Honey bee Temporal range: Oligocene–Recent PreꞒ Ꞓ O S D C P T J K Pg N
Western honey bee on the bars of a horizontal top-bar hive
Scientific classification
Domain:	Eukaryota
Kingdom:	Animalia
Phylum:	Arthropoda
Class:	Insecta
Order:	Hymenoptera
Family:	Apidae
Clade:	Corbiculata
Tribe:	Apini Latreille, 1802
Genus:	Apis Linnaeus, 1758
Type species
Apis mellifera Linnaeus, 1758
Species
†Apis lithohermaea †Apis nearctica Subgenus Micrapis: Apis andreniformis Apis florea Subgenus Megapis: Apis dorsata Apis laboriosa Subgenus Apis: Apis cerana Apis koschevnikovi Apis mellifera Apis nigrocincta

A honey bee (also spelled honeybee) is a eusocial flying insect within the genus Apis of the bee clade, all native to mainland Afro-Eurasia.^[1][2] After bees spread naturally throughout Africa and Eurasia, humans became responsible for the current cosmopolitan distribution of honey bees, introducing multiple subspecies into South America (early 16th century), North America (early 17th century), and Australia (early 19th century).^[1]

Honey bees are known for their construction of perennial colonial nests from wax, the large size of their colonies, and 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 8 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 was domesticated for honey production and crop pollination. The only other domesticated bee is the eastern honey bee (Apis cerana), which occurs in South, Southeast, and East Asia. Only members of the genus Apis are true honey bees,^[3] but some other types of bees produce and store honey and have been kept by humans for that purpose, including the stingless bees belonging to the genus Melipona and the Indian stingless or dammar bee Tetragonula iridipennis. Modern humans also use beeswax in making candles, soap, lip balms and various cosmetics, as a lubricant and in mould-making using the lost wax process.

Etymology and name

The genus name Apis is Latin for "bee".^[4][5] 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, because it is a kind or type of bee. It is incorrect to run the two words together, as in dragonfly or butterfly, which are appropriate because dragonflies and butterflies are not flies^[6] 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.^[7][8][9]

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.^[2]

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.^[10]

The close relatives of modern honey bees – e.g., bumblebees and stingless bees – are also social to some degree, and social behavior is considered to be a 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,^[11] only eight species of honey bee are recognized, with a total of 43 subspecies, although historically seven to 11 species are recognized:^[12] 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).^[13]

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).^[12][14]

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.^[15]

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.^[14] Apis florea have smaller wing spans than its sister species.^[16] Apis florea are also completely yellow with the exception of the scutellum of workers, which is black.^[16]

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.^[17]
• Apis laboriosa, the Himalayan giant honey bee, was initially described as a distinct species. Later, it was included in A. dorsata as a subspecies^[12] based on the biological species concept, though authors applying a genetic species concept have suggested it should be considered a separate species^[14] and more recent research has confirmed this classification.^[18] 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, A. 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 colour in the Malay Peninsula and Sumatra, Indonesia.^[19] 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.^[19]

Philippine honey bee

Apis nigrocincta is a cavity-nesting species. The species has rust-coloured scapes, legs, and clypeuses, with reddish-tan hair colour that covers most of the body.^[20]

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,^[14] though other researchers argue cerana is a single monophyletic species.^[21]

Western honey bee

A. mellifera, the most common domesticated^[22] species, was first domesticated before 2600 BC^[23] and 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 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, colour, and anatomy can be quite different from one subspecies or even strain to another.^[24]

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.^[24]

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.^[14]

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.^[25] 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.^[26]

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.^[27] 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.^{[citation needed]}

Living and fossil honey bees (Apini: Apis)

Tribe Apini Latreille^[28]

Genus Apis Linnaeus (sensu lato)

• henshawi species group (†Priorapis Engel, †Synapis Cockerell)
  • • †A. vetusta Engel
    • †A. henshawi Cockerell
    • †A. petrefacta (Říha)
    • †A. miocenica Hong
    • †A. "longtibia" Zhang
    • †A. "Miocene 1"
• armbrusteri species group (†Cascapis Engel)
  • • †A. armbrusteri Zeuner
    • †A. nearctica, species novus
• florea species group (Micrapis Ashmead)
  • • A. florea Fabricius
    • A. andreniformis Smith
• dorsata species group (Megapis Ashmead)
  • • †A. lithohermaea Engel
    • A. dorsata Fabricius
    • A. laboriosa Smith
• mellifera species group (Apis Linnaeus sensu stricto)
  • mellifera subgroup
    • A. mellifera Linnaeus (Apis Linnaeus sensu strictissimo)
  • cerana subgroup (Sigmatapis Maa)
    • A. cerana Fabricius
    • A. nigrocincta Smith
    • A. koschevnikovi Enderlein

Life cycle

As in a few other types of eusocial bees, a colony generally contains one queen bee, a female; seasonally up to a few thousand drone bees, or males;^[29] and tens of thousands of female worker bees. 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, produced and shaped by the worker bees. Using her spermatheca, the queen can choose to fertilize the egg she is laying, usually depending on which cell she is laying it into. 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 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 moultings before spinning a cocoon within the cell, and pupating.
2. Young worker bees, sometimes called "nurse 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 dances, which recruit receiver bees to collect nectar from returning foragers.
5. Virgin queens go on mating flights away from their home colony to a drone congregation area and mate with multiple drones before returning. The drones die in the act of mating. Queen honey bees do not mate with drones from their home colony.
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 which was scouted by worker bees beforehand and whose location is communicated with a special type of dance. Once the swarm arrives, 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 several groups of vespid wasps also found new nests by 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".

Gallery

• 
  Honey bee eggs shown in opened wax cells
• 
  Eggs and larvae
• 
  Drone pupae
• 
  Emergence of a European dark honey bee (A. m. mellifera)

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 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 15 to 50 kilograms (33 to 110 lb).^[30] In addition, certain bees, including the western honey bee as well as Apis cerana, are known to engage in effective methods of nest thermoregulation during periods of varying temperature in both summer and winter. During the summer, however, this is achieved through fanning and water evaporation from water collected in various fields.^[31][32]

Pollination

Zdroj:https://en.wikipedia.org?pojem=Honeybee
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