Editing Honey bee (section)

==Origin, systematics, and distribution==
[[File:Apis distribution map.svg|right|thumb|Distribution of honey bees around the world]]
[[File:HoneyBeeAnatomy.svg|right|thumb|[[Morphology (biology)|Morphology]] of a sterile female worker honey bee]]

Honey bees appear to have their center of origin in [[South Asia|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.<ref name="Han-2012" />

The first ''Apis'' bees appear in the [[fossil record]] at the [[Eocene]]–[[Oligocene]] boundary (34&nbsp;[[myr|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.&nbsp;mellifera'' by Europeans. Only one fossil species is documented from the New World, ''[[Apis nearctica]]'', known from a single 14&nbsp;million-year-old specimen from Nevada.<ref>{{cite journal |author1=Michael S. Engel |author2=I. A. Hinojosa-Diaz |author3=A. P. Rasnitsyn |year=2009 |title=A honey bee from the Miocene of Nevada and the biogeography of ''Apis'' (Hymenoptera: Apidae: Apini) |journal=[[Proceedings of the California Academy of Sciences]] |volume=60 |issue=3 |pages=23–38}}</ref>

The close relatives of modern honey bees&nbsp;– e.g., [[bumblebee]]s and [[stingless bee]]s – 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 (biology)|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,<ref name="Nicholls-2015">{{cite web | last=Nicholls | first=Henry | title=The truth about bees | website=BBC | date=15 June 2015 | url=https://www.bbc.com/earth/story/20150615-the-truth-about-bees | access-date=9 July 2020}}</ref> only eight [[species]] of honey bee are recognized, with a total of 43 [[subspecies]], although historically seven to 11 species are recognized:<ref name="Engel-1999">{{cite journal |author=Michael S. Engel |year=1999 |title=The taxonomy of recent and fossil honey bees (Hymenoptera: Apidae: ''Apis'') |journal=[[Journal of Hymenoptera Research]] |volume=8 |pages=165–196 |author-link=Michael S. Engel}}</ref> ''[[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); ''[[Western honey bee|Apis mellifera]]'' (the western honey bee); and ''[[Apis nigrocincta]]'' (the Philippine honey bee).<ref name="Encyclopedia of Life">{{cite web | title=Honey Bees | website=Encyclopedia of Life | url=https://eol.org/pages/104135 | access-date=9 July 2020}}</ref>

Honey bees are the only extant members of the [[Tribe (biology)|tribe]] Apini. Today's honey bees constitute three [[clade]]s: ''Micrapis'' (the dwarf honey bees), ''Megapis'' (the giant honey bee), and ''Apis'' (the western honey bee and its close relatives).<ref name="Engel-1999"/><ref name="Arias-2005">{{cite journal |author1=Maria C. Arias |author2=Walter S. Sheppard |year=2005 |title=Phylogenetic relationships of honey bees (Hymenoptera:Apinae:Apini) inferred from nuclear and mitochondrial DNA sequence data |journal=[[Molecular Phylogenetics and Evolution]] |volume=37 |issue=1 |pages=25–35 |doi=10.1016/j.ympev.2005.02.017 |pmid=16182149|bibcode=2005MolPE..37...25A }}<br />{{cite journal |author1=Maria C. Arias |author2=Walter S. Sheppard | year=2005 |title=Corrigendum to "Phylogenetic relationships of honey bees (Hymenoptera:Apinae:Apini) inferred from nuclear and mitochondrial DNA sequence data" |journal=[[Molecular Phylogenetics and Evolution]] |volume=40 |issue=1 |page=315 |doi=10.1016/j.ympev.2006.02.002|doi-access=free }}</ref>

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.&nbsp;mellifera'' has been cultivated at least since the time of the building of the [[Egyptian pyramid]]s, and only that species has been moved extensively beyond its native range.<ref>{{Cite book |last=Clark |first=Michael C. |url=https://books.google.com/books?id=ZPyFDwAAQBAJ&dq=A.+mellifera+has+been+cultivated+at+least+since+the+time+of+the+building+of+the+Egyptian+pyramids%2C+and+only+that+species+has+been+moved+extensively+beyond+its+native+range&pg=PT291 |title=Coexisting on Earth Homo sapiens Quagmire |date=2018-04-03 |publisher=Michael C. Clark |language=en}}</ref>

===''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 [[Beehive (beekeeping)|hive]] and [[swarm]]s can be handled with minimal protection. They occur largely [[sympatric]]ally, though they are very distinct [[evolution]]arily and are probably the result of [[allopatric speciation]], their distribution later converging.

