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This article refers collectively to all true honey bees; for the "common" domesticated honey bee, see .

Honey bees (or honeybees) are a subset of in the genus Apis, primarily distinguished by the production and storage of and the construction of , nests out of . Honey bees are the only extant members of the tribe Apini, all in the genus Apis. Currently, there are only seven recognised species of honey bee with a total of 44 subspecies, though historically, anywhere from six to eleven species have been recognised. Honey bees represent only a small fraction of the approximately 20,000 known species of bees. Some other types of related bees produce and store honey, but only members of the genus Apis are true honey bees.

The study of honey bees is known as .


Origin, systematics and distribution
Honey bees appear to have their center of origin in and (including the ), as all but one (i.e. Apis mellifera), of the extant species are native to that region. Notably, living representatives of the earliest lineages to diverge ( and ) have their center of origin there.

The first Apis bees appear in the at the (23–56 Mya) boundary, in European deposits. The origin of these prehistoric honey bees does not necessarily indicate that Europe is the place of origin of the genus, only that it occurred there at that time. There are few known fossil deposits in South Asia, the suspected region of honey bee origin, and fewer still have been thoroughly studied.

No Apis species existed in the New World during human times before the introduction of Apis mellifera by Europeans. There is only one fossil species documented from the New World, , known from a single 14-million-year old specimen from Nevada.

The close relatives of modern honey bees—e.g. and —are also social to some degree, and social behavior seems a trait that predates the origin of the genus. Among the extant members of Apis, the more species make single, exposed combs, while the more recently evolved species nest in cavities and have multiple combs, which has greatly facilitated their domestication.

Most species have historically been cultured or at least exploited for honey and by humans indigenous to their native ranges. Only two of these species have been truly , one ( ) at least since the time of the building of the , and only that species has been moved extensively beyond its native range.

Today's honey bees constitute three .


Genetics
The counts of female bees for the three clades are Micrapis 2N=16, Megapis 2N=16, Apis 2N=32. Drones of all species have 1N chromosome counts. The genome of Apis has been mapped.

Drones (males) are produced from unfertilized eggs and therefore represent only the of the queen that laid the eggs, i.e. have only a mother. Workers and queens (both female) result from fertilized eggs and therefore have both a mother and a father. A modified form of controls sex differentiation. The sex is and so long as two different variants are present, a female bee results. If both sex alleles are identical, diploid drones are produced. Honeybees detect and destroy diploid drones after the eggs hatch.

Queens typically mate with multiple drones on more than one mating flight. Once mated, they lay eggs and fertilize them as needed from sperm stored in the . Since the number of sex alleles is limited—about 18 are known in Apis—there is a high probability that a queen will mate with one or more drones having sex alleles identical with one of the sex alleles in the queen. It is therefore typical for a queen to produce a percentage of diploid drone eggs.


Micrapis
and are small honey bees of southern and southeastern Asia. They make very small, exposed nests in trees and shrubs. Their stings are often incapable of penetrating human skin, so the and can be handled with minimal protection. They occur largely though they are very distinct and are probably the result of , their distribution later converging. Given that A. florea is more widely distributed and A. andreniformis is considerably more aggressive, honey is—if at all—usually harvested from the former only. They are the most ancient extant lineage of honey bees, maybe diverging in the (some 40 million years ago or slightly later) from the other lineages, but among themselves do not seem to have diverged a long time before the .


Megapis
There is one recognised species in the subgenus Megapis. It usually builds single or a few exposed combs on high tree limbs, on cliffs, and sometimes on buildings. They can be very fierce. Periodically robbed of their honey by human "honey hunters", colonies are easily capable of stinging a human being to death if provoked.

  • , the giant honey bee, is native and widespread across most of South and Southeast Asia.
  • , the Indonesian honey bee, is classified as the subspecies of the giant honey bee or a distinct species; in the latter case, and/or other lineages would probably also have to be considered species.
  • , the Himalayan honey bee, was initially described as a distinct species. Later, it was included in A. dorsata as a subspecies based on the , though authors applying a genetic species concept have suggested it should be considered a species. Essentially restricted to the , it differs little from the giant honey bee in appearance, but has extensive behavioral that enable it to nest in the open at high altitudes despite low ambient temperatures. It is the largest living honey bee.


