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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; 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.
Meliponini stingless bees. 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, genes of ten corbiculate bee species supports the single origin of eusociality hypothesis in the corbiculate bees.
However, Romiguier et al. 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 incomplete. Around 11 specimens that might possibly be Bombini, some poorly documented, had been described by ; some such as Calyptapis florissantensis from Florissant , USA, and Oligoapis beskonakensis from Beskonak , Turkey dated from the Oligocene.
The genus Bombus , the only one extant genus in the tribe Bombini, comprises over 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 Examples of Bombus species include Bombus atratus , Bombus dahlbomii , Bombus fervidus , Bombus lapidarius , Bombus ruderatus , and Bombus rupestris.
Mendacibombus , 12 species. Kallobombus , 1 species. Orientalibombus , 3 species. Subterraneobombus , 10 species. Megabombus , 22 species.
Thoracobombus , 50 species. Alpinobombus , 5 species. Bombus subgenus , 5 species. Melanobombus , 17 species. Sibiricobombus , 7 species. Cullumanobombus , 23 species.
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 are typically found in temperate climates , and are often found at higher latitudes and altitudes than other bees, although a few lowland tropical species exist.
This is the northernmost occurrence of any eusocial insect. Other bees have similar physiology , but the mechanisms seem best developed and have been most studied in bumblebees.
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, that is, repeated dipping of the tongue into liquid.
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.
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 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.
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.
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. 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. Bumblebees are active under conditions during which honeybees stay at home, and can readily absorb heat from even weak sunshine.
The muscle temperature can be raised by shivering. 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. 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. 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.
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 study involving Bombus terrestris , bees were taught to complete an unnatural task of moving large objects to obtain a reward.
Bees that first observed another bee complete the task were significantly more successful in learning the task than bees that observed the same action performed by a magnet, indicating the importance of social information.
The bees did not copy each other exactly: in fact, the study suggested that the bees were instead attempting to emulate each other's goals.
Nest size depends on species of bumblebee. Many species nest underground, choosing old rodent 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 drones that are forcibly driven out of the colony.
They survive in a resting state diapause , generally below ground, until the weather warms up in the spring with the early bumblebee being the species that is among the first to emerge.
Bombus pensylvanicus is a species that follows this type of colony cycle. 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 to the vagina where there is a chamber called the spermatheca , 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 females and queens.
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. 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. 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.
Workers eventually begin to lay male eggs later in the season when the queen's ability to suppress their reproduction diminishes.
Although a large majority of bumblebees follow such monogynous colony cycles that only involve one queen, some select Bombus species such as Bombus atratus will spend part of their life cycle in a polygynous phase have multiple queens in one nest during these periods of polygyny.
Bumblebees use a combination of colour and spatial relationships to learn which flowers to forage from. They use this information to find out if a flower has been recently visited by another bee.
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 in the normal way, these bees bite directly through the base of the corolla 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. 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. 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.
They may visit quite different flowers from the workers because of their different nutritional needs. Bees beat their wings about 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 times per second. 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. 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.
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.
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.
Queen and worker bumblebees can sting. Unlike in honeybees, a bumblebee's stinger lacks barbs, so the bee can sting repeatedly without injuring itself; by the same token, the stinger is not left in the wound.
Female cuckoo bumblebees aggressively attack host colony members, and sting the host queen, but ignore other animals unless disturbed.
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.
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 B. Female bee moths Aphomia sociella prefer to lay their eggs in bumblebee nests. The A. Bumblebees are important pollinators of both crops and wildflowers.
Bumblebees are increasingly cultured for agricultural use as pollinators, among other reasons because they can pollinate plants such as tomato in greenhouses by buzz pollination whereas other pollinators cannot.
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 and the Canterbury Acclimatization Society brought in queens, of which 93 survived and quickly multiplied.
As planned, red clover was soon being produced from locally-grown seed. 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 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.
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. 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 s. The last native British short-haired bumblebee was captured near Dungeness in 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 impacted by the pesticides; their brood development has been reduced and their memory has been negatively affected.
Additionally, pesticide use negatively impacts colony development and size. 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.
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.
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.
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.
Some bumblebees native to North America are also vanishing, such as Bombus balteatus ,  Bombus terricola ,  Bombus affinis ,   and Bombus occidentalis , and one, Bombus franklini , may be extinct.
In the bumblebee researcher Dave Goulson founded a registered charity, the Bumblebee Conservation Trust , to prevent the extinction "of any of the UK's bumblebees.
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 hectares of additional flower-rich habitat for the bees.
By the summer of , 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 ; the brown-banded carder bee, Bombus humilis and the moss carder bee, Bombus muscorum.
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.
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.
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.
The following passage appears in the introduction to Le Vol des Insectes : . First prompted by what is done in aviation, I applied the laws of air resistance to insects, and I arrived, with Mr.
Others say Swiss gas dynamicist Jakob Ackeret — did the calculations. 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.
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 their size, and this reduces the power required to sustain flight by an order of magnitude.
The orchestral interlude Flight of the Bumblebee was composed c. 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 ,  although the music may reflect the flight of a bluebottle rather than a bumblebee.
This early attempt at " surround sound " was unsuccessful, and the music was excluded from the film's release. Where Watts wrote "How skilfully she builds her cell!
How neat she spreads the wax! Bumblebees appear as characters, often eponymously, in children's books. The surname Dumbledore in the Harry Potter series — is an old name for bumblebee.
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".
Tolkien , in his poem Errantry , also used the name Dumbledor, but for a large bee-like creature. Tittlemouse From Wikipedia, the free encyclopedia.
Genus of insect. For the film, see Bumblebee film. For other uses, see Bumblebee disambiguation. For other uses, see Bombus disambiguation.
Further information: List of bumblebee species. Further information: Characteristics of common wasps and bees. Further information: aposematism and mimicry.
Further information: haplodiploidy and worker policing. Further information: Bumblebee communication and nectar robbing. A bumblebee " nectar robbing " a flower.
Main article: Psithyrus. Further information: List of crop plants pollinated by bees. Further information: Insect flight.
Flight of the Bumblebee. DNA analysis was used to estimate how many colonies these individuals came from. Shorter Oxford English dictionary on historical principles.
Oxford [Oxfordshire]: Oxford University Press. Transactions of the Philological Society 6 : Bumblebee transformer.
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