Bumblebee
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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 genera (e.g., Calyptapis) are known from fossils. They are found primarily 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. Female bumblebees can sting repeatedly, but generally ignore humans and other animals.
Most bumblebees are eusocial insects that form colonies with a single queen. The colonies are smaller than those of honey bees, 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. 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 bumblebees 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, whereas in cuckoo bumblebees, the hind leg is hairy all around, 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 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, the mechanisation of agriculture, and pesticides.
Etymology
The English name bumblebee combines bumble, meaning to buzz or hum, with bee, both words imitative of the insect's sound.<ref name="OED"/> The generic name Bombus, introduced by Pierre André Latreille in 1802, derives from the Latin bombus ("buzzing" or "booming"), itself from Ancient Greek βόμβος (bómbos).<ref>Template:Cite book</ref>
Charles Darwin referred to bumblebees as "humble-bees" in his 1859 book On the Origin of Species, emphasising their importance in pollinating clovers and other wildflowers.<ref>Template:Cite book</ref> The term humblebee remained in use through the nineteenth century, but was gradually replaced by bumblebee in both scientific and common language during the twentieth century.<ref name="OED">Template:Cite dictionary</ref><ref name="Jones 2010">Template:Cite news</ref>
Evolution
Phylogeny
The tribe Bombini (bumblebees) is one of four lineages of corbiculate bees in the Apidae. It is distinguished by having pollen baskets (corbiculae) on their hind legs. The other three are the Apini (honeybees), Euglossini (orchid bees), and Meliponini (stingless bees). Together they form a monophyletic group.<ref name="Cardinal-2011">Template:Cite journal</ref>
Early analyses of morphology and behaviour suggested that complex eusociality evolved twice within the corbiculate bees, once in the ancestor of the Apini and once in the Meliponini. Molecular and morphological data indicate that the primitively eusocial Bombini are, despite appearances, closely related to the Meliponini, while the Apini and Euglossini form a separate branch. Cardinal and Danforth (2011) described this dual-origin model as consistent with earlier evidence from social and anatomical traits.<ref name="Cardinal-2011"/>
More recent phylogenomic studies based on the mRNA expression profile from more than 3,600 genes support instead a single origin of eusociality within the corbiculate bees. Romiguier et al. (2015) found that the Bombini, Meliponini, and Apini form a single clade, with Euglossini as their sister group, implying that advanced social organisation arose once in the common ancestor of these three tribes.<ref>Template:Cite journal</ref>
Revisiting the "corbiculate controversy",<ref name="Porto 2021"/> on the mismatch between morphological and molecular analyses in 2021, Diego Sasso Porto and Eduardo A. B. Almeida find that morphological analysis still suggests that Bombini are sister to a clade containing Apini and Meliponini, as shown in the phylogenetic tree. They propose "a more conciliatory"<ref name="Porto 2021"/> scenario involving "a diversification followed by several extinctions"<ref name="Porto 2021"/> so that Meliponini and Bombini shared "few apomorphic changes",<ref name="Porto 2021"/> while the similarities between Meliponini and Apini could be convergent evolution caused by "similar biology or similar eusocial behaviors".<ref name="Porto 2021">Template:Cite journal</ref>
The earliest divergence estimates suggest that the Bombini originated between 25 and 40 million years ago, whereas the Bombini–Meliponini clade dates to roughly 80–95 million years ago, around the time the corbiculate group as a whole diversified.<ref name="Cardinal-2011"/>
Fossil record
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Fossil evidence of Bombini is limited. By 2019, about 14 potential fossil species had been described.<ref name="Dehon-2019">Template:Cite journal</ref> The oldest known relatives include Calyptapis florissantensis from the Late Eocene Florissant Formation (United States) and Oligobombus cuspidatus from the Bembridge Marls (Isle of Wight).<ref>Template:Cite journal</ref>
Several Oligocene and Miocene species have been assigned to modern subgenera, including Bombus (Mendacibombus) beskonakensis and B. (Paraelectrobombus) patriciae from Turkey, B. randeckensis from Germany, B. cerdanyensis from Spain, and B. trophonius from the Czech Republic.<ref name="Wappler-2012">Template:Cite journal</ref><ref name="Prokop-2017">Template:Cite journal</ref><ref name="Dehon-2014">Template:Cite journal</ref>
These fossils indicate that the major subgenera of Bombus were already differentiated by the Miocene and that diversification of the tribe was well underway by the Oligocene.<ref name="Dehon-2019" />