"Apis"
Eastern species
These are three or four species. The reddish Koschevnikov's bee ( ) from is well distinct; it probably derives from the first colonization of the island by cave-nesting honey bees. , the Eastern honey bee proper, is the traditional honey bee of southern and eastern Asia, kept in hives in a similar fashion to , though on a much smaller and regionalised scale. It has not been possible yet to resolve its relationship to the Bornean and from the to satisfaction; the most recent hypothesis is that these are indeed distinct species but that A. cerana is still , consisting of several good species.
European/Western/Common honey bee

, the most commonly domesticated species, was the third insect to have its mapped. It seems to have originated in eastern tropical and spread from there to and eastwards into to the range. It is variously called the European, Western or Common honey bee in different parts of the world. There are many that have adapted to the local geographic and climatic environments, and in addition, hybrid strains such as the have been bred. Behavior, color and anatomy can be quite different from one subspecies or even strain to another.

Regarding , this is the most enigmatic honey bee species. It seems to have diverged from its Eastern relatives only during the . This would fit the hypothesis that the ancestral stock of cave-nesting honey bees was separated into the Western group of E Africa and the Eastern group of tropical Asia by in the and adjacent regions, which caused declines of foodplants and trees that provided nest sites, eventually causing to cease. The diversity of subspecies is probably the product of a largely aided by climate and habitat changes during the . That the Western honey bee has been intensively managed by humans for many millennia – including hybridization and introductions – has apparently increased the speed of its and confounded the DNA sequence data to a point where little of substance can be said about the exact relationships of many A. mellifera subspecies.

Apis mellifera are not native to the Americas and therefore were not present upon the arrival of the European explorers and colonists. There were, however, other native bee species kept and traded by indigenous peoples.Charles F. Calkins, "Beekeeping in Yucatán: A Study in Historical-Cultural Zoogeography (PhD diss., University of Nebraska, 1974), as quoted in Crane, World History of Beekeeping, p. 292. Calkins cites the original translated source as Hernán Cortés, Letters of Cortés: The Five Letters of Relation from Fernando Cortes to the Emperor Charles V, trans. and ed. Francis A. MacNutt (New York: Putnam, 1908), 1:145. In 1622, European colonists brought the dark bee ( ) to the Americas, followed later by ( A. m. ligustica) and others. Many of the crops that depend on honey bees for pollination have also been imported since colonial times. Escaped swarms (known as "wild" bees, but actually ) spread rapidly as far as the , usually preceding the colonists. Honey bees did not naturally cross the ; they were transported by the Mormon pioneersHorn, Bees in America, p. 80–81. http://maxwellinstitute.byu.edu/publications/review/?vol=20&num=1&id=694#_ednref30 to Utah in the late 1840s, and by ship to in the early 1850s.


Africanized bee
Africanized bees (known colloquially as "killer bees") are hybrids between European stock and one of the African subspecies ; they are often more aggressive than and do not create as much of a surplus as European bees, but are more resistant to disease and are better foragers. Originating by accident in Brazil, they have spread to North America and constitute a in some regions. However, these strains do not overwinter well, and so are not often found in the colder, more northern parts of North America. On the other hand, the original breeding experiment for which the African bees were brought to Brazil in the first place has continued (though not as intended). Novel hybrid strains of domestic and re-domesticated Africanized bees combine high resilience to tropical conditions and good yields. They are popular among beekeepers in Brazil.


Beekeeping
Two species of honey bee, A. mellifera and A. cerana indica, are often maintained, fed, and transported by beekeepers. Modern hives also enable beekeepers to transport bees, moving from field to field as the crop needs pollinating and allowing the beekeeper to charge for the pollination services they provide, revising the historical role of the self-employed beekeeper, and favoring large-scale commercial operations.


