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{{Short description|Colonial flying insect of genus Apis}}
{{Other uses}}
{{pp-pc}}
{{Use British English|date=June 2015}}
{{Automatic taxobox
| name = Honey bee
| fossil_range = {{fossil range|Oligocene|Recent}}
| image = The Lone Pollinator.jpg
| image_caption = [[Western honey bee]] on the bars of a [[horizontal top-bar hive]]
| parent_authority = [[Latreille]], 1802
| taxon = Apis
| authority = [[Linnaeus]], [[Systema Naturae|1758]]
| display_parents = 2
| type_species = ''[[Apis mellifera]]''
| type_species_authority = [[Linnaeus]], [[10th edition of Systema Naturae|1758]]
| subdivision_ranks = Species
| subdivision = *†''[[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]]''
}}
[[File:Honeybees.webm|thumb|thumbtime=94|Honeybees on [[Verbesina alternifolia|yellow ironweed]]. Followed by segment at one-tenth speed.]]
A '''honey bee''' (also spelled '''honeybee''') is a [[eusocial]] [[flying insect]] from the [[genus]] '''''Apis''''' of the largest [[bee]] [[family (biology)|family]], [[Apidae]].<ref>{{cite encyclopedia |title=Chapter 9 – Apis Species: (Honey Bees) |encyclopedia=Encyclopedia of Insects (Second Edition) |publisher=Academic Press |last=Crane |first=Eva |date=2009 |author-link=Eva Crane |editor1-last=Resh |editor1-first=Vincent H. |pages=31–32 |doi=10.1016/B978-0-12-374144-8.00009-6 |isbn=978-0-12-374144-8 |editor2-last=Cardé |editor2-first=Ring T.}}</ref> All honey bees are [[nectarivorous]] [[pollinator]]s native to mainland [[Afro-Eurasia]],<ref name="Whitfield-2006">{{cite journal|display-authors=3 |last1=Whitfield |first1=Charles W. |last2=Behura |first2=Susanta K. |last3=Berlocher |first3=Stewart H. |last4=Clark |first4=Andrew G. |last5=Johnston |first5=J. Spencer |last6=Sheppard |first6=Walter S. |last7=Smith |first7=Deborah R. |last8=Suarez |first8=Andrew V. |last9=Weaver |first9=Daniel |last10=Tsutsui |first10=Neil D. |title=Thrice Out of Africa: Ancient and Recent Expansions of the Honey Bee, Apis mellifera |journal=Science|pmid= 17068261|date=27 October 2006 |volume=314 |issue=5799 |pages=642–645 |doi=10.1126/science.1132772|bibcode=2006Sci...314..642W |s2cid=15967796 }}</ref><ref name="Han-2012">{{cite journal |last1=Han |first1=Fan |last2=Wallberg |first2=Andreas |last3=Webster |first3=Matthew T. |title=From where did the Western honeybee (''Apis mellifera'') originate? |journal=Ecology and Evolution |date=August 2012 |volume=2 |issue=8 |pages=1949–1957 |doi=10.1002/ece3.312|pmid=22957195 |pmc=3433997 |bibcode=2012EcoEv...2.1949H }}</ref> but [[human migration]]s and [[colonization]]s to the [[New World]] since the [[Age of Discovery]] have been responsible for the [[species introduction|introduction]] of multiple [[subspecies]] into South America (early 16th century), North America (early 17th century) and Australia (early 19th century), resulting in the current [[cosmopolitan distribution]] of honey bees in all continents except [[Antarctica]].<ref name="Whitfield-2006"/>
Honey bees are known for their construction of [[wiktionary:perennial|perennial]] [[honeycomb|hexagonally celled]] [[nest]]s made of [[beeswax|secreted wax]] (i.e. [[beehive]]s),<ref>{{cite encyclopedia |last=Smith |first=Michael L. |editor-last=Starr |editor-first=Christopher K. |encyclopedia=Encyclopedia of Social Insects |date=2021 |publisher=Springer, Cham |isbn=978-3-030-28102-1 |pages=626–632 |language=en |title=Nest Structure: Honey Bees |doi=10.1007/978-3-030-28102-1_85}}</ref> their large [[colony (biology)|colony]] sizes, and their routine [[regurgitation (digestion)|regurgitation]] of digested [[carbonhydrate|carbohydrate]]s as surplus [[food storage]] in the form of [[honey]], the lattermost of which distinguishes their hives as a prized [[foraging]] target of many [[mellivorous]] animals including [[honey badger]]s, bears and human [[hunter-gatherer]]s. Only 8 [[extant taxon|extant]] [[species]] of honey bees are recognized, with a total of 43 [[subspecies]], though historically 7 to 11 species are recognized. Although honey bees represent only a small fraction of the roughly 20,000 known species of bees, they are the bee [[clade]] most familiar to humans and are also the most valuable [[beneficial insect]]s to agriculture and horticulture.<ref>{{cite journal |last1=Papa |first1=Giulia |last2=Maier |first2=Roberto |last3=Durazzo |first3=Alessandra |last4=Lucarini |first4=Massimo |last5=Karabagias |first5=Ioannis K. |last6=Plutino |first6=Manuela |last7=Bianchetto |first7=Elisa |last8=Aromolo |first8=Rita |last9=Pignatti |first9=Giuseppe |last10=Ambrogio |first10=Andrea |last11=Pellecchia |first11=Marco |last12=Negri |first12=Ilaria |title=The Honey Bee Apis mellifera: An Insect at the Interface between Human and Ecosystem Health |journal=Biology |date=2022 |volume=11 |issue=2 |pages=233 |doi=10.3390/biology11020233 |doi-access=free}}</ref>
The best-known honey bee species is the [[western honey bee]] (''Apis mellifera''), which was [[domesticate]]d and [[insect farming|farmed]] (i.e. [[beekeeping]]) for honey production and crop [[pollination]]. The only other domesticated species is the [[eastern honey bee]] (''Apis cerana''), which are raised in [[South Asia|South]], [[Southeast Asia|Southeast]] and East Asia. Only members of the genus ''Apis'' are true honey bees,<ref>{{cite book |last=Buchmann |first=Stephen L. |author-link=Stephen L. Buchmann |title=Honey Bees: Letters from the Hive |date=8 June 2010 |publisher=Random House Children's Books |isbn=9780375895579 |edition=1st |___location=New York |page=157}}</ref> but some other bee species also produce and store honey and have been kept by humans for that purpose, including the [[stingless bee]]s belonging to the genus ''[[Melipona]]'' and the Indian stingless or dammar bee ''[[Tetragonula iridipennis]]''. In addition to harvesting honey, modern humans also use [[beeswax]] in making [[candle]]s, soap, [[lip balm]]s and various cosmetics, as a lubricant and in mould-making using the [[Lost-wax casting|lost wax process]]. Other honey bee [[secretion]]s such as [[royal jelly]] and [[bee venom]] are used [[pharmaceutic]]ally, especially in [[alternative medicine]].
==Etymology and name==
The [[genus]] name ''Apis'' is [[Latin]] for "bee".<ref>{{cite web |title=Apis |url=http://latindictionary.wikidot.com/noun:apis |website=The Latin Dictionary |access-date=23 November 2021}}</ref><ref>{{cite web |title=Honeybee |publisher=Online Etymology Dictionary, Douglas Harper |date=2019 |url=http://www.etymonline.com/index.php?search=Honeybee |access-date=27 February 2016 }}</ref> Although modern dictionaries may refer to ''Apis'' as either honey bee or honeybee, [[entomologist]] [[Robert Evans Snodgrass|Robert Snodgrass]] asserts that [[linguistic prescription|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 [[Dragonfly|dragonflies]] and [[Butterfly|butterflies]] are not flies.<ref name="Snodgrass-1984">{{cite book |last=Snodgrass |first=Robert E. |author-link=Robert Evans Snodgrass |url=https://books.google.com/books?id=IHGmkX1zDS8C |title=Anatomy of the Honey Bee |publisher=Cornell University Press |year=1984 |isbn=978-0-8014-9302-7 |page=vii}}</ref> 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]].<ref>{{cite web |title=Integrated Taxonomic Information System – Search, ''Apinae''|url=https://www.itis.gov/servlet/SingleRpt/SingleRpt?search_topic=TSN&search_value=633913#null |access-date=26 February 2016|date=2008}}</ref><ref name="Entomological Society of America">{{cite web |title=Common Names of Insects Database|publisher=Entomological Society of America |url=http://entsoc.org/common-names |access-date=21 February 2016}}</ref><ref>{{cite web |publisher=Tree of Life Web Project |title=Apinae|url=http://tolweb.org/Apinae|date=2004 |access-date=25 February 2016 }}</ref>
==Origin, systematics, and distribution==
[[File:Apis distribution map.svg|right|thumb|Distribution of honey bees around the world<ref>{{cite encyclopedia |last=Smith |first=Deborah R. |editor-last=Starr |editor-first=Christopher K. |encyclopedia=Encyclopedia of Social Insects |date=2021 |publisher=Springer, Cham |isbn=978-3-030-28102-1 |pages=117–131 |language=en |title=Biogeography of Honey Bees |doi=10.1007/978-3-030-28102-1_60}}</ref>]]
[[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 [[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. 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.<ref>{{cite journal |author1=Engel |first=Michael S. |author-link=Michael S. Engel |last2=Hinojosa-Díaz |first2=I. A. |last3=Rasnitsyn |first3=A. P. |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—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 |last=Engel |first=Michael S. |author-link=Michael S. Engel |year=1999 |title=The taxonomy of recent and fossil honey bees (Hymenoptera: Apidae: ''Apis'') |url=https://www.biodiversitylibrary.org/part/29302#/summary |journal=[[Journal of Hymenoptera Research]] |volume=8 |pages=165–196}}</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 bees), and ''Apis'' (the western honey bee and its close relatives).<ref name="Engel-1999"/><ref name="Arias-2005">{{cite journal |author1=Arias |first=Maria C. |last2=Sheppard |first2=Walter S. |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 |bibcode=2005MolPE..37...25A |doi=10.1016/j.ympev.2005.02.017 |pmid=16182149}}<br />{{cite journal |author1=Arias |first=Maria C. |last2=Sheppard |first2=Walter S. |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. 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. 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.<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 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.<ref>{{cite journal |author1=Lo |first=Nathan |last2=Gloag |first2=Rosalyn S. |last3=Anderson |first3=Denis L. |last4=Oldroyd |first4=Benjamin P. |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. 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 species, including ''[[Apis koschevnikovi|A. koschevnikovi]]'', ''[[Apis nigrocincta|A. nigrocincta]]'', and ''[[Apis cerana|A. cerana]]''. The genetics of the western honey bee (''[[Apis mellifera|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|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. c. indica]]''), was [[Domestication|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 ''[[Apis cerana nuluensis|A. c. nuluensis]]'' and ''[[Apis nigrocincta]]'' from the Philippines to satisfaction; some researchers argue that these are indeed distinct species, but that ''[[Apis cerana|A. 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=Hepburn |first3=H. Randall |last4=Fuchs |first4=Stefan |last5=Hadisoesilo |first5=Soesilawati |last6=Tan |first6=Ken |last7=Engel |first7=Michael S. |last8=Kuznetsov |first8=Viktor |date=15 March 2010 |title=Population structure and classification of ''Apis cerana'' |url=https://hal.archives-ouvertes.fr/hal-00892035/file/hal-00892035.pdf |journal=Apidologie |volume=41 |issue=6 |pages=589–601 |doi=10.1051/apido/2010008 |s2cid=32751472}}</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 |date=26 June 2014 |title=What's Happening To The Bees? – Part 5: Is There A Difference Between Domesticated And Feral Bees? |url=http://scientificbeekeeping.com/whats-happening-to-the-bees-part-5-is-there-a-difference-between-domesticated-and-feral-bees/ |website=scientificbeekeeping.com}}</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 |last=Reuber |first=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—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.<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 | last=Villanueva |first=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. m. mellifera'') to the Americas first, followed later by the [[Italian bee|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.<ref>{{cite web |last=Head |first=RJ |date=2008 |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 |url-status=dead |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 |access-date=16 January 2012 |publisher=The FARMS Review}}</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 African subspecies ''[[Apis mellifera scutellata|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 African 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>
== Living and fossil honey bees (Apini: ''Apis'') ==
'''Tribe Apini Latreille'''<ref>{{cite journal |last=Engel |first=Michael S. |author-link=Michael S. Engel |last2=Hinojosa-Díaz |first2=Ismael A. |last3=Rasnitsyn |first3=Alexandr P. |name-list-style=amp |year=2009 |title=A honey bee from the Miocene of Nevada and the biogeography of ''Apis'' (Hymenoptera: Apidae: Apini) |url=http://susquehannabeekeepers.com/pdfs/A_honey_bee_from_the_Miocene_of_Nevada_and_the_bio.pdf |url-status=dead |journal=[[Proceedings of the California Academy of Sciences]] |series=4 |volume=60 |issue=3 |pages=23–38 |archive-url=https://web.archive.org/web/20210227030258/http://susquehannabeekeepers.com/pdfs/A_honey_bee_from_the_Miocene_of_Nevada_and_the_bio.pdf |archive-date=2021-02-27 |access-date=2017-10-28}}</ref>
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 bee]]s, or males;<ref name="Gould-1995">{{cite book |author1=Gould |first=James L. |title=The Honey Bee |last2=Gould |first2=Carol Grant |publisher=Scientific American Library |year=1995 |isbn=978-0-7167-6010-8 |page=19}}</ref> and tens of thousands of female [[worker bee]]s. Details vary among the different species of honey bees, but common features include:
#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 [[Ploidy#Haploid and monoploid|haploid]], while females (queens and worker bees) develop from fertilised eggs and are [[Ploidy#Diploid|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 [[pupate|pupating]].
