Bee

The immediate ancestors of bees were stinging wasps in the family Crabronidae, which were 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 evolutionary scenario may have occurred within the vespoid wasps, where the pollen wasps 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 West Gondwana, just prior to its breakup into South America and Africa. The supercontinent is thought to have been a largely xeric environment at this time; modern bee diversity hotspots are also in xeric and seasonal temperate environments, suggesting strong niche conservatism among bees ever since their origins.{{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 }}

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 plants 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. 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.

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.{{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}}{{efn|Triassic nests in a petrified forest in Arizona, implying that bees evolved much earlier, are now thought to be beetle borings.{{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 }}}} A second extinction event among bees is thought to have occurred due to rapid climatic cooling around the 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.

The oldest non-compression bee fossil is Cretotrigona prisca, a corbiculate bee of Late Cretaceous age (~70 mya) found in New Jersey amber.{{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}} A fossil from the early Cretaceous (~100 mya), Melittosphex burmensis, was initially considered "an extinct lineage of pollen-collecting Apoidea sister to the modern bees",{{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}} 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.{{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}}

The Allodapini (within the Apidae) appeared around 53 Mya.{{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 }}

The Colletidae appear as fossils only from the late Oligocene (~25 Mya) to early Miocene.{{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}}

The Melittidae are known from Palaeomacropis eocenicus in the Early Eocene.{{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 }}

The Megachilidae are known from trace fossils (characteristic leaf cuttings) from the Middle Eocene.{{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 |doi=10.1111/j.1475-4983.2008.00787.x |url=https://www.researchgate.net/publication/229562255 |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|hdl=11336/100644 |hdl-access=free }}

The Andrenidae are known from the Eocene-Oligocene boundary, around 34 Mya, of the Florissant shale.{{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 }}

The Halictidae first appear in the Early Eocene{{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 }} with species{{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}}{{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 }} found in amber. The Stenotritidae are known from fossil brood cells of Pleistocene age.{{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}}

File:Amegilla on long tube of Acanthus ilicifolius flower.jpg, like this Amegilla species (Apidae) on Acanthus ilicifolius.]]

{{further|Coevolution}}

The earliest animal-pollinated flowers were shallow, cup-shaped blooms pollinated by insects such as beetles, so the 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{{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}} such as nectar and longer tubes, and bees developed longer tongues to extract the nectar. Bees also developed structures known as 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.{{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.}} The appearance of these structures drove the adaptive radiation of the angiosperms, and, in turn, bees themselves.{{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}} Bees 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 mutualistic.{{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|bibcode=2009ANat..173..841B |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 }}{{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 }}

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 paraphyletic. In their study, the placement of the monogeneric Heterogynaidae was uncertain. The small family Mellinidae was not included in this analysis.{{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}}

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 Pemphredoninae, was recovered as the most sister family to bees.{{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 |display-authors=5 |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 }}

{{clade| style=font-size:85%;line-height:85%

|label1=Apoidea

|1={{clade

|1=Ampulicidae (Cockroach wasps) 70px

|2={{clade

|1=Astatidae 70px

|2={{clade

|1={{clade

|1=Bembicidae 70px

|2={{clade

|1=Sphecidae (sensu stricto) 70px

|2={{clade

|1={{clade

|1=Crabronidae (sensu stricto) 70px

|2=Mellinidae 70px

}} }} }} }}

|2={{clade

|1={{clade

|1=Pemphredonidae 70px

|2=Philanthidae 70px}}

|2={{clade

|1=Psenidae 70px

|2={{clade

|1={{clade

|1=Ammoplanidae

|2=Anthophila (bees) 70px

}} }} }} }} }}

}}

}}

}}

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.{{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 }} 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 70px