Colony collapse disorder (CCD)
Beekeepers in Western countries have been reporting slow declines of stocks for many years, apparently due to impaired protein production, changes in agricultural practice, or unpredictable weather. In early 2007, abnormally high die-offs (30–70% of hives) of European honey bee colonies occurred in ; such a decline seems unprecedented in recent history. This has been dubbed "Colony collapse disorder" (CCD); it is unclear whether this is simply an accelerated phase of the general decline due to more adverse conditions in 2006, or a novel phenomenon. Research has so far failed to determine what causes it, but the weight of evidence is tentatively leaning towards CCD being a rather than a as it seems to be caused by a combination of various contributing factors rather than a single or .


Life cycle
As in a few other types of bees, a colony generally contains one , a fertile female; seasonally up to a few thousand or fertile males; ξ1 and a large seasonally variable population of sterile female . Details vary among the different species of honey bees, but common features include:

1. Eggs are laid singly in a cell in a wax , produced and shaped by the worker bees. Using her , the queen actually can choose to fertilize the egg she is laying, usually depending on what cell she is laying in. Drones develop from unfertilised eggs and are , while females (queens and worker bees) develop from fertilised eggs and are . Larvae are initially fed with produced by worker bees, later switching to honey and pollen. The exception is a larva fed solely on royal jelly, which will develop into a queen bee. The larva undergoes several moltings before spinning a within the cell, and .

2. Young worker bees clean the hive and feed the larvae. When their royal jelly producing glands begin to atrophy, they begin building comb cells. They progress to other within-colony tasks as they become older, such as receiving nectar and pollen from foragers, and guarding the hive. Later still, a worker takes her first orientation flights and finally leaves the hive and typically spends the remainder of her life as a forager.

3. Worker bees cooperate to find food and use a pattern of "dancing" (known as the ) to communicate information regarding resources with each other; this dance varies from species to species, but all living species of Apis exhibit some form of the behavior. If the resources are very close to the hive, they may also exhibit a less specific dance commonly known as the "Round Dance".

4. Honey bees also perform , which recruit receiver bees to collect nectar from returning foragers.

5. Virgin queens go on mating flights away from their home colony, and mate with multiple drones before returning. The drones die in the act of mating.

6. Colonies are established not by solitary queens, as in most bees, but by groups known as "", which consist of a mated queen and a large contingent of worker bees. This group moves en masse to a nest site that has been scouted by worker bees beforehand. Once they arrive, they immediately construct a new wax comb and begin to raise new worker brood. This type of nest founding is not seen in any other living bee genus, though there are several groups of wasps that also found new nests via swarming (sometimes including multiple queens). Also, will start new nests with large numbers of worker bees, but the nest is constructed before a queen is escorted to the site, and this worker force is not a true "swarm".


Winter survival
In cold climates honey bees stop flying when the temperature drops below about and crowd into the central area of the hive to form a "winter cluster". The worker bees huddle around the queen bee at the center of the cluster, shivering in order to keep the center between at the start of winter (during the broodless period) and once the queen resumes laying. The worker bees rotate through the cluster from the outside to the inside so that no bee gets too cold. The outside edges of the cluster stay at about 8–9 °C (46–48 °F). The colder the weather is outside, the more compact the cluster becomes. During winter, they consume their stored honey to produce body heat. The amount of honey consumed during the winter is a function of winter length and severity but ranges in temperate climates from 15 to 50 kg (30 to 100 pounds).http://www.bees-online.com/Winter.htm


Pollination
Species of Apis are generalist floral visitors, and will pollinate a large variety of plants, but by no means all plants. Of all the honey bee species, only has been used extensively for commercial pollination of crops and other plants. The value of these pollination services is commonly measured in the billions of dollars. Bees collect 66 pounds of pollen per year, per hive. Back Yard Beekeepers Association. "Facts about Honeybees." http://www.backyardbeekeepers.com/facts.html


Honey
Honey is the complex substance made when the nectar and sweet deposits from plants and trees are gathered, modified and stored in the honeycomb by honey bees as a food source for the colony. All living species of Apis have had their honey gathered by indigenous peoples for consumption, though for commercial purposes only and have been exploited to any degree. Honey is sometimes also gathered by humans from the nests of various .

