honey bee (also spelled honeybee) is a eusocial flying insect within the genus Apis of the bee clade, all native to Eurasia but spread to four other continents by human beings.

They are known for construction of perennial, colonial nests from wax, for the large size of their colonies, and for their surplus production and storage of honey, distinguishing their hives as a prized foraging target of many animals, including honey badgers, bears and human hunter-gatherers.

Only eight surviving species of honey bee are recognized, with a total of 43 subspecies, though historically 7 to 11 species are recognized. Honey bees represent only a small fraction of the roughly 20,000 known species of bees.

The best known honey bee is the western honey bee (Apis mellifera), which has been domesticated for honey production and crop pollination; the only other domesticated bee is the eastern honey bee (Apis cerana), which occurs in South Asia.

Some other types of related bees produce and store honey and have been kept by humans for that purpose, including the stingless honey bees, but only members of the genus Apis are true honey bees. Modern humans also value the wax for use in making candlessoaplip balms, and other products.

Etymology and name

The genus name Apis is Latin for “bee”.

Although modern dictionaries may refer to Apis as either honey bee or honeybee, entomologist Robert Snodgrass asserts that correct usage requires two words, i.e. honey bee, as it is a kind or type of bee, whereas it is incorrect to run the two words together, as in dragonfly or butterfly, because the latter are not flies, and have no connection with dragons or butter.

Honey bee, not honeybee, is the listed common name in the Integrated Taxonomic Information System, the Entomological Society of America Common Names of Insects Database, and the Tree of Life Web Project.

Origin, systematics, and distribution

Distribution of honey bees around the world
Morphology of a sterile female worker honey bee

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.

Species

While about 20,000 species of bees exist, only eight species of honey bee are recognized, with a total of 43 subspecies, although historically seven to 11 species are recognized: Apis andreniformis (the black dwarf honey bee); Apis cerana (the eastern honey bee); Apis dorsata (the giant honey bee); Apis florea (the red dwarf honey bee); Apis koschevnikovi (Koschevnikov’s honey bee); Apis laboriosa (the Himalayan giant honey bee); Apis mellifera (the western honey bee); and Apis nigrocincta (the Philippine honey bee).

Honey bees are the only extant members of the tribe Apini.

Today’s honey bees constitute three clades: Micrapis (the dwarf honey bees), Megapis (the giant honey bee), and Apis (the western honey bee and its close relatives).

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. 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.

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

Two species are recognized in the subgenus Megapis. They usually build 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.
    • 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.
  • Apis 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 and more recent research has confirmed this classification. 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

Apis nigrocincta is a cavity-nesting species. The species has rust-colored scapes, legs, and clypeuses, with reddish-tan hair color that covers most of the body.

Eastern honey bee

Apis cerana, the eastern honey bee proper, is the traditional honey bee of southern and eastern Asia. One of its subspecies, the Indian honey bee (A. c. indica), was domesticated 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 honey bee A. c. nuluensis and Apis nigrocincta from the Philippines to satisfaction; some researchers argue that these are indeed distinct species, but that A. cerana as defined is still paraphyletic, consisting of several separate species,though other researchers argue cerana is a single monophyletic species.

Western honey bee

The European honey bee may have originated from eastern Africa. This bee is pictured in Tanzania.

A. mellifera, the most common domesticated species, was the third insect whose genome was mapped. It seems to have originated in eastern tropical Africa and spread from there to Europe and eastwards into Asia to the Tian 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 radiation 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 German honey 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” honey bees, but actually feral) spread rapidly as far as the Great Plains, usually preceding the colonists.

Honey bees did not naturally cross the Rocky Mountains; they were transported by the Mormon pioneers to Utah in the late 1840s, and by ship to California in the early 1850s.

An Africanized honey bee (left) and a European honey bee on a honeycomb

Africanized honey bee

Africanized honey 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 honey 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 they are not often found in the colder, more northern parts of North America.

The original breeding experiment for which the East African lowland honey bees were brought to Brazil in the first place has continued (though not as originally intended).

Novel hybrid strains of domestic and re-domesticated Africanized honey bees combine high resilience to tropical conditions and good yields. They are popular among beekeepers in Brazil.

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