#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 flight and finally leaves the hive and typically spends the remainder of her life as a forager.
#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".
#Honey bees also perform [[tremble dance]]s, which recruit receiver bees to collect nectar from returning foragers.
#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.
#Colonies are established not by solitary queens, as in most bees, but by groups known as "[[Swarming (honey bee)|swarm]]s", which consist of a mated queen and a large contingent of worker bees. This group moves ''en masse'' to a nest site that 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 a 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 bee]]s 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===
<gallery mode="packed">
File:Bienenwabe mit Eiern 39.jpg|Honey bee eggs shown in opened wax cells
File:Bienenwabe mit Eiern und Brut 5.jpg|Eggs and larvae
File:Drohnenpuppen 79b.jpg|Drone pupae
File:Birth of black bee (Apis mellifera mellifera).jpg|Emergence of a European dark honey bee (''A. m. mellifera'')
File:HoneyBeeFlying.webm|thumbtime=115|[[Honey bee]] on [[Geranium]] (cultivar Rozanne) and [[insect flight|flying]]. Second portion shown at 12,000 frames per second with 60 frames shown per second
</gallery>
===Winter survival===
In cold climates, honey bees stop flying when the temperature drops below about {{cvt|10|C}} 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 {{cvt|27|C}} at the start of winter (during the broodless period) and {{cvt|34|C}} 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 {{cvt|8-9|C}}. 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 {{convert|15 to 50|kg}}.<ref>{{cite web|url=http://www.bees-online.com/Winter.htm|title=What do bees do in the winter?|access-date=12 March 2016|url-status=dead|archive-url=https://web.archive.org/web/20160304175741/http://www.bees-online.com/Winter.htm|archive-date=4 March 2016}}</ref> 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.<ref>Oldroyd, Benjamin P.; Wongsiri, Siriwat (2006). ''Asian Honey Bees (Biology, Conservation, and Human Interactions)''. Cambridge, Massachusetts and London, England: Harvard University Press.{{ISBN|0674021940}}.</ref><ref>{{cite web |title=Conservation Work for Honey Bees |url=https://www.nrcs.usda.gov/conservation-basics/natural-resource-concerns/plants |website=USDA |access-date=24 November 2021}}</ref>
<!-- Question: This paragraph is true for European honey bees. Is it applicable on this page?-->
==Pollination==
{{Main|Pollination management|List of crop plants pollinated by bees}}
[[File:Bee Leg with Pollen Bulb.jpg|thumb|Hind leg of a honey bee with pollen pellet stuck on the pollen basket or corbicula. When the worker bee is collecting pollen, their legs make the transfer of pollen from the inner basitarsal combs to the outer pollen basket (shown in the figure).]]
[[File:Buzzing bees.ogg|thumb|Buzzing bees on the flowering plum]]
Of all the honey bee species, only ''A. mellifera'' has been used extensively for commercial pollination of fruit and vegetable crops. The scale of these pollination services is commonly measured in the billions of dollars, credited with adding about 9% to the value of crops across the world. However, despite contributing substantially to crop pollination, there is debate about the potential spillover to natural landscapes and competition between managed honey bees and many of the ~20,000 species of wild pollinators.<ref name="Geldmann-2018">{{cite journal |last1=Geldmann |first1=Jonas |last2=González-Varo|first2=Juan P. |title=Conserving honey bees does not help wildlife |journal=Science |volume=359 |issue=6374 |year=2018 |pages=392–393 |doi=10.1126/science.aar2269|pmid=29371456 |bibcode=2018Sci...359..392G |s2cid=206665383 }}</ref>
Species of ''Apis'' are generalist floral visitors, and pollinate many species of flowering plants, but because of their "generalized" nature, they often do so inefficiently. Without specialized adaptations for specific flowers, their ability to reach pollen and nectar is often limited. This combined with their behavioural flexibility may be why they are the most commonly documented [[Pollen theft|pollen thieves]].<ref>{{Cite journal |last1=Hargreaves |first1=Anna L. |last2=Harder |first2=Lawrence D. |last3=Johnson |first3=Steven D. |date=2009 |title=Consumptive emasculation: the ecological and evolutionary consequences of pollen theft |url=https://onlinelibrary.wiley.com/doi/10.1111/j.1469-185X.2008.00074.x |journal=Biological Reviews |language=en |volume=84 |issue=2 |pages=259–276 |doi=10.1111/j.1469-185X.2008.00074.x|pmid=19382932 |s2cid=205599079 |url-access=subscription }}</ref> Indeed, for plant species with more specialized pollinators, experiments show that increased honeybee visitation can reduce pollination, both where honey bees are non-native<ref>{{Cite journal |last1=do Carmo |first1=Roselaini Mendes |last2=Franceschinelli |first2=Edivani Villaron |last3=da Silveira |first3=Fernando Amaral |date=2004 |title=Introduced Honeybees (Apis mellifera) Reduce Pollination Success without Affecting the Floral Resource Taken by Native Pollinators |url=https://www.jstor.org/stable/30043128 |journal=Biotropica |volume=36 |issue=3 |pages=371–376 |doi=10.1111/j.1744-7429.2004.tb00329.x |jstor=30043128 |bibcode=2004Biotr..36..371D |s2cid=86316179 |issn=0006-3606|url-access=subscription }}</ref> and even where they are native.<ref>{{Cite journal |last1=Hargreaves |first1=Anna L. |last2=Harder |first2=Lawrence D. |last3=Johnson |first3=Steven D. |date=2010 |title=Native pollen thieves reduce the reproductive success of a hermaphroditic plant, Aloe maculata |url=http://doi.wiley.com/10.1890/09-0792.1 |journal=Ecology |language=en |volume=91 |issue=6 |pages=1693–1703 |doi=10.1890/09-0792.1 |pmid=20583711 |bibcode=2010Ecol...91.1693H |issn=0012-9658|url-access=subscription }}</ref> What is more, their tendency to visit all species in a given area means that the pollen they carry for any one species is often very diluted. As such, they can provide some pollination to many plants, but most plants have some native pollinator that is more effective at pollinating that species.<ref>[https://www.decodedscience.org/plants-pollinators-and-the-price-of-almonds/28797/2 Plants, Pollinators, and the Price of Almonds] {{Webarchive|url=https://web.archive.org/web/20180919094322/https://www.decodedscience.org/plants-pollinators-and-the-price-of-almonds/28797/2 |date=2018-09-19 }}<br/>"Flowers set more seeds when visited by wild insects, and the more plants that were visited by wild insects, the more likely they were to set fruit. In some places the researchers considered, wild insects were pollinating most of the plants despite rented honey bees being present."</ref> When honey bees are present as an invasive species in an area, they compete for flowers with native pollinators, which can actually push out the native species.<ref>[https://www.popsci.com/honeybees-hog-limelight Saving pollinators is about more than just honeybees]<br/>The problem is that there are only so many flowers and places to nest. And once the numbers of honeybees have been artificially inflated (commercial-scale beekeeping would not exist without humans) the increased competition for these resources can push native non-''Apis'' pollinators out of their natural habitats. Honeybees also spread exotic plants and transmit pathogens, both of which have been shown to harm other pollinators.</ref>
===Claims of human dependency===
[[Western honey bee]]s have been described as essential to human food production, leading to claims that without their pollination humanity would starve or die out.<ref>{{Cite web |last=Haltiwanger |first=John |date=15 September 2014 |title=If All The Bees In The World Die, Humans Will Not Survive |url=https://www.elitedaily.com/news/world/humans-need-bees-to-survive/755737 |access-date=11 August 2025 |website=Elite Daily}}</ref><ref>{{Cite web |last=Spector |first=Dina |date=2013-06-22 |title=What Our World Would Look Like Without Honeybees |url=https://www.businessinsider.com/the-world-without-honeybees-2013-6 |access-date=2025-08-11 |website=Business Insider |language=en-US}}</ref> Apples, blueberries, and cherries, for example, are 90 percent dependent on honeybee pollination.<ref name="AgBioResearch-2018">{{cite web | title=Supporting pollination in agriculture | website=AgBioResearch, [[Michigan State University College of Agriculture & Natural Resources]] | date=2018-10-01 | url=https://www.canr.msu.edu/news/supporting-pollination-in-agriculture | access-date=2022-07-09}}</ref> [[Albert Einstein]] is sometimes misquoted as saying "If bees disappeared off the face of the earth, man would only have four years left to live"''.''<ref>[https://www.vice.com/en/article/what-would-happen-if-all-the-bees-died-tomorrow/ What Would Happen if All the Bees Went Extinct?]<br/>"First, the easy part: "I've never seen anything definitively link the quote to Einstein," says Mark Dykes, the chief inspector for Texas Apiary Inspection Service. Quote checkers like this one, and this one agrees. But debunking its message? That's more complicated."</ref> Einstein did not say this and there is no science to support this prediction.<ref>[https://www.nrdc.org/onearth/would-world-without-bees-be-world-without-us Would a World Without Bees Be a World Without Us?]<br/>"Albert Einstein is sometimes quoted as saying, "If the bee disappears from the surface of the earth, man would have no more than four years to live." It's highly unlikely that Einstein said that. For one thing, there's no evidence of him saying it. For another, the statement is hyperbolic and wrong (and Einstein was rarely wrong)."</ref>