|2={{clade

|label1={{nowrap|long-tongued bees}}

|1={{clade

|1=Apidae (inc. honeybees, cuckoo bees, carpenter bees) ≈87 Mya 70px

|2=Megachilidae (mason, leafcutter bees) ≈50 Mya 70px

}}

|label2={{nowrap|short-tongued bees}}

|2={{clade

|1=Andrenidae (mining bees) ≈34 Mya 70px

|2={{clade

|1=Halictidae (sweat bees) ≈50 Mya 70px

|2={{clade

|1=Colletidae (plasterer bees) ≈25 Mya 70px

|2=Stenotritidae (large Australian bees) ≈2 Mya 70px

}}

}}

}}

}}

}}

}}

}}

File:European Honeybee (Apis mellifera) lapping mouthparts, showing labium and maxillae..jpg of a honey bee, showing labium and maxillae]]

{{see also|Characteristics of common wasps and bees}}

Bees differ from closely related groups such as wasps by having branched or plume-like setae (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.{{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}}

Bees have the following characteristics:

• 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.
• The antennae usually have 13 segments in males and 12 in females, and are 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.
• The mouthparts are adapted for both chewing and sucking by having both a pair of mandibles and a long proboscis for sucking up nectar.{{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}}
• 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 synchronized 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.

File:Carpenter bee head and compound eyes.jpg, showing antennae, three ocelli, compound eyes, 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|sp=us}}.{{ 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 }} The smallest species may be dwarf stingless bees in the tribe Meliponini whose workers are less than {{convert|2|mm|inch|2|sp=us}} in length.{{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}}

{{further|Haplodiploidy}}

File:Wasp attack.jpgs, along with a dead yellowjacket. Such altruistic behaviour may be favoured by the haplodiploid 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.{{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 }}

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.{{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 }} 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.{{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}}

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.{{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 }} 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.{{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 | 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 }}

File:Bee swarm on fallen tree03.jpg swarm]]

File:Wildbienen.jpg nest in the trunk of a 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.{{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}} The most advanced of these are species with eusocial colonies; these are characterized by cooperative brood care and a division of labour into reproductive and non-reproductive adults, plus overlapping generations.{{Cite book|title=The Insect Societies|last=Wilson|first=Edward O|publisher=Belknap Press of Harvard University Press|year=1971|location=Cambridge, Mass}} This division of labour creates specialized groups within eusocial societies which are called 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) and her daughters (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 wasps; when the castes are morphologically discrete, the system is considered highly eusocial.{{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}}

True honey bees (genus Apis, of which eight species are currently recognized) are highly eusocial, and are among the best known insects. Their colonies are established by 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 bees are a hybrid strain of A. mellifera that escaped from experiments involving crossing European and African subspecies; they are extremely defensive.{{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}}

Stingless bees are also highly eusocial. They practice mass provisioning, with complex nest architecture and perennial colonies also established via swarming.{{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 }}

File:Bumblebee 05.JPG carrying pollen in its pollen baskets (corbiculae)]]

Many bumblebees are eusocial, similar to the eusocial Vespidae such as hornets 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.{{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}} 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.{{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}}

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.{{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 }} Some species are eusocial in parts of their range and solitary in others,{{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 }} or have a mix of eusocial and solitary nests in the same population.{{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 }} The orchid bees (Apidae) include some primitively eusocial species with similar biology. Some 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.{{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}}

File:Megachile rotundata.JPG, cutting circles from acacia leaves]]

Most other bees, including familiar insects such as carpenter bees, 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 wasps have similar behaviours, and a few species of bee scavenge from carcases to feed their offspring. 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 genera which are commonly known by their nesting behavior or preferences, namely: carpenter bees, sweat bees, mason bees, plasterer bees, squash bees, dwarf carpenter bees, leafcutter bees, alkali bees and digger bees.{{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}}

File:Anthidium February 2008-1.jpg (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 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.{{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}}{{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}}

File:Osmia cornifrons.5.1.08.w.jpg Osmia cornifrons nests in a hole in dead wood. Bee "hotels" are often sold for this purpose.]]