Taxonomy
The genus Bombus, the only one extant genus in the tribe Bombini, comprises over 250 species;<ref>Template:Cite journal</ref> 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).<ref name="Williams-2008">Template:Cite journal</ref><ref>Template:Cite journal</ref> The cuckoo bumblebees Psithyrus have sometimes been treated as a separate genus but are now considered to be part of Bombus.<ref name="Williams-2008"/>
- Subgenera of the genus Bombus
Description
Bumblebees vary in appearance, but are characteristically plump and densely furry. They are larger, broader and stouter-bodied than honeybees, and the tip of the abdomen is more rounded. Many species have broad bands of colour, whose patterns help 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.<ref>Template:Cite web</ref> 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.<ref>Template:Cite web</ref> Sizes are very variable even within species; the largest British species, B. terrestris, has queens up to Template:Cvt long, males up to Template:Cvt long, and workers between Template:Cvt long.<ref>Template:Cite web</ref> The largest bumblebee species in the world is B. dahlbomii of Chile, up to about Template:Cvt long, and described as "flying mice" and "a monstrous fluffy ginger beast".<ref>Template:Cite news</ref>
Distribution and habitat
Abundance and diversity
Bumblebees are most abundant across temperate and montane ecosystems of the Northern Hemisphere, with the greatest species richness found in Eurasia and North America.<ref>Template:Cite web</ref> Bumblebees have a largely cosmopolitan distribution but are absent from Australia (apart from Tasmania, where they have been introduced<ref name="Larkins-2012">Template:Cite news</ref>) and occur in Africa only north of the Sahara.<ref name="BugGuide.Net">Template:Cite web</ref> More than a hundred years ago they were also introduced to New Zealand, where they play an important role as efficient pollinators.<ref>Template:Cite journal</ref>
At the northern limits of their range, species such as Bombus polaris, B. alpinus, and the parasitic B. hyperboreus occupy Arctic tundra ecosystems that extend as far north as Greenland and Canada's Ellesmere Island. These populations endure some of the planet's shortest flowering seasons and coldest conditions, marking the northernmost boundary of eusocial insect life.<ref>Template:Cite journal</ref>
Physiological adaptations
One reason for bumblebee presence in such cold places is that they can regulate their body temperature through several physiological mechanisms. They use solar radiation to warm themselves, generate heat internally through "shivering" known as heterothermy, and employ countercurrent exchange to retain heat.<ref>Template:Cite journal</ref> Although other bees show similar forms of thermoregulation, these mechanisms are especially well developed and extensively studied in bumblebees.<ref name="Heinrich-19812">Template:Cite book</ref> They also adapt to high-elevation environments by increasing their wing stroke amplitude to sustain flight in thin air.<ref>Template:Cite journal</ref>
Biology
Feeding
File:Yellowfaced lBumblebee foraging.webm
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 or repeated dipping of the tongue into liquid.<ref>Template:Cite journal</ref> 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.<ref>Template:Cite book</ref> Bees with shorter proboscides, like Bombus bifarius, 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.<ref>Template:Cite journal</ref>
Wax production
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.<ref name="Bumblebee.org">Template:Cite web</ref>
Coloration
The black-and-yellow coloration of bumblebees acts as an aposematic (warning) signal to predators, indicating that the insects can inflict a painful sting.<ref>Poulton, E. B. (1890). The Colours of Animals: Their Meaning and Use Especially Considered in the Case of Insects. London: Kegan Paul, Trench, Trübner & Co. pp. 100–104.</ref> This visual warning serves as an anti-predator adaptation that discourages birds and other vertebrates, representing a common example of aposematism among members of the Hymenoptera.<ref>Williams, P. (2007). "The distribution of bumblebee colour patterns worldwide". Biological Journal of the Linnean Society. 92 (1): 97–118. doi:10.1111/j.1095-8312.2007.00878.x.</ref> Depending on the species and morph, the warning colours range from entirely black, to bright yellow, red, orange, white, and pink.<ref name="Williams-2007">Template:Cite journal</ref> Dipteran flies in the families Syrphidae (hoverflies), Asilidae (robber flies), Tabanidae (horseflies), Oestridae (bot or warble flies) and Bombyliidae (bee flies, such as Bombylius major) all include Batesian mimics of bumblebees, resembling them closely enough to deceive at least some predators.<ref name="Thorp-1983">Template:Cite book</ref>
