Honey bee

Honey bee

Overview
This article refers collectively to all true honey bees; for the "common" domesticated honey bee, see European honey bee
European honey bee
The Western honey bee or European honey bee is a species of honey bee. The genus Apis is Latin for "bee", and mellifera comes from Latin melli- "honey" and ferre "to bear"—hence the scientific name means "honey-bearing bee"...

.

Honey bees (or honeybees) are a subset of bee
Bee
Bees are flying insects closely related to wasps and ants, and are known for their role in pollination and for producing honey and beeswax. Bees are a monophyletic lineage within the superfamily Apoidea, presently classified by the unranked taxon name Anthophila...

s in the genus Apis, primarily distinguished by the production and storage of honey
Honey
Honey is a sweet food made by bees using nectar from flowers. The variety produced by honey bees is the one most commonly referred to and is the type of honey collected by beekeepers and consumed by humans...

 and the construction of perennial, colonial
Colony (biology)
In biology, a colony reference to several individual organisms of the same species living closely together, usually for mutual benefit, such as stronger defense or the ability to attack bigger prey. Some insects live only in colonies...

 nests out of wax
Beeswax
Beeswax is a natural wax produced in the bee hive of honey bees of the genus Apis. It is mainly esters of fatty acids and various long chain alcohols...

. Honey bees are the only extant members of the tribe Apini, all in the genus Apis.
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Encyclopedia
This article refers collectively to all true honey bees; for the "common" domesticated honey bee, see European honey bee
European honey bee
The Western honey bee or European honey bee is a species of honey bee. The genus Apis is Latin for "bee", and mellifera comes from Latin melli- "honey" and ferre "to bear"—hence the scientific name means "honey-bearing bee"...

.

Honey bees (or honeybees) are a subset of bee
Bee
Bees are flying insects closely related to wasps and ants, and are known for their role in pollination and for producing honey and beeswax. Bees are a monophyletic lineage within the superfamily Apoidea, presently classified by the unranked taxon name Anthophila...

s in the genus Apis, primarily distinguished by the production and storage of honey
Honey
Honey is a sweet food made by bees using nectar from flowers. The variety produced by honey bees is the one most commonly referred to and is the type of honey collected by beekeepers and consumed by humans...

 and the construction of perennial, colonial
Colony (biology)
In biology, a colony reference to several individual organisms of the same species living closely together, usually for mutual benefit, such as stronger defense or the ability to attack bigger prey. Some insects live only in colonies...

 nests out of wax
Beeswax
Beeswax is a natural wax produced in the bee hive of honey bees of the genus Apis. It is mainly esters of fatty acids and various long chain alcohols...

. 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.

Origin, systematics and distribution


Honey bees as a group appear to have their centre of origin in South
South Asia
South Asia, also known as Southern Asia, is the southern region of the Asian continent, which comprises the sub-Himalayan countries and, for some authorities , also includes the adjoining countries to the west and the east...

 and South East Asia (including the Philippines
Philippines
The Philippines , officially known as the Republic of the Philippines , is a country in Southeast Asia in the western Pacific Ocean. To its north across the Luzon Strait lies Taiwan. West across the South China Sea sits Vietnam...

), as all but one (i.e. Apis mellifera), of the extant species are native to that region. Notably the most plesiomorphic living species (Apis florea
Apis florea
The dwarf honey bee or the red dwarf honey bee, Apis florea, is one of two species of small, wild honey bees of southern and southeastern Asia. It has a much wider distribution than its sister species, Apis andreniformis....

and Apis andreniformis
Apis andreniformis
Apis andreniformis or the black dwarf honey bee is a species of honey bee whose native habitat is the tropical and subtropical regions of Asia....

) has the center of origin there.. The age of Apis melifera specie is estimated for only 1 million-year old.

The first Apis bees appear in the fossil record at the Eocene
Eocene
The Eocene Epoch, lasting from about 56 to 34 million years ago , is a major division of the geologic timescale and the second epoch of the Paleogene Period in the Cenozoic Era. The Eocene spans the time from the end of the Palaeocene Epoch to the beginning of the Oligocene Epoch. The start of the...

