View source for Bee

{{Short description|Clade of insects}}
{{About|the group of flying insects|other uses}}
{{Redirect|Anthophila|the moth genus|Anthophila (moth){{!}}''Anthophila'' (moth)}}
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{{Pp-semi-indef}}
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{{Use dmy dates|date=November 2022}}
{{Automatic taxobox
| name = Bees
| fossil_range = {{fossilrange|70|0}}[[Late Cretaceous]]&nbsp;– [[Holocene|Present]]
| image = Tetragonula carbonaria (14521993792).jpg
| image_caption = The sugarbag bee, ''[[Tetragonula carbonaria]]''
| display_parents = 3
| taxon = Anthophila
| authority =
| subdivision_ranks = Families
| subdivision = {{Plain list|
* [[Andrenidae]]
* [[Apidae]]
* [[Colletidae]]
* [[Halictidae]]
* [[Megachilidae]]
* [[Melittidae]]
* [[Stenotritidae]]}}
| synonyms = '''Apiformes''' (from Latin ''[[wikt:apis#Etymology|'apis']]'')
}}
'''Bees''' are winged [[insect]]s closely related to [[wasp]]s and [[ant]]s, known for their roles in [[pollination]] and, in the case of the best-known bee species, the [[western honey bee]], for producing [[honey]]. Bees are a [[monophyly|monophyletic]] lineage within the superfamily [[Apoidea]]. They are currently considered a [[clade]], called '''Anthophila'''.<ref>Engel, M. S. (2005) Family-group names for bees (Hymenoptera, Apoidea). ''American Museum Novitates'' 3476.</ref> There are over 20,000 known species of bees in seven recognized [[Family (biology)|biological families]].<ref name="Danforthetal2006">{{cite journal |author1=Danforth, B. N. |author2=Sipes, S. |author3=Fang, J. |author4=Brady, S. G. |title=The history of early bee diversification based on five genes plus morphology |journal=PNAS |volume=103 |issue=41 |pages=15118–15123 |date=October 2006 |pmid=17015826 |pmc=1586180 |doi=10.1073/pnas.0604033103|bibcode=2006PNAS..10315118D |doi-access=free }}</ref><ref name=Michener2000/><ref>{{cite journal |last1=Almeida |first1=Eduardo A.B. |last2=Bossert |first2=Silas |last3=Danforth |first3=Bryan N. |last4=Porto |first4=Diego S. |last5=Freitas |first5=Felipe V. |last6=Davis |first6=Charles C. |last7=Murray |first7=Elizabeth A. |last8=Blaimer |first8=Bonnie B. |last9=Spasojevic |first9=Tamara |last10=Ströher |first10=Patrícia R. |last11=Orr |first11=Michael C. |last12=Packer |first12=Laurence |last13=Brady |first13=Seán G. |last14=Kuhlmann |first14=Michael |last15=Branstetter |first15=Michael G. |last16=Pie |first16=Marcio R. |title=The evolutionary history of bees in time and space |journal=[[Current Biology]] |date=2023 |volume=33 |issue=16 |pages=3409–3422.e6 |doi=10.1016/j.cub.2023.07.005 |doi-access=free|pmid=37506702 |bibcode=2023CBio...33E3409A }}</ref> Some species{{snd}}including [[honey bee]]s, [[bumblebee]]s, and [[stingless bee]]s{{snd}}live [[eusociality|socially]] in colonies while most species (>90%){{snd}}including [[mason bee]]s, [[carpenter bee]]s, [[Megachile|leafcutter bees]], and [[Halictidae|sweat bees]]{{snd}}are solitary.
Bees are found on every continent except [[Antarctica]], in every habitat on the planet that contains insect-pollinated [[flowering plant]]s. The most common bees in the [[Northern Hemisphere]] are the [[Halictidae]], or sweat bees, but they are small and often mistaken for wasps or flies. Bees range in size from tiny stingless bee species, whose workers are less than {{convert|2|mm|inch|2}} long,<ref name=":0">{{Cite book|publisher = Springer New York|date = 2020|isbn = 978-3-030-60089-1|first = Christoph|last = Grüter|doi = 10.1007/978-3-030-60090-7|title = Stingless Bees: Their Behaviour, Ecology and Evolution|series = Fascinating Life Sciences|s2cid = 227250633|url = https://link.springer.com/book/10.1007%2F978-3-030-60090-7#toc|url-access = limited}}</ref> to the leafcutter bee ''[[Megachile pluto]]'', the largest species of bee, whose females can attain a length of {{convert|39|mm|inch|2}}.
Bees feed on [[nectar]] and [[pollen]], the former primarily as an energy source and the latter primarily for [[protein]] and other nutrients. Most pollen is used as food for their [[larva]]e. Vertebrate predators of bees include [[primates]] and birds such as [[bee-eater]]s; insect predators include [[beewolf|beewolves]] and [[dragonfly|dragonflies]].
Bee [[Pollination management|pollination]] is important both ecologically and [[List of crop plants pollinated by bees|commercially]], and the decline in wild bees has increased the value of pollination by commercially managed hives of honey bees. The analysis of 353 wild bee and hoverfly species across Britain from 1980 to 2013 found the insects have been lost from a quarter of the places they inhabited in 1980.<ref>{{cite news |url=https://www.theguardian.com/environment/2019/mar/26/widespread-losses-of-pollinating-insects-revealed-across-britain |title=Widespread losses of pollinating insects revealed across Britain |newspaper=[[The Guardian]] |date=26 March 2019}}</ref>
Human [[beekeeping]] or apiculture ([[meliponiculture]] for stingless bees) has been practised for millennia, since at least the times of [[Ancient Egypt]] and [[Ancient Greece]]. Bees have appeared in mythology and folklore, through all phases of art and literature from ancient times to the present day, although primarily focused in the [[Northern Hemisphere]] where beekeeping is far more common. In [[Mesoamerica]], the [[Mayans]] have practiced large-scale intensive meliponiculture since pre-Columbian times.<ref name=":0" />
== Evolution ==
The immediate ancestors of bees were [[Aculeata|stinging wasp]]s in the family [[Crabronidae]], which were [[Predation|predators]] of other insects. The switch from insect prey to pollen may have resulted from the consumption of prey insects which were flower visitors and were partially covered with pollen when they were fed to the wasp larvae. This same [[evolution]]ary scenario may have occurred within the [[vespoidea|vespoid]] wasps, where the [[pollen wasp]]s evolved from predatory ancestors.
Based on phylogenetic analysis, bees are thought to have originated during the [[Early Cretaceous]] (about 124 million years ago) on the supercontinent of [[Gondwana|West Gondwana]], just prior to its breakup into [[South America]] and [[Africa]]. The supercontinent is thought to have been a largely [[Deserts and xeric shrublands|xeric]] environment at this time; modern bee diversity hotspots are also in xeric and seasonal temperate environments, suggesting strong [[Phylogenetic niche conservatism|niche conservatism]] among bees ever since their origins.<ref name=":1">{{Cite journal |last1=Almeida |first1=Eduardo A. B. |last2=Bossert |first2=Silas |last3=Danforth |first3=Bryan N. |last4=Porto |first4=Diego S. |last5=Freitas |first5=Felipe V. |last6=Davis |first6=Charles C. |last7=Murray |first7=Elizabeth A. |last8=Blaimer |first8=Bonnie B. |last9=Spasojevic |first9=Tamara |last10=Ströher |first10=Patrícia R. |last11=Orr |first11=Michael C. |last12=Packer |first12=Laurence |last13=Brady |first13=Seán G. |last14=Kuhlmann |first14=Michael |last15=Branstetter |first15=Michael G. |date=2023-08-21 |title=The evolutionary history of bees in time and space |journal=Current Biology |volume=33 |issue=16 |pages=3409–3422.e6 |doi=10.1016/j.cub.2023.07.005 |issn=0960-9822|doi-access=free |pmid=37506702 |bibcode=2023CBio...33E3409A }}</ref>
Genomic analysis indicates that despite only appearing much later in the fossil record, all modern bee families had already diverged from one another by the end of the Cretaceous. The [[Melittidae]], [[Apidae]], and [[Megachilidae]] had already evolved on the supercontinent prior to its fragmentation. Further divergences were facilitated by West Gondwana's breakup around 100 million years ago, leading to a deep Africa-South America split within both the Apidae and Megachilidae, the isolation of the Melittidae in Africa, and the origins of the [[Colletidae]], [[Andrenidae]] and [[Halictidae]] in South America. The rapid radiation of the South American bee families is thought to have followed the concurrent radiation of [[flowering plant]]s in the same region. Later in the Cretaceous (80 million years ago), colletid bees colonized [[Australia]] from [[South America]] (with an offshoot lineage evolving into the [[Stenotritidae]]), and by the end of the Cretaceous, South American bees had also colonized North America.<ref name=":1" /> The North American fossil taxon ''[[Cretotrigona]]'' belongs to a group that is no longer found in North America, suggesting that many bee lineages went extinct during the [[Cretaceous–Paleogene extinction event|Cretaceous-Paleogene extinction event]].<ref name=":1" />
Following the K-Pg extinction, surviving bee lineages continued to spread into the Northern Hemisphere, colonizing [[Europe]] from Africa by the [[Paleocene]], and then spreading east to [[Asia]]. This was facilitated by the warming climate around the same time, allowing bees to move to higher latitudes following the spread of tropical and subtropical habitats. By the [[Eocene]] (~45 mya) there was already considerable diversity among eusocial bee lineages.<ref name="pnas3054932">{{cite journal |last=Engel |first=Michael S. |year=2001 |title=Monophyly and Extensive Extinction of Advanced Eusocial Bees: Insights from an Unexpected Eocene Diversity |journal=PNAS |publisher=National Academy of Sciences |volume=98 |issue=4 |pages=1661–1664 |bibcode=2001PNAS...98.1661E |doi=10.1073/pnas.041600198 |jstor=3054932 |pmc=29313 |pmid=11172007 |doi-access=free}}</ref>{{efn|[[Triassic]] nests in a petrified forest in Arizona, implying that bees evolved much earlier,<ref name=Buchmann2012/> are now thought to be beetle borings.<ref>{{cite journal |last1=Lucas |first1=Spencer G. |last2=Minter |first2=Nicholas J. |last3=Hunt |first3=Adrian P. |title=Re-evaluation of alleged bees' nests from the Upper Triassic of Arizona |journal=Palaeogeography, Palaeoclimatology, Palaeoecology |date=February 2010 |volume=286 |issue=3–4| pages=194–201 |doi=10.1016/j.palaeo.2010.01.010|bibcode=2010PPP...286..194L }}</ref>}} A second extinction event among bees is thought to have occurred due to rapid climatic cooling around the [[Eocene–Oligocene extinction event|Eocene-Oligocene boundary]], leading to the extinction of some bee lineages such as the tribe [[Melikertini]]. Over the Paleogene and [[Neogene]], different bee lineages continued to spread all over the world, and the shifting habitats and connectedness of continents led to the isolation and evolution of many new bee tribes.<ref name=":1" />
=== Fossils ===
The oldest non-compression bee fossil is ''[[Cretotrigona prisca]]'', a [[Apinae|corbiculate bee]] of [[Late Cretaceous]] age (~70 mya) found in [[New Jersey amber]].<ref>{{cite journal |author1=Cardinal, Sophie |author2=Danforth, Bryan N. |year=2011 |title=The Antiquity and Evolutionary History of Social Behavior in Bees |journal=PLOS ONE |volume=6 |issue=6 |pages=e21086 |bibcode=2011PLoSO...621086C |doi=10.1371/journal.pone.0021086 |pmc=3113908 |pmid=21695157 |doi-access=free}}</ref> A fossil from the early Cretaceous (~100 mya), ''[[Melittosphex burmensis]]'', was initially considered "an extinct lineage of pollen-collecting Apoidea [[Cladistics|sister]] to the modern bees",<ref name="Poinar">{{cite journal |author1=Poinar, G. O. |author2=Danforth, B. N. |year=2006 |title=A fossil bee from Early Cretaceous Burmese amber |url=http://fossilinsects.net/pdfs/Poinar_Danforth_2006_MelittosphexBurmese.pdf |url-status=dead |journal=Science |volume=314 |issue=5799 |page=614 |doi=10.1126/science.1134103 |pmid=17068254 |s2cid=28047407 |archive-url=https://web.archive.org/web/20121204122518/http://fossilinsects.net/pdfs/Poinar_Danforth_2006_MelittosphexBurmese.pdf |archive-date=4 December 2012}}</ref> but subsequent research has rejected the claim that ''Melittosphex'' is a bee, or even a member of the superfamily [[Apoidea]] to which bees belong, instead treating the lineage as ''[[incertae sedis]]'' within the [[Aculeata]].<ref name="Allommation">{{Cite journal |last1=Rosa |first1=B. B. |last2=Melo |first2=G. A. R. |year=2021 |title=Apoid wasps (Hymenoptera: Apoidea) from mid-Cretaceous amber of northern Myanmar |journal=Cretaceous Research |volume=122 |pages=Article 104770 |bibcode=2021CrRes.12204770R |doi=10.1016/j.cretres.2021.104770 |issn=0195-6671 |s2cid=234071940}}</ref>
The [[Allodapini]] (within the Apidae) appeared around 53 Mya.<ref name="danforth2012">{{cite journal |last1=Danforth |first1=Bryan |last2=Cardinal |first2=Sophie |last3=Praz |first3=Christophe |last4=Almeida |first4=Eduardo |last5=Michez |first5=Denis |s2cid=28274420 |title=The Impact of Molecular Data on Our Understanding of Bee Phylogeny and Evolution |journal=Annual Review of Entomology |date=28 August 2012 |volume= 58 |pages=57–78 |doi=10.1146/annurev-ento-120811-153633 |pmid=22934982 }}</ref>
The Colletidae appear as fossils only from the late [[Oligocene]] (~25 Mya) to early [[Miocene]].<ref>{{cite journal |last1=Almeida |first1=Eduardo A. B. |last2=Pie |first2=Marcio R. |last3=Brady |first3=Sean G. |last4=Danforth |first4=Bryan N. |title=Biogeography and diversification of colletid bees (Hymenoptera: Colletidae): emerging patterns from the southern end of the world |journal=Journal of Biogeography |year=2012 |volume=39 |issue=3 |pages=526–544 |url=http://entomology.si.edu/staffpages/Brady/2012_AlmeidaPieBradyDanforth_jBiogeog.pdf |doi=10.1111/j.1365-2699.2011.02624.x |bibcode=2012JBiog..39..526A |s2cid=34626231 |url-status=live |archive-url=https://web.archive.org/web/20130921025319/http://entomology.si.edu/StaffPages/Brady/2012_AlmeidaPieBradyDanforth_jBiogeog.pdf |archive-date=21 September 2013}}</ref>
The Melittidae are known from ''Palaeomacropis eocenicus'' in the [[Ypresian|Early Eocene]].<ref>{{cite journal |last1=Michez |first1=Denis |last2=Nel |first2=Andre |last3=Menier |first3=Jean-Jacques |last4=Rasmont |first4=Pierre |title=The oldest fossil of a melittid bee (Hymenoptera: Apiformes) from the early Eocene of Oise (France) |journal=Zoological Journal of the Linnean Society |year=2007 |volume=150 |issue=4 |pages=701–709 |url=http://www.atlashymenoptera.net/biblio/194_Michez_et_al_2007_Oldest_Melittid_from_Oise_Palaeomacropis.pdf |doi=10.1111/j.1096-3642.2007.00307.x |url-status=live |archive-url=https://web.archive.org/web/20150923175837/http://www.atlashymenoptera.net/biblio/194_Michez_et_al_2007_Oldest_Melittid_from_Oise_Palaeomacropis.pdf |archive-date=23 September 2015|doi-access=free }}</ref>
The Megachilidae are known from trace fossils (characteristic leaf cuttings) from the [[Eocene|Middle Eocene]].<ref>{{cite journal |last1=Sarzetti |first1=Laura C. |last2=Lanandeira |first2=Conrad C. |last3=Genise |first3=Jorge F. |title=A Leafcutter Bee Trace Fossil from the Middle Eocene of Patagonia, Argentina, and a Review of Megachilid (Hymenoptera) Ichnology |journal=Palaeontology |year=2008 |volume=51 |issue=4 |pages=933–994 |url=https://www.researchgate.net/publication/229562255 |doi=10.1111/j.1475-4983.2008.00787.x |bibcode=2008Palgy..51..933S |url-status=live |archive-url=https://web.archive.org/web/20150924123512/http://www.researchgate.net/profile/Conrad_Labandeira/publication/229562255_A_LEAFCUTTER_BEE_TRACE_FOSSIL_FROM_THE_MIDDLE_EOCENE_OF_PATAGONIA_ARGENTINA_AND_A_REVIEW_OF_MEGACHILID_%28HYMENOPTERA%29_ICHNOLOGY/links/53fe0d4b0cf23bb019bd1b59.pdf |archive-date=24 September 2015|doi-access=free |hdl=11336/100644 |hdl-access=free }}</ref>
The Andrenidae are known from the Eocene-Oligocene boundary, around 34 Mya, of the Florissant shale.<ref>{{cite journal |last1=Dewulf |first1=Alexandre |last2=De Meulemeester |first2=Thibaut |last3=Dehon |first3=Manuel |last4=Engel |first4=Michael S. |last5=Michez |first5=Denis |title=A new interpretation of the bee fossil Melitta willardi Cockerell (Hymenoptera, Melittidae) based on geometric morphometrics of the wing |journal=ZooKeys |year=2014 |issue=389 |pages=35–48 |doi=10.3897/zookeys.389.7076 |pmid=24715773 |pmc=3974431|doi-access=free |bibcode=2014ZooK..389...35D }}</ref>
The Halictidae first appear in the Early Eocene<ref>{{cite journal | last1=Engel | first1=M.S. | last2=Archibald | first2=S.B. | year=2003 | title=An Early Eocene bee (Hymenoptera: Halictidae) from Quilchena, British Columbia | doi=10.4039/n02-030| journal=The Canadian Entomologist | volume=135 | issue=1| pages=63–69| hdl=1808/16473 | s2cid=54053341 | url=https://kuscholarworks.ku.edu/bitstream/1808/16473/1/Engel_CE_135%281%2963.pdf |archive-url=https://web.archive.org/web/20170812132942/https://kuscholarworks.ku.edu/bitstream/1808/16473/1/Engel_CE_135%281%2963.pdf |archive-date=2017-08-12 |url-status=live | hdl-access=free }}</ref> with species<ref name="Engel1995">{{cite journal |last1=Engel |first1=M.S. |year=1995 |title=''Neocorynura electra'', a New Fossil Bee Species from Dominican Amber (Hymenoptera:Halictidae) |journal= Journal of the New York Entomological Society |volume=103 |pages=317–323 |jstor=25010174 |issue=3}}</ref><ref name="Engel2000">{{cite journal |last1=Engel |first1=M.S. |year=2000 |title=Classification of the bee tribe Augochlorini (Hymenoptera, Halictidae) |journal=Bulletin of the American Museum of Natural History |volume=250 |pages=1 |url=http://digitallibrary.amnh.org/dspace/bitstream/2246/1598/1/B250.pdf |url-status=live |archive-url=https://web.archive.org/web/20110110015144/http://digitallibrary.amnh.org/dspace/bitstream/2246/1598/1/B250.pdf |archive-date=10 January 2011|doi=10.1206/0003-0090(2000)250<0001:COTBTA>2.0.CO;2 |hdl=2246/1598 |s2cid=85810077 }}</ref> found in amber. The Stenotritidae are known from fossil brood cells of [[Pleistocene]] age.<ref>{{cite journal | last1=Houston | first1=T.F. | year=1987 | title=Fossil brood cells of stenotritid bees (Hymenoptera: Apoidea) from the Pleistocene of South Australia | url=http://eurekamag.com/research/001/840/001840568.php | journal=Transactions of the Royal Society of South Australia | volume=1111–2 | pages=93–97 | url-status=live | archive-url=https://web.archive.org/web/20150701102937/http://eurekamag.com/research/001/840/001840568.php | archive-date=1 July 2015}}</ref>
===Coevolution===
[[File:Amegilla on long tube of Acanthus ilicifolius flower.jpg|thumb|Long-tongued bees and long-tubed flowers [[Coevolution|coevolved]], like this ''[[Amegilla ]]'' species (Apidae) on ''[[Acanthus ilicifolius]]''.]]