Many important crops need no insect pollination at all. The ten most important crops,<ref>{{Cite web|url=https://www.businessinsider.com/10-crops-that-feed-the-world-2011-9|title=The 10 Most Important Crops In The World|first=Eric|last=Goldschein|website=Business Insider}}</ref> comprising 60% of all human food energy,<ref>{{Cite web|url=https://www.worldatlas.com/articles/most-important-staple-foods-in-the-world.html|title=What Are the World's Most Important Staple Foods?|website=WorldAtlas|date=7 June 2019}}</ref> fall into this category: [[Plantain (cooking)|plantain]]s are sterile and propagated by cuttings, as are [[cassava]]; potatoes, [[Yam (vegetable)|yam]]s, and [[sweet potato]]es are root vegetables propagated by [[tuber]]s; [[soybean]]s are [[Self-pollination|self-pollinated]]; and rice, wheat, [[sorghum]], and maize, are [[Anemophily|wind-pollinated]], as are most other grasses.<ref>{{Cite book| last=Gibson | first=D.J. | title=Grasses and grassland ecology | year=2009 | ___location=Oxford | publisher=Oxford University Press | page=82 | isbn=9780198529187 }}</ref>
No crops originating in the [[New World]] depend on the western honey bee (''Apis mellifera'') at all, as the bee is an [[invasive species]] brought over with colonists in the last few centuries.<ref>{{Cite web|url=https://www.usgs.gov/faqs/are-honey-bees-native-north-america|title=Are honey bees native to North America?|website=USGS|date=7 May 2024}}</ref> [[Tomato]]es, [[Capsicum annuum|pepper]]s, [[Cucurbita|squash]], and all other New World crops evolved with native pollinators such as [[squash bee]]s, [[bumble bee]]s, and other native bees.{{Citation needed|date=August 2021}} The [[stingless bee]]s mentioned by Jefferson{{Clarify|date=August 2021}} are distant relatives of the honey bees, in the genus ''[[Melipona]]''.{{Citation needed|date=August 2021}}
Still, honey bees are considered "crucial to the food supply, pollinating more than 100 of the crops we eat, including nuts, vegetables, berries, citrus and melons."<ref name="PBS News-2023">{{Cite web|url=https://www.pbs.org/newshour/economy/struggling-beekeepers-stabilize-u-s-honeybee-population-after-nearly-half-of-colonies-died-last-year|title=Struggling beekeepers stabilize U.S. honeybee population after nearly half of colonies died last year|date=June 22, 2023|website=PBS News}}</ref> The [[USDA]] reports "Three-fourths of the world's flowering plants and about 35 percent of the world's food crops depend on animal pollinators to reproduce"<ref>{{Cite web|url=https://www.usda.gov/about-usda/general-information/initiatives-and-highlighted-programs/peoples-garden/importance-pollinators|title=The Importance of Pollinators | USDA|website=usda.gov}}</ref> and honey bees "pollinate 80 percent of all flowering plants, including more than 130 types of fruits and vegetables."<ref>{{Cite web|url=https://www.farmers.gov/blog/value-birds-and-bees|title=The Value of Birds and Bees|date=June 22, 2020|website=Farmers.gov}}</ref>
==Nutrition==
[[File:A bee on a quince flower.jpg|thumb|A honey bee forager on a [[quince]] flower|alt=The foraging behavior of a honeybee]]
Honey bees obtain all of their nutritional requirements from a diverse combination of pollen and nectar. Pollen is the only natural protein source for honey bees. Adult worker honey bees consume 3.4–4.3 mg of pollen per day to meet a dry matter requirement of 66–74% protein.<ref name="Brodschneider-2010">{{Cite journal |last1=Brodschneider |first1=Robert |last2=Crailsheim|first2=Karl|date=1 May 2010 |title=Nutrition and health in honey bees |journal=Apidologie |language=en |volume=41|issue=3|pages=278–294|doi=10.1051/apido/2010012|s2cid=40046635 |issn=0044-8435|url=https://hal.archives-ouvertes.fr/hal-00892087/file/hal-00892087.pdf }}</ref> The rearing of one larva requires 125-187.5 mg pollen or 25–37.5 mg protein for proper development.<ref name="Brodschneider-2010"/> Dietary proteins are broken down into amino acids, ten of which are considered essential to honey bees: methionine, tryptophan, arginine, lysine, histidine, phenylalanine, isoleucine, threonine, leucine, and valine. Of these amino acids, honey bees require the highest concentrations of leucine, isoleucine, and valine; however, elevated concentrations of arginine and lysine are required for brood rearing.<ref name="Anderson-1976">{{Cite journal|last1=Anderson|first1=Leroy M|last2=Dietz|first2=A.|date=1976|title=Pyridoxine Requirement of the Honey Bee (Apis mellifera) For Brood Rearing|url=https://hal.archives-ouvertes.fr/hal-00890394/document|journal=Apidologie|doi=10.1051/apido:19760105|volume=7|pages=67–84|doi-access=free}}</ref> In addition to these amino acids, some B vitamins including biotin, folic acid, nicotinamide, riboflavin, thiamine, pantothenate, and most importantly, pyridoxine are required to rear larvae. Pyridoxine is the most prevalent B vitamin found in royal jelly and concentrations vary throughout the foraging season with the lowest concentrations found in May and the highest concentrations found in July and August. Honey bees lacking dietary pyridoxine were unable to rear brood.<ref name="Anderson-1976"/>
[[File:Bee Collecting Pollen 2004-08-14.jpg|thumb|left|250px|A forager collecting pollen]]
Pollen is also a lipid source for honey bees ranging from 0.8% to 18.9%.<ref name="Brodschneider-2010"/> Lipids are metabolized during the brood stage for precursors required for future biosynthesis. Fat-soluble vitamins A, D, E, and K are not considered essential but have been shown to significantly improve the number of brood reared.<ref name="Brodschneider-2010"/> Honey bees ingest phytosterols from pollen to produce 24-methylene cholesterol and other sterols as they cannot directly synthesize cholesterol from phytosterols. Nurse bees can selectively transfer sterols to larvae through brood food.<ref name="Brodschneider-2010"/>
Nectar is collected by foraging worker bees as a source of water and carbohydrates in the form of sucrose. The dominant monosaccharides in honey bee diets are fructose and glucose but the most common circulating sugar in hemolymph is trehalose which is a disaccharide consisting of two glucose molecules.<ref>{{Cite book|title=Physiological Ecology: How Animals Process Energy, Nutrients, and Toxins|last1=Karasov|first1=William H.|last2=Martinez del Rio|first2=Carlos|publisher=Princeton|year=2008|pages=63–66}}</ref> Adult worker honey bees require 4 mg of utilizable sugars per day and larvae require about 59.4 mg of carbohydrates for proper development.<ref name="Brodschneider-2010"/>
Honey bees require water to maintain osmotic homeostasis, prepare liquid brood food, and cool the hive through evaporation. A colony's water needs can generally be met by nectar foraging as it has high water content. Occasionally on hot days or when nectar is limited, foragers will collect water from streams or ponds to meet the needs of the hive.<ref>{{Cite journal|last=Kuhnholz|first=Susanne|date=1997|title=The Control of Water Collection in Honey Bee Colonies|journal=Behavioral Ecology and Sociobiology|doi=10.1007/s002650050402|volume=41|issue=6|pages=407–422|bibcode=1997BEcoS..41..407K |s2cid=41311766}}</ref>
==Beekeeping==
{{Main|Beekeeping}}
[[File:Beekeeper with moveable comb hive.jpg|thumb|right|A beekeeper inspecting a [[hive frame]] from a [[Langstroth hive]]. The modular design allows for easier management and honey harvesting.]]
[[File:Beekeeper in Pendro.jpg|thumb|right|A professional beekeeper inspects hives for breeding and selection, [[Pendro]].]]
[[File:Gurian Beekeeper.jpg|thumb|right|A professional beekeeper from [[Guria]] inspects a well-developed jumbo frame hive.]]
[[File:Honeybee hive entrance.webm|thumb|right|thumbtime=89|Honey bee hive entrance with audio. The last part is at one-fourth speed.]]