While solitary, females each make individual nests.{{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}} Some species, such as the European mason bee Hoplitis anthocopoides,{{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 }} and the Dawson's Burrowing bee, Amegilla dawsoni,{{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 }} 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 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.

File:Bee morning glory visitors.webm flower. A Tumbling flower beetle remains in the flower with a bee visitor.]]

{{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 pupation stage during which the insect undergoes 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.[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] 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 haplodiploid; the sex of a bee is determined by whether or not the egg is fertilized. After mating, a female stores the sperm, and determines which sex is required at the time each individual egg is laid, fertilized eggs producing female offspring and unfertilized eggs, males. Tropical bees may have several generations in a year and no diapause stage.{{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}}{{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}}{{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}}{{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}}

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.{{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}}

In most species, larvae are whitish grubs, roughly oval and bluntly-pointed at both ends. They have 15 segments and spiracles 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 exuviae and breaks out of the cell.

File:Apoidea.jpg|Nest of common carder bumblebee, wax canopy removed to show winged 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

File:Apis mellifera flying.jpg]]

{{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 insects 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".Ingram, Jay (2001) The Barmaid's Brain, Aurum Press, pp. 91–92, {{ISBN|0716741202}}. 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 helicopters.{{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}} In 1996 it was shown that vortices created by many insects' wings helped to provide lift.{{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}} High-speed cinematography{{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}} and robotic mock-up of a bee wing{{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 Dr. Michael H. Dickinson. 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, it flaps approximately 230 times per second, faster than a fruitfly (200 times per second) which is 80 times smaller.{{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 }}

File:Bee_dance.svg (1953) discovered that honey bee workers can 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 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 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.{{cite book |last=von Frisch |first=Karl |title=The Dancing Bees |year=1953 |publisher=Harcourt, Brace & World | pages=93–96}} Bees navigate using spatial memory with a "rich, map-like organization".{{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 }}

The gut of bees is relatively simple, but multiple metabolic strategies exist in the gut microbiota.{{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}} 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 polysaccharides: branching pectin and hemicellulose.{{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}} 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 clades 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 niches.{{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 }}

Although most bee species are nectarivorous and palynivorous, some are not. Particularly unusual are vulture bees in the genus Trigona, which consume carrion and wasp brood, turning meat into a honey-like substance.{{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 }} Drinking guttation drops from leaves is also a source of energy and nutrients.[https://royalsocietypublishing.org/doi/10.1098/rspb.2020.1080 Plant guttation provides nutrient-rich food for insects - Journals]

Most bees are polylectic (generalist) meaning they collect pollen from a range of flowering plants, but some are oligoleges (specialists), in that they only gather pollen from one or a few species or genera of closely related plants.{{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}} 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.{{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 }} Male orchid bees in some species gather aromatic compounds from orchids, 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,{{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}} and even electromagnetic fields.{{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}} Once landed, a bee then uses nectar quality and pollen taste{{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}} 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 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 creosote bush in the arid parts of the United States southwest is associated with some 40 oligoleges.{{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}}

File:Bombylius major on flower.jpg, a Batesian mimic of bees, taking nectar and pollinating a flower]]

{{main|Mimicry|Batesian mimicry|Müllerian mimicry}}

File:Ophrys apifera flower1.jpg lures male bees to attempt to mate with the flower's lip, which resembles a bee perched on a pink flower.]]

Many bees are aposematically colored, 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, robber flies and hoverflies,{{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, 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}} all of which gain a measure of protection by superficially looking and behaving like bees.

Bees are themselves Müllerian mimics of other aposematic insects with the same color scheme, including wasps, 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 recognized warning coloration.{{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}}

Bees are also mimicked by plants such as the 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.{{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}}

File:Bumblebee January 2008-4.jpg, a brood parasite of the bumblebee Bombus terrestris]]

{{main|Brood parasite|Nest usurpation}}

Brood parasites occur in several bee families including the apid subfamily Nomadinae.{{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}} Females of these species lack pollen collecting structures (the 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.{{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}} 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.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.

