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(Redirected from Honeybee)
:''This article refers collectively to all true honey bees; for the "common" domesticated honey bee, see Western honey bee''
'Honey bees' (or 'honeybees') are a subset of bees which represent a far smaller fraction of bee diversity than most people suspect; of the approximately 20,000 known species of bees, there are only seven presently-recognized species with a total of 44 subspecies (Engel, 1999; historically, anywhere from six to eleven species have been recognized). These bees are the only living members of the tribe Apini, all in the genus ''Apis'', and all of which produce and store liquefied sugar ("honey") to some degree, and construct colonial nests out of wax secreted by the workers in the colony. Other types of related bees produce and store honey, but only members of the genus ''Apis'' are considered true honey bees.
Honey bees as a group appear to have their center of origin in 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''). [1] 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 honeybees 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'''
★ ''Apis florea'' and ''Apis andreniformis'' are small honey bees of southern and southeastern Asia. The make very small, exposed nests in trees and shrubs. The stings often not capable 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. 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 Bartonian (some 40 mya or slightly later) from the other lineages, but among themselves do not seem to have diverged a long time before the Neogene.(Arias & Sheppard 2005)
'Giant honey bees - subgenus ''Megapis'''
There is one recognized species which 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 when provoked. Their origin as a distinct lineage is only slightly more recent than that of the dwarf honey bees.
★ ''Apis dorsata'', the Giant honey bee proper, is native and widespread across most of South and Southeast Asia.
★ ''Apis dorsata binghami'', the Indonesian honey bee, is classified as the Indonesian subspecies of the Giant honey bee or a distinct species; in the latter case, ''A. d. breviligula'' and/or other lineages would probably also have to be considered species.
★ ''Apis dorsata laboriosa'', the Himalayan honey bee, was initially described as a distinct species. Later, it was included in ''A. dorsata'' as a subspecies (Engel 1999), though genetic studies suggest it is divergent enough to merit species status (Arias & Sheppard 2005). Essentially restricted to the Himalayas, it differs little from the Giant honey bee in appearance, but has extensive behavioral adaptations which enable it to nest in the open at high altitudes despite low ambient temperatures. It is the largest living honey bee.
'Cave-nesting honey bees - subgenus ''Apis'''
★ Western (European, Common) honey bee
Main articles: Apis mellifera
:''Apis mellifera'', the most commonly domesticated species, was the third insect to have its genome mapped. 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 Western, European or Common honey bee in different parts of the world. There are many subspecies that have adapted to the local geographic and climatic environment, and 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.
:Regarding phylogeny, this is the most enigmatic honey bee species. 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 E Africa and the Eastern group of tropical Asia by desertification in the Middle East and adjacent regions, which caused declines of foodplants and trees which provided nest sites, eventually causing gene flow to cease. The diversity of subspecies is probably the product of a (largely) Early Pleistocene radiation aided by climate and habitat changes during the last ice age. That the Western honey bee has been intensively managed by humans since many millennia - including hybridization and introductions - has apparently increased the speed of its evolution 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.(Arias & Sheppard 2005)
:In 1622, European colonists brought the dark bee (''A. m. mellifera'') to the Americas, followed later by Italian bees (''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 feral) spread rapidly as far as the Great Plains, usually preceding the colonists. The Native Americans called the honey bee "the white man's fly". Honey bees did not naturally cross the Rocky Mountains; they were carried by ship to California in the early 1850s.
:Main articles: Africanized bee
:The so-called "killer bee" are highly aggressive hybrids between European stock and the African subspecies ''A. m. scutellata''; they are thus often called "Africanized bees". Originating by accident in Brazil, they have spread to North America and constitutes a pest in some regions. 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, and are popular among beekeepers in Brazil.
:Main articles: Colony Collapse Disorder
:Beekeepers in Western countries have been reporting slow declines of stocks since many years, apparently to changes in agricultural practice and perhaps climate change causing more unpredictable weather. In early 2007, abnormally high die-offs (30-70% of hives) of Western honey bee colonies occurred in the US and possibly Québec; 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 stochastically more adverse conditions in 2006, or a novel phenomenon. Research has hitherto failed to determine what causes it, but the weight of evidence is tentatively leaning towards CCD being a syndrome rather than a disease as it seems to be caused by a combination of various contributing factors rather than a single pathogen or poison.