The only domesticated species of honey bee are ''A. mellifera'' and ''[[Apis cerana|A. cerana]]'', and they are often maintained, fed, and transported by beekeepers. In Japan, where ''A. mellifera'' is vulnerable to local hornets and disease, the Japanese honey bee ''[[A. cerana japonica]]'' is used in its place. Modern hives also enable beekeepers to transport bees, moving from field to field as the crop needs pollinating and allowing the beekeeper to charge for the pollination services they provide, revising the historical role of the self-employed beekeeper, and favoring large-scale commercial operations. Bees of various types other than honey bees are also domesticated and used for pollination or other means around the world, including ''[[Tetragonula iridipennis]]'' in India, the [[blue orchard bee]] for tree nut and fruit pollination in the United States, and several species of ''[[Bombus]]'' (bumblebees) for pollination in various regions globally, such as tomatoes, which are not effectively pollinated by honey bees.<ref>{{Cite web|url=https://www.biobestgroup.com/en/biobest/pollination/things-to-know-about-bumblebees-7052/species-6674/|title=Bumblebee species | Biobest|website=biobestgroup.com}}</ref>
===Colony collapse disorder===
{{Main|Colony collapse disorder}}
Primarily in places where western honey bees were imported by humans, periodic collapses in western honey bee populations have occurred at least since the late 19th century.<ref>{{Cite news|url=https://www.megabeepestcontrol.com/wp-content/uploads/2024/03/Colony-Collapse-Disorder-The-Vanishing-Honeybee-Apis-Mellifera.pdf|title=Colony Collapse Disorder: The Vanishing Honeybee (Apis Mellifera)|date=12 March 2011|work=CU Scholar|access-date=26 March 2024|language=en-US}}</ref>
However, as humans continued to manipulate the western honey bee and deliberately transferred them on a global scale, diseases simultaneously spread and harmed managed colonies. Colony losses have occurred periodically throughout history. Fungus, mites, and starvation have all been thought to be the cause of the deaths. Limited occurrences resembling CCD were documented as early as 1869.<ref>{{cite news |author1=Underwood |first=Robyn M. |last2=van Engelsdorp |first2=Dennis |title=Colony Collapse Disorder: Have We Seen This Before? |url=http://ento.psu.edu/pollinators/publications/underwood |access-date=2010-05-02 |publisher=The Pennsylvania State University, Department of Entomology}}</ref><ref>{{cite news |last=Lester |first=Benjamin |date=7 March 2007 |title=Mystery of the dying bees |url=http://www.cosmosmagazine.com/node/1087?page=1 |url-status=dead |archive-url=https://web.archive.org/web/20080324131409/http://www.cosmosmagazine.com/node/1087?page=1 |archive-date=24 March 2008 |work=[[Cosmos (Australian magazine)|Cosmos]]}}</ref> Colony collapses were called "May Disease" in Colorado in 1891 and 1896.<ref>{{cite journal |last=vanEngelsdorp |first=Dennis |last2=Evans |first2=Jay D. |last3=Saegerman |first3=Claude |last4=Mullin |first4=Chris |last5=Haubruge |first5=Eric |last6=Nguyen |first6=Bach Kim |last7=Frazier |first7=Maryann |last8=Frazier |first8=Jim |last9=Cox-Foster |first9=Diana |last10=Chen |first10=Yanping |last11=Underwood |first11=Robyn |last12=Tarpy |first12=David R. |last13=Pettis |first13=Jeffery S. |date=August 2009 |title=Colony Collapse Disorder: A Descriptive Study |url=https://nature.berkeley.edu/garbelottoat/wp-content/uploads/vanengelsdorp2009.pdf |journal=PLOS ONE |volume=4 |issue=8 |pages=e6481 |bibcode=2009PLoSO...4.6481V |doi=10.1371/journal.pone.0006481 |pmc=2715894 |pmid=19649264 |doi-access=free}}</ref>
Starting in the first decade of the 21st century, abnormally high die-offs (30–70% of hives) of western honey bee colonies have occurred in North America. This has been dubbed "colony collapse disorder" (CCD) and was at first unexplained.<ref>{{cite journal|last=Bryony|first=Bonning|title=Honey Bee Disease Overview|journal=Journal of Invertebrate Pathology|date=11 November 2009|volume=103|pages=S2-4|doi=10.1016/j.jip.2009.07.015 |pmid=19909974}}</ref> It seems to be caused by a combination of factors rather than a single [[pathogen]] or [[poison]], possibly including [[neonicotinoid]] pesticides<ref>{{cite news|last=McDonald-Gibson|first=Charlotte|title='Victory for bees' as European Union bans neonicotinoid pesticides blamed for destroying bee population|url=https://www.independent.co.uk/environment/nature/victory-for-bees-as-european-union-bans-neonicotinoid-pesticides-blamed-for-destroying-bee-population-8595408.html|newspaper=The Independent|access-date=2 July 2014}}</ref> or [[Israeli acute paralysis virus]].<ref>{{cite web|title=Colony Collapse Disorder |url=http://www.beeologics.com/colony-health/colony-collapse-disorder/ |archive-url=https://web.archive.org/web/20130206024748/http://www.beeologics.com/colony-health/colony-collapse-disorder/ |url-status=dead |archive-date=6 February 2013 |website=Beeologics |access-date=23 October 2014 }}</ref>
A survey by the [[University of Maryland]] and [[Auburn University]] published in 2023 found the number of United States honeybee colonies "remained relatively stable" although 48% of colonies were lost in the year that ended April 1, 2023, with a 12-year average annual mortality rate of 39.6%. The previous year (2021–2022) the loss was 39% and the 2020–2021 loss was 50.8%. Beekeepers told the surveying scientists that a 21% loss over the winter is acceptable and more than three-fifths of beekeepers surveyed said their losses were higher than that in 2022–2023.<ref name="PBS News-2023"/>
=== Parasites ===
{{for| parasites affecting Apis cerana |Apis cerana#Pathogens and parasites affecting Apis cerana}}
====Acarapis woodi====
''[[Acarapis woodi]]'' (or "tracheal mites") are parasitic mites that live and reproduce in adult bees' tracheae, or respiratory tubes, piercing the tube walls with their mouthparts to feed on haemolymph. To infest new hosts, the mites must find newly emerged bees; after three days, the bristles (setae) guarding the [[Spiracle (arthropods)|spiracles]] are firm enough to prevent the mites' entry into the tracheae. Mite infestations are known as acarine and have been called "Isle of Wight disease".<ref>{{cite web|url=http://www.sel.barc.usdj.gov/acari/content/trachealmites.html |title="Tracheal mites" Tarsonemidae |publisher=[[Agricultural Research Service]], [[United States Department of Agriculture]] |date=February 18, 2005 |access-date=March 10, 2011 |url-status=dead |archive-url=https://web.archive.org/web/20110517030353/http://www.sel.barc.usda.gov/acari/content/trachealmites.html |archive-date=May 17, 2011 }}</ref>
====''Galleria mellonella''====
Larval stages of the moth ''[[Galleria mellonella]]'' parasitize both wild and cultivated honey bees, in particular ''Apis mellifera'' and ''Apis cerana''. Eggs are laid within the hive and the [[larva]]e that hatch tunnel through and destroy the honeycombs that contain bee larva and their honey stores. The tunnels they create are lined with silk, which entangles and starves emerging bees. Destruction of honeycombs also results in honey leaking and being wasted. Both ''G. mellonella'' adults and larvae are possible vectors for pathogens that can infect bees, including the [[Israeli acute paralysis virus]] and the [[black queen cell virus]].<ref name="Kwadha-2017">{{Cite journal|last1=Kwadha|first1=Charles A.|last2=Ong'amo|first2=George O.|last3=Ndegwa|first3=Paul N.|last4=Raina|first4=Suresh K.|last5=Fombong|first5=Ayuka T.|date=9 June 2017|title=The Biology and Control of the Greater Wax Moth, Galleria mellonella|journal=Insects|language=en|volume=8|issue=2|pages=61|doi=10.3390/insects8020061|pmid=28598383|pmc=5492075|doi-access=free}}</ref>
To manage the mites, temperature treatments are possible, but also distort the wax of the honeycombs. Chemical fumigants, particularly CO<sub>2</sub>, are also used.<ref name="Kwadha-2017"/>
====''Varroa'' mites====
[[Varroa destructor|''Varroa'' mites]] are arguably the biggest threat to honey bees in the United States.<ref name="PBS News-2023"/> These mites invade hives and reproduce by laying eggs on the pupa. The hatching mites eat away at the pupa, causing deformities as well as spreading disease. If not detected and treated early on, the mite population may increase to such an extent that the hive will succumb to the diseases and deformities caused by the mites. It was widely believed that the mites drank the blood of bees. However, a 2018 study Article in PNAS: "Linking pesticides and gut health in bees" showed that they feed on the fat body tissue of live bees, not the blood.
Mite treatment is accomplished by several methods, including treatment strips and acid vaporization.
==Bee products==
===Honey===
{{Main|Honey}}
Honey is a complex substance made when bees ingest nectar, process it, and store the substance in honeycombs.<ref name="Crane-1990">{{cite journal |last=Crane |first=E |author-link=Eva Crane |year=1990 |title=Honey from honeybees and other insects |journal=Ethology Ecology & Evolution |volume=3 |issue=sup1 |pages=100–105 |doi=10.1080/03949370.1991.10721919}}</ref> All living species of ''Apis'' have had their honey gathered by indigenous peoples for consumption. ''A. mellifera'' and ''A. cerana'' are the only species that have had their honey harvested for commercial purposes.
===Beeswax===
{{Main|Beeswax}}
Worker bees of a certain age secrete [[beeswax]] from a series of exocrine glands on their abdomens.<ref name="Sanford-1976">{{cite journal |last1=Sanford |first1=M.T. |last2=Dietz |first2=A. |year=1976 |title=The fine structure of the wax gland of the honey bee (Apis mellifera L.). |journal=Apidologie |volume=7 |issue=3 |pages=197–207 |doi=10.1051/apido:19760301|doi-access=free }}</ref> They use the wax to form the walls and caps of the comb. As with honey, beeswax is gathered by humans for various purposes such as candle making, waterproofing, soap and cosmetics manufacturing, pharmaceuticals, art, furniture polish, and more.<ref>{{Cite news|url=https://www.beeculture.com/wax-rendering/|title=Wax Rendering {{!}} Bee Culture|date=23 March 2016|work=Bee Culture|access-date=26 October 2018|language=en-US}}</ref>
===Bee bread===
{{Main|Bee pollen}}
Bees collect [[pollen]] in their [[pollen basket]]s called corbiculae and carry it back to the hive.<ref>{{cite book |last=Gillott |first=Cedric |title=Entomology |publisher=Springer |year=1995 |page=79}}</ref>
Worker bees combine pollen, honey, and glandular secretions and allow them to ferment in the comb to make bee bread. The fermentation process releases additional nutrients from the pollen and can produce antibiotics and fatty acids which inhibit spoilage.<ref>{{cite journal |last1=Anderson |first1=Kirk E. |last2=Carroll |first2=Mark J. |last3=Sheehan |first3=Tim |last4=Lanan |first4=Michele C. |last5=Mott |first5=Brendon M. |last6=Maes |first6=Patrick |last7=Corby-Harris |first7=Vanessa |date=5 November 2014 |title=Hive-stored pollen of honey bees: many lines of evidence are consistent with pollen preservation, not nutrient conversion |journal=Molecular Ecology |volume=23 |issue=23 |pages=5904–5917 |doi=10.1111/mec.12966 |pmc=4285803 |pmid=25319366|bibcode=2014MolEc..23.5904A }}</ref> Bee bread is eaten by [[Worker bee#Nurse bees (days 3–12)|nurse bees]] (younger workers) which produce the protein-rich royal jelly needed by the queen and developing larvae in their hypopharyngeal glands.
In the hive, pollen is used as a [[protein]] source necessary during brood-rearing. In certain environments, excess pollen can be collected from the hives of ''A. mellifera'' and ''A. cerana''. The product is used as a health supplement. It has been used with moderate success as a source of pollen for [[hand-pollination|hand pollination]].