{{further|Coevolution}}
The earliest animal-pollinated flowers were shallow, cup-shaped blooms [[pollination|pollinated]] by insects such as [[beetle]]s, so the [[Pollination syndrome|syndrome of insect pollination]] was well established before the first appearance of bees. The novelty is that bees are specialized as pollination agents, with behavioral and physical modifications that specifically enhance pollination, and are the most efficient pollinating insects. In a process of [[coevolution]], flowers developed floral rewards<ref name="Armbruster ch. 3 Evol. Plant-pollinator relationships">{{cite book |author=Armbruster, W. Scott |author-link=Evolution and ecological implications of 'specialized' pollinator rewards |editor1-last=Patiny |editor1-first=Sébastien |title=Evolution of Plant-Pollinator Relationships |date=2012 |publisher=Cambridge University Press |pages=45–67 |chapter=3}}</ref> such as [[nectar]] and longer tubes, and bees developed longer tongues to extract the nectar.<ref name=Michener1974/> Bees also developed structures known as [[scopa (biology)|scopal hairs]] and [[pollen baskets]] to collect and carry pollen. The location and type differ among and between groups of bees. Most species have scopal hairs on their hind legs or on the underside of their abdomens. Some species in the family Apidae have [[pollen baskets]] on their hind legs, while very few lack these and instead collect pollen in their crops.<ref name=Michener2000>{{cite book |title=The Bees of the World |date=2000 |publisher=Johns Hopkins University Press |isbn=0-8018--6133-0 |pages=19–25 |last1=Michener |first1=Charles D.}}</ref> The appearance of these structures drove the [[adaptive radiation]] of the [[Flowering plant|angiosperms]], and, in turn, bees themselves.<ref name=Buchmann2012>{{cite book |author1=Buchmann, Stephen L. |author2=Nabhan, Gary Paul |title=The Forgotten Pollinators |url=https://books.google.com/books?id=YWTZs5fSqb8C&pg=PA41 |year=2012 |publisher=Island Press |isbn=978-1-59726-908-7 |pages=41–42 |url-status=live |archive-url=https://web.archive.org/web/20160527145146/https://books.google.com/books?id=YWTZs5fSqb8C&pg=PA41 |archive-date=27 May 2016}}</ref> Bees [[Coevolution|coevolved]] not only with flowers but it is believed that some species coevolved with mites. Some provide tufts of hairs called [[acarinaria]] that appear to provide lodgings for mites; in return, it is believed that mites eat fungi that attack pollen, so the relationship in this case may be [[mutualism (biology)|mutualistic]].<ref name=":5">{{cite journal |last1=Biani |first1=Natalia B. |last2=Mueller |first2=Ulrich G.|last3=Wcislo |first3=William T. |title=Cleaner Mites: Sanitary Mutualism in the Miniature Ecosystem of Neotropical Bee Nests |journal=The American Naturalist |date=June 2009 |volume=173 |issue=6 |pages=841–847 |doi=10.1086/598497 |pmid=19371167|url=https://repositories.lib.utexas.edu/bitstream/2152/31261/1/CleanerMites.pdf |archive-url=https://web.archive.org/web/20180328155201/https://repositories.lib.utexas.edu/bitstream/2152/31261/1/CleanerMites.pdf |archive-date=2018-03-28 |url-status=live |hdl=2152/31261 |s2cid=4845087 |hdl-access=free }}</ref><ref name="Pavel B 2007">{{cite journal|last1=Klimov |first1=Pavel B. |last2=OConnor |first2=Barry M. |last3=Knowles |first3=L. Lacey |title=Museum Specimens And Phylogenies Elucidate Ecology's Role in Coevolutionary Associations Between Mites And Their Bee Hosts |journal=Evolution |date=June 2007 |volume=61 |issue=6 |pages=1368–1379 |doi=10.1111/j.1558-5646.2007.00119.x |pmid=17542846|url=https://deepblue.lib.umich.edu/bitstream/2027.42/74970/1/j.1558-5646.2007.00119.x.pdf |archive-url=https://web.archive.org/web/20190504194936/https://deepblue.lib.umich.edu/bitstream/2027.42/74970/1/j.1558-5646.2007.00119.x.pdf |archive-date=2019-05-04 |url-status=live |hdl=2027.42/74970 |s2cid=32318137 |doi-access=free }}</ref>
===Phylogeny===
====External====
Molecular phylogeny was used by Debevic ''et al'', 2012, to demonstrate that the bees ([[Anthophila]]) arose from deep within the [[Crabronidae]] ''[[sensu lato]]'', which was thus rendered [[paraphyly|paraphyletic]]. In their study, the placement of the monogeneric [[Heterogyna|Heterogynaidae]] was uncertain. The small family [[Mellinidae]] was not included in this analysis.<ref>{{cite journal |last1=Debevec |first1=Andrew H. |last2=Cardinal |first2=Sophie |last3=Danforth |first3=Bryan N. |title=Identifying the sister group to the bees: a molecular phylogeny of Aculeata with an emphasis on the superfamily Apoidea |journal=Zoologica Scripta |date=2012 |volume=41 |issue=5 |pages=527–535 |doi=10.1111/j.1463-6409.2012.00549.x |s2cid=33533180 |url=http://www.danforthlab.entomology.cornell.edu/files/all/debevec_etal_2012.pdf |url-status=live |archive-url=https://web.archive.org/web/20150923212548/http://www.danforthlab.entomology.cornell.edu/files/all/debevec_etal_2012.pdf |archive-date=23 September 2015}}</ref>
Further studies by Sann ''et al.'', 2018, elevated the subfamilies (plus one tribe and one subtribe) of [[Crabronidae]] ''sensu lato'' to family status. They also recovered the placement of ''Heterogyna'' within [[Nyssonini]] and sunk [[Heterogynaidae]]. The newly erected family, [[Ammoplanidae]], formerly a subtribe of [[Pemphredonidae|Pemphredoninae]], was recovered as the most sister family to bees.<ref name=Sann2018>{{cite journal |first1=Manuela |last1=Sann  |first2=Oliver |last2=Niehuis |first3=Ralph S. |last3=Peters |first4=Christoph |last4=Mayer |first5=Alexey |last5=Kozlov |first6=Lars |last6=Podsiadlowski |first7=Sarah |last7=Bank |first8=Karen |last8=Meusemann |first9=Bernhard |last9=Misof |first10=Christoph |last10=Bleidorn |first11=Michael |last11=Ohl |date=2018 |title=Phylogenomic analysis of Apoidea sheds new light on the sister group of bees. |journal=BMC Evolutionary Biology |volume= 18 |issue=71 |page=71 |doi=10.1186/s12862-018-1155-8 |doi-access=free|pmid=29776336 |pmc=5960199 |bibcode=2018BMCEE..18...71S }}</ref>
{{clade| style=font-size:85%;line-height:85%
|label1=Apoidea
|1={{clade
|1=[[Ampulicidae]] (Cockroach wasps) [[File:Emerald Cockroach Wasp.JPG|70px]]
|2={{clade
|1=[[Astatidae]] [[File:Astata boopsDSC05135.JPG|70px]]
|2={{clade
|1={{clade
|1=[[Bembicinae|Bembicidae]] [[File:Bembix sp2.jpg|70px]]
|2={{clade
|1=[[Sphecidae]] (''[[sensu stricto]]'') [[File:Sceliphron spirifex TZ edit1.jpg|70px]]
|2={{clade
|1={{clade
|1=[[Crabroninae|Crabronidae]] (''[[sensu stricto]]'') [[File:Ectemnius.lapidarius.-.lindsey.jpg|70px]]
|2=[[Mellinidae]] [[File:Mellinus arvensis - Flickr - gailhampshire (1).jpg|70px]]
}} }} }} }}
|2={{clade
|1={{clade
|1=[[Pemphredonidae]] [[File:Pemphredon ? (Crabronidae) - Zwerggrabwespe (11462528464).jpg|70px]]
|2=[[Philanthidae]] [[File:P. gibbosus57306787w.jpg|70px]]}}
|2={{clade
|1=[[Psenidae]] [[File:Aphid Wasp - Flickr - treegrow (2).jpg|70px]]
|2={{clade
|1={{clade
|1=[[Ammoplanidae]]
|2='''Anthophila (bees)''' [[File:Abeille butineuse et son pollen.JPG|70px]]
}} }} }} }} }}
}}
}}
}}
====Internal====
This cladogram of the bee families is based on Hedtke et al., 2013, which places the former families Dasypodaidae and Meganomiidae as subfamilies inside the Melittidae.<ref name=Hedtke2013>{{cite journal|last1=Hedtke|first1=Shannon M.|last2=Patiny|first2=Sébastien|last3=Danforth|first3=Bryan M.|title=The bee tree of life: a supermatrix approach to apoid phylogeny and biogeography|journal=BMC Evolutionary Biology|date=2013|volume=13|issue=138|pages=138|doi=10.1186/1471-2148-13-138|pmid=23822725|pmc=3706286 |doi-access=free |bibcode=2013BMCEE..13..138H }}</ref> English names, where available, are given in parentheses.
{{clade| style=font-size:85%;line-height:85%
|label1=[[Anthophila]] (bees)
|1={{clade
|1={{clade
|1=[[Melittidae]] (inc. [[Dasypodainae]], [[Meganomiinae]]) at least 50 Mya [[File:Macropis sp 01.jpg|70px]]
|2={{clade
|label1={{nowrap|long-tongued bees}}
|1={{clade
|1=[[Apidae]] (inc. honeybees, cuckoo bees, carpenter bees) ≈87 Mya [[File:Apis mellifera flying2.jpg|70px]]
|2=[[Megachilidae]] (mason, leafcutter bees) ≈50 Mya [[File:Leafcutter bee (Megachile sp.) collecting leaves (7519316920).jpg|70px]]
}}
|label2={{nowrap|short-tongued bees}}
|2={{clade
|1=[[Andrenidae]] (mining bees) ≈34 Mya [[File:Thomas Bresson - Hyménoptère sur une fleur de pissenlit (by).jpg|70px]]
|2={{clade
|1=[[Halictidae]] (sweat bees) ≈50 Mya [[File:Iridescent.green.sweat.bee1.jpg|70px]]
|2={{clade
|1=[[Colletidae]] (plasterer bees) ≈25 Mya [[File:Colletes cunicularius m1.JPG|70px]]
|2=[[Stenotritidae]] (large Australian bees) ≈2 Mya [[File:Stenotritus pubescens, f, side, australia 2014-07-05-12.18.33 ZS PMax.jpg|70px]]
}}
}}
}}
}}
}}
}}
}}
{{see also|Characteristics of common wasps and bees}}
==Characteristics==
[[File:European Honeybee (Apis mellifera) lapping mouthparts, showing labium and maxillae..jpg|thumb|upright|The lapping [[insect mouthparts|mouthparts]] of a honey bee, showing labium and maxillae]]
Bees differ from closely related groups such as wasps by having branched or plume-like [[seta]]e (hairs), combs on the forelimbs for cleaning their antennae, small anatomical differences in limb structure, and the venation of the hind wings; and in females, by having the seventh dorsal abdominal plate divided into two half-plates.<ref name=Grimaldi>{{cite book |author1=Grimaldi, David |author2=Engel, Michael S. |title=Evolution of the Insects |url=https://books.google.com/books?id=Ql6Jl6wKb88C&pg=PA454 |year=2005 |publisher=Cambridge University Press |isbn=978-0-521-82149-0 |page=454 |url-status=live |archive-url=https://web.archive.org/web/20180328155201/https://books.google.com/books?id=Ql6Jl6wKb88C&pg=PA454 |archive-date=28 March 2018}}</ref>
Bees have the following characteristics:<ref name=Extension/>
* A pair of large [[compound eyes]] which cover much of the surface of the head. Between and above these are three small simple eyes ([[ocelli]]) which provide information on light intensity.<ref name=Extension/>
* The [[Antenna (biology)|antennae]] usually have 13 segments in males and 12 in females, and are [[Insect morphology#Antennae|geniculate]], having an elbow joint part way along. They house large numbers of sense organs that can detect touch (mechanoreceptors), smell and taste; and small, hairlike mechanoreceptors that can detect air movement so as to "hear" sounds.<ref name=Extension/>
* The [[insect mouthparts|mouthparts]] are adapted for both chewing and sucking by having both a pair of [[Mandible (insect mouthpart)|mandibles]] and a long [[proboscis]] for sucking up nectar.<ref name=Extension>{{cite web |url=http://www.extension.org/pages/21754/anatomy-of-the-honey-bee |title=Anatomy of the Honey Bee |date=19 June 2014 |publisher=Extension |access-date=30 June 2015 |url-status=dead |archive-url=https://web.archive.org/web/20150701141426/http://www.extension.org/pages/21754/anatomy-of-the-honey-bee |archive-date=1 July 2015}}</ref>
* The thorax has three segments, each with a pair of robust legs, and a pair of membranous wings on the hind two segments. The front legs of corbiculate bees bear combs for cleaning the antennae, and in many species the hind legs bear pollen baskets, flattened sections with incurving hairs to secure the collected pollen. The wings are synchronised in flight, and the somewhat smaller hind wings connect to the forewings by a row of hooks along their margin which connect to a groove in the forewing.
