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Among the extant members of Apis , the more basal 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 beeswax by humans indigenous to their native ranges.

Only two species have been truly domesticated: Apis mellifera and Apis cerana indica. Honey bees are the only extant members of the tribe Apini.

Today's honey bees constitute three clades: Micrapis dwarf honey bees , Megapis giant honey bee , and Apis domestic honey bees and close relatives.

The chromosome counts of female bees for the three clades are:. Drones males are produced from unfertilized eggs, so they represent only the DNA of the queen that laid the eggs, i.

Workers and queens both female result from fertilized eggs, so have both a mother and a father.

Arrhenotokous parthenogenesis , a modified form of parthenogenesis , controls sex differentiation. The sex allele is polymorphic , and so long as two different variants are present, a female bee results.

If both sex alleles are identical, diploid drones are produced. Honey bees 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 spermatheca.

Since the number of sex alleles is limited at a regional level, a queen will most likely mate with one or more drones having sex alleles identical with one of the sex alleles in the queen.

The queen, then, typically produces a percentage of diploid drone eggs. Apis florea and Apis andreniformis 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 hive and swarms can be handled with minimal protection.

They occur largely sympatrically , though they are very distinct evolutionarily and are probably the result of allopatric speciation , their distribution later converging.

One species is recognized 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 genetics of the western honey bee A. The domesticated species are A. The reddish Koschevnikov's bee Apis koschevnikovi from Borneo is clearly a distinct species; it probably derives from the first colonization of the island by cave-nesting honey bees.

Apis cerana , the eastern honey bee proper, is the traditional honey bee of southern and eastern Asia. It was domesticated as subspecies A.

It has not been possible yet to resolve its relationship to the Bornean A. It seems to have originated in eastern tropical Africa and spread from there to Northern Europe and eastwards into Asia to the Tien Shan range.

It is variously called the European, western, or common honey bee in different parts of the world. Many subspecies have adapted to the local geographic and climatic environments; in addition, hybrid strains, such as the Buckfast bee , have been bred.

Behavior, color, and anatomy can be quite different from one subspecies or even strain to another. It seems to have diverged from its eastern relatives only during the Late Miocene.

This would fit the hypothesis that the ancestral stock of cave-nesting honey bees was separated into the western group of East Africa and the eastern group of tropical Asia by desertification in the Middle East and adjacent regions, which caused declines of food plants and trees that provided nest sites, eventually causing gene flow to cease.

The diversity of A. Apis mellifera is not native to the Americas , so was not present upon the arrival of the European explorers and colonists.

However, other native bee species were kept and traded by indigenous peoples. In , European colonists brought the dark bee A. 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 feral spread rapidly as far as the Great Plains , usually preceding the colonists. Honey bees did not naturally cross the Rocky Mountains ; they were transported by the Mormon pioneers to Utah in the late s, and by ship to California in the early s.

Africanized bees known colloquially as "killer bees" are hybrids between European stock and one of the African subspecies A. However, these strains do not overwinter well, so are not often found in the colder, more northern parts of North America.

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 redomesticated Africanized bees combine high resilience to tropical conditions and good yields. They are popular among beekeepers in Brazil.

Tribe Apini Latreille [16]. As in a few other types of eusocial bees, a colony generally contains one queen bee , a fertile female; seasonally up to a few thousand drone bees , or fertile males; [17] and tens of thousands of sterile female worker bees.

Details vary among the different species of honey bees, but common features include:. The worker bees rotate through the cluster from the outside to the inside so that no bee gets too cold.

The colder the weather is outside, the more compact the cluster becomes. During winter, they consume their stored honey to produce body heat.

During the summer, however, this is achieved through fanning and water evaporation from water collected in various fields.

Of all the honey bee species, only A. Species of Apis are generalist floral visitors, and pollinate many species of flowering plants, but because of their "generalized" nature, they do so inefficiently.

Without specialized adaptations for specific flowers, their ability to reach pollen and nectar is often limited. What's more, their tendency to visit all species in a given area means that the pollen they carry for any one species is often very diluted.

As such, they can provide some pollination to many plants, especially non-native crops, but most native plants have some native pollinator that is far more effective at pollinating that species.

European honey bees are often described as being essential to all human food production, leading to claims that without their pollination, all of humanity would starve, or even die out.

Plantains are sterile and propagated by cuttings, as are cassava. Potatoes , yams , and sweet potatoes are root vegetables propagated by tubers.

Rice , wheat , and corn are all wind-pollinated , because this is true of all grasses. Similarly, no crops originating in the New World depend on the domesticated honey bee Apis mellifera at all, as the insect is invasive, having been brought over with colonists in the last few centuries.

Thomas Jefferson mentioned this in his Notes on the State of Virginia:. Tomatoes , peppers , squash , and all other New World crops evolved with native pollinators like squash bees , bumble bees , other native bees.

The stingless bees mentioned by Jefferson are distant relatives of the honey bees, in the genus Melipona.

Honey bees obtain all of their nutritional requirements from a diverse combination of pollen and nectar. Pollen is the only natural protein source for honey bees.

Adult worker honey bees consume 3. Of these amino acids, honey bees require highest concentrations of leucine, isoleucine, and valine, however elevated concentrations of arginine and lysine are required for brood rearing.

Pyridoxine is the most prevalent B vitamin found in royal jelly and concentrations vary throughout the foraging season with lowest concentrations found in May and highest concentrations found in July and August.

Honey bees lacking dietary pyridoxine were unable to rear brood. Pollen is also a lipid source for honey bees ranging from 0. Fat-soluble vitamins A, D, E, and K are not considered essential but have shown to significantly improve the number of brood reared.

Nurse bees have the ability to selectively transfer sterols to larvae through brood food. Nectar is collected by foraging worker bees as a source of water and carbohydrates in the form of sucrose.