===Bees as food===
{{Main|Bee brood#As food}}
Bee brood—the eggs, larvae or [[pupa]]e of honey bees—is nutritious and seen as a delicacy in countries such as Indonesia,<ref name="Haris-2013">{{cite web |last=Haris |first=Emmaria |title=Sensasi Rasa Unik Botok Lebah yang Menyengat (Unique taste sensation botok with stinging bees) |url=http://www.sayangi.com/gayahidup1/kuliner/read/12669/sensasi-rasa-unik-botok-lebah-yang-menyengat |publisher=Sayangi.com |access-date=14 February 2018 |language=id |date=6 December 2013 |archive-url=https://web.archive.org/web/20150622203413/http://www.sayangi.com/gayahidup1/kuliner/read/12669/sensasi-rasa-unik-botok-lebah-yang-menyengat |archive-date=22 June 2015 |url-status=dead }}</ref> Mexico, Thailand, and many African countries; it has been consumed since ancient times by the Chinese and Egyptians.{{efn|The Mayans kept and collected honey and brood, but from stingless social bees such as ''[[Melipona beecheii]]'', not from ''Apis'' honeybees.<ref>{{cite magazine|last1=Pearson|first1=Gwen|title=Women Work to Save Native Bees of Mexico|url=https://www.wired.com/2014/03/women-work-save-native-bees-mexico/|magazine=Wired|access-date=15 May 2018|date=3 May 2014}}</ref>}}<ref name="UN FAO-2016">{{cite web|url=http://teca.fao.org/read/8775|title=How to collect drone larvae from the beehive|publisher=Home technologies and practices for small agricultural producers, UN Food and Agriculture Organization|date=29 August 2016|access-date=13 February 2018|archive-url=https://web.archive.org/web/20180213135222/http://teca.fao.org/read/8775|archive-date=13 February 2018|url-status=dead}}</ref><ref name="Holland-2013">{{cite web |url=http://news.nationalgeographic.com/news/2013/13/130514-edible-insects-entomophagy-science-food-bugs-beetles/ |archive-url=https://web.archive.org/web/20130606223425/http://news.nationalgeographic.com/news/2013/13/130514-edible-insects-entomophagy-science-food-bugs-beetles |url-status=dead |archive-date=June 6, 2013 |title=U.N. Urges Eating Insects: 8 Popular Bugs to Try |last=Holland |first=Jennifer |date=14 May 2013 |work=National Geographic}}</ref>
Adult wild honeybees are also consumed as food in parts of China, including [[Yunnan]]. According to a worker at a Yunnan-based specialty restaurant, the bees are best served "deep-fried with salt and pepper", and they are "naturally sweet and tasty". Kellie Schmitt of [[CNN]] described the dish as one of "Shanghai's weirdest foods".<ref name="Schmitt-2011">{{cite news|last=Schmitt|first=Kellie|date=26 December 2011|title=Shanghai's weirdest foods|newspaper=CNN|url=http://travel.cnn.com/shanghai/eat/shanghais-weird-eats-779548|url-status=dead|archiveurl=https://web.archive.org/web/20130328112430/http://travel.cnn.com/shanghai/eat/shanghais-weird-eats-779548|archive-date=28 March 2013}}</ref>
===Propolis===
{{Main|Propolis}}
Propolis is a resinous mixture collected by honey bees from tree buds, sap flows, or other botanical sources, which is used as a sealant for unwanted open spaces in the hive.<ref>{{cite journal |last1=Simone-Finstrom |first1=Michael |last2=Spivak |first2=Marla |doi=10.1051/apido/2010016 |title=Propolis and bee health: The natural history and significance of resin use by honey bees |journal=Apidologie |volume=41 |issue=3 |pages=295–311 |date=May–June 2010|doi-access=free |hdl=11299/182451 |hdl-access=free }}</ref> Propolis may cause severe [[allergic reaction]]s and have [[Drug interaction|adverse interactions]] with [[prescription drug]]s in some individuals.<ref>{{cite web|url=https://www.medlineplus.gov/druginfo/natural/390.html|title=Propolis|publisher=U.S. National Library of Medicine |date= 28 July 2022|access-date=5 February 2023}}</ref> Propolis is also used in wood finishes on [[string instrument]]s.<ref>{{cite journal |last=Gambichler |first=T |last2=Boms |first2=S |last3=Freitag |first3=M |date=April 2004 |title=Contact dermatitis and other skin conditions in instrumental musicians |journal=BMC Dermatol. |volume=4 |pages=3 |doi=10.1186/1471-5945-4-3 |pmc=416484 |pmid=15090069 |doi-access=free}}</ref>
===Royal jelly===
{{Main|Royal jelly}}
Royal jelly is a honey bee secretion made in the hypopharyngeal and mandibular glands of nurse bees used to nourish larvae and queen bees. It is marketed for its alleged but unsupported claims of health benefits.<ref>{{cite journal | journal = EFSA Journal | date = 2011 | volume = 9 | issue = 4| pages = 2083 | doi = 10.2903/j.efsa.2011.2083 | title = Scientific Opinion on the substantiation of health claims related to: anthocyanidins and proanthocyanidins (ID 1787, 1788, 1789, 1790, 1791); sodium alginate and ulva (ID 1873); vitamins, minerals, trace elements and standardised ginseng G115 extract (ID 8, 1673, 1674); vitamins, minerals, lysine and/or arginine and/or taurine (ID 6, 1676, 1677); plant-based preparation for use in beverages (ID 4210, 4211); Carica papaya L. (ID 2007); "fish protein" (ID 651); acidic water-based, non-alcoholic flavoured beverages containing calcium in the range of 0.3 to 0.8 mol per mol of acid with a pH not lower than 3.7 (ID 1170); royal jelly (ID 1225, 1226, 1227, 1228, 1230, 1231, 1326, 1328, 1329, 1982, 4696, 4697); foods low in cholesterol (ID 624); and foods low in trans-fatty acids (ID 672, 4333) pursuant to Article 13(1) of Regulation (EC) No 1924/2006|author=EFSA Panel on Dietetic Products, Nutrition and Allergies, European Food Safety Authority| doi-access = free }}</ref><ref>{{cite web | url = https://www.fda.gov/NewsEvents/Newsroom/PressAnnouncements/ucm207416.htm | archive-url = https://web.archive.org/web/20100407160606/http://www.fda.gov/NewsEvents/Newsroom/PressAnnouncements/ucm207416.htm | url-status = dead | archive-date = April 7, 2010 | title = Federal Government Seizes Dozens of Misbranded Drug Products: FDA warned company about making medical claims for bee-derived products | date = 5 April 2010 | publisher = [[Food and Drug Administration]]}}</ref> On the other hand, it may cause severe [[allergic reaction]]s in some individuals.<ref>{{cite journal | pmid = 9088660 | volume=27 | issue=3 | title=Royal jelly consumption and hypersensitivity in the community |date=March 1997 | journal=Clin. Exp. Allergy | pages=333–6 | doi = 10.1111/j.1365-2222.1997.tb00712.x | last1 = Leung | first1 = R | last2 = Ho | first2 = A | last3 = Chan | first3 = J | last4 = Choy | first4 = D | last5 = Lai | first5 = CK| s2cid=19626487 }}</ref>
==Sexes and castes==
Honey bees have three [[caste]]s: [[drone (bee)|drones]], [[Worker bee|workers]], and [[queen bee|queens]].<ref>{{cite encyclopedia |last1=Cervoni |first1=Mário Sérgio |last2=Hartfelder |first2=Klaus |editor-last=Starr |editor-first=Christopher K. |encyclopedia=Encyclopedia of Social Insects |date=2021 |publisher=Springer, Cham |isbn=978-3-030-28102-1 |pages=177–184 |language=en |title=Caste Differentiation: Honey Bees |doi=10.1007/978-3-030-28102-1_151}}</ref><ref>{{cite web|url=http://www.ikonet.com/en/visualdictionary/animal-kingdom/insects-and-arachnids/honeybee/castes.php|title=Bee castes|publisher=Visual Dictionary, QA International|date=2017|access-date=18 May 2017}}</ref><ref name="University of Georgia-2017">{{cite web|title=Getting Started: Honey Bee Biology|url=http://caes2.caes.uga.edu/bees/get-started/biology.html|publisher=University of Georgia College of Agricultural and Environmental Sciences|access-date=18 May 2017|date=2017|archive-date=24 January 2018|archive-url=https://web.archive.org/web/20180124075205/http://caes2.caes.uga.edu/bees/get-started/biology.html|url-status=dead}}</ref> Drones are male, while workers and queens are female.<ref name="University of Georgia-2017"/>
===Drones===
{{Main|Drone (bee)}}
[[File:Haplodiploid Sex Determination in Honey Bees.svg|thumb|upright=2.0|Honey bees have a [[haplodiploid]] system of sex determination.]]
Drones are typically [[haploid]], having only one set of [[chromosome]]s, and primarily exist for [[reproduction]].<ref name="University of Georgia-2017"/> They are produced by the queen if she chooses not to fertilize an egg or by an unfertilized laying worker. There are rare instances of diploid drone larvae. This phenomenon usually arises when there are more than two generations of brother-sister mating.<ref>Woyka, J.; Pszczelnictwa, Zaklad; [http://jerzy_woyke.users.sggw.pl/1963_dipldrlarvae.pdf Drone Larvae from Fertilized Eggs of the Honey Bee] {{Webarchive|url=https://web.archive.org/web/20141022232521/http://jerzy_woyke.users.sggw.pl/1963_dipldrlarvae.pdf |date=2014-10-22 }} Journal of Apiculture Research, (1963), pages 19–24</ref> Sex determination in honey bees is initially due to a single locus, called the complementary sex determiner (''csd'') gene. In developing bees, if the conditions are that the individual is heterozygous for the ''csd'' gene, they will develop into females. If the conditions are so that the individual is [[zygosity|hemizygous]] or homozygous for the ''csd'' gene, they will develop into males. The instances where the individual is homozygous at this gene are the instances of diploid males.<ref>Weinstock, George M.; Robinson, Gene E., & the Honeybee Genome Sequencing Consortium [http://www.nature.com/nature/journal/v443/n7114/full/nature05260.htm Insights into social insects from the genome of the honeybee Apis mellifera] ''[[Nature (journal)|Nature]]'', volume "'443'" (2006), pages 931–949</ref> Drones take 24 days to develop and may be produced from summer through to autumn, numbering as many as 500 per hive.<ref name="University of Georgia-2017"/> They are expelled from the hive during the winter months when the hive's primary focus is warmth and food conservation.<ref name="University of Georgia-2017"/> Drones have large eyes used to locate queens during mating flights. They do not defend the hive or kill intruders and do not have a [[stinger]].<ref name="PerfectBee-2017"/>
===Workers===
{{Main|Worker bee}}
Workers have two sets of chromosomes.<ref>{{cite web |last=Harbo |first=JR |last2=Rinderer |first2=TE |date=1980 |title=Breeding and Genetics of Honey Bees |url=http://beesource.com/resources/usda/breeding-and-genetics-of-honey-bees/ |access-date=18 May 2017 |publisher=Beesource Beekeeping}}</ref> They are produced from an egg that the queen has selectively fertilized from stored sperm. Workers typically develop in 21 days. A typical colony may contain as many as 60,000 worker bees.<ref name="University of Georgia-2017"/> Workers exhibit a wider range of behaviors than either queens or drones. Their duties change with age in the following order (beginning with cleaning out their cell after eating through their capped brood cell): feed brood, receive nectar, clean hive, guard duty, and forage.<ref name="University of Georgia-2017"/><ref name="PerfectBee-2017">{{cite web|url=https://www.perfectbee.com/learn-about-bees/types-of-bees/|title=Worker, drone and queen bees|publisher=PerfectBee LLC|date=2017|access-date=18 May 2017}}</ref> Some workers engage in other specialized behaviors, such as "undertaking" (removing corpses of their nestmates from inside the hive).<ref name="PerfectBee-2017"/>
Workers have morphological specializations, including the [[pollen basket]] ''(corbicula)'',<ref>{{cite web|url=http://www.ikonet.com/en/visualdictionary/animal-kingdom/insects-and-arachnids/honeybee/morphology-of-a-honeybee-worker.php|title=Morphology of a honeybee: worker|publisher=Visual Dictionary, QA International|date=2017|access-date=18 May 2017}}</ref> abdominal glands that produce beeswax, brood-feeding glands, and barbs on the sting. Under certain conditions (for example, if the colony becomes queenless), a worker may develop ovaries.