Oligocene
Oligocene
The Oligocene is a geologic epoch of the Paleogene Period and extends from about 34 million to 23 million years before the present . As with other older geologic periods, the rock beds that define the period are well identified but the exact dates of the start and end of the period are slightly...

 (23-56 Mya) boundary, in European deposits. 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 South Asia, the suspected region of honey bee origin, and fewer still have been thoroughly studied.

No Apis specie existed in New World in human time before introduction of Apis melifera by Europeans. There is only one fossil species documented from the New World, Apis nearctica
Apis nearctica
Apis nearctica is an extinct species of honey bee which existed in what is now the United States, during the middle Miocene period. It was described by Michael S. Engel, Ismael A. Hinojosa-Díaz, and Alexandr P. Rasnitsyn in 2009, and is the first fossil honey bee described from the New World....

, known from a single 14-million-year old specimen from Nevada.

The close relatives of modern honey bees –- e.g. bumblebee
Bumblebee
A bumble bee is any member of the bee genus Bombus, in the family Apidae. There are over 250 known species, existing primarily in the Northern Hemisphere although they are common in New Zealand and in the Australian state of Tasmania.Bumble bees are social insects that are characterised by black...

s and stingless bee
Stingless bee
Stingless bees, sometimes called stingless honey bees or simply meliponines, are a large group of bees, comprising the tribe Meliponini . They belong in the family Apidae, and are closely related to common honey bees, carpenter bees, orchid bees and bumblebees...

s –- are also social to some degree, and 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
Beeswax
Beeswax is a natural wax produced in the bee hive of honey bees of the genus Apis. It is mainly esters of fatty acids and various long chain alcohols...

 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 clade
Clade
A clade is a group consisting of a species and all its descendants. In the terms of biological systematics, a clade is a single "branch" on the "tree of life". The idea that such a "natural group" of organisms should be grouped together and given a taxonomic name is central to biological...

s.


Genetics


The chromosomes 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. Further information is available in .

Drones are produced from unfertilized eggs and therefore represent only the DNA
DNA
Deoxyribonucleic acid is a nucleic acid that contains the genetic instructions used in the development and functioning of all known living organisms . The DNA segments that carry this genetic information are called genes, but other DNA sequences have structural purposes, or are involved in...

of the queen that laid the eggs, i.e. have only a mother. Workers and queens result from fertilized eggs and therefore have both a mother and a father. A modified form of Parthenogenesis
Parthenogenesis
Parthenogenesis is a form of asexual reproduction found in females, where growth and development of embryos occur without fertilization by a male...

controls sex differentiation. The sex Allele
Allele
An allele is one of two or more forms of a gene or a genetic locus . "Allel" is an abbreviation of allelomorph. Sometimes, different alleles can result in different observable phenotypic traits, such as different pigmentation...

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. 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 Spermatheca
Spermatheca
The spermatheca , also called receptaculum seminis , is an organ of the female reproductive tract in insects, some molluscs, oligochaeta worms and certain other invertebrates and vertebrates...

. 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


Apis florea
Apis florea
The dwarf honey bee or the red dwarf honey bee, Apis florea, is one of two species of small, wild honey bees of southern and southeastern Asia. It has a much wider distribution than its sister species, Apis andreniformis....

and Apis andreniformis
Apis andreniformis
Apis andreniformis or the black dwarf honey bee is a species of honey bee whose native habitat is the tropical and subtropical regions of Asia....

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
Beehive (beekeeping)
A beehive is an enclosed structure in which some honey bee species of the subgenus Apis live and raise their young. Natural beehives are naturally occurring structures occupied by honeybee colonies, while domesticated honeybees live in man-made beehives, often in an apiary. These man-made...

 and swarm
Swarm
Swarm behaviour, or swarming, is a collective behaviour exhibited by animals of similar size which aggregate together, perhaps milling about the same spot or perhaps moving en masse or migrating in some direction. As a term, swarming is applied particularly to insects, but can also be applied to...

s can be handled with minimal protection. They occur largely sympatrically though they are very distinct evolution
Evolution
Evolution is any change across successive generations in the heritable characteristics of biological populations. Evolutionary processes give rise to diversity at every level of biological organisation, including species, individual organisms and molecules such as DNA and proteins.Life on Earth...

arily and are probably the result of allopatric speciation
Allopatric speciation
Allopatric speciation or geographic speciation is speciation that occurs when biological populations of the same species become isolated due to geographical changes such as mountain building or social changes such as emigration...