★ Eastern species
:These are 2-4 species. The reddish Koschevnikov's Bee (''Apis koschevnikovi'') from Borneo is well distinct; it probably derives from the first colonizaion 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, kept in hives in a similar fashion to ''Apis mellifera'', though on a much smaller and regionalized scale. It has not been possible yet to resolve its relationship to the Bornean ''Apis cerana nuluensis'' and ''Apis nigrocincta'' from the Philippines to satisfaction; the most recent hypothesis is that these are indeed distinct species but that ''A. cerana'' is still paraphyletic, consisting of several good species.(Arias & Sheppard 2005)
Main articles: Beekeeping
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.
Like other eusocial bees, a colony generally contains one breeding female, or "queen"; seasonally up to a few thousand males, or "drones"; and a large seasonally variable population of sterile female workers. Many minor details vary among the different species of honeybees, though there are some common features.
Eggs are laid singly in a cell in a wax honeycomb, produced and shaped by the workers. 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.
Young worker bees clean the hive and feed the larvae. After this, they begin building comb cells. They progress to other within-colony tasks as they become older, such as receiving nectar and pollen from foragers. Later still, a worker leaves the hive and typically spends the remainder of its life as a forager.
Workers cooperate to find food and use a pattern of "dancing" (known as ''the bee dance or waggle dance'') to communicate with each other; this dance varies from species to species, but all living species of ''Apis'' exhibit some form of the behavior.
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 workers. This group moves ''en masse'' to a nest site that has been scouted by workers 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 workers, but the nest is constructed before a queen is escorted to the site, which is not a true "swarm".
Main articles: Pollination management
Main articles: List of plants pollinated by bees
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.
Main articles: 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. All living species of ''Apis'' have had their honey gathered by indigenous peoples for consumption, though for commercial purposes only ''A. mellifera'' and ''A. cerana'' have been exploited to any degree. Honey is sometimes also gathered by humans from the nests of various stingless bees.
Main articles: Beeswax
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, most indigenous peoples will gather beeswax for various purposes.
Main articles: Pollen
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.
Main articles: Propolis
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 also used in some cosmetics.
All honey bees live in colonies where the workers 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 workers. 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 workers, 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.
Main articles: Bee learning and communication
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 workers orient the dance in the actual compass direction of the resource to which they are recruiting.
A community of honey bees has often been employed throughout history by political theorists as a model of human society:
Honeybees, signifying immortality and resurrection, were royal emblems of the Napoleonic Empire. [1]
★ Apidae
★ Apiology
★ Bee
★ Beekeeping
★ Bees and toxic chemicals
★ Colony Collapse Disorder
★ List of plants pollinated by bees
1. Smith, Deborah R.; Villafuerte, Lynn ; Otisc, Gard; Palmer,Michael R. Biogeography of Apis cerana F. and A. nigrocincta Smith: insights from mtDNA studies Apidologie 31 (2000) 265-279
★ (2005): Phylogenetic relationships of honey bees (Hymenoptera:Apinae:Apini) inferred from nuclear and mitochondrial DNA sequence data. ''Molecular Phylogenetics and Evolution'' '37'(1): 25–35. . Erratum in ''Molecular Phylogenetics and Evolution'' '40'(1): 315.
★ (1999): The taxonomy of recent and fossil honey bees (Hymenoptera: Apidae: ''Apis''). ''Journal of Hymenoptera Research'' '8': 165-196.
★ (2005): ''Evolution of the Insects''. Cambridge University Press.
★ (1991): The Honey Bee Dance Language Controversy. ''The Mankind Quarterly'' Summer 1991: 357-365. HTML fulltext
★ (1971): ''Communication among social bees''. Harvard University Press.