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.{{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}}

The cuckoo bees in the 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 nomadine apid, two species of which are cleptoparasites of the dasypodaid genus Hesperapis,{{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 }} while the other species in the same genus attacks halictid bees.{{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}}

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 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.{{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 |bibcode=2008JExpB.211.1737W }}

{{further|Diseases of the honey bee}}

File:Pair of Merops apiaster feeding detail.jpg, specializes in feeding on bees; here a male catches a nuptial gift for his mate.]]

Vertebrate predators of bees include bee-eaters, shrikes and flycatchers, which make short sallies to catch insects in flight. Swifts and swallows fly almost continually, catching insects as they go. The honey buzzard attacks bees' nests and eats the larvae.{{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}} 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.{{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 }} Among mammals, predators such as the badger dig up bumblebee nests and eat both the larvae and any stored food.{{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}}

File:Wasp and bee August 2008-2.jpg paralysing a bee with its sting]]

Specialist ambush predators of visitors to flowers include crab spiders, which wait on flowering plants for pollinating insects; predatory bugs, and praying mantises,{{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}} some of which (the flower mantises of the tropics) wait motionless, aggressive mimics camouflaged as flowers.{{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}} Beewolves are large wasps that habitually attack bees; 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.{{cite book | author=Tinbergen, Niko | title=Curious Naturalists | publisher=Methuen | year=1958 | page=21}} Other predatory insects that sometimes catch bees include robber flies and dragonflies. Honey bees are affected by parasites including tracheal and Varroa mites.{{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}} However, some bees are believed to have a mutualistic relationship with mites.

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.{{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}}

{{Main|Bees in mythology}}

File:Plaque bee-goddess BM GR1860.4-123.4.jpg, Rhodes. 7th century BC.]]

Homer's 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.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}} Honey, according to a Greek myth, was discovered by a nymph called Melissa ("Bee"); and honey was offered to the Greek gods from Mycenean times. Bees were also associated with the Delphic oracle and the prophetess was sometimes called a bee.{{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}}

The image of a community of honey bees has been used from ancient to modern times, in Aristotle and Plato; in Virgil and 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.{{cite book |last=Wilson |first=Bee |year=2004 |title=The Hive: the Story of the Honeybee |location=London |publisher=John Murray |isbn=0-7195-6598-7}} In English folklore, bees would be told of important events in the household, in a custom known as "Telling the bees".{{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}}

File:Mrs tittlemouse.jpg's illustration of Babbity Bumble in The Tale of Mrs Tittlemouse, 1910]]

Some of the oldest examples of bees in art are rock paintings in Spain which have been dated to 15,000 BC.{{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}}

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 in the West of London.{{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}} Beatrix Potter's illustrated book The Tale of Mrs Tittlemouse (1910) features Babbity Bumble and her brood (pictured). Kit Williams' 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 (2004), and the 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.

Bees have appeared in films such as Jerry Seinfeld's animated Bee Movie,{{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}} or Eugene Schlusser's A Sting in the Tale (2014). The playwright Laline Paull's fantasy The Bees (2015) tells the tale of a hive bee named Flora 717 from hatching onwards.{{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 }}

{{main|Beekeeping}}

File:Beekeeper.jpg

File:Western honey bee on a honeycomb.jpg

Humans have kept honey bee colonies, commonly in hives, for millennia. 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.{{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}} Simple hives and smoke were used.{{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 }}{{Cite book |title=Animal and Man in Bible Lands |first=F. S. |last=Bodenheimer |publisher=Brill Archive |year=1960 |page=79}}

Among Classical Era authors, beekeeping with the use of smoke is described in Aristotle's History of Animals Book 9.{{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}} The account mentions that bees die after stinging; that workers remove corpses from the hive, and guard it; castes including workers and non-working drones, 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).{{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 }}{{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".}} Beekeeping is described in detail by Virgil in his Georgics; it is mentioned in his Aeneid, and in Pliny's Natural History.