* The abdomen has nine segments, the hindermost three being modified into the sting.<ref name=Extension/>
[[File:Carpenter bee head and compound eyes.jpg|thumb|Head-on view of a male [[carpenter bee]], showing antennae, three [[ocelli]], [[compound eye]]s, and mouthparts]]
The largest species of bee is thought to be Wallace's giant bee ''[[Megachile pluto]]'', whose females can attain a length of {{convert|39|mm|inch|2}}.<ref>{{ cite journal | author=Messer, A. C. | title=''Chalicodoma pluto'': The World's Largest Bee Rediscovered Living Communally in Termite Nests (Hymenoptera: Megachilidae) | journal=Journal of the Kansas Entomological Society | year=1984 | volume=57 | issue=1 | pages=165–168 | jstor=25084498 }}</ref> The smallest species may be dwarf stingless bees in the tribe [[Meliponini]] whose workers are less than {{convert|2|mm|inch|2}} in length.<ref>{{cite journal |url=http://eprints.lib.hokudai.ac.jp/dspace/bitstream/2115/27567/1/19(2)_P361-421.pdf |title=Oviposition Behavior of Two Dwarf Stingless Bees, ''Hypotrigona'' (''Leurotrigona'') ''muelleri'' and ''H.'' (''Trigonisca'') ''duckei'', with Notes on the Temporal Articulation of Oviposition Process in Stingless Bees |author1=Sakagami, Shôichi F. |author2=Zucchi, Ronaldo |journal=Journal of the Faculty of Science Hokkaido University Series Vi. Zoology |volume=19 |issue=2 |pages=361–421 |year=1974 |url-status=live |archive-url=https://web.archive.org/web/20160304063836/http://eprints.lib.hokudai.ac.jp/dspace/bitstream/2115/27567/1/19(2)_P361-421.pdf |archive-date=4 March 2016}}</ref>
==Sociality==
===Haplodiploid breeding system===
{{further|Haplodiploidy}}
[[File:Wasp attack.jpg|thumb|left|upright=1.3<!--size for low image-->|Willing to die for their sisters: worker honey bees killed defending their hive against [[yellowjacket]]s, along with a dead yellowjacket. Such [[altruism|altruistic]] behaviour may be favoured by the [[haplodiploid]] [[Sex-determination system|sex determination]] system of bees.]]
According to [[inclusive fitness]] theory, organisms can gain fitness not just through increasing their own reproductive output, but also that of close relatives. In evolutionary terms, individuals should help relatives when ''Cost < Relatedness * Benefit''. The requirements for eusociality are more easily fulfilled by [[haplodiploid]] species such as bees because of their unusual relatedness structure.<ref name=Hughes2008>{{Cite journal | last1=Hughes | first1=W. O. H. | last2=Oldroyd | first2=B. P. | last3=Beekman | first3=M. | last4=Ratnieks | first4=F. L. W. | title=Ancestral Monogamy Shows Kin Selection is Key to the Evolution of Eusociality | doi=10.1126/science.1156108 | journal=Science | volume=320 | issue=5880 | pages=1213–1216 | year=2008 | pmid= 18511689| bibcode=2008Sci...320.1213H | s2cid=20388889 }}</ref>
In haplodiploid species, females develop from fertilized eggs and males from unfertilized eggs. Because a male is [[haploid]] (has only one copy of each gene), his daughters (which are [[diploid]], with two copies of each gene) share 100% of his genes and 50% of their mother's. Therefore, they share 75% of their genes with each other. This mechanism of sex determination gives rise to what [[W. D. Hamilton]] termed "supersisters", more closely related to their sisters than they would be to their own offspring.<ref name="Hamilton1964II">{{cite journal | last=Hamilton | first=W. D. | title=The Genetical Evolution of Social Behaviour II | journal=Journal of Theoretical Biology | date=20 March 1964 | volume=7 | issue=1 | pages=17–52 | doi=10.1016/0022-5193(64)90039-6 | pmid=5875340| bibcode=1964JThBi...7...17H }}</ref> Workers often do not reproduce, but they can pass on more of their genes by helping to raise their sisters (as queens) than they would by having their own offspring (each of which would only have 50% of their genes), assuming they would produce similar numbers. This unusual situation has been proposed as an explanation of the multiple (at least nine) evolutions of eusociality within [[Hymenoptera]].<ref name="Hughes" /><ref>{{cite book |last1=Gullan |first1=P. J. |last2=Cranston |first2=P. S. |title=The Insects: An Outline of Entomology |date=2014 |publisher=Wiley Blackwell |isbn=978-1-118-84615-5 |edition=5th |pages=328, 348–350}}</ref>
Haplodiploidy is neither necessary nor sufficient for eusociality. Some eusocial species such as [[termites]] are not haplodiploid. Conversely, all bees are haplodiploid but not all are eusocial, and among eusocial species many queens mate with multiple males, creating half-sisters that share only 25% of each other's genes.<ref name="NTW">{{cite journal | last=Nowak | first=Martin | author2=Tarnita, Corina | author3=Wilson, E.O. | title=The evolution of eusociality | journal=Nature | year=2010 | volume=466 | pmid=20740005 | issue=7310 | pages=1057–1062 | doi=10.1038/nature09205 | pmc=3279739| bibcode=2010Natur.466.1057N }}</ref> But, monogamy (queens mating singly) is the ancestral state for all eusocial species so far investigated, so it is likely that haplodiploidy contributed to the evolution of eusociality in bees.<ref name="Hughes">{{cite journal | author1=Hughes, William O. H. | author2=Oldroyd, Benjamin P. | author3=Beekman, Madeleine | author4=Ratnieks, Francis L. W. | title=Ancestral Monogamy Shows Kin Selection Is Key to the Evolution of Eusociality | journal=[[Science (journal)|Science]] | volume=320 | issue=5880 | pages=1213–1216 | publisher=American Association for the Advancement of Science | date=May 2008 | doi=10.1126/science.1156108 | pmid=18511689| bibcode=2008Sci...320.1213H | s2cid=20388889 }}</ref>
===Eusociality===
[[File:Bee swarm on fallen tree03.jpg|thumb|upright|A [[Western honey bee]] swarm]]
[[File:Wildbienen.jpg|thumb|[[Western honey bee]] nest in the [[Trunk (botany)|trunk]] of a [[Picea abies|spruce]]]]
{{further|Eusociality}}
Bees may be solitary or may live in various types of communities. [[Eusociality]] appears to have originated from at least three independent origins in halictid bees.<ref>{{cite journal |last1=Brady |first1=Seán G. |last2=Sipes |first2=Sedonia |last3=Pearson |first3=Adam |last4=Danforth |first4=Bryan N. |date=2006 |title=Recent and simultaneous origins of eusociality in halictid bees |journal=Proceedings of the Royal Society of London B: Biological Sciences |volume=273 |issue=1594 |pages=1643–1649 |doi=10.1098/rspb.2006.3496 |issn=0962-8452 |pmc=1634925 |pmid=16769636}}</ref> The most advanced of these are species with [[Eusociality|eusocial]] colonies; these are characterised by cooperative brood care and a [[division of labour]] into reproductive and non-reproductive adults, plus overlapping generations.<ref>{{Cite book|title=The Insect Societies|last=Wilson|first=Edward O|publisher=Belknap Press of Harvard University Press|year=1971|location=Cambridge, Mass}}</ref> This division of labour creates specialized groups within eusocial societies which are called [[Eusociality#In insects|castes]]. In some species, groups of cohabiting females may be sisters, and if there is a division of labour within the group, they are considered [[semisocial]]. The group is called eusocial if, in addition, the group consists of a mother (the [[Queen bee|queen]]) and her daughters ([[Worker bee|workers]]). When the castes are purely behavioural alternatives, with no morphological differentiation other than size, the system is considered primitively eusocial, as in many [[paper wasp]]s; when the castes are morphologically discrete, the system is considered highly eusocial.<ref name=Michener1974>{{cite book|author=Michener, Charles Duncan |title=The Social Behavior of the Bees: A Comparative Study |url=https://books.google.com/books?id=aordrL_D-30C&pg=PA78 |year=1974 |publisher=Harvard University Press |isbn=978-0-674-81175-1 |pages=22–78}}</ref>
True honey bees (genus ''[[Apis (genus)|Apis]]'', of which eight species are currently recognized) are highly eusocial, and are among the best known insects. Their colonies are established by [[Swarming (honey bee)|swarms]], consisting of a queen and several thousand workers. There are 29 subspecies of one of these species, ''[[Apis mellifera]]'', native to Europe, the Middle East, and Africa. [[Africanized bee]]s are a hybrid strain of ''A. mellifera'' that escaped from experiments involving crossing European and African subspecies; they are extremely defensive.<ref>{{cite web | last1=Sanford | first1=Malcolm T. | title=The Africanized Honey Bee in the Americas: A Biological Revolution with Human Cultural Implications | url=http://apisenterprises.com/papers_htm/Misc/AHB%20in%20the%20Americas.htm | publisher=Apis Enterprises | access-date=29 March 2015 | year=2006 | url-status=live | archive-url=http://archive.wikiwix.com/cache/20150329231615/http://apisenterprises.com/papers_htm/Misc/AHB%20in%20the%20Americas.htm | archive-date=29 March 2015}}</ref>
[[Stingless bee]]s are also highly [[eusocial]]. They practise [[mass provisioning]], with complex nest architecture and perennial colonies also established via swarming.<ref name=":0" /><ref>{{cite journal | last1=Roubik | first1=D. W. | year=2006 | title=Stingless bee nesting biology | journal=Apidologie | volume=37 | issue=2 | pages=124–143 | doi=10.1051/apido:2006026| url=http://www.apidologie.org/articles/apido/pdf/2006/02/m6034sp.pdf |archive-url=https://ghostarchive.org/archive/20221009/http://www.apidologie.org/articles/apido/pdf/2006/02/m6034sp.pdf |archive-date=9 October 2022 |url-status=live | doi-access=free }}</ref>
[[File:Bumblebee 05.JPG|thumb|left|A [[bumblebee]] carrying pollen in its [[pollen basket]]s (corbiculae)]]
Many bumblebees are eusocial, similar to the eusocial [[Vespidae]] such as [[hornet]]s in that the queen initiates a nest on her own rather than by swarming. Bumblebee colonies typically have from 50 to 200 bees at peak population, which occurs in mid to late summer. Nest architecture is simple, limited by the size of the pre-existing nest cavity, and colonies rarely last more than a year.<ref name="BBCT-nest">{{cite web | title=Bumblebee nests | url=http://bumblebeeconservation.org/about-bees/habitats/bumblebee-nests/ | publisher=Bumblebee Conservation Trust | access-date=26 June 2015 | archive-url=https://web.archive.org/web/20170922230805/https://bumblebeeconservation.org/about-bees/habitats/bumblebee-nests/ | archive-date=22 September 2017}}</ref> In 2011, the [[International Union for Conservation of Nature]] set up the Bumblebee Specialist Group to review the threat status of all bumblebee species worldwide using the [[IUCN Red List]] criteria.<ref>{{cite news | url=http://cmsdata.iucn.org/downloads/bumblebee_sg_proofed.pdf | title=Bumblebee Specialist Group: 2011 Update | publisher=[[IUCN]] | access-date=7 October 2012 | url-status=live | archive-url=https://web.archive.org/web/20121203034020/http://cmsdata.iucn.org/downloads/bumblebee_sg_proofed.pdf | archive-date=3 December 2012}}</ref>
There are many more species of primitively eusocial than highly eusocial bees, but they have been studied less often. Most are in the family [[Halictidae]], or "sweat bees". Colonies are typically small, with a dozen or fewer workers, on average. Queens and workers differ only in size, if at all. Most species have a single season colony cycle, even in the tropics, and only mated females hibernate. A few species have long active seasons and attain colony sizes in the hundreds, such as ''[[Halictus hesperus]]''.<ref>{{cite journal | last1=Brooks | first1=R. W. | last2=Roubik | first2=D. W. | year=1983 | title=A Halictine bee with distinct castes: ''Halictus hesperus'' (Hymenoptera: Halictidae) and its bionomics in Central Panama | journal=Sociobiology | volume=7 | pages=263–282 }}</ref> Some species are eusocial in parts of their range and solitary in others,<ref>{{Cite journal|last1=Eickwort |first1=G. C. |last2=Eickwort |first2=J. M. |last3=Gordon|first3=J. |last4=Eickwort |first4=M. A. |last5=Wcislo |first5=W. T. |title=Solitary behavior in a high-altitude population of the social sweat bee Halictus rubicundus (Hymenoptera: Halictidae) |journal=Behavioral Ecology and Sociobiology |volume=38 |issue=4 |pages=227–233 |doi=10.1007/s002650050236 |year=1996|bibcode=1996BEcoS..38..227E |s2cid=12868253 }}</ref> or have a mix of eusocial and solitary nests in the same population.<ref>{{cite journal | last1=Yanega | first1=D. | year=1993 | title=Environmental effects on male production and social structure in ''Halictus rubicundus'' (Hymenoptera: Halictidae) | journal=Insectes Sociaux | volume=40 | pages=169–180 | doi=10.1007/bf01240705| s2cid=44934383 }}</ref> The [[orchid bee]]s (Apidae) include some primitively eusocial species with similar biology. Some [[Allodapini|allodapine]] bees (Apidae) form primitively eusocial colonies, with [[progressive provisioning]]: a larva's food is supplied gradually as it develops, as is the case in honey bees and some bumblebees.<ref>{{cite book |author=Michener, Charles Duncan |title=The Social Behavior of the Bees: A Comparative Study |url=https://books.google.com/books?id=aordrL_D-30C&pg=PA308 |year=1974 |publisher=Harvard University Press |isbn=978-0-674-81175-1 |page=308 |url-status=live |archive-url=https://web.archive.org/web/20161224104244/https://books.google.com/books?id=aordrL_D-30C&pg=PA308 |archive-date=24 December 2016}}</ref>
===Solitary and communal bees===
[[File:Megachile rotundata.JPG|thumb|A leafcutting bee, ''[[Megachile rotundata]]'', cutting circles from acacia leaves]]
Most other bees, including familiar insects such as [[carpenter bee]]s, [[leafcutter bees]] and [[mason bees]] are solitary in the sense that every female is fertile, and typically inhabits a nest she constructs herself. There is no division of labor so these nests lack queens and ''worker'' bees for these species. Solitary bees typically produce neither honey nor [[beeswax]].