★ (2004): ''The Hive: The Story Of The Honeybee''. John Murray, London, UK. ISBN 0-7195-6598-7
:''This article refers collectively to all true honey bees; for the "common" domesticated honey bee, see Western honey bee''
'Honey bees' (or 'honeybees') are a subset of bees which represent a far smaller fraction of bee diversity than most people suspect; of the approximately 20,000 known species of bees, there are only seven presently-recognized species with a total of 44 subspecies (Engel, 1999; historically, anywhere from six to eleven species have been recognized). These bees are the only living members of the tribe Apini, all in the genus ''Apis'', and all of which produce and store liquefied sugar ("honey") to some degree, and construct colonial nests out of wax secreted by the workers in the colony. Other types of related bees produce and store honey, but only members of the genus ''Apis'' are considered true honey bees.
| Contents |
| Origin, systematics and distribution |
| Beekeeping |
| Life cycle |
| Pollination |
| Honey |
| Beeswax |
| Pollen |
| Propolis |
| Defense |
| Communication |
| Symbolism |
| See also |
| References |
Origin, systematics and distribution
Morphology of a female honey bee.
Honey bees as a group appear to have their center of origin in 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''). [1] 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 honeybees 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'''
★ ''Apis florea'' and ''Apis andreniformis'' are small honey bees of southern and southeastern Asia. The make very small, exposed nests in trees and shrubs. The stings often not capable 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. 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 Bartonian (some 40 mya or slightly later) from the other lineages, but among themselves do not seem to have diverged a long time before the Neogene.(Arias & Sheppard 2005)
'Giant honey bees - subgenus ''Megapis'''
There is one recognized species which 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 when provoked. Their origin as a distinct lineage is only slightly more recent than that of the dwarf honey bees.
★ ''Apis dorsata'', the Giant honey bee proper, is native and widespread across most of South and Southeast Asia.
★ ''Apis dorsata binghami'', the Indonesian honey bee, is classified as the Indonesian subspecies of the Giant honey bee or a distinct species; in the latter case, ''A. d. breviligula'' and/or other lineages would probably also have to be considered species.
★ ''Apis dorsata laboriosa'', the Himalayan honey bee, was initially described as a distinct species. Later, it was included in ''A. dorsata'' as a subspecies (Engel 1999), though genetic studies suggest it is divergent enough to merit species status (Arias & Sheppard 2005). Essentially restricted to the Himalayas, it differs little from the Giant honey bee in appearance, but has extensive behavioral adaptations which enable it to nest in the open at high altitudes despite low ambient temperatures. It is the largest living honey bee.
'Cave-nesting honey bees - subgenus ''Apis'''
★ Western (European, Common) honey bee
Main articles: Apis mellifera
:''Apis mellifera'', the most commonly domesticated species, was the third insect to have its genome mapped. 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 Western, European or Common honey bee in different parts of the world. There are many subspecies that have adapted to the local geographic and climatic environment, and 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.
:Regarding phylogeny, this is the most enigmatic honey bee species. 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 E Africa and the Eastern group of tropical Asia by desertification in the Middle East and adjacent regions, which caused declines of foodplants and trees which provided nest sites, eventually causing gene flow to cease. The diversity of subspecies is probably the product of a (largely) Early Pleistocene radiation aided by climate and habitat changes during the last ice age. That the Western honey bee has been intensively managed by humans since many millennia - including hybridization and introductions - has apparently increased the speed of its evolution 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.(Arias & Sheppard 2005)
:In 1622, European colonists brought the dark bee (''A. m. mellifera'') to the Americas, followed later by Italian bees (''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 feral) spread rapidly as far as the Great Plains, usually preceding the colonists. The Native Americans called the honey bee "the white man's fly". Honey bees did not naturally cross the Rocky Mountains; they were carried by ship to California in the early 1850s.
:Main articles: Africanized bee
:The so-called "killer bee" are highly aggressive hybrids between European stock and the African subspecies ''A. m. scutellata''; they are thus often called "Africanized bees". Originating by accident in Brazil, they have spread to North America and constitutes a pest in some regions. 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, and are popular among beekeepers in Brazil.