Given that ''A.&nbsp;florea'' is more widely distributed and ''A.&nbsp;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&nbsp;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]].<ref name="Arias-2005"/> ''[[Apis florea]]'' have smaller wing spans than its sister species.<ref name="Wongsiri-1997">{{cite journal | last1 = Wongsiri | first1 = S. | display-authors = et al | year = 1997 | title = Comparative biology of Apis andreniformis and Apis florea in Thailand | journal = Bee World | volume = 78 | issue = 1| pages = 23–35 | doi = 10.1080/0005772X.1997.11099328 }}</ref> ''Apis florea'' are also completely yellow except the scutellum of workers, which is black.<ref name="Wongsiri-1997" />

===''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 [[bee sting|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 [[Indonesia]]n subspecies of the giant honey bee or a distinct species; in the latter case, ''A.&nbsp;d. breviligula'' and/or other lineages would probably also have to be considered species.<ref>{{cite journal |author1=Nathan Lo |author2=Rosalyn S. Gloag |author3=Denis L. Anderson |author4=Benjamin P. Oldroyd |year=2009 |title=A molecular phylogeny of the genus ''Apis'' suggests that the Giant Honey Bee of the Philippines, ''A. breviligula'' Maa, and the Plains Honey Bee of southern India, ''A. indica'' Fabricius, are valid species |journal=[[Systematic Entomology]] |volume=35 |issue=2 |pages=226–233 |doi=10.1111/j.1365-3113.2009.00504.x|s2cid=84531938 }}</ref>
*''[[Apis laboriosa]]'', the Himalayan giant honey bee, was initially described as a distinct species. Later, it was included in ''A.&nbsp;dorsata'' as a subspecies<ref name="Engel-1999"/> based on the [[biological species concept]], though authors applying a genetic species concept have suggested it should be considered a separate species<ref name="Arias-2005"/> and more recent research has confirmed this classification.<ref name="Kitnya-2020">Kitnya N, Prabhudev MV, Bhatta CP, Pham TH, Nidup T, Megu K, Chakravorty J, Brockmann A, Otis GW (2020) Geographical distribution of the giant honey bee ''Apis laboriosa'' Smith, 1871 (Hymenoptera, Apidae). ZooKeys 951: 67–81. https://doi.org/10.3897/zookeys.951.49855</ref> Essentially restricted to the [[Himalayas]], it differs little from the giant honey bee in appearance but has extensive behavioral [[adaptation (biology)|adaptations]] that enable it to nest in the open at high altitudes despite low ambient temperatures. It is the largest living honey bee.

===''Apis''===
[[File:Western honey bee on a honeycomb.jpg|thumb|Western honey bee on a honeycomb]]
Eastern ''Apis'' species include three or four&nbsp;species, including ''[[Apis koschevnikovi|A.&nbsp;koschevnikovi]]'', ''[[Apis nigrocincta|A.&nbsp;nigrocincta]]'', and ''[[Apis cerana|A.&nbsp;cerana]]''. The genetics of the western honey bee (''[[Apis mellifera|A.&nbsp;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.&nbsp;koschevnikovi'' is pale reddish in [[Sabah|Sabah State]], [[Borneo]], [[Malaysia]], but a dark, coppery colour in the [[Malay Peninsula]] and [[Sumatra]], [[Indonesia]].<ref name="Hadisoesilo-2008">{{Cite journal|title = Morphometric analysis and biogeography of Apis koschevnikovi Enderlein (1906)|journal = Apidologie|date = 1 September 2008|issn = 0044-8435|pages = 495–503|volume = 39|issue = 5|doi = 10.1051/apido:2008029|first1 = S.|last1 = Hadisoesilo|first2 = Rika|last2 = Raffiudin|first3 = Wirian|last3 = Susanti|first4 = Tri|last4 = Atmowidi|first5 = Colleen|last5 = Hepburn|first6 = Sarah E.|last6 = Radloff|first7 = Stefan|last7 = Fuchs|first8 = H. Randall|last8 = Hepburn|s2cid = 6605920}}</ref> 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]].<ref name="Hadisoesilo-2008"/>

====Philippine honey bee====
''[[Apis nigrocincta]]'' is a cavity-nesting species. The species has rust-coloured [[Antenna (biology)#Insects|scapes]], legs, and [[Clypeus (arthropod anatomy)|clypeuses]], with reddish-tan hair colour that covers most of the body.<ref name="Hadisoesilo-1995">{{cite journal |last1=Hadisoesilo |first1=S. |last2=Otis |first2=G. W. |last3=Meixner |first3=M. |title=Two distinct populations of cavity-nesting honey bees (Hymenoptera: Apidae) in South Sulawesi, Indonesia |journal=Journal of the Kansas Entomological Society |volume=68 |issue=4 |year=1995 |pages=399–407 |jstor=25085613 }}</ref>

====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 (''[[Apis cerana indica|A.&nbsp;c.&nbsp;indica]]''), was [[Domestication|domesticated]] and kept in hives in a fashion similar to ''A.&nbsp;mellifera'', though on a more limited, regional scale.