Many species of Bombus, including the Psithyrus group of cuckoo bumblebees, have evolved a form of Müllerian mimicry. This is a system in which different bumblebee species within the same region share similar warning color patterns, so that a predator only has to learn once to avoid any of them. In California, for example, several largely black species like B. californicus, B. caliginosus, B. vandykei, B. vosnesenskii, B. insularis, and B. fernaldae, form one such mimicry group. Another group in the same region includes species banded in black and yellow. In both cases, the shared coloration gives all members of the group a selective advantage.<ref name="Williams2007">Template:Cite journal</ref>
Parasitic, or cuckoo, bumblebees also tend to resemble their hosts more closely than would be expected by chance, especially in regions such as Europe where parasite and host species have evolved together. This resemblance is likely another case of Müllerian mimicry, rather than a strategy to deceive the host through aggressive mimicry.<ref>Fisher, R. M. & Sampson, J. (1992). "Mimetic coevolution between cuckoo bumblebees and their hosts". Evolution. 46 (3): 775–783.</ref>
Temperature control
Bumblebees are active under conditions during which honeybees and other smaller bees stay at home, and can readily absorb heat from even weak sunshine.<ref name="Macdonald-2003">Template:Harvnb</ref> 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.<ref>Template:Cite journal</ref> The temperature of the flight muscles, which occupy much of the thorax, needs to be at least Template:Convert 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 Template:Convert.<ref>Template:Cite web</ref>
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 Template:Convert. 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.<ref>Template:Cite journal</ref>
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 vibrations transmitted through substrates such as wood or other materials.<ref name="BeeCulture2016">Template:Cite web</ref>
Unlike the well-known "dance" communication of honeybees, bumblebees do not perform a spatial waggle or round dance to indicate the location of food sources. Instead, when a forager returns from a successful trip to the nest, it often moves rapidly and excitedly around the nest for several minutes before departing again to forage.<ref name="XercesBumbleBees">Template:Cite web</ref>
Bumblebees have also been observed to engage in social learning. In experiments with Bombus terrestris, workers were trained to solve a novel task (string-pulling) to obtain a reward, and naïve individuals that observed trained demonstrators were significantly more successful than those that did not.<ref name="Alem 2016">Template:Cite journal</ref> Furthermore, subsequent work showed that novel problem-solving behaviours can spread through bumblebee colonies via social learning under open-diffusion conditions.<ref name="Bridges2023">Template:Cite journal</ref>
Reproduction and nesting
Nest size depends on species of bumblebee. Most form colonies of between 50 and 400 individuals,<ref name="Bumblebee Conservation Trust-2"/> but colonies have been documented as small as ~20 individuals and as large as 1700.<ref>Template:Cite journal</ref> These nests are 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 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.<ref name="Bumblebee Conservation Trust-2">Template:Cite web</ref>
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.<ref name="Goulson-2013a"/> 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 being the species that is among the first to emerge.<ref name="Goulson-2013a"/><ref>Template:Cite journal</ref><ref name="Bumblebee Conservation Trust">Template:Cite web</ref> Many species of bumblebee follow this general trend within the year. Bombus pensylvanicus is a species that follows this type of colony cycle.<ref name="University of Wisconsin La Crosse-2013">Template:Cite web</ref> 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.<ref name="Di Trani de la Hoz-2006">Template:Cite journal</ref>
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.<ref name="Biobees Bumblebee Pollination">Template:Cite web</ref> The hormones that stimulate the development of the ovaries are suppressed in female worker bees, while the queen remains dominant.<ref name="Goulson-2013a"/>
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.<ref name="Evans-2007">Template:Cite book</ref>
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.<ref name="Goulson-2013a">Template:Harvnb</ref> Bumblebee workers 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.<ref>Template:Harvnb</ref>
In a young colony, the queen minimises reproductive competition from workers by suppressing their egg-laying through physical aggression and pheromones.<ref name="Van Honk-1980">Template:Cite journal</ref> Worker policing leads to nearly all eggs laid by workers being eaten.<ref name="Zanette-2012"/> 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.<ref name="Fletcher-1985">Template:Cite journal</ref> Because of the reproductive competition between workers and the queen, bumblebees are considered "primitively eusocial".<ref name="Cardinal-2011"/><ref name="Zanette-2012">Template:Cite journal</ref>
