Genus:Apis Linnaeus, 17
Subgenus Micrapis: Apis andreniformis Apis florea, or dwarf honey bee
Subgenus Megapis: Apis dorsata, or giant honey bee
Subgenus Apis: Apis cerana, or eastern honey bee Apis koschevnikovi Apis mellifera,or
western honey bee Apis nigrocincta
Honey bees (or honeybees) are a subset of bees, primarily distinguished by the
production and storage of honey and the construction of perennial, 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.
Origin, systematics and distribution
Morphology of a female honey bee.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 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
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):
Two views of an Apis florea dwarf honey bee nest in Thailand. Dwarf honey
bees – subgenus Micrapis
Apis florea and Apis andreniformis are small honey bees of southern and
southeastern Asia. They make very small, exposed nests in trees and shrubs.
Their stings are often incapable of penetrating human skin, so the hive and
swarms can be handled with minimal protection. They occur largely
sympatrically though they are very distinct evolutionarily and are probably
the result of allopatric speciation, their distribution later converging.
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 nest, Thailand. The comb is approximately 1m across.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 on combApis 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) based on the biological species concept,
though authors applying a genetic species concept have suggested it should be
considered a species (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
Cavity-nesting honey bees – subgenus Apis
Eastern honey bee (Apis cerana) from Hong Kong.Eastern species
These are 3 or 4 species. The reddish Koschevnikov's Bee (Apis koschevnikovi)
from Borneo is well distinct; it probably derives from the first colonization
of the island by cave-nesting honey bees. Apis cerana, the Eastern honey bee
proper, is the traditional honey bee of southern and eastern Asia, 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).
European (Western, Common) honey bee
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 European, Western 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 &
There are no honey bees native to the Americas. 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
Widely known as the "killer bee", Africanized bees 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. 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, and are popular among beekeepers in Brazil.
Beekeeping: Frame removed from Langstroth hive.Main article: Colony Collapse
Beekeepers in Western countries have been reporting slow declines of stocks for
many years, apparently due 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 European 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.
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
Queen bee. Yellow dot is added to aid beekeeper.
Honey bee eggs shown in cut open wax cells
Emergence of a black bee (Apis mellifera mellifera).
Eggs and larvae
Foragers coming in loaded with pollen on the hive landing board.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 her 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 bee drinkingHoney 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 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 combsHoney 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 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.
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
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
(and virtually all other Hymenoptera) 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. The worker bee dies after
the sting is torn out of its body.
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.
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 worker bees orient the dance
in the actual compass direction of the resource to which they are recruiting.
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
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 and Plato; in Virgil and Seneca; in Erasmus
and Shakespeare; in Marx and Tolstoy." (Wilson 2004: p.4)
Honey bees, signifying immortality and resurrection, were royal emblems of the
Merovingians, revived by Napoleon. The bee is the heraldic emblem too of the