, 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
Bartonian
The Bartonian is, in the ICS's geological timescale, a stage or age in the middle Eocene epoch or series. The Bartonian age spans the time between and . It is preceded by the Lutetian and is followed by the Priabonian age.-Stratigraphic definition:...

 (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 Neogene
Neogene
The Neogene is a geologic period and system in the International Commission on Stratigraphy Geologic Timescale starting 23.03 ± 0.05 million years ago and ending 2.588 million years ago...

.

Megapis


There is one recognised species in 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 when provoked.


  • Apis dorsata
    Apis dorsata
    Apis dorsata, the giant honey bee, is a honey bee of southern and southeastern Asia mainly in forested areas like the Terai of Nepal...

    , the giant honey bee, is native and widespread across most of South and Southeast Asia.
  • Apis dorsata binghami, the Indonesian honey bee, is classified as the Indonesia
    Indonesia
    Indonesia , officially the Republic of Indonesia , is a country in Southeast Asia and Oceania. Indonesia is an archipelago comprising approximately 13,000 islands. It has 33 provinces with over 238 million people, and is the world's fourth most populous country. Indonesia is a republic, with an...

    n 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
    Apis dorsata laboriosa
    Apis dorsata laboriosa, the Himalayan honey bee, is the world’s largest honey bee; single adults can measure up to in length. Before 1980, Apis dorsata laboriosa was considered to be a subspecies of the widespread Apis dorsata, the giant honey bee, but in 1980 and for almost 20 years thereafter...

    , the Himalayan honey bee, was initially described as a distinct species. Later, it was included in A. dorsata as a subspecies based on the biological species concept, though authors applying a genetic species concept have suggested it should be considered a species. Essentially restricted to the Himalayas
    Himalayas
    The Himalaya Range or Himalaya Mountains Sanskrit: Devanagari: हिमालय, literally "abode of snow"), usually called the Himalayas or Himalaya for short, is a mountain range in Asia, separating the Indian subcontinent from the Tibetan Plateau...

    , 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.

Apis


Eastern species

These are three or four species. The reddish Koschevnikov's bee (Apis koschevnikovi
Apis koschevnikovi
Koschevnikov's bee, or Apis koschevnikovi, is a species of honey bee which inhabits Malaysian and Indonesian Borneo, where it lives conspecifically with other honey bee species such as Apis cerana Koschevnikov's bee, or Apis koschevnikovi, is a species of honey bee which inhabits Malaysian and...

) from Borneo
Borneo
Borneo is the third largest island in the world and is located north of Java Island, Indonesia, at the geographic centre of Maritime Southeast Asia....

 is well distinct; it probably derives from the first colonization of the island by cave-nesting honey bees. Apis cerana
Apis cerana
Apis cerana, or the Asiatic honey bee , is a species of honey bee found in southern and southeastern Asia, such as China, India, Japan, Malaysia, Nepal, Bangladesh and Papua New Guinea. This species is the sister species of Apis koschevnikovi, and both are in the same subgenus as the Western honey...

, 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 regionalised scale. It has not been possible yet to resolve its relationship to the Bornean Apis cerana nuluensis
Apis cerana nuluensis
Apis cerana nuluensis is a subspecies of honey bee described in 1996 by Tingek, Koeniger & Koeniger. The geographic distribution of the subspecies is the south-east Asian island of Borneo, politically divided between Indonesia, Malaysia, and Brunei....

and Apis nigrocincta
Apis nigrocincta
Apis nigrocincta is a species of honey bee that inhabits the Philippine island of Mindanao as well as the Indonesian islands of Sangihe and the Celebes or Sulawesi.The species builds nests in cavities like the closely related Apis cerana...

from the Philippines
Philippines
The Philippines , officially known as the Republic of the Philippines , is a country in Southeast Asia in the western Pacific Ocean. To its north across the Luzon Strait lies Taiwan. West across the South China Sea sits Vietnam...