Although a large majority of bumblebees follow such monogynous colony cycles that only involve one queen, some select Bombus species (such as Bombus pauloensis) will spend part of their life cycle in a polygynous phase (have multiple queens in one nest during these periods of polygyny).<ref name="Cameron-1998">Template:Cite journal</ref>
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.<ref name="Walther-Hellwig-2000">Template:Cite journal</ref> They tend to visit the same patches of flowers every day, as long as they continue to find nectar and pollen there,<ref name="Dramstad-2003">Template:Cite journal</ref> a habit known as pollinator or flower constancy. While foraging, bumblebees can reach ground speeds of up to Template:Convert.<ref>Template:Cite journal</ref>
Bumblebees use a combination of colour and spatial relationships to learn which flowers to forage from.<ref>Template:Cite journal</ref> 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.<ref>Template:Cite journal
- Template:Cite journal</ref> Bumblebees can detect the temperature of flowers,<ref>Template:Cite journal</ref> as well as which parts of the flower are hotter or cooler<ref>Template:Cite journal</ref> 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 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.<ref name="Maloof-2001">Template:Cite journal</ref>
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.<ref>Template:Cite web</ref> Bumblebees are also capable of buzz pollination, in which they dislodge pollen from the anthers by creating a resonant vibration with their flight muscles.<ref>Template:Cite web</ref>
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.<ref name="Goulson-1998">Template:Cite journal</ref> This scent mark is a general chemical bouquet that bumblebees leave behind in different locations (e.g. nest, neutral, and food sites),<ref name="Saleh-2007">Template:Cite journal</ref> and they learn to use this bouquet to identify both rewarding and unrewarding flowers,<ref>Template:Cite journal</ref> and may be able to identify who else has visited a flower.<ref name="Pearce-2017">Template:Cite journal</ref> 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.<ref>Template:Cite journal</ref>
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.<ref>Template:Cite web</ref> 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.<ref>Template:Harvnb</ref>
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.<ref name="Scherer">Template:Cite web</ref> These types of muscles are called asynchronous muscles<ref>Template:Cite web</ref> and are found in the insect wing systems in families such as Hymenoptera, Diptera, Coleoptera, and Hemiptera.<ref name="Scherer"/> 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.<ref name="Heinrich-1981">Template:Cite book</ref><ref name="Schmidt-Nielsen-1997">Template:Cite book</ref>
Cuckoo bumblebees
Bumblebees of the subgenus Psithyrus (known as 'cuckoo bumblebees', and formerly considered a separate genus) are brood parasites,<ref name="Erler-2010">Template:Cite journal</ref> sometimes called kleptoparasites,<ref name="Slater-2005">Template:Cite book</ref> 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,<ref>Pierre Rasmont. "Bombus (Psithyrus) sylvestris (Lepeletier, 1832)". Université de Mons. Retrieved 6 January 2013.</ref> 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.<ref name="Zimma-2003">Template:Cite journal</ref> 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,<ref name="Erler-2010"/> though some species, such as B. bohemicus, actually enjoy increased success when they leave the host queen alive.<ref name="Fisher-1988">Template:Cite journal</ref>
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.<ref name="Fisher-1992">Template:Cite journal</ref> 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 bumblebee has a specific host species, which it may physically resemble.<ref>Template:Harvnb</ref> 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 locationTemplate:Efn had "[lost] their fight against their parasite".<ref name="Erler-2010"/>
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.<ref>Template:Cite web</ref><ref>Template:Cite web</ref> 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.<ref>Template:Harvnb</ref>
Predators, parasites, and pathogens
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.<ref>Template:Harvnb</ref> Adults are preyed upon by robber flies and beewolves in North America.<ref>Template:Harvnb</ref> 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.<ref>Template:Harvnb</ref>
The great grey shrike is able to detect flying bumblebees up to Template:Cvt away; once captured, the sting is removed by repeatedly squeezing the insect with the mandibles and wiping the abdomen on a branch.<ref>Template:Cite book</ref> 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.<ref>Template:Cite web</ref>