Bees collect pollen to feed their young, and have the necessary adaptations to do this. However, certain wasp species such as [[pollen wasp]]s have similar behaviours, and a few species of bee [[Scavenger|scavenge]] from carcases to feed their offspring.<ref name=Grimaldi/> Solitary bees are important pollinators; they gather pollen to provision their nests with food for their brood. Often it is mixed with nectar to form a paste-like consistency. Some solitary bees have advanced types of pollen-carrying structures on their bodies. Very few species of solitary bee are being cultured for commercial pollination. Most of these species belong to a distinct set of [[genus|genera]] which are commonly known by their nesting behavior or preferences, namely: carpenter bees, [[Halictidae|sweat bees]], mason bees, [[Colletes inaequalis|plasterer bees]], [[squash bee]]s, [[Ceratina|dwarf carpenter bees]], leafcutter bees, [[alkali bee]]s and [[Anthophorini|digger bees]].<ref>{{cite web |url=http://www.beesource.com/resources/usda/management-of-wild-bees/ |author1=Parker, Frank D. |author2=Torchio, Philip F. |title=Management of Wild Bees |publisher=Beesource Beekeeping Community |date=1 October 1980 |access-date=26 June 2015 |url-status=live |archive-url=https://web.archive.org/web/20150626235657/http://www.beesource.com/resources/usda/management-of-wild-bees/ |archive-date=26 June 2015}}</ref>
[[File:Anthidium February 2008-1.jpg|thumb|A solitary bee, ''[[Anthidium florentinum]]'' (family [[Megachilidae]]), visiting ''[[Lantana]]'']]
Most solitary bees are [[fossorial]], digging nests in the ground in a variety of soil textures and conditions, while others create nests in hollow [[Phragmites|reeds]] or twigs, or holes in [[wood]]. The female typically creates a compartment (a "cell") with an egg and some provisions for the resulting larva, then seals it off. A nest may consist of numerous cells. When the nest is in wood, usually the last (those closer to the entrance) contain eggs that will become males. The adult does not provide care for the brood once the egg is laid, and usually dies after making one or more nests. The males typically emerge first and are ready for mating when the females emerge. Solitary bees are very unlikely to sting (only in self-defense, if ever), and some (esp. in the family [[Andrenidae]]) are stingless.<ref>{{cite web |title=Solitary Bees (Hymenoptera) |url=http://www.royensoc.co.uk/insect_info/what/solitary_bees.htm |publisher=Royal Entomological Society |access-date=12 October 2015 |archive-url=https://web.archive.org/web/20170702005458/http://www.royensoc.co.uk/insect_info/what/solitary_bees.htm |archive-date=2 July 2017 |url-status=dead}}</ref><ref>{{cite web |title=Other bees |url=http://bumblebeeconservation.org/about-bees/identification/other-bees/ |publisher=Bumblebee Conservation Trust |access-date=12 October 2015 |url-status=dead |archive-url=https://web.archive.org/web/20150905113303/http://bumblebeeconservation.org/about-bees/identification/other-bees/ |archive-date=5 September 2015}}</ref>
[[File:Osmia cornifrons.5.1.08.w.jpg|thumb|The [[mason bee]] ''[[Osmia cornifrons]]'' nests in a hole in dead wood. [[Insect hotel#Solitary bees and wasps|Bee "hotels"]] are often sold for this purpose.]]
While solitary, females each make individual nests.<ref>{{Cite web|last=Westreich|first=Lila|title=Spring signals female bees to lay the next generation of pollinators|url=http://theconversation.com/spring-signals-female-bees-to-lay-the-next-generation-of-pollinators-134852|access-date=8 October 2020|website=The Conversation|date=May 2020 |language=en}}</ref> Some species, such as the European mason bee ''[[Hoplitis anthocopoides]]'',<ref name=Eickwort>{{cite journal |last1=Eickwort |first1=George C. |title=Gregarious Nesting of the Mason Bee Hoplitis anthocopoides and the Evolution of Parasitism and Sociality Among Megachilid Bees |journal=Evolution |date=1975 |volume=29 |issue=1 |pages=142–150 |doi=10.2307/2407147|jstor=2407147 |pmid=28563288 }}</ref> and the [[Amegilla dawsoni|Dawson's Burrowing bee]], ''Amegilla dawsoni,''<ref>{{Cite journal |title=The Nesting Behavior of Dawson's Burrowing Bee, Amegilla dawsoni (Hymenoptera: Anthophorini), and the Production of Offspring of Different Sizes |journal=Journal of Insect Behavior |date=1 May 1999 |issn=0892-7553 |pages=363–384 |volume=12 |issue=3 |doi=10.1023/A:1020843606530 |first=John |last=Alcock|bibcode=1999JIBeh..12..363A |s2cid=24832889 }}</ref> are gregarious, preferring to make nests near others of the same species, and giving the appearance of being social. Large groups of solitary bee nests are called ''aggregations'', to distinguish them from [[colony (biology)|colonies]]. In some species, multiple females share a common nest, but each makes and provisions her own cells independently. This type of group is called "communal" and is not uncommon. The primary advantage appears to be that a nest entrance is easier to defend from predators and parasites when multiple females use that same entrance regularly.<ref name=Eickwort/>
==Biology==
[[File:Bee morning glory visitors.webm|thumb|thumbtime=26|Various bees visit a [[morning glory]] flower. A [[Mordellidae|Tumbling flower beetle]] remains in the flower with a bee visitor.]]
===Life cycle===
{{further|Honey bee life cycle}}
The life cycle of a bee, be it a solitary or social species, involves the laying of an egg, the development through several moults of a legless [[larva]], a [[pupa]]tion stage during which the insect undergoes [[Holometabolism|complete metamorphosis]], followed by the emergence of a winged adult. The number of eggs laid by a female during her lifetime can vary from eight or less in some solitary bees, to more than a million in highly social species.<ref>[https://books.google.com/books?id=bu_1gmY13FIC&dq=eggs+laid+during+her+lifetime+eight+or+fewer+solitary+million+queens&pg=PA8-IA12 The Bees of the World, Volum 1]</ref> Most solitary bees and bumble bees in temperate climates overwinter as adults or pupae and emerge in spring when increasing numbers of flowering plants come into bloom. The males usually emerge first and search for females with which to mate. Like the other members of Hymenoptera bees are [[Haplodiploidy|haplodiploid]]; the sex of a bee is determined by whether or not the egg is fertilised. After mating, a female stores the sperm, and determines which sex is required at the time each individual egg is laid, fertilised eggs producing female offspring and unfertilised eggs, males. Tropical bees may have several generations in a year and no [[diapause]] stage.<ref name=Roubik1992>{{cite book |author=Roubik, David W. |title=Ecology and Natural History of Tropical Bees |url=https://books.google.com/books?id=ljlaYMeI6noC |year=1992 |publisher=Cambridge University Press |isbn=978-0-521-42909-2 |page=15 |url-status=live |archive-url=https://web.archive.org/web/20160617144437/https://books.google.com/books?id=ljlaYMeI6noC |archive-date=17 June 2016}}</ref><ref>{{cite web|title=The bumblebee lifecycle|url=http://bumblebeeconservation.org/about-bees/lifecycle/|publisher=Bumblebee Conservation Trust|access-date=1 July 2015|url-status=live|archive-url=https://web.archive.org/web/20150629043439/http://bumblebeeconservation.org/about-bees/lifecycle/|archive-date=29 June 2015}}</ref><ref>{{cite web |title=Learning About Honey Bees |url=http://www.scmidstatebeekeepers.org/honeybeelifecycle.htm |publisher=The South Carolina Mid-State Beekeepers Association |access-date=1 July 2015 |url-status=live |archive-url=https://web.archive.org/web/20150701180422/http://www.scmidstatebeekeepers.org/honeybeelifecycle.htm |archive-date=1 July 2015}}</ref><ref>{{cite web |title=Solitary Bees |url=http://www.nationalbeeunit.com/downloadDocument.cfm?id=901 |publisher=National Bee Unit |access-date=1 July 2015 |url-status=live |archive-url=https://web.archive.org/web/20150701202809/http://www.nationalbeeunit.com/downloadDocument.cfm?id=901 |archive-date=1 July 2015}}</ref>
The egg is generally oblong, slightly curved and tapering at one end. Solitary bees, lay each egg in a separate cell with a supply of mixed pollen and nectar next to it. This may be rolled into a pellet or placed in a pile and is known as mass provisioning. Social bee species provision progressively, that is, they feed the larva regularly while it grows. The nest varies from a hole in the ground or in wood, in solitary bees, to a substantial structure with wax combs in bumblebees and honey bees.<ref name=Shuckard>{{cite book |author=Shuckard, William Edward |title=British bees: an introduction to the study of the natural history and economy of the bees indigenous to the British Isles |url=https://archive.org/details/britishbeesanin01shucgoog |year=1866 |publisher=L. Reeve & Co. |pages=[https://archive.org/details/britishbeesanin01shucgoog/page/n38 18]–23}}</ref>
In most species, larvae are whitish grubs, roughly oval and bluntly-pointed at both ends. They have 15 segments and [[Spiracle (arthropods)|spiracle]]s in each segment for breathing. They have no legs but move within the cell, helped by tubercles on their sides. They have short horns on the head, jaws for chewing food and an appendage on either side of the mouth tipped with a bristle. There is a gland under the mouth that secretes a viscous liquid which solidifies into the silk they use to produce a cocoon. The cocoon is semi-transparent and the pupa can be seen through it. Over the course of a few days, the larva undergoes metamorphosis into a winged adult. When ready to emerge, the adult splits its skin dorsally and climbs out of the [[exuvia]]e and breaks out of the cell.<ref name=Shuckard/>
<gallery mode="packed" heights="125px">
File:Apoidea.jpg|Nest of [[Bombus pascuorum|common carder]] [[bumblebee]], wax canopy removed to show winged [[worker (bee)|workers]] and [[pupae]] in irregularly placed wax cells
File:Carpenter Bee Galleries.jpeg|[[Carpenter bee]] nests in a cedar wood beam (sawn open)
File:Bienen mit Brut 2.jpg|Honeybees on [[brood comb]] with eggs and [[larvae]] in cells
</gallery>
===Flight===
[[File:Apis mellifera flying.jpg|thumb|Honeybee in flight carrying pollen in [[pollen basket]]]]
{{further|Insect flight}}
[[Antoine Magnan]]'s 1934 book {{Lang|fr|Le vol des insectes}} says that he and [[André Sainte-Laguë]] had applied the equations of [[air resistance]] to [[insect]]s and found that their flight could not be explained by fixed-wing calculations, but that "One shouldn't be surprised that the results of the calculations don't square with reality".<ref>Ingram, Jay (2001) ''The Barmaid's Brain'', Aurum Press, pp. 91–92, {{ISBN|0716741202}}.</ref> This has led to a common misconception that bees "violate aerodynamic theory". In fact it merely confirms that bees do not engage in fixed-wing flight, and that their flight is explained by other mechanics, such as those used by [[helicopter]]s.<ref>{{cite web|author=Adams, Cecil |url=http://www.straightdope.com/columns/read/1076/is-it-aerodynamically-impossible-for-bumblebees-to-fly |title=Is it aerodynamically impossible for bumblebees to fly? |publisher=The Straight Dope |date=4 May 1990 |access-date=7 March 2009| archive-url= https://web.archive.org/web/20090303140245/http://www.straightdope.com/columns/read/1076/is-it-aerodynamically-impossible-for-bumblebees-to-fly| archive-date= 3 March 2009 | url-status=live}}</ref> In 1996 it was shown that vortices created by many insects' wings helped to provide lift.<ref>{{cite journal |url=https://www.newscientist.com/channel/life/dn8382-secrets-of-bee-flight-revealed.html |title=Life, animal and plant news, articles and features |journal=[[New Scientist]] |date=9 March 2016 |access-date=16 March 2016 |url-status=live |archive-url=https://web.archive.org/web/20081007163926/http://www.newscientist.com/channel/life/dn8382-secrets-of-bee-flight-revealed.html |archive-date=7 October 2008}}</ref> High-speed [[cinematography]]<ref>{{cite journal |url=https://www.newscientist.com/data/images/ns/av/dn8382.avi |title=Images of flight |journal=[[New Scientist]] |access-date=16 March 2016 |url-status=live |archive-url=https://web.archive.org/web/20160323022150/https://www.newscientist.com/data/images/ns/av/dn8382.avi |archive-date=23 March 2016}}</ref> and robotic mock-up of a bee wing<ref name=caltech>{{cite web|url=https://www.caltech.edu/news/deciphering-mystery-bee-flight-1075|title=Deciphering the Mystery of Bee Flight|publisher=[[California Institute of Technology]]|date=29 November 2005|access-date=8 September 2016|url-status=live|archive-url=https://web.archive.org/web/20160917023603/https://www.caltech.edu/news/deciphering-mystery-bee-flight-1075|archive-date=17 September 2016}} Re: work of [[Michael Dickinson (biologist)|Dr. Michael H. Dickinson]].</ref> showed that lift was generated by "the unconventional combination of short, choppy wing strokes, a rapid rotation of the wing as it flops over and reverses direction, and a very fast wing-beat frequency". Wing-beat frequency normally increases as size decreases, but as the bee's wing beat covers such a small [[arc (geometry)|arc]], it flaps approximately 230 times per second, faster than a [[Drosophilidae|fruitfly]] (200 times per second) which is 80 times smaller.<ref>{{cite journal |author1=Altshuler, Douglas L. |author2=Dickson, William B. |author3=Vance, Jason T. |author4=Roberts, Stephen P. |author5=Dickinson, Michael H. |title=Short-amplitude high-frequency wing strokes determine the aerodynamics of honeybee flight |journal=Proceedings of the National Academy of Sciences |volume=102 |issue=50 |pages=18213–18218 |year=2005 |doi=10.1073/pnas.0506590102 |pmid=16330767 |pmc=1312389|bibcode=2005PNAS..10218213A |doi-access=free }}</ref>
<!--temperature regulation - bees need to warm up for flight...-->
===Navigation, communication, and finding food===
[[File:Bee_dance.svg|thumb|upright|[[Karl von Frisch]] (1953) discovered that honey bee workers can [[animal navigation|navigate]], indicating the range and direction to food to other workers with a [[waggle dance]].]]