 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.

European/Western/Common honey bee
Apis mellifera, the most commonly domesticated species, was the third insect to have its genome
Genome
In modern molecular biology and genetics, the genome is the entirety of an organism's hereditary information. It is encoded either in DNA or, for many types of virus, in RNA. The genome includes both the genes and the non-coding sequences of the DNA/RNA....

 mapped. It seems to have originated in eastern tropical Africa
Africa
Africa is the world's second largest and second most populous continent, after Asia. At about 30.2 million km² including adjacent islands, it covers 6% of the Earth's total surface area and 20.4% of the total land area...

 and spread from there to Northern Europe
Northern Europe
Northern Europe is the northern part or region of Europe. Northern Europe typically refers to the seven countries in the northern part of the European subcontinent which includes Denmark, Estonia, Latvia, Lithuania, Norway, Finland and Sweden...

 and eastwards into Asia
Asia
Asia is the world's largest and most populous continent, located primarily in the eastern and northern hemispheres. It covers 8.7% of the Earth's total surface area and with approximately 3.879 billion people, it hosts 60% of the world's current human population...

 to the Tien Shan range. It is variously called the European, Western or Common honey bee in different parts of the world. There are many subspecies
Subspecies
Subspecies in biological classification, is either a taxonomic rank subordinate to species, ora taxonomic unit in that rank . A subspecies cannot be recognized in isolation: a species will either be recognized as having no subspecies at all or two or more, never just one...

 that have adapted to the local geographic and climatic environment, and in addition, hybrid strains such as the Buckfast bee
Buckfast bee
The Buckfast bee is a honey bee a cross of many species of bees into a stable race developed by "Brother Adam", , who was in charge of beekeeping at Buckfast Abbey, where the bees are still bred today.-Origin:...

 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
Late Miocene
The Late Miocene is a sub-epoch of the Miocene Epoch made up of two stages. The Tortonian and Messinian stages comprise the Late Miocene sub-epoch....

. 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
Desertification
Desertification is the degradation of land in drylands. Caused by a variety of factors, such as climate change and human activities, desertification is one of the most significant global environmental problems.-Definitions:...

 in the Middle East
Middle East
The Middle East is a region that encompasses Western Asia and Northern Africa. It is often used as a synonym for Near East, in opposition to Far East...

 and adjacent regions, which caused declines of foodplants and trees which provided nest sites, eventually causing gene flow
Gene flow
In population genetics, gene flow is the transfer of alleles of genes from one population to another.Migration into or out of a population may be responsible for a marked change in allele frequencies...

 to cease. The diversity of subspecies is probably the product of a largely Early Pleistocene
Early Pleistocene
Calabrian is a subdivision of the Pleistocene Epoch of the Geologic time scale. ~1.8 Ma.—781,000 years ago ± 5,000 years, a period of ~.The end of the stage is defined by the last magnetic pole reversal and plunge in to an ice age and global drying possibly colder and drier than the late Miocene ...

 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
Evolution
Evolution is any change across successive generations in the heritable characteristics of biological populations. Evolutionary processes give rise to diversity at every level of biological organisation, including species, individual organisms and molecules such as DNA and proteins.Life on Earth...

 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.

There are no honey bees native to the Americas
Americas
The Americas, or America , are lands in the Western hemisphere, also known as the New World. In English, the plural form the Americas is often used to refer to the landmasses of North America and South America with their associated islands and regions, while the singular form America is primarily...

. In 1622, European colonists brought the dark bee (A. m. mellifera) to the Americas, followed later by Italian bee
Italian bee
Apis mellifera ligustica is the Italian bee which is a sub-species of the western honey bee .- Origin :The Italian honey bee is thought to originate from the continental part of Italy, South of the Alps, and North of Sicily...

s (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
Feral
A feral organism is one that has changed from being domesticated to being wild or untamed. In the case of plants it is a movement from cultivated to uncultivated or controlled to volunteer. The introduction of feral animals or plants to their non-native regions, like any introduced species, may...

