Origin, Systematics and Distribution of honey bees

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

      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.^[14]

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.