In 1911 a bee culturist estimated that a (approx. 1 litre) of honey represented bees flying over an estimated 48,000 miles to gather the nectar needed to produce the honey. Mechanics Science installing linoleum&hl=en&sa=X&ei=y4zsT9OSGMriqAGU1PW8BQ&sqi=2&ved=0CDYQ6AEwAA#v=onepage&q&f=true "A Quart of Honey Means 48,000 miles of flight" Popular Mechanics, December 1911, p. 889.


Nectar
Nectar is a liquid high in sucrose that is produced in plant glands known as nectaries. It is an important energetic resource for honeybees and plays a significant role in foraging economics and evolutionary differentiation between different subspecies. It was proposed through an experiment conducted with the African honey bee, , that nectar temperature impacts the foraging decisions of honey bees.


Beeswax
Worker bees of a certain age will secrete from a series of glands on their . They use the wax to form the walls and caps of the comb. As with honey, beeswax is gathered by humans for various purposes.


Pollen
Bees collect pollen in the and carry it back to the hive. In the hive, pollen is used as a source necessary during brood-rearing. In certain environments, excess pollen can be collected from the hives of A. mellifera and A. cerana. It is often eaten as a health supplement.


Propolis
or bee glue is created from resins, balsams and . Those species of honey bees that nest in tree cavities use propolis to seal cracks in the hive. use propolis to defend against ants by coating the branch from which their nest is suspended to create a sticky moat. Propolis is consumed by humans as a health supplement in various ways and also used in some cosmetics.


Sexes and castes
There are two sexes of honey bee: females (workers and queens); and males (or drones). A is a different form, morphologically or reproductively, within the same sex of a species. In sum, there are three types of honey bees: drones, workers and queens; two sexes: male and female; and two female castes: queens and workers.


Drones
Males or drones are typically haploid, having only one set of chromosomes. They are produced by the queen if she chooses not to fertilize an egg; or by a non-fertilized laying worker. Diploid drones may be produced if an egg is fertilized but is homozygous for the sex-determination allele. Drones take 24 days to develop and may be produced anywhere from summer through autumn. Drones have large eyes used to locate queens during mating flights. Drones do not have a .


Workers
Workers are female and have two sets of chromosomes. They are produced from an egg that the queen has selectively fertilized from stored sperm. Workers typically develop in 21 days. A typical colony may contain as many as 60,000 worker bees. Workers exhibit a wider range of behaviors than either queens or drones. Their duties change upon the age of the bee in the following order (beginning with cleaning out their own cell after eating through their capped brood cell): feed brood; receive nectar; clean hive; guard duty; and foraging. Some workers engage in other specialized behaviors, such as "undertaking" (removing corpses of their nestmates from inside the hive).

Workers have morphological specializations: including the or pollen basket, abdominal glands that produce , brood-feeding glands, and barbs on the . Under certain conditions (for example, if the colony becomes queenless), a worker may develop ovaries.


Queens
Queen honey bees, like workers, are female. They are created at the decision of the worker bees by feeding a larva only royal jelly throughout its development, rather than switching from royal jelly to pollen once the larva grows past a certain size. Queens are produced in oversized cells and develop in only 16 days. Queens have a different morphology and behavior from worker bees. In addition to the greater size of the queen, she has a functional set of ovaries, and a spermatheca, which stores and maintains sperm after she has mated. The sting of queens is not barbed like a worker's sting, and queens lack the glands that produce beeswax. Once mated, queens may lay up to 2,000 eggs per day. Queens produce a variety of pheromones that regulate behavior of workers, and helps swarms track the queen's location during the migratory phase.


Defense
All honey bees live in colonies where the workers will intruders as a form of defense, and alarmed bees will release a that stimulates the attack response in other bees. The different species of honey bees are distinguished from all other bee species (and virtually all other ) by the possession of small barbs on the , but these barbs are found only in the worker bees. The sting and associated sac of honey bees are also modified so as to pull free of the body once lodged (), and the sting apparatus has its own musculature and , which allow it to keep delivering venom once detached. The worker dies after the sting becomes lodged and is subsequently torn loose from the bee's abdomen.