) spread rapidly as far as the Great Plains
Great Plains
The Great Plains are a broad expanse of flat land, much of it covered in prairie, steppe and grassland, which lies west of the Mississippi River and east of the Rocky Mountains in the United States and Canada. This area covers parts of the U.S...

, usually preceding the colonists. Honey bees did not naturally cross the Rocky Mountains
Rocky Mountains
The Rocky Mountains are a major mountain range in western North America. The Rocky Mountains stretch more than from the northernmost part of British Columbia, in western Canada, to New Mexico, in the southwestern United States...

; they were carried by ship to California
California
California is a state located on the West Coast of the United States. It is by far the most populous U.S. state, and the third-largest by land area...

 in the early 1850s.

Africanized bee



Africanized bees (known colloquially as "killer bees") are hybrids between European stock and one of the African subspecies A. m. scutellata; 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 pest
Pest (animal)
A pest is an animal which is detrimental to humans or human concerns. It is a loosely defined term, often overlapping with the related terms vermin, weeds, parasites and pathogens...

 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 North America
North America
North America is a continent wholly within the Northern Hemisphere and almost wholly within the Western Hemisphere. It is also considered a northern subcontinent of the Americas...

; 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 stochastic
Stochastic
Stochastic refers to systems whose behaviour is intrinsically non-deterministic. A stochastic process is one whose behavior is non-deterministic, in that a system's subsequent state is determined both by the process's predictable actions and by a random element. However, according to M. Kac and E...

ally 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 syndrome
Syndrome
In medicine and psychology, a syndrome is the association of several clinically recognizable features, signs , symptoms , phenomena or characteristics that often occur together, so that the presence of one or more features alerts the physician to the possible presence of the others...

 rather than a disease
Disease
A disease is an abnormal condition affecting the body of an organism. It is often construed to be a medical condition associated with specific symptoms and signs. It may be caused by external factors, such as infectious disease, or it may be caused by internal dysfunctions, such as autoimmune...

 as it seems to be caused by a combination of various contributing factors rather than a single pathogen
Pathogen
A pathogen gignomai "I give birth to") or infectious agent — colloquially, a germ — is a microbe or microorganism such as a virus, bacterium, prion, or fungus that causes disease in its animal or plant host...

 or poison
Poison
In the context of biology, poisons are substances that can cause disturbances to organisms, usually by chemical reaction or other activity on the molecular scale, when a sufficient quantity is absorbed by an organism....

, though the Israel acute paralysis virus has recently emerged as a significant candidate.

Recent research (2009) has found that an indicator for an impaired protein
Protein
Proteins are biochemical compounds consisting of one or more polypeptides typically folded into a globular or fibrous form, facilitating a biological function. A polypeptide is a single linear polymer chain of amino acids bonded together by peptide bonds between the carboxyl and amino groups of...

 production is common among all bees affected by CCD. It is conjectured that Dicistroviridae
Dicistroviridae
The Dicistroviridae are a family of Group IV insect-infecting viruses. Some of the insects commonly infected by dicistroviruses include aphids, leafhoppers, flies, bees, ants, silkworms.-Taxonomy:...

 like the IAPV may influence the genetic material of the ribosome
Ribosome
A ribosome is a component of cells that assembles the twenty specific amino acid molecules to form the particular protein molecule determined by the nucleotide sequence of an RNA molecule....

s, which are responsible for protein production of cells
Cell (biology)
The cell is the basic structural and functional unit of all known living organisms. It is the smallest unit of life that is classified as a living thing, and is often called the building block of life. The Alberts text discusses how the "cellular building blocks" move to shape developing embryos....

.

Life cycle




As in a few other types of eusocial bees, a colony generally contains one queen bee
Queen bee
The term queen bee is typically used to refer to an adult, mated female that lives in a honey bee colony or hive; she is usually the mother of most, if not all, the bees in the hive. The queens are developed from larvae selected by worker bees and specially fed in order to become sexually mature...