{{further|Animal navigation|Waggle dance}}
The ethologist [[Karl von Frisch]] studied [[animal navigation|navigation]] in the honey bee. He showed that honey bees communicate by the [[waggle dance]], in which a worker indicates the location of a food source to other workers in the hive. He demonstrated that bees can recognize a desired compass direction in three different ways: by the Sun, by the [[Polarization (waves)|polarization]] pattern of the blue sky, and by the Earth's magnetic field. He showed that the Sun is the preferred or main compass; the other mechanisms are used under cloudy skies or inside a dark [[beehive]].<ref name=vonFrisch93>{{cite book |last=von Frisch |first=Karl |title=The Dancing Bees |year=1953 |publisher=Harcourt, Brace & World | pages=93–96}}</ref> Bees navigate using [[spatial memory]] with a "rich, map-like organization".<ref>{{cite journal |author1=Menzel, Randolf |author2=Greggers, Uwe |author3=Smith, Alan |author4=Berger, Sandra |author5=Brandt, Robert |author6=Brunke, Sascha |author7=Bundrock, Gesine |author8=Hülse, Sandra |author9=Plümpe, Tobias |author10=Schaupp, Schaupp |author11=Schüttler, Elke |author12=Stach, Silke |author13=Stindt, Jan |author14=Stollhoff, Nicola |author15=Watzl, Sebastian |title=Honey bees Navigate According to a Map-Like Spatial Memory |journal=PNAS |year=2005 |volume=102 |issue=8 |pages=3040–3045 |doi=10.1073/pnas.0408550102 |pmid=15710880 |pmc=549458|bibcode=2005PNAS..102.3040M |doi-access=free }}</ref>
=== Digestion ===
The gut of bees is relatively simple, but multiple metabolic strategies exist in the gut [[microbiota]].<ref>{{Cite web |url=https://www.sciencedaily.com/releases/2017/12/171212141506.htm  |title=How honey bee gut bacteria help to digest their pollen-rich diet |website=ScienceDaily |access-date=2 January 2020}}</ref> Pollinating bees consume nectar and pollen, which require different digestion strategies by somewhat specialized bacteria. While nectar is a liquid of mostly [[monosaccharide]] sugars and so easily absorbed, pollen contains complex [[polysaccharide]]s: branching [[pectin]] and [[hemicellulose]].<ref>{{Cite web |url=https://phys.org/news/2019-12-bee-gut-microbes-division-labor.html |title=Bee gut microbes have a division of labor when it comes to metabolizing complex polysaccharides |website=phys.org |access-date=2 January 2020}}</ref> Approximately five groups of bacteria are involved in digestion. Three groups specialize in simple sugars (''[[Snodgrassella]]'' and two groups of ''[[Lactobacillus]]''), and two other groups in complex sugars (''[[Gilliamella]]'' and ''[[Bifidobacterium]]''). Digestion of pectin and hemicellulose is dominated by bacterial [[clade]]s ''Gilliamella'' and ''[[Bifidobacterium]]'' respectively. Bacteria that cannot digest polysaccharides obtain enzymes from their neighbors, and bacteria that lack certain amino acids do the same, creating multiple [[ecological niche]]s.<ref>{{Cite journal |last1=Zheng |first1=Hao |last2=Perreau |first2=Julie |last3=Powell |first3=J. Elijah |last4=Han |first4=Benfeng |last5=Zhang |first5=Zijing |last6=Kwong |first6=Waldan K. |last7=Tringe |first7=Susannah G. |last8=Moran |first8=Nancy A. |date=December 2019 |title=Division of labor in honey bee gut microbiota for plant polysaccharide digestion |journal=Proceedings of the National Academy of Sciences |volume=116 |issue=51 |pages=25909–25916 |doi=10.1073/pnas.1916224116 |issn=0027-8424 |pmid=31776248|pmc=6926048 |bibcode=2019PNAS..11625909Z |doi-access=free }}</ref>
Although most bee species are [[Nectarivore|nectarivorous]] and [[Palynivore|palynivorous]], some are not. Particularly unusual are [[vulture bee]]s in the genus ''[[Trigona]],'' which consume carrion and wasp brood, turning meat into a honey-like substance.<ref>{{Cite journal |last1=Mateus |first1=Sidnei |last2=Noll |first2=Fernando B. |date=February 2004 |title=Predatory behavior in a necrophagous bee Trigona hypogea (Hymenoptera; Apidae, Meliponini) |journal=Naturwissenschaften |volume=91 |issue=2 |pages=94–96 |doi=10.1007/s00114-003-0497-1 |pmid=14991148 |issn=1432-1904|bibcode=2004NW.....91...94M |s2cid=26518321 }}</ref> Drinking [[guttation]] drops from leaves is also a source of energy and nutrients.<ref>[https://royalsocietypublishing.org/doi/10.1098/rspb.2020.1080 Plant guttation provides nutrient-rich food for insects - Journals]</ref>
==Ecology==
=== Floral relationships ===
Most bees are [[polylectic]] (generalist) meaning they collect pollen from a range of flowering plants, but some are [[oligolege]]s (specialists), in that they only gather pollen from one or a few species or genera of closely related plants.<ref name=Waser2006>{{cite book |author=Waser, Nickolas M. |title=Plant-Pollinator Interactions: From Specialization to Generalization |url=https://books.google.com/books?id=Fbl5c9fUxTIC&pg=PA110 |year=2006 |publisher=University of Chicago Press |isbn=978-0-226-87400-5 |pages=110– |url-status=live |archive-url=https://web.archive.org/web/20180328155201/https://books.google.com/books?id=Fbl5c9fUxTIC&pg=PA110 |archive-date=28 March 2018}}</ref> In Melittidae and Apidae we also find a few genera that are highly specialized for collecting plant oils both in addition to, and instead of, nectar, which is mixed with pollen as larval food.<ref>{{cite journal | pmc=2838259 | year=2010 | last1=Renner | first1=S. S. | last2=Schaefer | first2=H. | title=The evolution and loss of oil-offering flowers: New insights from dated phylogenies for angiosperms and bees | journal=Philosophical Transactions of the Royal Society of London. Series B, Biological Sciences | volume=365 | issue=1539 | pages=423–435 | doi=10.1098/rstb.2009.0229 | pmid=20047869 }}</ref> Male orchid bees in some species gather aromatic compounds from [[orchid]]s, which is one of the few cases where male bees are effective pollinators. Bees are able to sense the presence of desirable flowers through ultraviolet patterning on flowers, floral odors,<ref name=Dafni>{{cite book |author1=Dafni, Amots |author2=Hesse, Michael |author3=Pacini, Ettore |title=Pollen and Pollination |url=https://books.google.com/books?id=-M7yCAAAQBAJ&pg=PA80 |year=2012 |publisher=Springer Science & Business Media |isbn=978-3-7091-6306-1 |page=80 |url-status=live |archive-url=https://web.archive.org/web/20180328155201/https://books.google.com/books?id=-M7yCAAAQBAJ&pg=PA80 |archive-date=28 March 2018}}</ref> and even electromagnetic fields.<ref>{{cite journal|last1=Suttona|first1=Gregory P.|last2=Clarkea|first2=Dominic|last3=Morleya|first3=Erica L.|last4=Robert|first4=Daniel|title=Mechanosensory hairs in bumblebees (Bombus terrestris) detect weak electric fields|journal=PNAS|date=2016|volume=113|issue=26|pages=7261–7265|doi=10.1073/pnas.1601624113|pmid=27247399|pmc=4932954|bibcode=2016PNAS..113.7261S |doi-access=free}}</ref> Once landed, a bee then uses nectar quality<ref name=Dafni /> and pollen taste<ref>{{cite journal|last1=Muth|first1=Felicity|last2=Francis|first2=Jacob S.|last3=Leonard|first3=Anne S.|title=Bees use the taste of pollen to determine which flowers to visit|journal=Biology Letters|date=2016|volume=12|issue=7|page=20160356|doi=10.1098/rsbl.2016.0356|pmid=27405383|pmc=4971173}}</ref> to determine whether to continue visiting similar flowers.
In rare cases, a [[plant]] species may only be effectively pollinated by a single bee species, and some plants are [[endangered species|endangered]] at least in part because their pollinator is also threatened. But, there is a pronounced tendency for oligolectic bees to be associated with common, widespread plants visited by multiple pollinator species. For example, the [[Larrea tridentata|creosote bush]] in the arid parts of the United States southwest is associated with some 40 oligoleges.<ref>{{cite journal |author=Hurd, P.D. Jr. |author2=Linsley, E.G. |year=1975 |title=The principal ''Larrea'' bees of the southwestern United States |journal=Smithsonian Contributions to Zoology |volume=193 |pages=1–74 |doi=10.5479/si.00810282.193 |issue=193}}</ref>
===As mimics and models===
[[File:Bombylius major on flower.jpg|thumb|left|The bee-fly ''[[Bombylius major]]'', a [[Batesian mimic]] of bees, taking nectar and pollinating a flower]]
{{main|Mimicry|Batesian mimicry|Müllerian mimicry}}
[[File:Ophrys apifera flower1.jpg|thumb|upright|[[Ophrys apifera|Bee orchid]] lures male bees to attempt to mate with the flower's lip, which resembles a bee perched on a pink flower.]]
Many bees are [[aposematic]]ally coloured, typically orange and black, warning of their ability to defend themselves with a powerful sting. As such they are models for [[Batesian mimicry]] by non-stinging insects such as [[bee-flies]], [[Asilidae|robber flies]] and [[hoverflies]],<ref name="ThorpHorning1983">{{cite book |last1=Thorp |first1=Robbin W. |last2=Horning |first2=Donald S. |last3=Dunning |first3=Lorry L. |title=Bumble Bees and Cuckoo Bumble Bees of California (Hymenoptera, Apidae) |url=https://books.google.com/books?id=v1eJ3fWwshIC&pg=PA9 |year=1983 |publisher=University of California Press |isbn=978-0-520-09645-5 |page=9 |quote=Of the forms of mimicry, two relate to [[Bombini]]. Batesian mimicry .. is exemplified by members of several families of flies: [[Syrphidae]], [[Asilidae]], [[Horse-fly|Tabanidae]], [[Oestridae]], and [[Bombyliidae]] (Gabritschevsky, 1926). |url-status=live |archive-url=https://web.archive.org/web/20170105171749/https://books.google.com/books?id=v1eJ3fWwshIC&pg=PA9 |archive-date=5 January 2017}}</ref> all of which gain a measure of protection by superficially looking and behaving like bees.<ref name="ThorpHorning1983"/>
Bees are themselves [[Müllerian mimicry|Müllerian mimics]] of other aposematic insects with the same colour scheme, including [[wasp]]s, [[Lycidae|lycid]] and other beetles, and many butterflies and moths ([[Lepidoptera]]) which are themselves distasteful, often through acquiring bitter and poisonous chemicals from their plant food. All the Müllerian mimics, including bees, benefit from the reduced risk of predation that results from their easily recognised warning coloration.<ref>{{cite book |last1=Cott |first1=Hugh |author-link=Hugh B. Cott |title=Adaptive Coloration in Animals |title-link=Adaptive Coloration in Animals |date=1940 |publisher=Oxford University Press |pages=196, 403 and passim}}</ref>
Bees are also mimicked by plants such as the [[Ophrys apifera|bee orchid]] which imitates both the appearance and the scent of a female bee; male bees attempt to mate ([[pseudocopulation]]) with the furry lip of the flower, thus pollinating it.<ref>{{cite web |title=Bee Orchids and Insect Mimicry |url=http://www.nhm.ac.uk/nature-online/life/plants-fungi/bee-orchids/ |publisher=Natural History Museum |access-date=1 July 2015 |url-status=live |archive-url=https://web.archive.org/web/20150708203045/http://www.nhm.ac.uk/nature-online/life/plants-fungi/bee-orchids/ |archive-date=8 July 2015}}</ref>
===As brood parasites===
[[File:Bumblebee January 2008-4.jpg|thumb|left|''[[Bombus vestalis]]'', a [[brood parasite]] of the bumblebee ''[[Bombus terrestris]]'']]
{{main|Brood parasite|Nest usurpation}}
[[Brood parasite]]s occur in several bee families including the [[apidae|apid]] subfamily [[Nomadinae]].<ref>{{cite web|title=Obligate Brood Parasitism|url=http://www.aculeataresearch.com/index.php/cuckoo-behavior/52-obligate-brood-parasitism|publisher=Aculeata Research Group|access-date=30 June 2015|url-status=live|archive-url=https://web.archive.org/web/20150707053542/http://www.aculeataresearch.com/index.php/cuckoo-behavior/52-obligate-brood-parasitism|archive-date=7 July 2015}}</ref> Females of these species lack pollen collecting structures (the [[scopa (biology)|scopa]]) and do not construct their own nests. They typically enter the nests of pollen collecting species, and lay their eggs in cells provisioned by the host bee. When the "cuckoo" bee larva hatches, it consumes the host larva's pollen ball, and often the host egg also.<ref>{{cite web |title=Brood Parasitism |url=http://www.amentsoc.org/insects/glossary/terms/brood-parasitism |publisher=Amateur Entomologists' Society |access-date=30 June 2015 |url-status=live |archive-url=https://web.archive.org/web/20150702165918/http://www.amentsoc.org/insects/glossary/terms/brood-parasitism |archive-date=2 July 2015}}</ref> In particular, the Arctic bee species, ''[[Bombus hyperboreus]]'' is an aggressive species that attacks and enslaves other bees of the same subgenus. However, unlike many other bee brood parasites, they have pollen baskets and often collect pollen.<ref>Gjershaug, Jan Ove (5 June 2009). "The social parasite bumblebee Bombus hyperboreus Schönherr, 1809 usurp nest of Bombus balteatus Dahlbom, 1832 (Hymenoptera, Apidae) in Norway" (PDF). ''Norwegian Journal of Entomology'' '''56'''(1): 28–31. Retrieved 26 September 2015.</ref>
In Southern Africa, hives of African honeybees (''A. mellifera scutellata'') are being destroyed by parasitic workers of the Cape honeybee, ''A. m. capensis''. These lay [[diploid]] eggs ("[[thelytoky]]"), escaping normal [[worker policing]], leading to the colony's destruction; the parasites can then move to other hives.<ref>{{cite book |last1=Gullan |first1=P. J. |last2=Cranston |first2=P. S. |title=The Insects: An Outline of Entomology |date=2014 |publisher=Wiley Blackwell |isbn=978-1-118-84615-5 |edition=5th |page=347}}</ref>
The [[cuckoo bee]]s in the ''[[List of bumblebee species|Bombus]]'' subgenus ''Psithyrus'' are closely related to, and resemble, their hosts in looks and size. This common pattern gave rise to the ecological principle "[[Emery's rule]]". Others parasitize bees in different families, like ''[[Townsendiella]]'', a [[nomadinae|nomadine]] [[apidae|apid]], two species of which are cleptoparasites of the [[Dasypodaidae|dasypodaid]] genus ''[[Hesperapis]]'',<ref>{{cite journal |author1=Rozen, Jerome George |author2=McGinley, Ronald J. |year=1991 |title=Biology and Larvae of the Cleptoparasitic Bee ''Townsendiella pulchra'' and Nesting Biology of its Host ''Hesperapis larreae'' (Hymenoptera, Apoidea) |journal=American Museum Novitates |issue=3005  |hdl=2246/5032 }}</ref> while the other species in the same genus attacks [[halictidae|halictid]] bees.<ref>{{cite book|author1=Moure, Jesus S.|author2=Hurd, Paul David |title=An Annotated Catalog of the Halictid Bees of the Western Hemisphere (Hymenoptera, Halictidae) |url=https://books.google.com/books?id=2iUlSfQt8vEC |year=1987 |publisher=Smithsonian Institution Press |pages=28–29}}</ref>
=== Nocturnal bees ===
Four bee families ([[Andrenidae]], [[Colletidae]], [[Halictidae]], and [[Apidae]]) contain some species that are [[crepuscular]]. Most are tropical or subtropical, but some live in arid regions at higher latitudes. These bees have greatly enlarged [[ocellus|ocelli]], which are extremely sensitive to light and dark, though incapable of forming images. Some have refracting superposition compound eyes: these combine the output of many elements of their compound eyes to provide enough light for each retinal photoreceptor. Their ability to fly by night enables them to avoid many predators, and to exploit flowers that produce nectar only or also at night.<ref>{{cite journal |last1=Warrant |first1=Eric J. |title=Seeing in the dark: vision and visual behaviour in nocturnal bees and wasps |journal=Journal of Experimental Biology |date=June 2008 |volume=211 |issue=11 |pages=1737–1746 |doi=10.1242/jeb.015396 |pmid=18490389|doi-access=free }}</ref>
===Predators, parasites and pathogens===
{{further|Diseases of the honey bee}}
[[File:Pair of Merops apiaster feeding detail.jpg|thumb|right|The bee-eater, ''[[Merops apiaster]]'', specialises in feeding on bees; here a male catches a nuptial gift for his mate.]]