Worker honey bees perform different behavioural tasks that cause them to be exposed to different local environments.<ref name="Jones-2018a">{{cite journal|doi=10.1002/ece3.3597|pmid=29321884|pmc=5756847|title=Gut microbiota composition is associated with environmental landscape in honey bees|journal=Ecology and Evolution|volume=8|issue=1|pages=441–451|year=2018|last1=Jones|first1=Julia C|last2=Fruciano|first2=Carmelo|last3=Hildebrand|first3=Falk|last4=Al Toufalilia|first4=Hasan|last5=Balfour|first5=Nicholas J|last6=Bork|first6=Peer|last7=Engel|first7=Philipp|last8=Ratnieks|first8=Francis LW|last9=Hughes|first9=William OH|bibcode=2018EcoEv...8..441J }}</ref><ref name="Jones-2018b">{{cite journal|doi=10.1007/s00040-018-0624-9|pmid=30100619|pmc=6061168|title=The gut microbiome is associated with behavioural task in honey bees|journal=Insectes Sociaux|volume=65|issue=3|pages=419–429|year=2018|last1=Jones|first1=J. C|last2=Fruciano|first2=C|last3=Marchant|first3=J|last4=Hildebrand|first4=F|last5=Forslund|first5=S|last6=Bork|first6=P|last7=Engel|first7=P|last8=Hughes|first8=W. O. H}}</ref> The [[Gut flora|gut microbial]] composition of workers varies according to the landscape and plant species they [[foraging|forage]], such as differences in [[rapeseed]] crops,<ref name="Jones-2018a"/> and with different hive tasks, such as nursing or food processing.<ref name="Jones-2018b"/>
===Queens===
{{Main|Queen bee}}
Queen honey bees are created when worker bees feed a single female larva an exclusive diet of a food called "[[royal jelly]]".<ref name="University of Georgia-2017"/><ref name="PerfectBee-2017"/> Queens are produced in oversized cells and develop in only 16 days; they differ in physiology, morphology, and behavior from worker bees. In addition to the greater size of the queen, she has a functional set of ovaries, and a spermatheca, which stores and maintains sperm after she has mated. ''Apis'' queens practice [[Polyandry in animals|polyandry]], with one female mating with multiple males. The highest documented mating frequency for an ''Apis'' queen is in ''[[Apis nigrocincta]]'', where queens mate with an extremely high number of males with observed numbers of different matings ranging from 42 to 69 drones per queen.<ref>Hadisoesilo, Soesilawati. "The Comparative Study of Two Species of Cavity-Nesting Honey Bees of Sulawesi, Indonesia" (PDF).</ref> The sting of queens is not barbed like a worker's sting, and queens lack the glands that produce beeswax. Once mated, queens may lay up to 2,000 eggs per day.<ref name="PerfectBee-2017"/> They produce a variety of pheromones that regulate the behavior of workers and help swarms track the queen's ___location during the swarming.<ref name="PerfectBee-2017"/>
=== Queen-worker conflict ===
{{Main|Worker policing}}
When a fertile female worker produces drones, a conflict arises between her interests and those of the queen. The worker shares half her genes with the drone and one-quarter with her brothers, favouring her offspring over those of the queen. The queen shares half her genes with her sons and one-quarter with the sons of fertile female workers.<ref>{{cite journal | last1 = Wenseleers | first1 = T. | last2 = Helanterä | first2 = H. | last3 = Hart | first3 = A. | last4 = Ratnieks | first4 = F. L. W. | year = 2004 | title = Worker reproduction and policing in insect societies: an ESS analysis | journal = Journal of Evolutionary Biology | volume = 17 | issue = 5| pages = 1035–1047 | doi = 10.1111/j.1420-9101.2004.00751.x | pmid = 15312076 | s2cid = 7239058 | doi-access = free }}</ref> This pits the worker against the queen and other workers, who try to maximize their [[reproductive fitness]] by rearing the offspring most related to them. This relationship leads to a phenomenon known as "worker policing". In these rare situations, other worker bees in the hive who are genetically more related to the queen's sons than those of the fertile workers will patrol the hive and remove worker-laid eggs. Another form of worker-based policing is aggression toward fertile females.<ref>{{cite journal | last1 = Ratnieks | first1 = F. | last2 = Visscher | first2 = P. Kirk | year = 1989 | title = Worker policing in the honeybee | journal = Nature | volume = 342 | issue = 6251| pages = 796–797 | doi = 10.1038/342796a0 | bibcode=1989Natur.342..796R| s2cid = 4366903 }}</ref> Some studies have suggested a queen pheromone which may help workers distinguish worker- and queen-laid eggs, but others indicate egg viability as the key factor in eliciting the behavior.<ref>{{cite journal | last1 = Pirk | first1 = C. | last2 = Neumann | first2 = P. | last3 = Hepburn | first3 = R. | last4 = Moritz | first4 = R. | last5 = Tautz | first5 = J. | year = 2003 | title = Egg viability and worker policing in honey bees | journal = PNAS | volume = 101 | issue = 23| pages = 8649–8651 | doi = 10.1073/pnas.0402506101 | pmid = 15169961 | pmc = 423249 | bibcode = 2004PNAS..101.8649P | doi-access = free }}</ref><ref>{{cite journal | last1 = Oldroyd | first1 = B. | last2 = Ratnieks | first2 = Francis | year = 2002 | title = Egg-marking pheromones in honey-bees Apis mellifera | journal = Behavioral Ecology and Sociobiology | volume = 51 | issue = 6| pages = 590–591 | doi = 10.1007/s00265-002-0480-4 | bibcode = 2002BEcoS..51..590O | s2cid = 30446742 }}</ref> Worker policing is an example of [[coercion|forced altruism]], where the benefits of worker reproduction are minimized and that of rearing the queen's offspring maximized.
In very rare instances workers subvert the policing mechanisms of the hive, laying eggs which are removed at a lower rate by other workers; this is known as anarchic syndrome. Anarchic workers can activate their ovaries at a higher rate and contribute a greater proportion of males to the hive. Although an increase in the number of drones would decrease the overall productivity of the hive, the reproductive fitness of the drones' mother would increase. The anarchic syndrome is an example of selection working in opposite directions at the [[Unit of selection|individual and group levels]] for the stability of the hive.<ref>{{cite journal | last1 = Barron | first1 = A. | last2 = Oldroyd | first2 = B | last3 = Ratnieks | first3 = F.L.W. | year = 2001 | title = Worker reproduction in honey-bees (Apis) and the anarchic syndrome: a review | journal = Behavioral Ecology and Sociobiology | volume = 50 | issue = 3| pages = 199–208 | doi = 10.1007/s002650100362 | bibcode = 2001BEcoS..50..199B | s2cid = 17246102 }}</ref>
Under ordinary circumstances, the death (or removal) of a queen increases reproduction in workers, and a significant proportion of workers will have active ovaries in the absence of a queen. The workers of the hive produce the last batch of drones before the hive eventually collapses. Although during this period worker policing is usually absent, in certain groups of bees it continues.<ref>{{cite journal | last1 = Châline | first1 = N. | last2 = Martin | first2 = S.J. | last3 = Ratnieks | first3 = F.L.W. | title = Worker policing persists in a hopelessly queenless honey bee colony (Apis mellifera) | journal = Insectes Soc | year = 2004 | volume = 51 | issue = 2 | pages = 1–4 | doi = 10.1007/s00040-003-0708-y | s2cid = 11988371 }}</ref>
According to the strategy of [[kin selection]], worker policing is not favored if a queen does not mate multiple times. Workers would be related by three-quarters of their genes, and the difference in [[Coefficient of relationship|relationship]] between the sons of the queen and those of the other workers would decrease. The benefit of policing is negated, and policing is less favored. Experiments confirming this hypothesis have shown a correlation between higher mating rates and increased rates of worker policing in many species of social [[hymenoptera]].<ref>Davies, N.R., Krebs, J.R., and West, S.A. An Introduction to Behavioral Ecology. 4th ed. West Sussex: Wiley-Blackwell, 2012. Print. pp. 387–388</ref>
== Timeline of reproduction ==
For ''Apis mellifera'', queens are the central reproducers among their colonies. Although reproduction may occur around the calendar, it may stop in the late fall due to falling temperatures. If a colony does not have a queen or she is unable to reproduce, workers can lay unfertilized eggs that may develop into males. The queens, however, do not reach this point immediately. Typically, it takes a queen 16 days to reach adulthood, with an additional week to begin developing and laying eggs.<ref name="Hammond-2009">{{Cite web |last1=Hammond |first1=George |last2=Blankenship |first2=Madison |title=''Apis mellifera'' (honey bee) |url=https://animaldiversity.org/accounts/Apis_mellifera/ |access-date=2023-04-17 |website=Animal Diversity Web |language=en}}</ref> To begin the process of reproduction in a honeybee colony, workers begin to produce queen larvae while simultaneously finding a place to create a new hive.<ref name="Hammond-2009"/> The queen larvae will then hatch at the old hive, and the queens will fight one another until there is only a single queen left to begin reproducing.<ref name="Hammond-2009"/>
== Reproductive strategies ==
Once a queen matures and is ready to begin reproducing, she will begin making flights to orient to mating in free flight and finding mates before actually beginning to mate. Queens that are ready to mate take between 1 and 6 flights across multiple consecutive days, called nuptial flights.<ref>{{Cite journal |last1=Heidinger |first1=Ina Monika Margret |last2=Meixner |first2=Marina Doris |last3=Berg |first3=Stefan |last4=Büchler |first4=Ralph |date=2014-07-01 |title=Observation of the Mating Behavior of Honey Bee (Apis mellifera L.) Queens Using Radio-Frequency Identification (RFID): Factors Influencing the Duration and Frequency of Nuptial Flights |journal=Insects |volume=5 |issue=3 |pages=513–527 |doi=10.3390/insects5030513 |issn=2075-4450 |pmc=4592583 |pmid=26462822 |doi-access=free }}</ref> Throughout their nuptial flights, queens engage with multiple mates and have little control over the number of times they do so.<ref name="Hammond-2009"/>
The process of queens engaging with their mates is not widely understood because the process takes place in free flight, so it is difficult to observe despite various advances in technology and observation techniques. It begins with drones flying in the same area where they know the queen will soon arrive, waiting for her to join them.<ref>{{Cite journal |last1=Barron |first1=Andrew B. |last2=Oldroyd |first2=Benjamin P. |last3=Ratnieks |first3=Francis L. |date=2001-08-01 |title=Worker reproduction in honey-bees (Apis) and the anarchic syndrome: a review |url=https://doi.org/10.1007/s002650100362 |journal=Behavioral Ecology and Sociobiology |language=en |volume=50 |issue=3 |pages=199–208 |doi=10.1007/s002650100362 |bibcode=2001BEcoS..50..199B |s2cid=17246102 |issn=1432-0762|url-access=subscription }}</ref> When the queen arrives, she is crowded immediately by the drones who are eager to mate with her. The drones receive a signal from the queen that her "sting chamber" is open, which induces the drones to mate with her and bring forward their physical contact which warrants reproduction. A successful drone clasps onto the queen and releases seminal fluid and spermatozoa into the queen. After this process is complete, the drone typically remains inside of the queen, which is indicative of the drone's desire to deter other drones from engaging with the queen and reproducing.<ref>{{Cite web |title=Chapter 8: Most Spectacular Mating {{!}} The University of Florida Book of Insect Records {{!}} Department of Entomology & Nematology {{!}} UF/IFAS |url=https://entnemdept.ufl.edu/walker/ufbir/chapters/chapter_08.shtml |access-date=2023-04-17 |website=entnemdept.ufl.edu}}</ref> This behavior also indicates that if the drone blocks other drones from mating with the queen, it will allow the mating drone to fertilize a greater number of the queen's eggs. If the drone does not remain within the queen and removes itself from her, the drone can reproduce again with slim chances. Finally, the drone will die after mating with the queen within minutes or hours after reproduction is complete.
==Defense==
{{Main|Bee sting}}
[[File:Honeybee thermal defence01.jpg|thumb|''[[Apis cerana japonica]]'' forming a ball around two [[Asian giant hornet|hornets]]: The body heat trapped by the ball will overheat and kill the hornets.]]