, a fertile female; seasonally up to a few thousand drone bees or fertile males; and a large seasonally variable population of sterile female worker bee
Worker bee
A Worker bee is any female eusocial bee that lacks the full reproductive capacity of the colony's queen bee; under most circumstances, this is correlated to an increase in certain non-reproductive activities relative to a queen, as well...

s. Details vary among the different species of honey bees, but common features include:

1. Eggs are laid singly in a cell in a wax honeycomb
Honeycomb
A honeycomb is a mass of hexagonal waxcells built by honey bees in their nests to contain their larvae and stores of honey and pollen.Beekeepers may remove the entire honeycomb to harvest honey...

, produced and shaped by the worker bees. Using her spermatheca
Spermatheca
The spermatheca , also called receptaculum seminis , is an organ of the female reproductive tract in insects, some molluscs, oligochaeta worms and certain other invertebrates and vertebrates...

, 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 haploid, while females (queens and worker bees) develop from fertilised eggs and are diploid. Larvae are initially fed with royal jelly
Royal jelly
Royal jelly is a honey bee secretion that is used in the nutrition of larvae, as well as adult queens. It is secreted from the glands in the hypopharynx of worker bees, and fed to all larvae in the colony....

 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.

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 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".

4. Honey bees also perform tremble dance
Tremble dance
A tremble dance is a dance performed by forager honey bees of the species Apis mellifera to recruit more receiver honey bees to collect nectar from the workers...

s 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 "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
Vespid
The Vespidae are a large , diverse, cosmopolitan family of wasps, including nearly all the known eusocial wasps and many solitary wasps. Each social wasp colony includes a queen and a number of female workers with varying degrees of sterility relative to the queen. In temperate social species,...

 wasps which also found new nests via swarming (sometimes including multiple queens). Also, stingless bee
Stingless bee
Stingless bees, sometimes called stingless honey bees or simply meliponines, are a large group of bees, comprising the tribe Meliponini . They belong in the family Apidae, and are closely related to common honey bees, carpenter bees, orchid bees and bumblebees...

s 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 10 °C (50 °F) 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 27 °C (81 °F) at the start of winter (during the broodless period) and 34 °C (93 °F) 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 30 to 100 lbs.

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 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


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 Apis mellifera and Apis cerana
Apis cerana
Apis cerana, or the Asiatic honey bee , is a species of honey bee found in southern and southeastern Asia, such as China, India, Japan, Malaysia, Nepal, Bangladesh and Papua New Guinea. This species is the sister species of Apis koschevnikovi, and both are in the same subgenus as the Western honey...

have been exploited to any degree. Honey is sometimes also gathered by humans from the nests of various stingless bee
Stingless bee
Stingless bees, sometimes called stingless honey bees or simply meliponines, are a large group of bees, comprising the tribe Meliponini . They belong in the family Apidae, and are closely related to common honey bees, carpenter bees, orchid bees and bumblebees...

s.

Beeswax



Worker bees of a certain age will secrete beeswax
Beeswax
Beeswax is a natural wax produced in the bee hive of honey bees of the genus Apis. It is mainly esters of fatty acids and various long chain alcohols...

 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.

Pollen



Bees collect pollen in the pollen basket
Pollen basket
The pollen basket or corbicula is part of the tibia on the hind legs of the four related lineages of apid bees that used to comprise the family Apidae: the honey bees, bumblebees, stingless bees, and orchid bees...

 and carry it back to the hive. In the hive, pollen is used as a protein
Protein
Proteins are biochemical compounds consisting of one or more polypeptides typically folded into a globular or fibrous form, facilitating a biological function. A polypeptide is a single linear polymer chain of amino acids bonded together by peptide bonds between the carboxyl and amino groups of...

 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



Propolis
Propolis
Propolis is a resinous mixture that honey bees collect from tree buds, sap flows, or other botanical sources. It is used as a sealant for unwanted open spaces in the hive. Propolis is used for small gaps , while larger spaces are usually filled with beeswax. Its color varies depending on its...