The dominant monosaccharides in honey bee diets are fructose and glucose but the most common circulating sugar in hemolymph is trehalose which is a disaccharide consisting of two glucose molecules.

Honey bees require water to maintain osmotic homeostasis, prepare liquid brood food, and to cool the hive through evaporation. A colony's water needs can generally be met by nectar foraging as it has high water content.

Occasionally on hot days or when nectar is limited, foragers will collect water from streams or ponds to meet the needs of the hive.

Larval stages of the G. Eggs are laid within the hive, and the larva that hatch tunnel through and destroy the honeycombs that contain bee larva and their honey stores.

The tunnels they create are lined with silk, which entangles and starves emerging bees. Destruction of honeycombs also result in honey leaking and being wasted.

Temperature treatments are possible, but also distorts wax of the honeycombs. Chemical fumigants, particularly CO 2 , are also used. The only domesticated species of honey bee are A.

In Japan, where mellifera is vulnerable to local hornets and disease, the Japanese honey bee a. 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.

Bees of various types other than honey bees are also domesticated and used for pollination or other means around the world, including Tetragonula iridipennis in India, the blue orchard bee for tree nut and fruit pollination in the United States, and a number of species of Bombus bumblebees for pollination in various regions globally, such as tomatoes , which are not effectively pollinated by honey bees.

Primarily in places where the bee was imported by humans, periodic collapses in honeybee populations have occurred at least since the late 19th century.

This has been dubbed "colony collapse disorder" CCD and was at first unexplained. Honey is the complex substance made when bees ingest nectar, process it, and store the substance into honey combs.

Worker bees of a certain age secrete beeswax from a series of exocrine glands on their abdomens. As with honey, beeswax is gathered by humans for various purposes such as candle making, waterproofing, soap and cosmetics manufacturing, pharmaceuticals, art, furniture polish and more.

Bees collect pollen in their pollen baskets and carry it back to the hive. Worker bees combine pollen, honey and glandular secretions and allow it to ferment in the comb to make bee bread.

The fermentation process releases additional nutrients from the pollen and can produce antibiotics and fatty acids which inhibit spoilage.

In the hive, pollen is used as a protein source necessary during brood-rearing. In certain environments, excess pollen can be collected from the hives of A.

The product is used as a health supplement. It has been used with moderate success as a source of pollen for hand pollination.

Bee brood — the eggs, larvae or pupae of honeybees — is nutritious and seen as a delicacy in countries such as Australia, Indonesia, [45] Mexico, Thailand, and many African countries; it has been consumed since ancient times by the Chinese and Egyptians.

Propolis is a resinous mixture collected by honey bees from tree buds, sap flows or other botanical sources, which is used as a sealant for unwanted open spaces in the hive.

Royal jelly is a honey-bee secretion used to nourish the larvae. Honey bees have three castes: Males, or drones, are typically haploid , having only one set of chromosomes , and primarily exist for the purpose of reproduction.

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 from summer through to autumn, numbering as many as per hive.

They do not defend the hive or kill intruders, and do not have a stinger. Workers have two sets of chromosomes.

Workers typically develop in 21 days. A typical colony may contain as many as 60, worker bees. 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: Workers have morphological specializations, including the pollen basket corbicula , [60] abdominal glands that produce beeswax, brood-feeding glands, and barbs on the sting.

Under certain conditions for example, if the colony becomes queenless , a worker may develop ovaries. Worker honey bees perform different behavioural tasks that cause them to be exposed to different local environments.

Queen honey bees are created when worker bees feed a single female larvae an exclusive diet of a food called " royal jelly ". 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.

Apis queens practice polyandry , with one female mating with multiple males. The highest documented mating frequency for an Apis queen is in Apis nigrocincta , where queens mate with an extremely high number of males with observed numbers of different matings ranging from 42 to 69 drones per queen.

Once mated, queens may lay up to 2, eggs per day. When a fertile female worker produces drones, a conflict arises between her interests and those of the queen.

The worker shares half her genes with the drone and one-quarter with her brothers, favouring her offspring over those of the queen. The queen shares half her genes with her sons and one-quarter with the sons of fertile female workers.

This relationship leads to a phenomenon known as "worker policing". In these rare situations, other worker bees in the hive who are genetically more related to the queen's sons than those of the fertile workers will patrol the hive and remove worker-laid eggs.

Another form of worker-based policing is aggression toward fertile females. In very rare instances workers subvert the policing mechanisms of the hive, laying eggs which are removed at a lower rate by other workers; this is known as anarchic syndrome.

Anarchic workers can activate their ovaries at a higher rate and contribute a greater proportion of males to the hive. Although an increase in the number of drones would decrease the overall productivity of the hive, the reproductive fitness of the drones' mother would increase.

Anarchic syndrome is an example of selection working in opposite directions at the individual and group levels for the stability of the hive.

Under ordinary circumstances the death or removal of a queen increases reproduction in workers, and a significant proportion of workers will have active ovaries in the absence of a queen.

The workers of the hive produce a last batch of drones before the hive eventually collapses. Although during this period worker policing is usually absent, in certain groups of bees it continues.

According to the strategy of kin selection , worker policing is not favored if a queen does not mate multiple times. Workers would be related by three-quarters of their genes, and the difference in relationship between sons of the queen and those of the other workers would decrease.

The benefit of policing is negated, and policing is less favored. Experiments confirming this hypothesis have shown a correlation between higher mating rates and increased rates of worker policing in many species of social hymenoptera.

All honey bees live in colonies where the workers sting intruders as a form of defense, and alarmed bees release a pheromone 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 Hymenoptera by the possession of small barbs on the sting, but these barbs are found only in the worker bees.