bees fly through the air (see 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.[42] After arriving at a flower, they extract nectar using their long tongues ("glossae") and store it in their crops. Many species of bumblebees also exhibit "nectar robbing": instead of inserting the mouthparts into the flower normally, these bees bite directly through the base of the corolla to extract nectar, avoiding pollen transfer.[43] These bees obtain pollen from other species of flowers which they "legitimately" visit.

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. The bumblebee's body hairs receive a dusting of pollen from the anthers, which is then groomed into the corbicula ("pollen basket"). Bumblebees are also capable of buzz pollination.[44]

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.[45] This scent mark is a general chemical bouquet that bumblebees leave behind in different locations (e.g. nest, neutral, and food sites),[46] and they learn to use this bouquet to identify both rewarding and unrewarding flowers.[47] Bumblebees rely on this chemical bouquet more when the flower has a high handling time (i.e. where it takes a longer time for the bee to find the nectar once inside the flower).[48]

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 honey bees, bumblebees only store a few days' worth of food, so are much more vulnerable to food shortages.[49] 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.[50]

Cuckoo bumblebees[edit]

The cuckoo bumblebee B. vestalis, a parasite of B. terrestris

Bumblebees of the subgenus Psithyrus (known as 'cuckoo bumblebees', and formerly considered a separate genus) are a lineage that live parasitically in the colonies of other bumblebees and have lost the ability to collect pollen. Before finding and invading a host colony, a Psithyrus female (there is no caste system in these species) feeds directly from flowers. Once she has infiltrated a host colony, the Psithyrus female kills or subdues the queen of that colony and forcibly (using pheromones and/or physical attacks) "enslaves" the workers of that colony to feed her and her young.[51] The female Psithyrus also 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.[52] 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 then their host species. Each species of cuckoo bee has a specific host species, which it may physically resemble.[53]

Sting[edit]

Queen and worker bumblebees can sting. Unlike in honey bees, a bumblebee's stinger lacks barbs, so it can sting repeatedly without injuring itself; by the same token, the stinger is not left in the wound.[54][55] 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,[56] but ignore other animals (e.g. humans) unless disturbed.

Predators, parasites and pathogens[edit]

Bumblebee nest dug up and destroyed by a predator, probably a badger

Bumblebees, despite their ability to sting, are eaten by certain predators. Nests may be dug up by badgers and eaten whole, including any adults present.[57] Adults are preyed upon by robber flies and beewolves in North America.[58] In Europe, birds including bee-eaters and shrikes capture adult bumblebees on the wing; smaller birds such as great tits also occasionally learn to take bumblebees, while camouflaged crab spiders catch them as they visit flowers.[59]

Bumblebees are parasitised by tracheal mites, Locustacarus buchneri; protozoans including Crithidia bombi and Apicystis bombi; and microsporidians including Nosema bombi and Nosema ceranae. The Tree bumblebee Bombus hypnorum has spread into the United Kingdom despite hosting high levels of a nematode that normally interferes with queen bees' attempts to establish colonies.[60]

Deformed wing virus has been found to affect 11% of bumblebees in Great Britain.[61]

Bumblebees and people[edit]

Bumblebees and human culture: Bombus anachoreta on a Russian postage stamp, 2005

Agricultural use[edit]

Bumblebees are important pollinators of both crops and wildflowers.[62] Because bumblebees do not overwinter the entire colony, they do not stockpile honey, so 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 in greenhouses that other pollinators cannot by buzz pollination.[63]

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 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.[64] Bumblebees are also reared commercially to pollinate tomatoes grown in greenhouses.[31] The New Zealand population of buff-tailed bumblebees naturally colonised Tasmania, 1,500 miles away, in 1992.[65]

Some concerns exist about the impact of the international trade in mass-produced bumblebee colonies. Evidence from Japan[66] and South America[67] indicates bumblebees can escape and naturalise in new environments, causing damage to native pollinators. In addition, mounting evidence indicates mass-produced bumblebees may also carry diseases, harmful to wild bumblebees[68][69] and honeybees.[69]

Endangered status[edit]

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 Bombus str. in North America.[70] Bumblebees are in danger in many developed countries due to habitat destruction and collateral pesticide damage. The European Food Safety Authority ruled that three neonicotinoid pesticides (clothianidin, imidacloprid, and thiamethoxam) presented a high risk for bees.[71] In Britain, until relatively recently, 19 species of native nestmaking bumblebees were recognised along with six species of cuckoo bumblebees.[72] Of these, three have been extirpated,[73][74] eight are in serious decline, and only six remain widespread.[75] Similar declines in bumblebees have been reported in Ireland, with four species being designated endangered, and another two species considered vulnerable to extinction.[76] A decline in bumblebee numbers could cause large-scale changes to the countryside, resulting from inadequate pollination of certain plants. The world's first bumblebee sanctuary was established at Vane Farm in the Loch Leven National Nature Reserve in Scotland in 2008.[77]