It is hypothesized that this complex apparatus, including the barbs on the sting, evolved specifically in response to predation by vertebrates, as the barbs do not usually function (and the sting apparatus does not detach) unless the sting is embedded in fleshy tissue. While the sting can also penetrate the membranes between joints in the exoskeleton of other insects (and is used in fights between queens), in the case of , defense against larger insects such as predatory wasps is usually performed by surrounding the intruder with a mass of defending worker bees, who vibrate their muscles so vigorously that it raises the temperature of the intruder to a lethal level. It was previously thought that the heat alone was responsible for killing intruding wasps, but recent experiments have demonstrated that it is the increased temperature in combination with increased carbon dioxide levels within the ball that produces the lethal effect. This phenomenon is also used to kill a queen perceived as intruding or defective, an action known to beekeepers as balling the queen, named for the ball of bees formed.

In the case of those honey bee species with open combs (e.g., A. dorsata), would-be predators are given a warning signal that takes the form of a "" that spreads as a ripple across a layer of bees densely packed on the surface of the comb when a threat is perceived, and consists of bees momentarily arching their bodies and flicking their wings.http://www.sciencedaily.com/releases/2008/09/080909204550.htm


Communication
Honey bees are known to communicate through many different chemicals and odors, as is common in insects, but also using specific behaviours that convey information about the quality and type of resources in the environment, and where these resources are located. The details of the signalling being used vary from species to species; for example, the two smallest species, and , dance on the upper surface of the comb, which is horizontal (not vertical, as in other species), and worker bees orient the dance in the actual compass direction of the resource to which they are recruiting.

A study has found that honey bees use their antenna asymmetrically for social interactions with a strong lateral preference to use their right antenna. Honeybees use right antennae to tell friend from foe July 1, 2013


Symbolism
Both the "O Asvins, lords of brightness, anoint me with the honey of the bee, that I may speak forceful speech among men! Atharva Veda 91-258, quoted in Maguelonne Toussaint-Samat (Anthea Bell, tr.) The History of Food, 2nd ed. 2009:14. and the ancient Greeks associated lips anointed with honey with the gift of eloquence and even of prescience. The priestess at was the "Delphic Bee". The Quran has a chapter titled .

A community of honey bees has often been employed throughout history by political theorists as a model of human society:

This image occurs in Aristotle and Plato; in VirgilVirgil, , book IV. and Seneca; in Erasmus and Shakespeare; in Marx and Tolstoy. ξ2

Honey bees, signifying immortality and resurrection, were royal emblems of the , revived by . The bee also is the heraldic emblem of the .


See also


Further reading
  • Adam, Brother. In Search of the Best Strains of Bees. Hebden Bridge, W. Yorks: Northern Bee Books, 1983.
  • Adam, Brother. Bee-keeping at Buckfast Abbey. Geddington, Northants: British Bee Publications, 1975.
  • Aldersey-Williams, H. Zoomorphic: New Animal Architecture. London: Laurence King Publishing, 2003.
  • Alexander, P. Rough Magic: A Biography of Sylvia Plath. New York: Da Capo Press, 2003.
  • Allan, M. Darwin and his flowers. London: Faber & Faber, 1977.
  • Alston, F. Skeps, their History, Making and Use. Hebden Bridge, W. Yorks: Northern Bee Books, 1987.
  • Barrett, P. The Immigrant Bees 1788 to 1898, 1995.
  • Barrett, P. William Cotton.
  • Beuys, J. Honey is Flowing in All Directions. Heidelberg: Edition Staeck, 1997.
  • Bevan, E. The Honey-bee: Its Natural History, Physiology and Management. London: Baldwin, Cradock & Joy, 1827.
  • Bill, L. For the Love of Bees. Newton Abbot, Devon: David & Charles, 1989.
  • Bodenheimer, F.S. Insects as Human Food The Hague: Dr. W. Junk, 1951.
  • Brothwell, D., Brothwell, P. Food in Antiquity. London: Thames & Hudson, 1969.
  • Kak, Subhash C. (1991): The Honey Bee Dance Language Controversy. The Mankind Quarterly Summer 1991: 357–365. HTML fulltext
  • Lindauer, Martin (1971): Communication among social bees. .
  • & (2005): Evolution of the Insects. .


External links


References
    ^ (1995). 9780716760108, Scientific American Library.
    ^ (2020). 9780719565984, .

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