Origin, Systematics and Distribution
Honey
bees appear to have their center of origin in South and Southeast
Asia (including the Philippines), as all the extant species
except Apis mellifera are
native to that region. Notably, living representatives of the earliest lineages
to diverge (Apis florea and Apis andreniformis) have their center
of origin there.
The
first Apis bees
appear in the fossil record at the Eocene-Oligocene boundary
(34 mya),
in European deposits. The origin of these prehistoric honey bees does not
necessarily indicate Europe as the place of origin of the genus, only that the
bees were present in Europe by that time. Few fossil deposits are known from
South Asia, the suspected region of honey bee origin, and fewer still have been
thoroughly studied.
No Apis species existed in the New
World during human times before the introduction of A. mellifera by Europeans. Only
one fossil species is documented from the New World, Apis nearctica, known from a single
14 million-year-old specimen from Nevada.
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 their
domestication.
Most
species have historically been cultured or at least exploited for honey
and beeswax by humans indigenous to their native ranges. Only two
species have been truly domesticated: Apis mellifera and Apis cerana indica. A.
mellifera has been cultivated at least since the time of the
building of the Egyptian pyramids, and only that species has been moved
extensively beyond its native range.
Honey
bees are the only extant members of the tribe Apini. Today's honey
bees constitute three clades: Micrapis (dwarf
honey bees), Megapis (giant
honey bee), and Apis (domestic
honey bees and close relatives).
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 million years ago or slightly later) from the other lineages, but do
not seem to have diverged from each other a long time before the Neogene. Apis florea have smaller wing
spans than its sister species. Apis
florea are also completely yellow with the exception of the
scutellum of workers, which is black.
Megapis
One species is recognized in the
subgenus Megapis. It
usually builds single or a few exposed combs on high tree limbs, on cliffs, and
sometimes on buildings. They can be very fierce. Periodically robbed of their
honey by human "honey hunters", colonies are easily capable of stinging a
human being to death if provoked.
·
Apis
dorsata, the giant honey bee, is native and
widespread across most of South and Southeast Asia.
o A. d. binghami, the Indonesian giant 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.
o A. d. laboriosa, the Himalayan giant 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 separate species. Essentially
restricted to the Himalayas, it differs little from the giant honey bee in
appearance, but has extensive behavioral adaptations that enable it
to nest in the open at high altitudes despite low ambient temperatures. It is
the largest living honey bee.
Apis
Eastern Apis species include three or four species,
including A. koschevnikovi, Apis nigrocincta, and A. cerana. The genetics of the
western honey bee A. mellifera are
unclear.
Koschevnikov's honey bee
Koschevnikov's honey bee (Apis koschevnikovi) is often referred
to in the literature as the "red bee of Sabah"; however, A. koschevnikovi is pale reddish
in Sabah State, Borneo, Malaysia, but a dark, coppery color in
the Malay Peninsula and Sumatra, Indonesia. Its
habitat is limited to the tropical evergreen forests of the Malay
Peninsula, Borneo and Sumatra and they do not live in
tropical evergreen rain forests which extend into Thailand, Myanmar, Cambodia and Vietnam.
Philippine honey bee
Eastern honey bee
Apis
cerana, the eastern honey bee proper, is
the traditional honey bee of southern and eastern Asia. It was domesticated as
subspecies A. c. indica and
kept in hives in a fashion similar to A. mellifera, though on a more limited, regional scale.
It has not been possible yet to resolve
its relationship to the Bornean A.
c. 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 parphyletic,
consisting of several separate species.
Western honey bee
A.
mellifera, the most common 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 Europe and eastwards into Asia to
the Tien Shan range. It is variously called the European, western, or
common honey bee in different parts of the world. Many subspecies have
adapted to the local geographic and climatic environments; in addition breeds
such as the Buckfast Bee, have been bred. Behavior, color, and anatomy can
be quite different from one subspecies or even strain to another.
A.
mellifera phylogeny is the most
enigmatic of all 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 East Africa and the eastern group of tropical Asia by desertification in
the Middle East and adjacent regions, which caused declines of food
plants and trees that provided nest sites, eventually causing gene flow to
cease.
The diversity of A. mellifera subspecies is
probably the product of a largely Early Pleistocene radition aided by
climate and habitat changes during the Last ice age. That the western honey bee
has been intensively managed by humans for 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.
Apis mellifera is not native to the Americas, so it was not
present when the European explorers and colonists arrived. However, other
native bee species were kept and traded by indigenous peoples. In 1622,
European colonists brought the European dark bee (A. m. mellifera) to the Americas
first, followed later by the Italian honey bee (A. m. ligustica) and others. Many of the crops that depend on
western honey bees for pollination have also been imported since colonial
times. Escaped swarms (known as "wild" bees, but actually feral)
spread rapidly as far as the Great Plains, usually preceding the
colonists. Honey bees did not naturally cross the Rocky Mountain; they
were transported by the Mormon pioneers to Utah in the late 1840s, and by
ship to California in the early 1850s.
Africanized bee
Africanized bees (known colloquially as
"killer bees") are hybrids between European stock and the
East African lowland subspecies A.
m. scutellata; they are often more aggressive than European bees and do
not create as much of a honey surplus, but are more resistant to disease and
are better foragers. Accidentally released from quarantine in Brazil,
they have spread to North America and constitute a pest in some
regions. However, these strains do not overwinter well, so are not often found
in the colder, more northern parts of North America. The original breeding
experiment for which the African bees were brought to Brazil in the first place
has continued (though not as originally intended). Novel hybrid strains of
domestic and redomesticated Africanized bees combine high resilience to tropical
conditions and good yields. They are popular among beekeepers in Brazil.