All honey bees live in colonies where the workers [[Bee sting|sting]] intruders as a form of defense, and alarmed bees release a [[pheromone]] that stimulates the attack response in other bees. The different species of honey bees are distinguished from all other bee species by the possession of small barbs on the sting, but these barbs are found only in the worker bees.<ref>{{cite web |title=Bee Bonanza. The story of Honey Bees. |url=https://askabiologist.asu.edu/bee-honey |website=askabiologist.asu.edu |date=13 June 2017 |publisher=Arizona State University |access-date=15 March 2022}}</ref>
The sting apparatus, including the barbs, may have evolved specifically in response to predation by vertebrates, as the barbs do not usually function (and the sting apparatus does not detach) unless the sting is embedded in fleshy tissue. While the sting can also penetrate the membranes between joints in the exoskeleton of other insects (and is used in fights between queens), in the case of ''Apis cerana japonica'', defense against larger insects such as predatory wasps (e.g. [[Asian giant hornet]]) is usually performed by surrounding the intruder with a mass of defending worker bees, which vibrate their muscles vigorously to raise the temperature of the intruder to a lethal level ("balling").<ref>{{cite web |url=http://www.bio.davidson.edu/people/midorcas/animalphysiology/websites/2001/Thawley/defense.htm |title=Heat tolerance as a weapon |last=Thawley |first=C. H. |publisher=[[Davidson College]] |access-date=1 June 2010 |url-status=dead |archive-url=https://web.archive.org/web/20100718201646/http://www.bio.davidson.edu/people/midorcas/animalphysiology/websites/2001/Thawley/defense.htm |archive-date=18 July 2010 }}</ref> Previously, heat alone was thought to be responsible for killing intruding wasps, but recent experiments have demonstrated the increased temperature in combination with increased carbon dioxide levels within the ball produce the lethal effect.<ref>{{cite journal|title=Heat and carbon dioxide generated by honeybees jointly act to kill hornets|journal=[[Naturwissenschaften]] |year=2009 |author1=Michio Sugahara |last2=Fumio Sakamoto |volume=96 |issue=9 |pages=1133–6 |doi=10.1007/s00114-009-0575-0 |pmid=19551367|bibcode=2009NW.....96.1133S |s2cid=22080257 }}</ref><ref>{{cite news |last=Gill |first=Victoria |date=3 July 2009 |title=Honeybee mobs overpower hornets |url=http://news.bbc.co.uk/2/hi/science/nature/8129536.stm |access-date=5 July 2009 |work=BBC News}}</ref> This phenomenon is also used to kill a queen perceived as intruding or defective, an action known to beekeepers as 'balling the queen', named for the ball of bees formed.
Defense can vary based on the habitat of the bee. In the case of those honey bee species with open combs (e.g., ''A. dorsata''), would-be predators are given a warning signal that takes the form of a "[[wave (audience)|wave]]" that spreads as a ripple across a layer of bees densely packed on the surface of the comb when a threat is perceived and consists of bees momentarily arching their bodies and flicking their wings.<ref>{{Cite web |date=15 September 2008 |title=Giant Honeybees Use Shimmering 'Mexican Waves' To Repel Predatory Wasps |url=https://www.sciencedaily.com/releases/2008/09/080909204550.htm |access-date=11 August 2025 |website=ScienceDaily}}</ref> In cavity dwelling species such as ''[[Apis cerana]]'', ''[[Apis mellifera]]'', and ''[[Apis nigrocincta]]'', entrances to these cavities are guarded and checked for intruders in incoming traffic. Another act of defense against nest invaders, particularly wasps, is "body shaking", a violent and pendulum-like swaying of the abdomen, performed by worker bees.<ref>Radloff, Sara E.; Hepburn, H. Randall; Engel, Michael S. (2011). ''Honeybees of Asia''. Berlin: Springer Science & Business Media. {{ISBN|978-3642164217}}.</ref>
A 2020 study of ''[[Apis cerana]]'' in Vietnam found that they use [[feces]] and even human urine to defend their hives against raids by hornets (''[[Vespa soror]]''), a strategy not replicated by their European and North American counterparts,<ref name="Mattila-2020">{{cite journal | last1=Mattila | first1=Heather R. | last2=Otis | first2=Gard W. | last3=Nguyen | first3=Lien T. P. | last4=Pham | first4=Hanh D. | last5=Knight | first5=Olivia M. | last6=Phan | first6=Ngoc T. | editor-last=Blenau | editor-first=Wolfgang | title=Honey bees (Apis cerana) use animal feces as a tool to defend colonies against group attack by giant hornets (''Vespa soror'') | journal=PLOS ONE | volume=15 | issue=12 | date=9 December 2020 | issn=1932-6203 | doi=10.1371/journal.pone.0242668 | page=e0242668| pmid=33296376 | pmc=7725375 | bibcode=2020PLoSO..1542668M | s2cid=228087051 | doi-access=free }}</ref> though collection and use of feces in nest construction is well-known in [[stingless bee]]s.<ref>{{Cite journal |last1=Basari |first1=Norasmah |last2=Ramli |first2=Sarah Najiah |last3=Mohd Khairi |first3=Nur 'Aina Syakirah |date=2018-10-11 |title=Food Reward and Distance Influence the Foraging Pattern of Stingless Bee, Heterotrigona itama |journal=Insects |language=en |volume=9 |issue=4 |pages=138 |doi=10.3390/insects9040138 |doi-access=free |issn=2075-4450 |pmc=6315735 |pmid=30314344}}</ref><ref>{{cite book |last=Jalil |first=Abu Hassan |url=https://books.google.com/books?id=fUh9BAAAQBAJ |title=Beescape for Meliponines: Conservation of Indo-Malayan Stingless Bees |date=8 July 2014 |publisher=Partridge Publishing Singapore |isbn=978-1-4828-2362-2}}</ref>
===Venom===
The stings of honey bees are barbed and therefore embed themselves into the sting site, and the sting apparatus has its own musculature and ganglion which keep delivering venom even after detachment.<ref name="Biller-2014">{{cite book |last1=Biller |first1=Jose |last2=Ferro |first2=Jose M |title=Neurologic Aspects of Systemic Disease, Part II |date=7 February 2014 |publisher=Elsevier |isbn=9780702040870 |page=995 |edition=1st}}</ref> The gland which produces the alarm pheromone is also associated with the sting apparatus. The embedded stinger continues to emit additional alarm pheromones after it has torn loose; other defensive workers are thereby attracted to the sting site. The worker dies after the sting becomes lodged and is subsequently torn loose from the bee's abdomen. The honey bee's venom, known as [[apitoxin]], carries several active components, the most abundant of which is [[melittin]],<ref name="Chen-2016">{{cite journal | vauthors = Chen J, Guan SM, Sun W, Fu H | title = Melittin, the Major Pain-Producing Substance of Bee Venom | journal = Neuroscience Bulletin | volume = 32 | issue = 3 | pages = 265–72 | year = 2016 | pmid = 26983715 | pmc = 5563768 | doi = 10.1007/s12264-016-0024-y }}</ref> and the most biologically active are [[enzyme]]s, particularly [[phospholipase A2]].<ref>{{cite journal|pmc=4548770|year=2015|last1=Ramanadham|first1=S|title=Calcium-independent phospholipases A2 and their roles in biological processes and diseases|journal=Journal of Lipid Research|volume=56|issue=9|pages=1643–1668|last2=Ali|first2=T|last3=Ashley|first3=J. W|last4=Bone|first4=R. N|last5=Hancock|first5=W. D|last6=Lei|first6=X|doi=10.1194/jlr.R058701 |doi-access=free |pmid=26023050}}</ref>
[[File:Honey Bee Colony Count by Number of United States' States (Jan 1, 2015).png|thumb|342x342px|Shows active colonies on January 1, 2015 with U.S. state count.<ref>{{Cite report |url=https://downloads.usda.library.cornell.edu/usda-esmis/files/rn301137d/p2676z253/gq67jv05t/BeeColonies-05-12-2016.pdf |title=Honey Bee Colonies (May 2016) |date=2016-05-12 |publisher=[[National Agricultural Statistics Service]] |issn=2470-993X |access-date=2025-06-07}}</ref> The highest states being those like California and Florida with the most colonies.]]
[[File:Honey Bee Colony Count by Number of States (Jan 1, 2023).png|thumb|341x341px|Shows active colonies on January 1, 2023 with state count.<ref>{{Cite report |url=https://downloads.usda.library.cornell.edu/usda-esmis/files/rn301137d/4j03fq210/8w32sw99n/hcny0824.pdf |title=Honey Bee Colonies (August 2024) |date=2024-08-01 |publisher=[[National Agricultural Statistics Service]] |issn=2470-993X |access-date=2025-06-07}}</ref> The highest states being those like California and Texas with the most production.]]
Honey bee venom is under laboratory and [[clinical research]] for its potential properties and uses in reducing risks for [[adverse event]]s from bee venom therapy,<ref>{{cite journal|pmc=4440710|year=2015|last1=Park|first1=J. H|title=Risk Associated with Bee Venom Therapy: A Systematic Review and Meta-Analysis|journal=PLOS ONE|volume=10|issue=5|pages=e0126971|last2=Yim|first2=B. K|last3=Lee|first3=J. H|last4=Lee|first4=S|last5=Kim|first5=T. H|doi=10.1371/journal.pone.0126971|pmid=25996493|bibcode=2015PLoSO..1026971P|doi-access=free}}</ref> [[rheumatoid arthritis]],<ref>{{cite journal|pmc=4225238|year=2014|last1=Lee|first1=J. A|title=Bee venom acupuncture for rheumatoid arthritis: A systematic review of randomised clinical trials|journal=BMJ Open|volume=4|issue=11|pages=e006140|last2=Son|first2=M. J|last3=Choi|first3=J|last4=Jun|first4=J. H|last5=Kim|first5=J. I|last6=Lee|first6=M. S|doi=10.1136/bmjopen-2014-006140|pmid=25380812}}</ref> and use as an [[immunotherapy]] for protection against [[allergy|allergies]] from insect stings.<ref>{{cite journal|pmid=23076950|year=2012|last1=Boyle|first1=R. J|title=Venom immunotherapy for preventing allergic reactions to insect stings|journal=Cochrane Database of Systematic Reviews|volume=10|pages=CD008838|last2=Elremeli|first2=M|last3=Hockenhull|first3=J|last4=Cherry|first4=M. G|last5=Bulsara|first5=M. K|last6=Daniels|first6=M|last7=Oude Elberink|first7=J. N|issue=2|doi=10.1002/14651858.CD008838.pub2|pmc=8734599 |url=https://pure.rug.nl/ws/files/67436563/Boyle_et_al_2012_Cochrane_Database_of_Systematic_Reviews.pdf}}</ref> Bee venom products are marketed in many countries, but, as of 2018, there are no approved clinical uses for these products which carry various warnings for potential allergic reactions.<ref name="Drugs.com-2018">{{cite web |title=Wasp and Bee venom |url=https://www.drugs.com/pro/wasp-and-bee-venom.html |publisher=Drugs.com |access-date=29 June 2018 |date=2018}}</ref>
==Competition==
{{further|Western honey bee#As an environmental threat}}
With an increasing population of honey bees in specific areas due to beekeeping, Western honey bees (as an [[invasive species]]) and native wild bees often have to compete for the limited habitat and food sources available,<ref>{{Cite journal|last1=Hudewenz|first1=Anika|last2=Klein|first2=Alexandra-Maria|date=1 December 2013|title=Competition between honey bees and wild bees and the role of nesting resources in a nature reserve|journal=Journal of Insect Conservation|language=en|volume=17|issue=6|pages=1275–1283|doi=10.1007/s10841-013-9609-1|bibcode=2013JICon..17.1275H |s2cid=16268870|issn=1366-638X}}</ref><ref>{{Cite journal |last1=MacInnis |first1=Gail |last2=Normandin |first2=Etienne |last3=Ziter |first3=Carly D. |date=2023-02-03 |title=Decline in wild bee species richness associated with honey bee ( Apis mellifera L.) abundance in an urban ecosystem |journal=PeerJ |language=en |volume=11 |pages=e14699 |doi=10.7717/peerj.14699 |doi-access=free |issn=2167-8359 |pmc=9901307 |pmid=36755869}}</ref><ref>{{Cite journal |last1=Geldmann |first1=Jonas |last2=González-Varo |first2=Juan P. |date=2018-01-26 |title=Conserving honey bees does not help wildlife |url=https://txmn.org/gulfcoast/files/2018/02/Science-Vol.-359-pp.-392-393.pdf |journal=Science |language=en |volume=359 |issue=6374 |pages=392–393 |doi=10.1126/science.aar2269 |pmid=29371456 |bibcode=2018Sci...359..392G |issn=0036-8075}}</ref> and Western honey bees may become defensive in response to the seasonal arrival of competition from other colonies, particularly Africanized bees which may be on the offence and defence year round due to their tropical origin.<ref>{{Cite journal|last1=Johnson|first1=Brian R.|last2=Nieh|first2=James C.|date=1 November 2010|title=Modeling the Adaptive Role of Negative Signaling in Honey Bee Intraspecific Competition|journal=Journal of Insect Behavior|language=en|volume=23|issue=6|pages=459–471|doi=10.1007/s10905-010-9229-5|issn=0892-7553|pmc=2955239|pmid=21037953|bibcode=2010JIBeh..23..459J }}</ref>
==Communication==
{{Main|Bee learning and communication}}
Honey bees are known to communicate through many different chemicals and odors, as is common in insects. They also rely on a sophisticated dance language that conveys information about the distance and direction to a specific ___location (typically a nutritional source, e.g., flowers or water). The dance language is also used during the process of colony fission, or swarming, when scouts communicate the ___location and quality of nesting sites.<ref>{{Cite web|url=https://content.ces.ncsu.edu/honey-bee-dance-language|title=The Honey Bee Dance Language|last=Tarpy|first=David|date=2016|website=NC State Extension}}</ref>
The details of the signalling being used vary from species to species; for example, the two smallest species, ''[[Apis andreniformis]]'' and ''A. florea'', dance on the upper surface of the comb, which is horizontal (not vertical, as in other species), and worker bees orient the dance in the actual compass direction of the resource to which they are recruiting.