 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
Apis florea
The dwarf honey bee or the red dwarf honey bee, Apis florea, is one of two species of small, wild honey bees of southern and southeastern Asia. It has a much wider distribution than its sister species, Apis andreniformis....

 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.

Defense



All honey bees live in colonies where the worker bees will sting
Bee sting
A bee sting is strictly a sting from a bee . In the vernacular it can mean a sting of a bee, wasp, hornet, or yellow jacket. Some people may even call the bite of a horse-fly a bee sting...

 intruders as a form of defense, and alarmed bees will release a pheromone
Pheromone
A pheromone is a secreted or excreted chemical factor that triggers a social response in members of the same species. Pheromones are chemicals capable of acting outside the body of the secreting individual to impact the behavior of the receiving individual...

 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
Hymenoptera
Hymenoptera is one of the largest orders of insects, comprising the sawflies, wasps, bees and ants. There are over 130,000 recognized species, with many more remaining to be described. The name refers to the heavy wings of the insects, and is derived from the Ancient Greek ὑμήν : membrane and...

) by the possession of small barbs on the sting, but these barbs are found only in the worker bees. The sting and associated venom
Apitoxin
Apitoxin, or honey bee venom, is a bitter colorless liquid. The active portion of the venom is a complex mixture of proteins, which causes local inflammation and acts as an anticoagulant. The venom is produced in the abdomen of worker bees from a mixture of acidic and basic secretions. Apitoxin...

 sac are also modified so as to pull free of the body once lodged (autotomy
Autotomy
Autotomy or self amputation is the act whereby an animal severs one or more of its own appendages, usually as a self-defense mechanism designed to elude a predator's grasp...

), and the sting apparatus has its own musculature and ganglion
Ganglion
In anatomy, a ganglion is a biological tissue mass, most commonly a mass of nerve cell bodies. Cells found in a ganglion are called ganglion cells, though this term is also sometimes used to refer specifically to retinal ganglion cells....

 which allow it to keep delivering venom once detached. The worker bee dies after the stinger
Stinger
-Biology:* Stinger, an organ or body part found in various animals that usually delivers some kind of venom.* Stinger , a minor neurological injury suffered by athletes.-Sports and entertainment:...

 is torn from its body. As with other forms of life, warnings are given before an attack is launched. In the case of some honey bees species in the wild, this takes the form of a 'Mexican wave' which spreads as a ripple across a layer of bees densely packed on the surface of a comb when a threat is perceived, and consists of bees momentarily arching their bodies and flicking their wings.

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
Apis cerana
Apis cerana, or the Asiatic honey bee , is a species of honey bee found in southern and southeastern Asia, such as China, India, Japan, Malaysia, Nepal, Bangladesh and Papua New Guinea. This species is the sister species of Apis koschevnikovi, and both are in the same subgenus as the Western honey...

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. 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.

Communication



Honey bees are known to communicate through many different chemicals and odours, 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, Apis andreniformis
Apis andreniformis
Apis andreniformis or the black dwarf honey bee is a species of honey bee whose native habitat is the tropical and subtropical regions of Asia....

and Apis florea
Apis florea
The dwarf honey bee or the red dwarf honey bee, Apis florea, is one of two species of small, wild honey bees of southern and southeastern Asia. It has a much wider distribution than its sister species, Apis andreniformis....

, 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.

Symbolism



Both the Atharva Veda and the ancient Greeks associated lips anointed with honey with the gift of eloquence and even of prescience. The priestess at Delphi
Delphi
Delphi is both an archaeological site and a modern town in Greece on the south-western spur of Mount Parnassus in the valley of Phocis.In Greek mythology, Delphi was the site of the Delphic oracle, the most important oracle in the classical Greek world, and a major site for the worship of the god...

 was the "Delphic Bee".

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
Aristotle
Aristotle was a Greek philosopher and polymath, a student of Plato and teacher of Alexander the Great. His writings cover many subjects, including physics, metaphysics, poetry, theater, music, logic, rhetoric, linguistics, politics, government, ethics, biology, and zoology...

 and Plato
Plato
Plato , was a Classical Greek philosopher, mathematician, student of Socrates, writer of philosophical dialogues, and founder of the Academy in Athens, the first institution of higher learning in the Western world. Along with his mentor, Socrates, and his student, Aristotle, Plato helped to lay the...

