Apis dorsata, or giant honey bee
Honey bees (or honeybees) are a subset of bees, primarily distinguished by the production and storage of honey and the construction of colonial nests out of wax. Honey bees are the only extant members of the tribe Apini, all in the genus Apis. Currently, there are only seven recognized species of honey bee with a total of 44 subspecies (Engel, 1999) though historically, anywhere from six to eleven species have been recognized. 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.
Honey bees as a group appear to have their center of origin in South and Southeast Asia (including the Philippines), as all but one of the extant species are native to that region, notably the most plesiomorphic living species (Apis florea and A. andreniformis). The first Apis bees appear in the fossil record at the Eocene-Oligocene boundary, in European deposits dating about 35 million years ago. The origin of these prehistoric honey bees does not necessarily indicate that Europe is where the genus originated, only that it occurred there at that time. There are few known fossil deposits in the suspected region of honeybee origin, and fewer still have been thoroughly studied; moreover, the tropical conditions are generally not ideal for fossilization of small land animals.
The close relatives of modern honey bees - e.g. bumblebees and stingless bees - are also social to some degree, and thus social behavior seems a plesiomorphic trait that predates the origin of the genus. 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 of these species have been truly domesticated, one (Apis mellifera) at least since the time of the building of the Egyptian pyramids, and only that species has been moved extensively beyond its native range.
Today's honey bees constitute three clades (Engel 1999, Arias & Sheppard 2005):
Dwarf honey bees - subgenus Micrapis
Cave-nesting honey bees - subgenus Apis
Two species of honey bee, A. mellifera and A. cerana, 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. For further information see the main article, or the articles for these species.
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; and a large seasonally variable population of sterile female worker bees. Details vary among the different species of honey bees, but common features include:
Eggs are laid singly in a cell in a wax honeycomb, produced and shaped by the worker bees. Larvae are initially fed with royal jelly 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 cocoon within the cell, and pupating. Drones hatch from unfertilized eggs, females (Queens and worker bees) hatch from fertilized eggs. The queen actually can choose to fertilize the egg she is laying, usually depending on what cell she is laying in.
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 its life as a forager.
Worker bees cooperate to find food and use a pattern of "dancing" (known as the bee dance or waggle dance) 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".
Honey bees also perform tremble dances which recruit receiver bees to collect nectar from returning foragers .
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.
Colonies are established not by solitary queens, as in most bees, but by groups known as "swarms", 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 Vespid wasps which also found new nests via swarming (sometimes including multiple queens). Also, stingless bees 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".
Species of Apis are generalist floral visitors, and will pollinate a large variety of plants, but by no means all plants. Of all the honeybee species, only Apis mellifera 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.
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. All living species of Apis have had their honey gathered by indigenous peoples for consumption, though for commercial purposes only Apis mellifera and Apis cerana have been exploited to any degree. Honey is sometimes also gathered by humans from the nests of various stingless bees.
Worker bees of a certain age will secrete beeswax from a series of glands on their abdomens. They use the wax to form the walls and caps of the comb. As with honey, beeswax is gathered for various purposes.
Bees collect pollen in the pollen basket and carry it back to the hive. 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. mellifera and A. cerana. It is often eaten as a health supplement.
Propolis (or bee glue) is created from resins, balsams and tree saps. Those species of honey bees which nest in tree cavities use propolis to seal cracks in the hive. Dwarf honey bees use propolis to defend against ants by coating the branch from which their nest is suspended to create a sticky moat. Propolis is consumed as a health supplement in various ways and also used in some cosmetics.
All honey bees live in colonies where the worker bees will sting intruders as a form of defense, and alarmed bees will release a pheromone that stimulates the attack response in other bees. The different species of honey bees are distinguished from all other bee species by the possession of small barbs on the sting, but these barbs are found only in the worker bees. The sting and associated venom sac are also modified so as to pull free of the body once lodged (autotomy), and the sting apparatus has its own musculature and ganglion which allow it to keep delivering venom once detached.
It is presumed 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 flexible exoskeletal joints in appendages of other insects (and is used in fights between queens), in the case of Apis cerana defense against other 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. This 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.
Honey bees are known to communicate through many different chemicals and odors, as is common in insects, but also using specific behaviors that convey information about the quality and type of resources in the environment, and where these resources are located. The details of the signaling being used vary from species to species; for example, the two smallest species, Apis andreniformis and Apis florea, 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.
Western honey bees (Apis mellifera) in Serbia
A Bee and a Rose
Worker bees (left) emerging from hive push away drones (right)
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 Virgil and Seneca; in Erasmus and Shakespeare; in Marx and Tolstoy." (Wilson 2004: p.4)
Honeybees, signifying immortality and resurrection, were royal emblems of the Napoleonic Empire.