It has not been possible yet to resolve its relationship to the Bornean honey bee ''[[Apis cerana nuluensis|A.&nbsp;c. nuluensis]]'' and ''[[Apis nigrocincta]]'' from the Philippines to satisfaction; some researchers argue that these are indeed distinct species, but that ''[[Apis cerana|A.&nbsp;cerana]]'' as defined is still [[paraphyletic]], consisting of several separate species,<ref name="Arias-2005" /> though other researchers argue ''cerana'' is a single monophyletic species.<ref name="Radloff-2010">{{cite journal |last1=Radloff |first1=Sarah E. |last2=Hepburn |first2=Colleen |last3=Randall Hepburn |first3=H. |last4=Fuchs |first4=Stefan |last5=Hadisoesilo |first5=Soesilawati |last6=Tan |first6=Ken |last7=Engel |first7=Michael S. |last8=Kuznetsov |first8=Viktor |title=Population structure and classification of ''Apis cerana'' |journal=Apidologie |date=15 March 2010 |volume=41 |issue=6 |pages=589–601 |doi=10.1051/apido/2010008|s2cid=32751472 |url=https://hal.archives-ouvertes.fr/hal-00892035/file/hal-00892035.pdf }}</ref>

====Western honey bee====
{{Main|Apis mellifera}}
[[File:Apis mellifera Tanzania.jpg|thumb|The European honey bee may have originated from eastern Africa. This bee is pictured in [[Tanzania]].]]

''A. mellifera'', the most common domesticated<ref>{{Cite web|url=http://scientificbeekeeping.com/whats-happening-to-the-bees-part-5-is-there-a-difference-between-domesticated-and-feral-bees/|title=What's Happening To The Bees? – Part 5: Is There A Difference Between Domesticated And Feral Bees?|date=26 June 2014}}</ref> species, was first domesticated before 2600 BC<ref>{{Cite web|url=https://www.fondazioneslowfood.com/en/ark-of-taste-slow-food/egyptian-honeybee/|title=Egyptian honeybee - Arca del Gusto|website=Slow Food Foundation}}</ref> 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 [[List of Apis mellifera subspecies|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.<ref name="Reuber-2015"/>

''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.<ref name="Reuber-2015">{{cite book |last1=Reuber |first1=Brant |title=21st Century Homestead: Beekeeping |date=21 February 2015 |publisher=lulu.com |isbn=978-1-312-93733-8 |page=116 |edition=First}}</ref>

The diversity of ''A. mellifera'' subspecies is probably the product of a largely [[Early Pleistocene]] [[Radiation (biology)|radiation]] aided by climate and habitat changes during the [[Quaternary glaciation|last ice age]]. That the western honey bee has been intensively managed by humans for many millennia&nbsp;– including hybridization and introductions&nbsp;– 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.&nbsp;mellifera'' subspecies.<ref name="Arias-2005" />

''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.<ref>{{cite journal | author=Villanueva, Rogel |display-authors=etal | title=Extinction of ''Melipona beecheii'' and traditional beekeeping in the Yucatán peninsula| journal=Bee World| year=2005 | volume=86 | issue=2 | pages=35–41 | doi=10.1080/0005772X.2005.11099651 |s2cid=31943555 }}</ref> In 1622, European colonists brought the [[European dark bee|German honey bee]] (''A.&nbsp;m. mellifera'') to the Americas first, followed later by the [[Italian bee|Italian honey bee]] (''A.&nbsp;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.<ref>{{cite web|author=Head RJ|title=A Brief Survey of Ancient Near Eastern Beekeeping; A Final Note|url=http://maxwellinstitute.byu.edu/publications/review/?vol=20&num=1&id=694#_ednref30|date=2008|publisher=The FARMS Review|access-date=16 January 2012|archive-url=https://web.archive.org/web/20130730162629/http://maxwellinstitute.byu.edu/publications/review/?vol=20&num=1&id=694#_ednref30|archive-date=30 July 2013|url-status=dead}}</ref>

[[File:Apis mellifera scutellata 1355020.jpg|thumb|right|An Africanized honey bee (left) and a European honey bee on a honeycomb]]

====Africanized honey bee====
{{Main|Africanized bee}}

Africanized honey bees (known colloquially as "killer bees") are [[Hybrid (biology)|hybrid]]s between European stock and the East African lowland subspecies ''[[African honey bee|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.<ref>{{Cite web|url=http://entnemdept.ufl.edu/creatures/misc/bees/ahb.htm|title=Africanized honey bee – Apis mellifera scutellata Lepeletier|website=entnemdept.ufl.edu|access-date=1 May 2019}}</ref> Accidentally released from quarantine in [[Brazil]], they have spread to North America and constitute a [[Pest (animal)|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.<ref>{{Cite web |others=Department of Systematic Biology |title=Africanized Bees |url=https://www.si.edu/spotlight/buginfo/killbee |access-date=2024-10-28 |website=Smithsonian Institution}}</ref>