; in Virgil
Virgil
Publius Vergilius Maro, usually called Virgil or Vergil in English , was an ancient Roman poet of the Augustan period. He is known for three major works of Latin literature, the Eclogues , the Georgics, and the epic Aeneid...

 and Seneca
Seneca the Younger
Lucius Annaeus Seneca was a Roman Stoic philosopher, statesman, dramatist, and in one work humorist, of the Silver Age of Latin literature. He was tutor and later advisor to emperor Nero...

; in Erasmus
Desiderius Erasmus
Desiderius Erasmus Roterodamus , known as Erasmus of Rotterdam, was a Dutch Renaissance humanist, Catholic priest, and a theologian....

 and Shakespeare
William Shakespeare
William Shakespeare was an English poet and playwright, widely regarded as the greatest writer in the English language and the world's pre-eminent dramatist. He is often called England's national poet and the "Bard of Avon"...

; in Marx
Karl Marx
Karl Heinrich Marx was a German philosopher, economist, sociologist, historian, journalist, and revolutionary socialist. His ideas played a significant role in the development of social science and the socialist political movement...

 and Tolstoy
Leo Tolstoy
Lev Nikolayevich Tolstoy was a Russian writer who primarily wrote novels and short stories. Later in life, he also wrote plays and essays. His two most famous works, the novels War and Peace and Anna Karenina, are acknowledged as two of the greatest novels of all time and a pinnacle of realist...

."


Honey bees, signifying immortality and resurrection, were royal emblems of the Merovingians, revived by Napoleon
Napoleon I of France
Napoleon Bonaparte was a French military and political leader during the latter stages of the French Revolution.As Napoleon I, he was Emperor of the French from 1804 to 1815...

. The bee is the heraldic emblem too of the Barberini
Barberini
The Barberini are a family of the Italian nobility that rose to prominence in 17th century Rome. Their influence peaked with the election of Cardinal Maffeo Barberini to the papal throne in 1623, as Pope Urban VIII...

.

See also



  • Apiology
    Apiology
    Apiology is the scientific study of honey bees, a subdiscipline of melittology, which is itself a branch of entomology...

  • Bees and toxic chemicals
    Bees and toxic chemicals
    Bees can suffer serious effects from toxic chemicals in their environments. These include various synthetic chemicals, such as insecticides and fertilizers, as well as a variety of naturally occurring chemicals from plants, such as ethanol resulting from the fermentation of organic material...

  • Honey bee life cycle
  • Bee sting
    Bee sting
    A bee sting is strictly a sting from a bee . In the vernacular it can mean a sting of a bee, wasp, hornet, or yellow jacket. Some people may even call the bite of a horse-fly a bee sting...



Further reading

  • Engel, Michael S.
    Michael S. Engel
    Michael S. Engel is an American paleontologist and entomologist. He has undertaken field work in Central Asia, Asia Minor, and the Western Hemisphere, and published more than 300 papers in scientific journals. He was trained at the University of Kansas where in 1993 he received a B.S. in Cellular...

     & Grimaldi, David
    David Grimaldi
    David A. Grimaldi is an entomologist and Curator of Invertebrate Zoology at the American Museum of Natural History in New York. He received his graduate training at Cornell University, where he earned his doctorate in Entomology in 1986. Dr. Grimaldi is an authority in many fields of insect...

     (2005): Evolution of the Insects. Cambridge University Press
    Cambridge University Press
    Cambridge University Press is the publishing business of the University of Cambridge. Granted letters patent by Henry VIII in 1534, it is the world's oldest publishing house, and the second largest university press in the world...

    .
  • 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. Harvard University Press
    Harvard University Press
    Harvard University Press is a publishing house established on January 13, 1913, as a division of Harvard University, and focused on academic publishing. In 2005, it published 220 new titles. It is a member of the Association of American University Presses. Its current director is William P...

    .