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. hypnorum has spread into the United Kingdom despite hosting high levels of a nematode that normally interferes with queen bees' attempts to establish colonies.<ref>"Parasites fail to halt European bumblebee invasion of the UK Template:Webarchive", Bumblebee Conservation Trust (retrieved 6 February 2015)</ref> Deformed wing virus has been found to affect 11% of bumblebees in Great Britain.<ref>"New study shows how bumblebees can be infected by honeybee diseases Template:Webarchive", Bumblebee Conservation Trust (retrieved 6 February 2015)</ref>
Female bee moths (Aphomia sociella) 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.<ref>Template:Cite journal</ref>
Relationship to humans
Agricultural use
Bumblebees are important pollinators of both crops and wildflowers.<ref>Template:Cite web</ref> 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 in greenhouses by buzz pollination whereas other pollinators cannot.<ref>Template:Cite web</ref> 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.<ref>Template:Harvnb</ref>
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.<ref name="Macdonald-2003"/> Bumblebees are also reared commercially to pollinate tomatoes grown in greenhouses.<ref name="Biobees Bumblebee Pollination"/> The New Zealand population of buff-tailed bumblebees began colonising Tasmania, Template:Convert away, after being introduced there in 1992 under unclear circumstances.<ref>Template:Harvnb</ref>
Some concerns exist about the impact of the international trade in mass-produced bumblebee colonies. Evidence from Japan<ref>Template:Cite journal</ref> and South America<ref>Template:Cite journal</ref> 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.<ref>Template:Cite journal</ref> In addition, mounting evidence indicates mass-produced bumblebees may also carry diseases, harmful to wild bumblebees<ref>Template:Cite journal</ref><ref name="Graystock-2013">Template:Cite journal
- Template:Cite news</ref> and honeybees.<ref name="Graystock-2013" />
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.<ref name="Goulson-2013b">Template:Harvnb</ref>
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.<ref>Template:Cite journal</ref> 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.<ref>Template:Harvnb</ref> 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.<ref>Template:Cite journal</ref>
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.<ref>Template:Cite news</ref> 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.<ref name="Whitehorn-2012">Template:Cite journal</ref> Another study on B. terrestris had results suggesting that use of neonicotinoid pesticides can affect how well bumblebees are able to forage and pollinate. Foragers from bee colonies that had been affected by the pesticide took longer to learn to manipulate flowers and visited flowers with less nutritious pollen.<ref>Template:Cite journal</ref> In another study, chronic exposure in a laboratory setting to field-realistic levels of the neonicotinoid pesticide thiamethoxam did not affect colony weight gain or the number or mass of sexuals produced.<ref>Template:Cite journal</ref> 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."<ref>Template:Cite web Research was published in the Journal of the Federation of American Societies for Experimental Biology by Chris Connolly and others.</ref>
Of 19 species of native nestmaking bumblebees and six species of cuckoo bumblebees formerly widespread in Britain,<ref>Template:Cite book</ref> three have been extirpated,<ref>Template:Cite web</ref><ref>Template:Cite journal</ref> eight are in serious decline, and only six remain widespread.<ref>Template:Cite journal</ref> Similar declines have been reported in Ireland, with four species designated endangered, and another two considered vulnerable to extinction.<ref>Template:Cite webTemplate:Dead link</ref> A decline in bumblebee numbers could cause large-scale changes to the countryside, resulting from inadequate pollination of certain plants.<ref name="Geographical-2008">Template:Cite journal</ref>
Some bumblebees native to North America are also vanishing, such as Bombus balteatus,<ref>Template:Cite journal</ref> Bombus terricola,<ref>Template:Cite journal</ref> Bombus affinis,<ref name="Flesher-2017">Template:Cite news</ref><ref>Template:Cite web</ref> and Bombus occidentalis; one, Bombus franklini, may be extinct.<ref>Template:Cite web</ref> 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.<ref>Template:Cite journal</ref>
Conservation efforts
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."<ref>Template:Cite web</ref><ref name="Barkham-2013">Template:Cite news</ref> In 2009 and 2010, the Trust attempted to reintroduce the short-haired bumblebee, Bombus subterraneus, which had become extinct in Britain, from the British-derived populations surviving in New Zealand from their introduction there a century earlier.<ref>Template:Harvnb</ref> From 2011, the Trust, in partnership with Natural England, Hymettus and the RSPB, has reintroduced short-haired bumblebee queens from 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; the brown-banded carder bee, Bombus humilis and the moss carder bee, Bombus muscorum.<ref>Template:Cite web</ref>