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.Thomas Wildman, A Treatise on the Management of Bees (London, 1768, 2nd edn 1770).{{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}}

{{See also|List of crop plants pollinated by bees|Pollinator decline|Pesticide toxicity to bees}}

Bees play an important role in pollinating flowering plants, and are the major type of pollinator in many ecosystems 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.{{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}}{{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}}

Since the 1970s, 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 decrease in the number of wild flowers. Climate change probably exacerbates the problem.{{cite journal |last1=Goulson |first1=Dave |author1-link=Dave Goulson |author2=Nicholls, Elizabeth |author3=Botías, Cristina |author4=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 }} This is a major cause of concern, as it can cause biodiversity loss and ecosystem degradation as well as increase climate change.{{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}}

Contract pollination has overtaken the role of honey production for beekeepers in many countries. After the introduction of Varroa mites, feral honey bees declined dramatically in the US, though their numbers have since recovered.{{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}}{{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 }} The number of colonies kept by beekeepers declined slightly, through urbanization, systematic pesticide use, 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.{{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}} In 2010 invertebrate iridescent virus and the fungus Nosema ceranae were shown to be in every killed colony, and deadly in combination.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.{{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}}{{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 }}[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. Winter losses increased to about 1/3.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).{{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}} Varroa mites were thought to be responsible for about half the losses.{{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}}

Apart from colony collapse disorder, losses outside the US have been attributed to causes including pesticide seed dressings, using neonicotinoids such as clothianidin, imidacloprid and thiamethoxam.{{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 }}{{cite web |url=http://www.efsa.europa.eu/en/press/news/130116 |title=EFSA identifies risks to bees from neonicotinoids | 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}} From 2013 the European Union restricted some pesticides to stop bee populations from declining further.{{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}} In 2014 the Intergovernmental Panel on Climate Change report warned that bees faced increased risk of extinction because of global warming.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 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.{{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}}

Farmers have focused on alternative solutions to mitigate these problems. By raising native plants, they provide food for native bee pollinators like Lasioglossum vierecki{{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 }} and L. leucozonium,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. leading to less reliance on honey bee populations.

File:Peponapis pruinosaCane-12.JPG|Squash bees (Apidae) are important pollinators of squashes and cucumbers.

File:A bee covered with pollen.jpg|Bee covered in pollen

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 rock paintings in Spain dated to 15,000 BC.

Honey bees are used commercially to produce honey.{{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 |last1=Hunt |first1=C.L. |last2=Atwater |first2=H.W. |date=7 April 1915 |title=Honey and Its Uses in the Home |access-date=14 July 2015}}

Bees are considered edible insects. People in some countries 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.{{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}} In the Indonesian dish 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.{{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){{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}}

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 and monounsaturated fatty acids with 2.0% being polyunsaturated fatty acids.{{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 }}{{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 }}

File:Botoktawon.jpg|Bee larvae as food in the Javanese dish botok tawon

File:Fried bees dish.jpg|Fried whole bees served in a Ukrainian restaurant

Apitherapy is a branch of alternative medicine that uses honey bee products, including raw honey, royal jelly, pollen, propolis, beeswax and apitoxin (Bee venom).{{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}} The claim that apitherapy treats cancer, which some proponents of apitherapy make, remains unsupported by evidence-based medicine.{{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}}{{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 }}

The painful 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,{{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}} and 22-docosanolide.{{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}}

• Australian native bees
• Fear of bees (apiphobia)
• Superorganism
• World Bee Day

{{notelist}}

{{Reflist}}

{{Wikiquote|Bees}}

{{Commons category|Anthophila|Anthophila
(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

{{Hymenoptera|2}}

{{Heraldic creatures}}

{{Insects in culture}}

{{Taxonbar|from=Q7391}}

{{Authority control}}

Category:Extant Early Cretaceous first appearances