Vertebrate predators of bees include [[bee-eater]]s, [[shrikes]] and [[Old World flycatcher|flycatcher]]s, which make short sallies to catch insects in flight.<ref name="ChittkaThomson2001"/> Swifts and swallows<ref name="ChittkaThomson2001"/> fly almost continually, catching insects as they go. The [[Pernis (bird)|honey buzzard]] attacks bees' nests and eats the larvae.<ref>{{cite web | url=https://www.theguardian.com/world/2013/sep/26/hornet-attacks-kill-18-china | work=The Guardian | title=Hornet attacks kill dozens in China | date=26 September 2013 | access-date=18 June 2015 | url-status=live | archive-url=https://web.archive.org/web/20150906191229/http://www.theguardian.com/world/2013/sep/26/hornet-attacks-kill-18-china | archive-date=6 September 2015}}</ref> The [[greater honeyguide]] interacts with humans by guiding them to the nests of wild bees. The humans break open the nests and take the honey and the bird feeds on the larvae and the wax.<ref>{{cite journal |author=Friedmann, Herbert |year=1955 |title=The Honey-Guides |journal=Bulletin of the United States National Museum |doi=10.5479/si.03629236.208.1 |pages=1–292 |issue=208|hdl=10088/10101 }}</ref> Among mammals, predators such as the [[badger]] dig up bumblebee nests and eat both the larvae and any stored food.<ref>{{cite web |title=What predators do bumblebees have? |url=http://bumblebeeconservation.org/about-bees/faqs/bumblebee-predators/ |publisher=Bumblebee Conservation Trust |access-date=29 June 2015 |url-status=dead |archive-url=https://web.archive.org/web/20150629025444/http://bumblebeeconservation.org/about-bees/faqs/bumblebee-predators/ |archive-date=29 June 2015}}</ref>
[[File:Wasp and bee August 2008-2.jpg|thumb|left|The beewolf ''[[Philanthus triangulum]]'' paralysing a bee with its sting]]
Specialist ambush predators of visitors to flowers include [[crab spiders]], which wait on flowering plants for pollinating insects; [[Hemiptera|predatory bugs]], and [[praying mantis]]es,<ref name="ChittkaThomson2001">{{cite book |last1=Chittka |first1=Lars |last2=Thomson |first2=James D. |title=Cognitive Ecology of Pollination: Animal Behaviour and Floral Evolution |url=https://books.google.com/books?id=g2Km4B6n-mQC&pg=PA215 |date=28 May 2001 |publisher=Cambridge University Press |isbn=978-1-139-43004-3 |pages=215–216 |url-status=live |archive-url=https://web.archive.org/web/20161224104844/https://books.google.com/books?id=g2Km4B6n-mQC&pg=PA215 |archive-date=24 December 2016}}</ref> some of which (the [[flower mantis]]es of the tropics) wait motionless, [[aggressive mimicry|aggressive mimics]] camouflaged as flowers.<ref>{{cite web |last1=Choi |first1=Charles Q. |title=Found! First Known Predator To Lure Prey By Mimicking Flowers |url=http://www.livescience.com/41605-predator-lures-prey-by-mimicking-flowers.html |publisher=LiveScience |access-date=2 July 2015 |date=30 November 2013 |quote=the color of the orchid mantis was indistinguishable from 13 species of wild flowers in the areas the predator lived. ... The orchid mantis is unique in that the mantis itself is the attractive stimulus. |url-status=live |archive-url=https://web.archive.org/web/20150630181114/http://www.livescience.com/41605-predator-lures-prey-by-mimicking-flowers.html |archive-date=30 June 2015}}</ref> [[Beewolf|Beewolves]] are large wasps that habitually attack bees;<ref name="ChittkaThomson2001"/> the [[ethologist]] [[Niko Tinbergen]] estimated that a single colony of the beewolf ''[[Philanthus triangulum]]'' might kill several thousand honeybees in a day: all the prey he observed were honeybees.<ref>{{cite book | author=Tinbergen, Niko | title=Curious Naturalists | publisher=Methuen | year=1958 | page=21}}</ref> Other predatory insects that sometimes catch bees include [[Asilidae|robber flies]] and [[dragonfly|dragonflies]].<ref name="ChittkaThomson2001"/> Honey bees are affected by parasites including [[Acarapis woodi|tracheal]] and ''[[Varroa]]'' [[mite]]s.<ref>{{cite web |title=Honey Bee Disorders: Honey Bee Parasites |url=http://www.ent.uga.edu/bees/disorders/honey-bee-parasites.html |publisher=University of Georgia |access-date=29 June 2015 |url-status=live |archive-url=https://web.archive.org/web/20150701225054/http://www.ent.uga.edu/bees/disorders/honey-bee-parasites.html |archive-date=1 July 2015}}</ref> However, some bees are believed to have a mutualistic relationship with mites.<ref name="Pavel B 2007"/>
Some mites of genus ''[[Tarsonemus]]'' are associated with bees. They live in bee nests and ride on adult bees for dispersal. They are presumed to feed on fungi, nest materials or pollen. However, the impact they have on bees remains uncertain.<ref>{{Cite web |title=Tarsonemus {{!}} Bee Mite ID |url=http://idtools.org/id/mites/beemites/factsheet.php?name=15293 |access-date=25 August 2022 |website=idtools.org}}</ref>
=== Symbiosis of Mycelium and Bees ===
==== Fungus properties ====
Recent studies have shown that [[mycelium]] provides [[honey bee]]s and [[stingless bee]]s with vital nutrients. Specific fungi, such as [[Zygosaccharomyces]] sp, [[Candida (fungus)|Candida sp]]., and [[Monascus]] ruber, produce chemicals that fight against [[bacteria]], [[fungal infection]]s from different species, and [[virus]]es. Recently these types of bees have been observed eating [[mycelium]], suggesting that honey bees have been “foraging [[mushroom]]s to collect [[antimicrobial]] medicine to boost their collective [[Immunity (medical)|immunity]]”.<ref>{{Cite web |last=White |first=Katie |date=2022-02-02 |title=Curious about Fungi and Honey Bees? |url=https://bestbees.com/2022/02/02/fungi-and-honey-bees/ |access-date=2023-04-25 |website=bestbees.com |language=en-US}}</ref> Without these vital nutrients, honey bee morbidity rates rise, and the possibility of [[fungal infection]]s can spike, leading to unhealthy bee hives and honey shortage. [[Fungal infection]]s can also lead to [[colony collapse disorder]], so the ingestion of [[mycelium]] lowers the morbidity rate of honey bees by preventing those [[fungal infection]]s from happening. [[Colony collapse disorder]] (CCD) is when worker bees abandon the [[queen bee]] and leave behind the brood and a few nurse bees. This however is not enough to sustain a hive as workers are required to construct and maintain the hive structure as well as produce honey. [[Colony collapse disorder]] can also happen when [[Varroa Mites|varroa mites]] infiltrate a hive. These mites will attack and eat bees inside a hive, making it impossible for them to continue to reproduce and make [[honey]]. The presence of [[Varroa destructor|varroa mites]] results in a decrease in bee population, deformed bees, an inability to reproduce on the bees part, and overall weakening of the colony. [[Varroa Mites|Varroa mites]] are only capable of reproducing inside of a honey bee colony, posing an even greater threat if they are able to infiltrate because it will destroy their home. [[Mycelium]] has been shown to [[Germination|germinate]] inside of [[Varroa Mites|varroa mites]] and grow from the inside out, killing the mites and protecting the bees. The extermination of mites by [[mycelium]] is a better alternative to [[pesticide]]s that have shown to be toxic towards the bee colony. [[Mycelium]] also plays a role in boosting [[anti-inflammatory]] and [[Antimicrobial resistance|antibacterial resistance]] in bees due to the [[ecdysteroid]]s and [[Zygosaccharomyces]] found in [[mycelium]], which are then fed to [[larva]]e, boosting the next generations immunity and improving overall hive health. [[Zygosaccharomyces]] are “spoilage yeasts that have an extreme resistance to acids and preservatives” and can “tolerate high concentrations of sugars and salts”.<ref>{{Cite web |title=Zygosaccharomyces - an overview {{!}} ScienceDirect Topics |url=https://www.sciencedirect.com/topics/agricultural-and-biological-sciences/zygosaccharomyces#:~:text=Zygosaccharomyces%20bailii%20is%20characterized%20by,delbrueckii%20of%20diverse%20physiological%20properties.%20(Accessed:%20March%2020,%202023). |access-date=2023-04-25 |website=www.sciencedirect.com}}</ref> Honey bees depend on this source of steroids to allow them to develop properly during insect [[Pupa|pupation.]]
==== Bee broods ====
The [[Symbiosis|symbiotic]] relationship between bees and [[mycelium]] is found primarily in Brazilian stingless bees and Malaysian stingless bees - or more commonly honey bees. [[Bee brood]]s are the [[larva]]e of honeybees. They can typically be found inside of a bee hive, and in man made hives especially, the honeybees can be found developing at different stages (eggs, larvae, and [[pupa]]e) inside a [[Honeycomb|hexagonal shape]]. Bee larvae are incapable of producing [[steroid]]s at birth, so they ingest [[mycelium]] to receive vital nutrients they cannot create on their own such as [[ecdysteroid]]s and Zygosaccharomyces sp.<ref name="ReferenceA">{{Cite journal |last1=Paludo |first1=Camila Raquel |last2=Pishchany |first2=Gleb |last3=Andrade-Dominguez |first3=Andres |last4=Silva-Junior |first4=Eduardo Afonso |last5=Menezes |first5=Cristiano |last6=Nascimento |first6=Fabio Santos |last7=Currie |first7=Cameron R. |last8=Kolter |first8=Roberto |last9=Clardy |first9=Jon |last10=Pupo |first10=Mônica Tallarico |date=2019-07-25 |title=Microbial community modulates growth of symbiotic fungus required for stingless bee metamorphosis |journal=PLOS ONE |language=en |volume=14 |issue=7 |pages=e0219696 |doi=10.1371/journal.pone.0219696 |issn=1932-6203 |pmc=6657851 |pmid=31344052 |bibcode=2019PLoSO..1419696P |doi-access=free }}</ref> Once the honey bee eggs hatch, a white microbial film starts to grow on the boundary between the brood cell and the larval food supply, and is then ingested by the [[larva]]e to complete their development.<ref name="ReferenceA"/>
[[Gut microbiota]] play an immense role in the health of the entire bee colony. Three studies were recently conducted and each introduced a new organism to the bees [[gut microbiota]]. The bees were fed aged [[pollen]], the assembly of the [[gut microbiota]] was disturbed, and antibiotic tetracycline entered their diet. All three studies showed that the honey bees' ability to survive decreased drastically and they became more likely to contract [[Parasitism|parasites]] and [[fungal infection]]s.<ref>{{Cite journal |last1=Bonilla-Rosso |first1=Germán |last2=Engel |first2=Philipp |date=2018-06-01 |title=Functional roles and metabolic niches in the honey bee gut microbiota |journal=Current Opinion in Microbiology |series=Environmental Microbiology * The New Microscopy |language=en |volume=43 |pages=69–76 |doi=10.1016/j.mib.2017.12.009 |pmid=29309997 |s2cid=19058556 |issn=1369-5274|doi-access=free }}</ref> The introduction of certain [[mycelium]] to the honey bees [[gut microbiota]] has the opposite effect to what took place in these three studies, highlighting the importance of what bees ingest and the impacts it has on their survivability during both the development and adult stages.
==== Bee-fungus symbiosis ====
As mentioned above, honey bees cannot produce [[steroid]]s themselves, they must be ingested through their diet, specifically in the early development process. Larvae eat the fungus and the [[ecdysteroid]]s and [[Zygosaccharomyces]] produced by the [[mycelium]] benefit the [[larva]]e. [[Ecdysteroid]]s are naturally occurring [[steroid]]s found in [[mycelium]] and they help enhance performance and reproduction, boosting honey production and keeping the hive population running at a stable rate. “[[Zygosaccharomyces]] sp. is essential for S. depilis larvae”.<ref name="ReferenceA"/> These [[sterol]]s thus have a high impact on the survival rate of honey bees. Their ingestion determines whether the honey bees will be able to protect themselves against [[fungal infection]]s, [[virus]]es, and whether or not they will have sufficient strength to increase honey production and the ability to [[Pollination|pollinate]] a larger area and more frequently.
Knowledge of how [[mycelium]] boosts honey bees immunity could be pivotal to the increase of a honey bee's lifespan and boost reproduction by helping implement new policies to prevent the use of harmful [[pesticide]]s.<ref name="ReferenceA" />
==== Impact of pesticides ====
[[Pesticide]]s have been diminishing the bee population recently due to a lack of regulations regarding what can and cannot be sprayed on produce to protect it from being damaged during growth. When honey bees collect [[pollen]] and [[nectar]] for nutrition and to make [[honey]], they are also ingesting harmful chemicals. Such chemicals take a toll on the honey bees' already sensitive [[Gut microbiota|gut microbiome]] and lead to a higher morbidity rate in honey bees. "These [[Microorganism|microbes]] can suffer with toxic [[pesticide]]s applied in agriculture, causing dangerous changes in the colony fitness and perturbing bees' health.” (Yordanova, M. et al., 2022)<ref>{{Cite journal |last1=Yordanova |first1=Monika |last2=Evison |first2=Sophie E. F. |last3=Gill |first3=Richard J. |last4=Graystock |first4=Peter |date=2022-04-01 |title=The threat of pesticide and disease co-exposure to managed and wild bee larvae |journal=International Journal for Parasitology: Parasites and Wildlife |language=en |volume=17 |pages=319–326 |doi=10.1016/j.ijppaw.2022.03.001 |pmid=35342713 |pmc=8943340 |bibcode=2022IJPPW..17..319Y |issn=2213-2244}}</ref>
With the research provided on the positive impact of [[mycelium]] on bees, the relationship between [[mycelium]] and honey bees is [[Symbiosis|symbiotic]] in that the survival of bees and the mycelium's ability to help boost bee pollination, boosts the ability of the [[Fungus|fungi]] to grow because bee pollination improves air and soil quality, thus boosting plant life. Allowing for a higher survivability rate for both bees and [[mycelium]] if they are able to perform their environmental roles properly without the interruption of harmful government approved [[pesticide]]s. The recent studies done on the [[Symbiosis|symbiotic]] relationship between [[mycelium]] and honey bees will prove to be vital in the argument towards lessening the types of chemicals legally allowed to be sprayed on produce. The use of [[pesticide]]s on lawns and for other agricultural uses destroy the livelihood of [[mycelium]] by killing the soil it grows in, inhibiting bees from ingesting the necessary nutrients [[mycelium]] provides to survive.
==Relationship with humans==
===In mythology and folklore===
{{Main|Bees in mythology}}
[[File:Plaque bee-goddess BM GR1860.4-123.4.jpg|thumb|Gold plaques embossed with winged bee goddesses. [[Kameiros|Camiros]], [[Rhodes]]. 7th century BC.]]
[[Homer]]'s ''[[Homeric Hymns|Hymn to Hermes]]'' describes three bee-maidens with the power of [[divination]] and thus speaking truth, and identifies the food of the gods as honey. Sources associated the bee maidens with [[Apollo]] and, until the 1980s, scholars followed Gottfried Hermann (1806) in incorrectly identifying the bee-maidens with the [[Thriae]].<ref>Susan Scheinberg, "The Bee Maidens of the Homeric ''Hymn to Hermes''", in Albert Heinrichs, ed., ''Harvard Studies in Classical Philology'' (Cambridge MA: Harvard University Press, 1980), 11. {{ISBN|0674379306}}; and many others since questioning Gottfried Hermann's 1806 equation of the ''Thriae'' with bee-maidens. Heinrich Gottfried, ''Homeri nomine dignissimum''/Homeric Hymns (Leipzig: 1806), 346 and cxiii. Many took Hermann's unfounded identification at face-value, repeating it ''ad nauseam'', e.g. Hilda M. Ransome, ''The Sacred Bee in Ancient Times and Folklore'' (NY: Courier, 1937; reprinted as recently as NY: Dover, 2012), 97. {{ISBN|0486122980}}</ref> Honey, according to a Greek myth, was discovered by a nymph called [[Melissa]] ("Bee"); and honey was offered to the Greek gods from [[Helladic period|Mycenean times]]. Bees were also associated with the [[Delphic oracle]] and the prophetess was sometimes called a bee.<ref>{{cite journal | author=Scheinberg, Susan | year=1979 | title=The Bee Maidens of the Homeric Hymn to Hermes | journal=Harvard Studies in Classical Philology | volume=83 | pages=1–28 | doi=10.2307/311093 | jstor=311093}}</ref>
The image of a community of honey bees has been used from ancient to modern times, in [[Aristotle]] and [[Plato]]; in [[Virgil]] and [[Seneca the Younger|Seneca]]; in [[Erasmus]] and [[Shakespeare]]; [[Tolstoy]], and by political and social theorists such as [[Bernard Mandeville]] and [[Karl Marx]] as a model for human [[society]].<ref>{{cite book |last=Wilson |first=Bee |year=2004 |title=The Hive: the Story of the Honeybee |location=London |publisher=[[John Murray (publishing house)|John Murray]] |isbn=0-7195-6598-7}}</ref> In English folklore, bees would be told of important events in the household, in a custom known as "[[Telling the bees]]".<ref name="Roud2006">{{cite book |author=Steve Roud |title=The Penguin Guide to the Superstitions of Britain and Ireland |url=https://books.google.com/books?id=1Mc4qPiICvcC&pg=PT128 |date=6 April 2006 |publisher=Penguin Books |isbn=978-0-14-194162-2 |page=128 |url-status=live |archive-url=https://web.archive.org/web/20161128210444/https://books.google.com/books?id=1Mc4qPiICvcC&pg=PT128 |archive-date=28 November 2016}}</ref>
===In art and literature===
[[File:Mrs tittlemouse.jpg|thumb|upright|[[Beatrix Potter]]'s illustration of Babbity Bumble in ''[[The Tale of Mrs Tittlemouse]]'', 1910]]
Some of the oldest examples of bees in art are [[Cave painting|rock paintings]] in [[Spain]] which have been dated to 15,000 BC.<ref name=EHSTMNWC>{{cite book |title=Encyclopaedia of the History of Science, Technology, and Medicine in Non-Western Cultures |url=https://books.google.com/books?id=kt9DIY1g9HYC&pg=PA1074 |publisher=Springer Science & Business Media |isbn=978-1-4020-4559-2 |page=1074 |url-status=live |archive-url=https://web.archive.org/web/20140704171807/http://books.google.com/books?id=kt9DIY1g9HYC |archive-date=4 July 2014|date = 12 March 2008}}</ref>
[[W. B. Yeats]]'s poem ''[[The Lake Isle of Innisfree]]'' (1888) contains the couplet "Nine bean rows will I have there, a hive for the honey bee, / And live alone in the bee loud glade." At the time he was living in [[Bedford Park, London|Bedford Park]] in the West of London.<ref>{{cite web |last1=Deering |first1=Chris |title=Yeats in Bedford Park |url=http://www.chiswickw4.com/default.asp?section=info&page=conyeats.htm |publisher=ChiswickW4.com |access-date=28 June 2015 |url-status=live |archive-url=https://web.archive.org/web/20150630131250/http://www.chiswickw4.com/default.asp?section=info&page=conyeats.htm |archive-date=30 June 2015}}</ref> [[Beatrix Potter]]'s illustrated book ''[[The Tale of Mrs Tittlemouse]]'' (1910) features Babbity Bumble and her brood ''(pictured)''. [[Kit Williams]]' [[Treasure hunt (game)|treasure hunt]] book ''[[The Bee on the Comb]]'' (1984) uses bees and beekeeping as part of its story and puzzle. [[Sue Monk Kidd]]'s ''[[The Secret Life of Bees (novel)|The Secret Life of Bees]]'' (2004), and the [[The Secret Life of Bees (film)|2009 film]] starring [[Dakota Fanning]], tells the story of a girl who escapes her abusive home and finds her way to live with a family of beekeepers, the Boatwrights.