The world's first bumblebee sanctuary was established at Vane Farm in the Loch Leven National Nature Reserve in Scotland in 2008.<ref name="Geographical-2008"/> In 2011, London's Natural History Museum led the establishment of an International Union for Conservation of Nature Bumblebee Specialist Group, chaired by Dr. Paul H. Williams,<ref name="Natural History Museum">Template:Cite web</ref> to assess the threat status of bumblebee species worldwide using Red List criteria.<ref>Template:Cite web</ref>
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.<ref name="Goulson-2013b"/>
Misconception about flight
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.<ref name="The secrets of bee flight"/>
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.<ref>Template:Cite journal cited in Template:Cite book</ref>
In later years, McMasters backed away from this origin, suggesting there could be multiple sources, and stating that the earliest he had found was a reference in the 1934 book Template:Lang by French entomologist Antoine Magnan (1881–1938). Magnan had applied the equations of air resistance to insects, giving the result that their flight would be impossible, but "One shouldn't be surprised that the results of the calculations don't square with reality".<ref>Template:Cite book</ref> The following passage appears in the introduction to Le Vol des Insectes:<ref name="Magnan-1934">Template:Cite book</ref>
Magnan refers to his assistant André Sainte-Laguë.<ref>"The bumblebee story can be traced back to a 1934 book by entomologist Antoine Magnan, who refers to a calculation by his assistant André Sainte-Laguë, who was an engineer. The conclusion was presumably based on the fact that the maximum possible lift produced by aircraft wings as small as a bumblebee's wings and traveling as slowly as a bee in flight would be much less than the weight of a bee."Template:Cite journal</ref> Some credit physicist Ludwig Prandtl (1875–1953) of the University of Göttingen in Germany with popularizing the idea. Others say Swiss gas dynamicist Jakob Ackeret (1898–1981) did the calculations.<ref>Template:Cite web</ref>
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.<ref>Template:Cite news</ref>
The evolutionary biologist John Maynard Smith 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.<ref>Template:Cite web</ref>
In music and literature
The orchestral interlude Flight of the Bumblebee was composed (c. 1900) by Nikolai Rimsky-Korsakov. 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,<ref>Template:Cite book</ref> although the music may reflect the flight of a bluebottle rather than a bumblebee.<ref>Template:Cite book</ref> 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 excluded from the film in later showings.<ref>Template:Cite book</ref>
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".<ref>Template:Cite book</ref> This was one of nine books censored under the Bishops' Ban issued by the Archbishop of Canterbury John Whitgift and the Bishop of London Richard Bancroft.<ref>Template:Cite web</ref>
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!",<ref>Template:Cite web</ref> 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 is said to have enclosed a dead bee.<ref name="Morgan-2010">Template:Cite book</ref><ref>Template:Cite book</ref>
In 1847, Ralph Waldo Emerson published his poem "The Humble-Bee".<ref>Template:Cite book</ref>
The entomologist Otto Plath wrote Bumblebees and Their Ways in 1934.<ref>Template:Cite web</ref> His daughter, the poet Sylvia Plath, wrote a group of poems about bees late in 1962, within four months of her suicide,<ref>Template:Cite web</ref> transforming her father's interest into her poetry.<ref name="Kirk-2004">Template:Cite book</ref>
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).<ref>Template:Cite book</ref>
The surname Dumbledore in the Harry Potter series (1997–2007) is an old name for bumblebee.<ref name="Dumbledor-1913">Template:Cite encyclopedia</ref> 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".<ref>Template:Cite book</ref> J. R. R. Tolkien, in his poem Errantry, used the term Dumbledor for a bumblebee.<ref>Template:Cite book</ref> Among Beatrix Potter's "little books", Babbity Bumble and other members of her nest appear in her 1910 The Tale of Mrs. Tittlemouse.<ref name="Jones 2010"/>
See also
- Ophrys bombyliflora – the bumblebee orchid
- Melittology (also known as apiology) – the study of bees
Notes
References
Sources
- Goulson, Dave. Bumblebees: Their Behaviour and Ecology, 2003. Oxford University Press. Template:ISBN.
- Template:Cite book
- Template:Cite web
- Macdonald, Murdo & Nisbet, G. Highland Bumblebees: Distribution, Ecology and Conservation. HBRG, 2006. Template:ISBN. – 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, Discover Life: List of Species, Worldwide Species Map.
- Deciphering the Mystery of Bee Flight
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