[[Carniolan honey bee]]s (''Apis mellifera carnica'') use their antennae asymmetrically for social interactions, with a strong lateral preference to use their right antennae.<ref name="Rogers-2013">{{cite journal |last=Rogers |first=Lesley J. |last2=Rigosi |first2=Elisa |last3=Frasnelli |first3=Elisa |last4=Vallortigara |first4=Giorgio |author-link4=Giorgio Vallortigara |date=27 June 2013 |title=A right antenna for social behaviour in honeybees |journal=Scientific Reports |volume=3 |pages=2045 |bibcode=2013NatSR...3.2045R |doi=10.1038/srep02045 |pmc=3694496 |pmid=23807465}}</ref><ref>{{cite web |last=Shugart |first=Jessica |title=Honeybees use right antennae to tell friend from foe |url=http://www.sciencenews.org/view/generic/id/351355/description/Honeybees_use_right_antennae_to_tell_friend_from_foe |access-date=12 March 2016 |work=Science News}}</ref>
There has been speculation as to honey bee [[consciousness]].<ref>{{cite news|url=https://www.nytimes.com/2016/04/19/science/honeybees-insects-consciousness-brains.html|title=Do Honeybees Feel? Scientists Are Entertaining the Idea|first=James|last=Gorman|newspaper=The New York Times|date=18 April 2016}}</ref> While honey bees lack the parts of the brain that a human being uses for consciousness like the cerebral cortex or even the cerebrum itself, when those parts of a human brain are damaged, the midbrain seems able to provide a small amount of consciousness. Honey bees have a tiny structure that appears similar to a human midbrain, so if it functions the same way they may be able to achieve a small amount of simple awareness of their bodies.
==Symbolism==
{{Main|Bee (mythology)}}
The bee was used as a symbol of government by Emperor [[Napoleon I of France]].<ref>{{cite web |url=http://www.napoleon.org/en/essential_napoleon/symbols/index.asp |title=The symbols of empire |publisher=Napoleon.org |access-date=1 June 2010}}</ref> Both the [[Hindu]] ''[[Atharva Veda]]''<ref>"O Asvins, lords of brightness, anoint me with the honey of the bee, that I may speak forceful speech among men! ''Atharva Veda'' 91–258, quoted in Maguelonne Toussaint-Samat (Anthea Bell, tr.) ''The History of Food'', 2nd ed. 2009:14.</ref> and the ancient Greeks associated lips anointed with honey with the gift of eloquence and even of prescience. The priestess at [[Delphi]] was the "Delphic Bee".
The ''[[Quran]]'' has a [[Sura]] (chapter) titled "[[An-Nahl|The Bee]]". It is named after honey bees and contains a comparison of the industry and adaptability of honey bees to the industry of man.<ref>{{Cite web|url=http://www.islamicity.com/science/quranandscience/animals/generatedfiles/thehoneybee.htm|title=THE HONEY BEE|website=islamicity.com}}</ref>
{{blockquote|And your Lord inspired the bees: "Make [your] homes in the mountains, the trees, and in what people construct, and feed from [the flower of] any fruit [you please] and follow the ways your Lord has made easy for you." From their bellies comes forth liquid of varying colours, in which there is healing for people. Surely in this is a sign for those who reflect.|{{qref|16|68–69|c=y}}<ref>{{Cite web|url=http://www.miracles-of-quran.com/honey_bees.htm|title=Honey Bees in Quran|website=miracles-of-quran.com|access-date=2019-08-25|archive-date=2019-08-25|archive-url=https://web.archive.org/web/20190825160604/http://www.miracles-of-quran.com/honey_bees.htm|url-status=dead}}</ref>}}
In ancient Egyptian mythology, honey bees were believed to be born from the tears of the [[Sun god]], [[Ra]].<ref>{{cite news |last=Norton |first=Holly |date=24 May 2017 |title=Honey, I love you: our 40,000-year relationship with the humble bee |url=https://www.theguardian.com/science/2017/may/24/honey-i-love-you-our-40000-year-relationship-with-the-humble-bee |access-date=29 June 2018 |newspaper=The Guardian}}</ref> Because of their divine origin, they were used to represent the Pharaoh. They were also used as a symbol of Lower Egypt in conjunction with the sedge, which represented Upper Egypt.<ref>{{Cite book |last=Haynes |first=Dawn |url=https://scholar.sun.ac.za/bitstream/10019.1/79920/1/haynes_symbolism_2013.pdf |title=The Symbolism and Significance of the Butterfly in Ancient Egypt}}</ref>
In Joseph and Asenath, a work composed by ancient Egyptian Jews who may have been affiliated with the [[Leontopolis (Heliopolis)|Leontopolis temple]], bee and honey imagery appears when Asenath converts and is visited by an angel. If the work was indeed connected to the Leontopolis temple, the bees likely represent Levite priests, and the imagery intends to signify the legitimacy of a Jewish temple in Egypt.<ref>{{Cite book |last=Piotrkowski |first=Meron M. |url=https://www.worldcat.org/title/on1076801783 |title=Priests in exile: the history of the temple of Onias and its community in the Hellenistic Period |date=2019 |publisher=De Gruyter |isbn=978-3-11-059107-1 |series=Studia Judaica |___location=Berlin |oclc=on1076801783}}</ref>
A community of honey bees has often been employed by political theorists as a model of human society, from Aristotle and Plato to Virgil.<ref>Virgil, ''[[Georgics]]'', book IV.</ref><ref>{{cite book |last=Wilson |first=Bee |title=The Hive: The Story of the Honeybee |publisher=[[John Murray (publishing house)|John Murray]] |year=2004 |isbn=978-0-7195-6598-4 |___location=London |page=14}}</ref> Honey bees, signifying immortality and resurrection, were royal emblems of the [[Merovingians]]. The U.S. state of [[Utah]] is called the "Beehive State", the state emblem is the beehive, the state insect is the honey bee, and a beehive and the word "industry" appear on both the state flag and seal.<ref>{{cite web |title=Utah State Motto and Emblem |url=http://onlinelibrary.utah.gov/research/utah_symbols/motto.html |publisher=Utah State Library |access-date=13 October 2017 |archive-date=21 October 2017 |archive-url=https://web.archive.org/web/20171021011044/http://onlinelibrary.utah.gov/research/utah_symbols/motto.html |url-status=dead }}</ref>
==Gallery==
<gallery mode="packed" heights="150">
File:Bienenkoenigin.JPG|A coloured dot applied by a beekeeper identifies a [[queen bee|queen]] [[Western honey bee]].
File:Honeybee02.jpg| [[Western honey bee]] foragers loaded with pollen on the hive landing board
File:Cerana.jpg|[[Eastern honey bee]] (''A. cerana'') in Hong Kong
File:ApisDorsataHive.jpg|[[Apis dorsata|Giant honey bee]] (''A. dorsata'')
File:Pollinationn.jpg|[[Western honey bee]] visiting flowers
File:Natural Beehive and Honeycombs.jpg|A colony of giant honey bees (''A. dorsata'') on their comb
File:Bee-apis.jpg|[[Western honey bee]]
File:Honey bee on rock rose.jpg|[[Western honey bee]] on rock rose (''[[Cistus]]'') in Oakland, California
File:Honeybee pollen turnips.jpg|[[Western honey bee]] collecting pollen from turnip blossoms in [[Eastern Oklahoma]]
File:Green Bee-eater Merops orientalis feeding on Honey-bee by Dr. Raju Kasambe DSCN1640 (1).jpg|A predator, the [[Asian green bee-eater]]
File:A natural beehive in Chandigarh.jpg|A wild colony in Chandigarh
File:Honeybee on Lavender.jpg|alt=Honeybee on Lavender|[[Western honey bee]] on lavender (''[[Lavandula]]'') in [[Lompoc, California]]
</gallery>
==See also==
* [[Bees and toxic chemicals]]
* [[Honey bee life cycle]]
* [[Honey bee starvation]]
* ''[[More than Honey]]'' – a 2012 Swiss documentary film about honey bees
*[[Melittology]] (also known as apiology) – the study of bees
*{{annotated link |Brachygastra}} (honey wasps) known to collect and store nectar as honey
*{{annotated link|Pollen wasp}} unique among wasps in feeding their larvae exclusively with pollen and nectar, in a fashion quite similar to many solitary bees
* [[Honeypot ant]]
==Notes==
{{notelist}}
==References==
{{Reflist|30em}}
==External links==
{{Commons category|Apis (insect)}}
{{Wiktionary}}
{{Wikispecies|Apis|''Apis''}}
* ''[http://watch.opb.org/video/2365583178/ Could a Mushroom Save a Honeybee?] {{Webarchive|url=https://web.archive.org/web/20160407145124/http://watch.opb.org/video/2365583178/ |date=2016-04-07 }}'' Documentary produced by [[Oregon Field Guide]], 14 October 2015
{{BeeColonyMemberTypes}}
{{Insects in culture}}
{{Eusociality}}
{{Animal domestication}}
{{Taxonbar|from=Q102857}}
{{Authority control}}
{{DEFAULTSORT:Honey Bee}}
[[Category:Apis (genus)| ]]
[[Category:Articles containing video clips]]
[[Category:Beekeeping]]
[[Category:Bees]]
[[Category:Extant Oligocene first appearances]]
[[Category:Insects in culture]]
[[Category:Insects described in 1758]]
[[Category:Animal taxa named by Carl Linnaeus]]
[[Category:Pollination]]
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