The 2007 animated comedy film ''[[Bee Movie]]'' used [[Jerry Seinfeld]]'s first script and was his first work for children; he starred as a bee named Barry B. Benson, alongside [[Renée Zellweger]]. Critics found its premise awkward and its delivery tame.<ref>{{cite web |url=http://www.rottentomatoes.com/m/bee_movie/ |title=Bee Movie |website=Rotten Tomatoes |access-date=30 June 2015 |url-status=live |archive-url=https://web.archive.org/web/20150623154308/http://www.rottentomatoes.com/m/bee_movie/ |archive-date=23 June 2015}}</ref> [[Dave Goulson]]'s ''[[A Sting in the Tale]]'' (2014) describes his efforts to save bumblebees in Britain, as well as much about their biology. The playwright [[Laline Paull]]'s [[fantasy]] ''The Bees'' (2015) tells the tale of a hive bee named Flora 717 from hatching onwards.<ref>{{cite news |last1=Jones |first1=Gwyneth |title=The Bees by Laline Paull review – a fantasy with a sting in its tail |url=https://www.theguardian.com/books/2014/may/21/bees-laline-paull-fantasy-novel-review |newspaper=The Guardian |access-date=28 June 2015 |url-status=live |archive-url=https://web.archive.org/web/20150701010722/http://www.theguardian.com/books/2014/may/21/bees-laline-paull-fantasy-novel-review |archive-date=1 July 2015|date=21 May 2014 }}</ref>
===Beekeeping===
{{main|Beekeeping}}
[[Image:Beekeeper.jpg|thumb|right|A commercial beekeeper at work]]
[[File:Western honey bee on a honeycomb.jpg|thumb|Western honey bee on a honeycomb]]
Humans have kept honey bee colonies, commonly in [[beehive|hives]], for millennia. [[Beekeeper]]s collect [[honey]], [[beeswax]], [[propolis]], [[pollen]], and [[royal jelly]] from hives; bees are also kept to [[pollination|pollinate]] crops and to produce bees for sale to other beekeepers.
Depictions of humans collecting honey from wild bees date to 15,000 years ago; efforts to domesticate them are shown in Egyptian art around 4,500 years ago.<ref>{{cite web |url=http://reshafim.org.il/ad/egypt/timelines/topics/beekeeping.htm |title=Ancient Egypt: Bee-keeping |website=Reshafim.org.il |date=6 April 2003 |access-date=16 March 2016 |url-status=live |archive-url=https://web.archive.org/web/20160309203227/http://www.reshafim.org.il/ad/egypt/timelines/topics/beekeeping.htm |archive-date=9 March 2016}}</ref> Simple hives and smoke were used;<ref>{{cite web |url=http://beelore.com/2008/02/23/beekeeping-in-ancient-egypt/ |title=Beekeeping in Ancient Egypt |publisher=Bee Lore |access-date=16 March 2016 |url-status=live |archive-url=https://web.archive.org/web/20160322224443/http://beelore.com/2008/02/23/beekeeping-in-ancient-egypt/ |archive-date=22 March 2016|date=23 February 2008 }}</ref><ref name="Bodenheimer1960">{{Cite book | title=Animal and Man in Bible Lands | first=F. S. | last=Bodenheimer | publisher=Brill Archive |year=1960 | page=79}}</ref> jars of honey were found in the tombs of [[pharaoh]]s such as [[Tutankhamun]].
Among Classical Era authors, beekeeping with the use of smoke is described in Aristotle's ''[[History of Animals]]'' Book 9.<ref name=Aristotle>{{cite book |last1=Aristotle |author1-link=Aristotle |last2=Thompson |first2=D'Arcy (trans.) |author2-link=D'Arcy Wentworth Thompson |title=The Works of Aristotle |date=1910 |publisher=Clarendon Press |pages=Book 9, Section 40 |url=https://archive.org/stream/worksofaristotle04arisuoft#page/n443/mode/2up}}</ref> The account mentions that bees die after stinging; that workers remove corpses from the hive, and guard it; castes including workers and non-working [[drone (bee)|drone]]s, but "kings" rather than queens; predators including toads and bee-eaters; and the [[waggle dance]], with the "irresistible suggestion" of {{lang|grc|άροσειονται}} ("{{lang|grc-Latn|aroseiontai}}", it waggles) and {{lang|grc|παρακολουθούσιν}} ("{{lang|grc-Latn|parakolouthousin}}", they watch).<ref name=Whitfield>{{cite journal |last1=Whitfield |first1=B. G. |title=Cambridge University Press and The Classical Association are collaborating with JSTOR to digitize, preserve and extend access to Greece & Rome. Virgil and the Bees: A Study in Ancient Apicultural Lor |journal=Greece and Rome |date=October 1956 |volume=3 |issue=2 |pages=99–117 |jstor=641360|doi=10.1017/S0017383500015126 |s2cid=161643666 }}</ref>{{efn|In D'Arcy Thompson's translation: "At early dawn they make no noise, until some one particular bee makes a buzzing noise two or three times and thereby awakes the rest; hereupon they all fly in a body to work. By and by they return and at first are noisy; ... until at last some one bee flies round about, making a buzzing noise, and apparently calling on the others to go to sleep".<ref name=Aristotle/>}}
Beekeeping is described in detail by [[Virgil]] in his ''[[Georgics]]''; it is also mentioned in his ''[[Aeneid]]'', and in [[Pliny the Elder|Pliny's]] ''[[Natural History (Pliny)|Natural History]]''.<ref name=Whitfield/>
From the 18th century, European understanding of the colonies and biology of bees allowed the construction of the moveable comb hive so that honey could be harvested without destroying the colony.<ref>Thomas Wildman, ''A Treatise on the Management of Bees'' (London, 1768, 2nd edn 1770).</ref><ref>{{cite journal |last1=Harissis |first1=H. V. |last2=Mavrofridis |first2=G. |date=2012 |title=A 17th Century Testimony on the Use of Ceramic Top-bar Hives |url=https://www.academia.edu/1929792 |url-status=live |journal=Bee World |volume=89 |issue=3 |pages=56–57 |doi=10.1080/0005772x.2012.11417481 |s2cid=85120138 |archive-url=https://web.archive.org/web/20151019113156/http://www.academia.edu/1929792/A_17th_Century_Testimony_On_The_Use_Of_Ceramic_Top-bar_Hives._2012 |archive-date=19 October 2015}}</ref>
===As commercial pollinators===
{{See also|List of crop plants pollinated by bees|Pollinator decline|Pesticide toxicity to bees}}
==== Role of bees ====
Bees play an important role in [[pollination|pollinating]] [[flowering plant]]s, and are the major type of [[pollinator]] in many [[ecosystem]]s that contain flowering plants. It is estimated that one third of the human food supply depends on [[pollination]] by insects, birds and bats, most of which is accomplished by bees, whether wild or domesticated.<ref>{{cite web |last1=Yang |first1=Sarah |title=Pollinators help one-third of world's crop production, says new study |url=http://www.berkeley.edu/news/media/releases/2006/10/25_pollinator.shtml |publisher=UC Berkeley |access-date=29 June 2015 |date=25 October 2006 |url-status=live |archive-url=https://web.archive.org/web/20150709060248/http://www.berkeley.edu/news/media/releases/2006/10/25_pollinator.shtml |archive-date=9 July 2015}}</ref><ref>{{cite news |url=https://www.independent.co.uk/news/science/wild-bees-found-to-be-just-as-important-as-honeybees-for-pollinating-food-crops-10324450.html |title=Wild bees just as important as domesticated bees for pollinating food crops |author=Connor, Steve |date=16 June 2015 |newspaper=The Independent |quote=Wild bees have become as important as domesticated honeybees in pollinating food crops around the world due to the dramatic decline in number of healthy honeybee colonies over the past half century, a study has found. |url-status=live |archive-url=https://web.archive.org/web/20170906052404/http://www.independent.co.uk/news/science/wild-bees-found-to-be-just-as-important-as-honeybees-for-pollinating-food-crops-10324450.html |archive-date=6 September 2017}}</ref>
==== In the United States ====
[[pollination management|Contract pollination]] has overtaken the role of honey production for [[beekeeper]]s in many countries. After the introduction of [[Varroa mites]], [[feral]] honey bees declined dramatically in the US, though their numbers have since recovered.<ref>{{cite journal |last1=Loper |first1=Gerald M. |last2=Sammataro |first2=Diana |last3=Finley |first3=Jennifer |last4=Cole |first4=Jerry |title=Feral honey bees in southern Arizona, 10 years after varroa infestation |journal=American Bee Journal |date=2006 |volume=146 |pages=521–524}}</ref><ref>{{cite journal |last1=Rangel |first1=Juliana |last2=Giresi |first2=Melissa |last3=Pinto |first3=Maria Alice |last4=Baum |first4=Kristen A. |last5=Rubink |first5=William L. |last6=Coulson |first6=Robert N. |last7=Johnston |first7=John Spencer |title=Africanization of a feral honey bee (Apis mellifera) population in South Texas: does a decade make a difference? |journal=Ecology and Evolution |date=2016 |volume=6 |issue=7 |pages=2158–2169 |doi=10.1002/ece3.1974|pmid=27069571 |pmc=4782243 |bibcode=2016EcoEv...6.2158R }}</ref> The number of colonies kept by beekeepers declined slightly, through [[urbanization]], systematic pesticide use, [[Acarapis woodi|tracheal]] and ''[[Varroa]]'' mites, and the closure of beekeeping businesses. In 2006 and 2007 the rate of attrition increased, and was described as [[colony collapse disorder]].<ref name="Penn">{{cite news|title=Honey Bee Die-Off Alarms Beekeepers, Crop Growers and Researchers|url=http://www.aginfo.psu.edu/News/07Jan/HoneyBees.htm|publisher=Pennsylvania State University College of Agricultural Sciences|date=29 January 2007|url-status=live|archive-url=https://web.archive.org/web/20080517154313/http://www.aginfo.psu.edu/News/07Jan/HoneyBees.htm|archive-date=17 May 2008}}</ref> In 2010 invertebrate iridescent virus and the fungus ''[[Nosema ceranae]]'' were shown to be in every killed colony, and deadly in combination.<ref>Johnson, Kirk (6 October 2010) [https://www.nytimes.com/2010/10/07/science/07bees.html Scientists and Soldiers Solve a Bee Mystery] {{webarchive|url=https://web.archive.org/web/20101007221509/http://www.nytimes.com/2010/10/07/science/07bees.html |archive-url=https://ghostarchive.org/archive/20220101/http://www.nytimes.com/2010/10/07/science/07bees.html |archive-date=1 January 2022 |url-access=limited |date=7 October 2010 }}{{cbignore}}. ''[[The New York Times]]''.</ref><ref>{{cite web |last=Eban |first=Katherine |url=https://money.cnn.com/2010/10/08/news/honey_bees_ny_times.fortune/index.htm |title=What a scientist didn't tell the New York Times about his study on bee deaths |publisher=CNN |date=8 October 2010 |access-date=20 August 2012 |url-status=live |archive-url=https://web.archive.org/web/20121019235428/https://money.cnn.com/2010/10/08/news/honey_bees_ny_times.fortune/index.htm |archive-date=19 October 2012}}</ref><ref>{{cite journal |title=Iridovirus and Microsporidian Linked to Honey Bee Colony Decline |author1=Jerry J. Bromenshenk |author2=Colin B. Henderson |author3=Charles H. Wick |author4=Michael F. Stanford |author5=Alan W. Zulich |author6=Rabih E. Jabbour |author7=Samir V. Deshpande |author8=Patrick E. McCubbin |author9=Robert A. Seccomb |author10=Phillip M. Welch |author11=Trevor Williams |author12=David R. Firth |author13=Evan Skowronski |author14=Margaret M. Lehmann |author15=Shan L. Bilimoria |author16=Joanna Gress |author17=Kevin W. Wanner |author18=Robert A. Cramer Jr |date=6 October 2010 |journal=PLOS ONE |doi=10.1371/journal.pone.0013181 |volume=5 |issue = 10|pages=e13181 |pmid=20949138 |pmc=2950847|bibcode=2010PLoSO...513181B |doi-access=free }}</ref><ref>[https://www.telegraph.co.uk/news/1545516/Honey-bees-in-US-facing-extinction.html "Honey bees in US facing extinction"] {{webarchive|url=https://web.archive.org/web/20080906233434/http://www.telegraph.co.uk/news/1545516/Honey-bees-in-US-facing-extinction.html |date=6 September 2008 }}, ''[[The Daily Telegraph]]'' (London), 14 March 2007.</ref> Winter losses increased to about 1/3.<ref>Benjamin, Alison (2 May 2010) [https://www.theguardian.com/environment/2010/may/02/food-fear-mystery-beehives-collapse Fears for crops as shock figures from America show scale of bee catastrophe] {{webarchive|url=https://web.archive.org/web/20131204232310/http://www.theguardian.com/environment/2010/may/02/food-fear-mystery-beehives-collapse |date=4 December 2013 }}. ''[[The Observer]]'' (London).</ref><ref>{{cite web|url=https://www.sciencedaily.com/releases/2008/05/080509111955.htm |title=Beekeepers Report Continued Heavy Losses From Colony Collapse Disorder |publisher=Sciencedaily.com |date=12 May 2008 |access-date=22 June 2010| archive-url= https://web.archive.org/web/20100731084827/https://www.sciencedaily.com/releases/2008/05/080509111955.htm| archive-date= 31 July 2010 | url-status=live}}</ref> ''Varroa'' mites were thought to be responsible for about half the losses.<ref>{{cite news | url=http://www.rts.ch/info/sciences-tech/4011954-hiver-fatal-pour-la-moitie-des-colonies-d-abeilles-en-suisse.html | title=Hiver fatal pour la moitié des colonies d'abeilles en Suisse | publisher=Radio Télévision Suisse | date=22 May 2012 | access-date=22 May 2012 | url-status=live | archive-url=https://web.archive.org/web/20121112095201/http://www.rts.ch/info/sciences-tech/4011954-hiver-fatal-pour-la-moitie-des-colonies-d-abeilles-en-suisse.html | archive-date=12 November 2012}}</ref>
==== In the European Union ====
Apart from colony collapse disorder, losses outside the US have been attributed to causes including pesticide seed dressings, using [[neonicotinoid]]s such as [[clothianidin]], [[imidacloprid]] and [[thiamethoxam]].<ref>{{cite journal |author=Storkstad, Erik |s2cid=206597443 |title=Field Research on Bees Raises Concern About Low-Dose Pesticides |journal=Science |page=1555 |volume= 335 |date=30 March 2012|doi= 10.1126/science.335.6076.1555|issue= 6076 |pmid=22461580|bibcode=2012Sci...335.1555S }}</ref><ref>{{cite web |url=http://www.efsa.europa.eu/en/press/news/130116 |title=EFSA identifies risks to bees from neonicotinoids &#124; European Food Safety Authority |website=Efsa.europa.eu |date=20 September 2012 |access-date=16 March 2016 |url-status=live |archive-url=https://web.archive.org/web/20150728203252/http://www.efsa.europa.eu/en/press/news/130116 |archive-date=28 July 2015}}</ref> From 2013 the [[European Union]] restricted some pesticides to stop bee populations from declining further.<ref>{{cite news | url= http://www.3news.co.nz/EU-moves-to-protect-bees-from-pesticides/tabid/1160/articleID/296028/Default.aspx | work= 3 News NZ | title= EU moves to protect bees | date= 30 April 2013 | url-status=live | archive-url= https://web.archive.org/web/20130729145145/http://www.3news.co.nz/EU-moves-to-protect-bees-from-pesticides/tabid/1160/articleID/296028/Default.aspx | archive-date= 29 July 2013}}</ref> In 2014 the [[Intergovernmental Panel on Climate Change]] report warned that bees faced increased risk of extinction because of [[global warming]].<ref>Gosden, Emily (29 March 2014) [https://www.telegraph.co.uk/earth/earthnews/10730667/Bees-and-the-crops-they-pollinate-are-at-risk-from-climate-change-IPCC-report-to-warn.html Bees and the crops they pollinate are at risk from climate change, IPCC report to warn] {{webarchive |url=https://web.archive.org/web/20140829051520/http://www.telegraph.co.uk/earth/earthnews/10730667/Bees-and-the-crops-they-pollinate-are-at-risk-from-climate-change-IPCC-report-to-warn.html |date=29 August 2014 }} ''The Daily Telegraph'' (London). Retrieved 30 March 2014</ref> In 2018 the European Union decided to ban field use of all three major neonicotinoids; they remain permitted in veterinary, greenhouse, and vehicle transport usage.<ref>{{cite news |last1=Carrington |first1=Damian |title=EU agrees total ban on bee-harming pesticides |url=https://www.theguardian.com/environment/2018/apr/27/eu-agrees-total-ban-on-bee-harming-pesticides |work=[[The Guardian]] |date=27 April 2018}}</ref>
==== Raising native plants ====
Farmers have focused on alternative solutions to mitigate these problems. By raising native plants, they provide food for native bee pollinators like ''[[Lasioglossum vierecki]]''<ref name="Farming for native bees">{{cite web |url=http://mysare.sare.org/mySARE/ProjectReport.aspx?do=viewRept&pn=LNE07-261&y=2011&t=1 |title=Farming for native bees |date=2011|website=Sustainable Agriculture Research & Education |access-date=4 November 2015 |archive-url=https://web.archive.org/web/20150930233624/http://mysare.sare.org/mySARE/ProjectReport.aspx?do=viewRept&pn=LNE07-261&y=2011&t=1 |archive-date=30 September 2015|last=Kuehn|first=Faith }}</ref> and ''[[Lasioglossum leucozonium|L. leucozonium]]'',<ref name="An Assessment of Non-Apis Bees as Fruit and Vegetable Crop Pollinators in Southwest Virginia">Adamson, Nancy Lee. [http://www.step-project.net/NPDOCS/Adamson_NL_D_2011.pdf An Assessment of Non-Apis Bees as Fruit and Vegetable Crop Pollinators in Southwest Virginia] {{webarchive|url=https://web.archive.org/web/20151120230411/http://www.step-project.net/NPDOCS/Adamson_NL_D_2011.pdf |date=20 November 2015}}. Diss. 2011. Web. 15 October 2015.</ref> leading to less reliance on honey bee populations.