:Main articles: Colony Collapse Disorder
:Beekeepers in Western countries have been reporting slow declines of stocks since many years, apparently to changes in agricultural practice and perhaps climate change causing more unpredictable weather. In early 2007, abnormally high die-offs (30-70% of hives) of Western honey bee colonies occurred in the US and possibly Québec; 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 stochastically more adverse conditions in 2006, or a novel phenomenon. Research has hitherto failed to determine what causes it, but the weight of evidence is tentatively leaning towards CCD being a syndrome rather than a disease as it seems to be caused by a combination of various contributing factors rather than a single pathogen or poison.
★ Eastern species
:These are 2-4 species. The reddish Koschevnikov's Bee (''Apis koschevnikovi'') from Borneo is well distinct; it probably derives from the first colonizaion 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, kept in hives in a similar fashion to ''Apis mellifera'', though on a much smaller and regionalized scale. It has not been possible yet to resolve its relationship to the Bornean ''Apis cerana nuluensis'' and ''Apis nigrocincta'' from the Philippines to satisfaction; the most recent hypothesis is that these are indeed distinct species but that ''A. cerana'' is still paraphyletic, consisting of several good species.(Arias & Sheppard 2005)
Beekeeping: Frame removed from Langstroth hive.
Beekeeping
Main articles: Beekeeping
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.
Life cycle
Queen bee. Yellow dot is added to aid beekeeper.
Honey bee eggs shown in cut open wax cells
Eggs and larvae
Foragers coming in loaded with pollen on the hive landing board.
Like other eusocial bees, a colony generally contains one breeding female, or "queen"; seasonally up to a few thousand males, or "drones"; and a large seasonally variable population of sterile female workers. Many minor details vary among the different species of honeybees, though there are some common features.
Eggs are laid singly in a cell in a wax honeycomb, produced and shaped by the workers. 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.
Young worker bees clean the hive and feed the larvae. After this, they begin building comb cells. They progress to other within-colony tasks as they become older, such as receiving nectar and pollen from foragers. Later still, a worker leaves the hive and typically spends the remainder of its life as a forager.
Workers cooperate to find food and use a pattern of "dancing" (known as ''the bee dance or waggle dance'') to communicate with each other; this dance varies from species to species, but all living species of ''Apis'' exhibit some form of the behavior.
Honey bee drinking
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 workers. This group moves ''en masse'' to a nest site that has been scouted by workers 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 workers, but the nest is constructed before a queen is escorted to the site, which is not a true "swarm".
Pollination
Main articles: Pollination management
Main articles: List of plants pollinated by bees
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
Main articles: Honey
Honey combs
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 ''A. mellifera'' and ''A. cerana'' have been exploited to any degree. Honey is sometimes also gathered by humans from the nests of various stingless bees.
Beeswax
Main articles: Beeswax
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, most indigenous peoples will gather beeswax for various purposes.
A forager collecting pollen.
Pollen
Main articles: Pollen
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
Main articles: Propolis
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 also used in some cosmetics.
Defense
''Apis cerana'' forming a ball around two hornets. The body heat trapped by the ball will overheat and kill the hornets.
All honey bees live in colonies where the workers 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 workers. 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 workers, 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.
Communication
Main articles: Bee learning and communication
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 workers orient the dance in the actual compass direction of the resource to which they are recruiting.
Symbolism
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. [1]
See also
★ Apidae
★ Apiology
★ Bee
★ Beekeeping
★ Bees and toxic chemicals
★ Colony Collapse Disorder
★ List of plants pollinated by bees
References
1. Smith, Deborah R.; Villafuerte, Lynn ; Otisc, Gard; Palmer,Michael R. Biogeography of Apis cerana F. and A. nigrocincta Smith: insights from mtDNA studies Apidologie 31 (2000) 265-279
★ (2005): Phylogenetic relationships of honey bees (Hymenoptera:Apinae:Apini) inferred from nuclear and mitochondrial DNA sequence data. ''Molecular Phylogenetics and Evolution'' '37'(1): 25–35. . Erratum in ''Molecular Phylogenetics and Evolution'' '40'(1): 315.
★ (1999): The taxonomy of recent and fossil honey bees (Hymenoptera: Apidae: ''Apis''). ''Journal of Hymenoptera Research'' '8': 165-196.
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