Some bumblebees native to North America are also vanishing, such as Bombus terricola, Bombus affinis, and Bombus occidentalis, and one, Bombus franklini, may even be extinct.[78] 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.[79]

In 2011, London's Natural History Museum led the establishment of an International Union for the Conservation of Nature Bumblebee Specialist Group, chaired by Dr. Paul H. Williams,[80] to assess the threat status of bumblebee species worldwide using Red List criteria.[81]

In 2006 the bumblebee researcher Dave Goulson founded a registered charity, the Bumblebee Conservation Trust, to prevent the extinction "of any of the UK's bumblebees."[82][83] He and the Trust have attempted to reintroduce the short-haired bumblebee, which had become extinct in Britain, from the populations surviving in New Zealand from their introduction there a century earlier.[84]

Misconception about flight[edit]

A bumblebee landing on a purple flower
A widely believed falsehood holds that scientists proved that bumblebees are incapable of flight[85]

According to 20th-century folklore, the laws of aerodynamics prove the bumblebee should be incapable of flight, 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.

Supposedly someone did a back of the envelope calculation, taking the weight of a bumblebee and its wing area into account, and worked out that if it only flies at a couple of metres per seconds, the wings wouldn’t produce enough lift to hold the bee up,’ explains Charlie Ellington, Professor of Animal Mechanics at Cambridge University.[86]

The origin of this claim has been difficult to pin down with any certainty. John H. McMasters 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.[87] In later years, McMasters has backed away from this origin, suggesting there could be multiple sources, and the earliest he has found was a reference in the 1934 book Le Vol des Insectes 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".[88]

The following passage appears in the introduction to Le Vol des Insectes:[89]

Tout d'abord poussé par ce qui se fait en aviation, j'ai appliqué aux insectes les lois de la résistance de l'air, et je suis arrivé avec M. Sainte-Laguë à cette conclusion que leur vol est impossible.

This translates to:

First prompted by what is done in aviation, I applied the laws of air resistance to insects, and I arrived, with Mr. Sainte-Laguë, at this conclusion that their flight is impossible.

Magnan refers to his assistant André Sainte-Laguë.[90] Some credit physicist Ludwig Prandtl (1875–1953) of the University of Göttingen in Germany with popularizing the idea. Others say Swiss gas dynamicist Jacob Ackeret (1898–1981) did the calculations.[91]

Bumblebee in flight. It has its tongue extended and a laden pollen basket.

The calculations that purported to show that bumblebees cannot fly are based upon a simplified linear treatment of oscillating aerofoils. The method assumes small amplitude oscillations without flow separation. This ignores the effect of dynamic stall (an airflow separation inducing a large vortex 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 cycle.[92]

Additionally, John Maynard Smith, 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 at the scale of small insects meant even their small wings can move a very large volume of air relative to the size, and this reduces the power required to sustain flight by an order of magnitude.[93]

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 nerves cannot fire 200 times per second. These types of muscles are called asynchronous muscles[94] and are often found in insect wings. Bumblebees must warm up their bodies considerably to get airborne at low ambient temperatures.[16] Bumblebees have been known to reach an internal thoracic temperature of 30 °C (86 °F) using this method.

In music and literature[edit]

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Flight of the Bumblebee performed by the US Army Band

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The Russian composer Nikolai Rimsky-Korsakov wrote the Flight of the Bumblebee, c. 1900

The orchestral interlude Flight of the Bumblebee was composed (c. 1900) by Nikolai Rimsky-Korsakov to represent 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,[95] although the music may reflect the flight of a bluebottle rather than a bumblebee.[96] The music inspired Walt Disney 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 unsuccessful, and the music was excluded from the film's release.[97]

In 1599, during the reign of Queen Elizabeth I, someone, possibly Tailboys Dymoke, published Caltha Poetarum: Or The Bumble Bee, under the pseudonym "T. Cutwode".[98] 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.[99]

Emily Dickinson wrote a poem about a bumblebee. Daguerreotype, c. 1848

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!",[100] 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.[101][102]

Bumblebees appear as characters, often eponymously, in children's books. The surname Dumbledore in the Harry Potter series (1997–2007) is an old name for bumblebee.[7] 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".[103] 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's "little books", Babbity Bumble and other members of her nest appear in The Tale of Mrs Tittlemouse (1910).

See also[edit]

References[edit]

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