<gallery widths="200px" heights="200px">
File:Peponapis pruinosaCane-12.JPG|[[Squash bee]]s (Apidae) are important pollinators of [[Cucurbita|squashes]] and [[cucumber]]s.
File:A bee covered with pollen.jpg|Bee covered in pollen
</gallery>
===As food producers===
Honey is a natural product produced by bees and stored for their own use, but its sweetness has always appealed to humans. Before domestication of bees was even attempted, humans were raiding their nests for their honey. Smoke was often used to subdue the bees and such activities are depicted in [[Cave painting|rock paintings]] in Spain dated to 15,000 BC.<ref name=EHSTMNWC/>
Honey bees are used commercially to produce [[honey]].<ref>{{cite book |url=http://babel.hathitrust.org/cgi/pt?id=wu.89094204153;view=1up;seq=3 |publisher=US Department of Agriculture, Farmers' Bulletin, No. 653 |author1=Hunt, C.L. |author2=Atwater, H.W. |date=7 April 1915 |title=Honey and Its Uses in the Home |access-date=14 July 2015}}</ref> They also produce some substances used as dietary supplements with possible health benefits, pollen,<ref>{{cite web| url=http://edis.ifas.ufl.edu/AA158| archive-url=https://web.archive.org/web/20070113100544/http://edis.ifas.ufl.edu/AA158| archive-date=13 January 2007| title=Producing Pollen | author=Sanford, Malcolm T. | publisher=University of Florida, Institute of Food and Agricultural Sciences | access-date=15 July 2015}}</ref> [[propolis]],<ref>{{cite web |url=https://www.nlm.nih.gov/medlineplus/druginfo/natural/390.html |title=Propolis:MedlinePlus Supplements |publisher=U.S. National Library of Medicine |date=19 January 2012 |url-status=live |archive-url=https://web.archive.org/web/20160630040550/https://www.nlm.nih.gov/medlineplus/druginfo/natural/390.html |archive-date=30 June 2016}}</ref> and [[royal jelly]],<ref>{{cite journal | author=European Food Safety Authority (EFSA) Panel on Dietetic Products, Nutrition and Allergies | journal=EFSA Journal | year=2011 | volume=9 | issue=4 | page=2083 | title=Scientific Opinion | doi=10.2903/j.efsa.2011.2083 | doi-access=free }}</ref> though all of these can also cause [[allergy|allergic reactions]].
===As food===
Bees are considered [[edible insects]]. People in some countries [[entomophagy|eat insects]], including the larvae and pupae of bees, mostly stingless species. They also gather larvae, pupae and surrounding cells, known as [[bee brood]], for consumption.<ref name=Holland2013>{{cite web |url=http://news.nationalgeographic.com/news/2013/13/130514-edible-insects-entomophagy-science-food-bugs-beetles/ |title=U.N. Urges Eating Insects: 8 Popular Bugs to Try |author=Holland, Jennifer |date=14 May 2013 |work=National Geographic |access-date=16 July 2015 |url-status=dead |archive-url=https://web.archive.org/web/20150716110446/http://news.nationalgeographic.com/news/2013/13/130514-edible-insects-entomophagy-science-food-bugs-beetles/ |archive-date=16 July 2015}}</ref> In the [[Indonesia]]n dish ''[[botok|botok tawon]]'' from Central and East [[Java]], bee larvae are eaten as a companion to [[rice]], after being mixed with shredded [[coconut]], wrapped in [[banana]] leaves, and steamed.<ref name=Tasty2015>{{cite web | title=Botok Tempe Tahu Teri (Botok Tempe Tofu Anchovy) | publisher=Tasty Indonesian Food | url=http://tasty-indonesian-food.com/indonesian-food-recipes/tahu-tempe-vegetables/botok-tempe-tahu-teri/ | access-date=22 June 2015 | url-status=live | archive-url=https://web.archive.org/web/20150626145342/http://tasty-indonesian-food.com/indonesian-food-recipes/tahu-tempe-vegetables/botok-tempe-tahu-teri/ | archive-date=26 June 2015}} (This particular Botok recipe uses anchovies, not bees)</ref><ref name=Sayangi2015>{{cite web |last1=Haris |first1=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=22 June 2015 |language=id |date=6 December 2013 |url-status=dead |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}}</ref>
Bee brood (pupae and larvae) although low in [[calcium]], has been found to be high in [[protein]] and [[carbohydrate]], and a useful source of [[phosphorus]], [[magnesium]], [[potassium]], and trace minerals [[iron]], [[zinc]], [[copper]], and [[selenium]]. In addition, while bee brood was high in fat, it contained no fat soluble vitamins (such as A, D, and E) but it was a good source of most of the water-soluble [[B vitamins]] including [[choline]] as well as vitamin C. The fat was composed mostly of [[Saturated fat|saturated]] and [[monounsaturated fatty acids]] with 2.0% being [[polyunsaturated fatty acids]].<ref>{{cite journal |title=Nutrient Composition of Bee Brood and its Potential as Human Food |author=Fink, Mark D. |publisher=Taylor & Francis, Ecology Food and Nutrition|date=2007| doi=10.1080/03670240500187278 |volume=44 |issue = 4|journal=Ecology of Food and Nutrition|pages=257–270|s2cid=84191573 }}</ref><ref>{{cite journal |title=Standard methods for Apis mellifera brood as human food |publisher=Taylor & Francis, Journal of Apicultural Research |author=Annette Bruun Jensen |date=2016 |doi=10.1080/00218839.2016.1226606 |journal=Journal of Apicultural Research |volume = 58|issue = 2|pages=1–28|doi-access=free }}</ref>
<gallery widths="200px" heights="145px">
File:Botoktawon.jpg|Bee larvae as food in the [[Java]]nese dish ''[[botok|botok tawon]]''
File:Fried bees dish.jpg|Fried whole bees served in a Ukrainian restaurant
</gallery>
=== As alternative medicine ===
[[Apitherapy]] is a branch of [[alternative medicine]] that uses honey bee products, including [[raw honey]], [[royal jelly]], pollen, [[propolis]], [[beeswax]] and [[apitoxin]] (Bee [[venom]]).<ref name=MedicineWorld>{{cite web |url=http://medicineworld.org/alternative/apitherapy/what-is-apitherapy.html |title=What is apitherapy? |publisher=MedicineWorld.Org |access-date=20 January 2018 |url-status=live |archive-url=https://web.archive.org/web/20150618182834/http://medicineworld.org/alternative/apitherapy/what-is-apitherapy.html |archive-date=18 June 2015}}</ref> The claim that apitherapy treats cancer, which some proponents of apitherapy make, remains unsupported by [[evidence-based medicine]].<ref name=Cass>{{cite book |last1=Barry R. |first1=Cassileth |author-link=Barrie R. Cassileth |title=The Complete Guide to Complementary Therapies in Cancer Care: Essential Information for Patients, Survivors and Health Professionals |chapter-url=https://books.google.com/books?id=J6kLNKw5baYC&pg=PA221 |year=2011 |publisher=World Scientific |isbn=978-981-4335-66-9 |pages=221–224 |chapter=Chapter 36: Apitherapy |url-status=live |archive-url=https://web.archive.org/web/20170307071311/https://books.google.com/books?id=J6kLNKw5baYC&pg=PA221 |archive-date=7 March 2017}}</ref><ref name=Ades2009>{{cite book |chapter-url=https://books.google.com/books?id=E8aHAAAACAAJ&pg=704 |title=American Cancer Society Complete Guide to Complementary and Alternative Cancer Therapies |publisher=American Cancer Society |year=2009 |isbn=978-0-944235-71-3 |editor1-last=Ades |editor1-first=Terri B. |edition=2nd |pages=[https://archive.org/details/americancancerso0000unse/page/704 704–708] |chapter=Chapter 9: Pharmacologic and Biologic Therapies |editor2-last=Russel |editor2-first=Jill |url=https://archive.org/details/americancancerso0000unse/page/704 }}</ref>
=== Stings ===
The painful [[Bee sting|stings of bees]] are mostly associated with the poison gland and the [[Dufour's gland]] which are abdominal exocrine glands containing various chemicals. In ''[[Lasioglossum leucozonium]]'', the Dufour's Gland mostly contains [[octadecanolide]] as well as some [[eicosanolide]]. There is also evidence of n-triscosane, n-[[heptacosane]],<ref name=":4" /> and 22-docosanolide.<ref name=":7">{{cite journal|title=Systematic relationship of halictinae bees based on the pattern of macrocyclic lactones in the Dufour gland secretion|doi=10.1016/0020-1790(82)90004-X|volume=12|issue=2|pages=161–170|journal=Insect Biochemistry|year=1982 | last1=Johansson | first1=Ingela}}</ref> However, the secretions of these glands could also be used for nest construction.<ref name=":4">{{Cite journal|title=Chemistry of the dufour's gland secretion of halictine bees|last1=Hefetz|first1=Abraham|date=1978|journal=Comparative Biochemistry and Physiology B|doi=10.1016/0305-0491(78)90229-8|last2=Blum|first2=Murray|last3=Eickwort|first3=George|last4=Wheeler|first4=James|issue=1|volume=61|pages=129–132}}</ref>
== Bees population decline ==
{{Further|Pollinator decline}}
Over the last half century, there has been a general decline in the species richness of wild bees and other pollinators, probably attributable to stress from increased parasites and disease, the use of pesticides, and a general decrease in the number of wild flowers. Climate change probably exacerbates the problem.<ref>{{cite journal |author=Goulson, Dave; Nicholls, Elizabeth; Botías, Cristina; Rotheray, Ellen L. |year=2015 |title=Bee declines driven by combined stress from parasites, pesticides, and lack of flowers |journal=Science |volume=347 |issue=6229 |pages=1255957 |doi=10.1126/science.1255957|pmid=25721506 |s2cid=206558985 |doi-access=free }}</ref> This is a major cause of concern, as it can cause biodiversity loss and ecosystem degradation as well as increase climate change.<ref>{{cite web |title=Why bees are climate heroes |url=https://www.worldwildlife.org/stories/why-bees-are-climate-heroes |website=World Wildlife Fund |access-date=3 June 2024}}</ref>
==See also==
* [[Australian native bees]]
* [[Fear of bees | Fear of bees (apiphobia)]]
* [[Superorganism]]
* [[World Bee Day]]
== Explanatory notes ==
{{notelist}}
==References==
{{Reflist}}
==External links==
{{Wikiquote|Bees}}
{{Commons category|Anthophila|Anthophila<br>(Bees)}}
{{Wikispecies|Apoidea}}
{{Wikibooks|Beekeeping}}
* {{cite web | title=Bees  | website=Encyclopedia of Life | url=https://eol.org/pages/677 }}
* [https://www.discoverlife.org/20/q?search=Apoidea "Apoidea"] at All Living Things{{snd}}images, identification guides, and maps of bees
* [https://web.archive.org/web/20190411210512/http://cache.ucr.edu/~heraty/beepage.html Bee Genera of the World]
* [https://bugguide.net/node/view/8267 Anthophila (Apoidea) – Bees]{{snd}}North American species of bees at [[BugGuide]]
* [https://bugguide.net/node/view/475348 Native Bees of North America] at [[BugGuide]]
* [https://www.science.org/doi/10.1126/science.1255957 "Bee declines driven by combined stress from parasites, pesticides, and lack of flowers"]{{snd}}''[[Science (journal)|Science]]''
{{Hymenoptera|2}}
{{Heraldic creatures}}
{{Insects in culture}}
{{Taxonbar|from=Q7391}}
{{Authority control}}
[[Category:Bees| ]]
<!-- [[Category:Pollinator insects]] "category:Bees" is already listed here -->
[[Category:Extant Early Cretaceous first appearances]]