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==Ecology== === Floral relationships === Most bees are [[polylectic]] (generalist) meaning they collect pollen from a range of flowering plants, but some are [[oligolege]]s (specialists), in that they only gather pollen from one or a few species or genera of closely related plants.<ref name=Waser2006>{{cite book |author=Waser, Nickolas M. |title=Plant-Pollinator Interactions: From Specialization to Generalization |url=https://books.google.com/books?id=Fbl5c9fUxTIC&pg=PA110 |year=2006 |publisher=University of Chicago Press |isbn=978-0-226-87400-5 |pages=110– |url-status=live |archive-url=https://web.archive.org/web/20180328155201/https://books.google.com/books?id=Fbl5c9fUxTIC&pg=PA110 |archive-date=28 March 2018}}</ref> In Melittidae and Apidae we also find a few genera that are highly specialized for collecting plant oils both in addition to, and instead of, nectar, which is mixed with pollen as larval food.<ref>{{cite journal | pmc=2838259 | year=2010 | last1=Renner | first1=S. S. | last2=Schaefer | first2=H. | title=The evolution and loss of oil-offering flowers: New insights from dated phylogenies for angiosperms and bees | journal=Philosophical Transactions of the Royal Society of London. Series B, Biological Sciences | volume=365 | issue=1539 | pages=423–435 | doi=10.1098/rstb.2009.0229 | pmid=20047869 }}</ref> Male orchid bees in some species gather aromatic compounds from [[orchid]]s, which is one of the few cases where male bees are effective pollinators. Bees are able to sense the presence of desirable flowers through ultraviolet patterning on flowers, floral odors,<ref name=Dafni>{{cite book |author1=Dafni, Amots |author2=Hesse, Michael |author3=Pacini, Ettore |title=Pollen and Pollination |url=https://books.google.com/books?id=-M7yCAAAQBAJ&pg=PA80 |year=2012 |publisher=Springer Science & Business Media |isbn=978-3-7091-6306-1 |page=80 |url-status=live |archive-url=https://web.archive.org/web/20180328155201/https://books.google.com/books?id=-M7yCAAAQBAJ&pg=PA80 |archive-date=28 March 2018}}</ref> and even electromagnetic fields.<ref>{{cite journal|last1=Suttona|first1=Gregory P.|last2=Clarkea|first2=Dominic|last3=Morleya|first3=Erica L.|last4=Robert|first4=Daniel|title=Mechanosensory hairs in bumblebees (Bombus terrestris) detect weak electric fields|journal=PNAS|date=2016|volume=113|issue=26|pages=7261–7265|doi=10.1073/pnas.1601624113|pmid=27247399|pmc=4932954|bibcode=2016PNAS..113.7261S |doi-access=free}}</ref> Once landed, a bee then uses nectar quality<ref name=Dafni /> and pollen taste<ref>{{cite journal|last1=Muth|first1=Felicity|last2=Francis|first2=Jacob S.|last3=Leonard|first3=Anne S.|title=Bees use the taste of pollen to determine which flowers to visit|journal=Biology Letters|date=2016|volume=12|issue=7|page=20160356|doi=10.1098/rsbl.2016.0356|pmid=27405383|pmc=4971173}}</ref> to determine whether to continue visiting similar flowers. In rare cases, a [[plant]] species may only be effectively pollinated by a single bee species, and some plants are [[endangered species|endangered]] at least in part because their pollinator is also threatened. But, there is a pronounced tendency for oligolectic bees to be associated with common, widespread plants visited by multiple pollinator species. For example, the [[Larrea tridentata|creosote bush]] in the arid parts of the United States southwest is associated with some 40 oligoleges.<ref>{{cite journal |author=Hurd, P.D. Jr. |author2=Linsley, E.G. |year=1975 |title=The principal ''Larrea'' bees of the southwestern United States |journal=Smithsonian Contributions to Zoology |volume=193 |pages=1–74 |doi=10.5479/si.00810282.193 |issue=193}}</ref> ===As mimics and models=== [[File:Bombylius major on flower.jpg|thumb|left|The bee-fly ''[[Bombylius major]]'', a [[Batesian mimic]] of bees, taking nectar and pollinating a flower]] {{main|Mimicry|Batesian mimicry|Müllerian mimicry}} [[File:Ophrys apifera flower1.jpg|thumb|upright|[[Ophrys apifera|Bee orchid]] lures male bees to attempt to mate with the flower's lip, which resembles a bee perched on a pink flower.]] Many bees are [[aposematic]]ally 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]], [[Asilidae|robber flies]] and [[hoverflies]],<ref name="ThorpHorning1983">{{cite book |last1=Thorp |first1=Robbin W. |last2=Horning |first2=Donald S. |last3=Dunning |first3=Lorry L. |title=Bumble Bees and Cuckoo Bumble Bees of California (Hymenoptera, Apidae) |url=https://books.google.com/books?id=v1eJ3fWwshIC&pg=PA9 |year=1983 |publisher=University of California Press |isbn=978-0-520-09645-5 |page=9 |quote=Of the forms of mimicry, two relate to [[Bombini]]. Batesian mimicry .. is exemplified by members of several families of flies: [[Syrphidae]], [[Asilidae]], [[Horse-fly|Tabanidae]], [[Oestridae]], and [[Bombyliidae]] (Gabritschevsky, 1926). |url-status=live |archive-url=https://web.archive.org/web/20170105171749/https://books.google.com/books?id=v1eJ3fWwshIC&pg=PA9 |archive-date=5 January 2017}}</ref> all of which gain a measure of protection by superficially looking and behaving like bees.<ref name="ThorpHorning1983"/> Bees are themselves [[Müllerian mimicry|Müllerian mimics]] of other aposematic insects with the same color scheme, including [[wasp]]s, [[Lycidae|lycid]] and other beetles, and many butterflies and moths ([[Lepidoptera]]) which are themselves distasteful, often through acquiring bitter and poisonous chemicals from their plant food. All the Müllerian mimics, including bees, benefit from the reduced risk of predation that results from their easily recognized warning coloration.<ref>{{cite book |last1=Cott |first1=Hugh |author-link=Hugh B. Cott |title=Adaptive Coloration in Animals |title-link=Adaptive Coloration in Animals |date=1940 |publisher=Oxford University Press |pages=196, 403 and passim}}</ref> Bees are also mimicked by plants such as the [[Ophrys apifera|bee orchid]] which imitates both the appearance and the scent of a female bee; male bees attempt to mate ([[pseudocopulation]]) with the furry lip of the flower, thus pollinating it.<ref>{{cite web |title=Bee Orchids and Insect Mimicry |url=http://www.nhm.ac.uk/nature-online/life/plants-fungi/bee-orchids/ |publisher=Natural History Museum |access-date=1 July 2015 |url-status=live |archive-url=https://web.archive.org/web/20150708203045/http://www.nhm.ac.uk/nature-online/life/plants-fungi/bee-orchids/ |archive-date=8 July 2015}}</ref> ===As brood parasites=== [[File:Bumblebee January 2008-4.jpg|thumb|left|''[[Bombus vestalis]]'', a [[brood parasite]] of the bumblebee ''[[Bombus terrestris]]'']] {{main|Brood parasite|Nest usurpation}} [[Brood parasite]]s occur in several bee families including the [[apidae|apid]] subfamily [[Nomadinae]].<ref>{{cite web|title=Obligate Brood Parasitism|url=http://www.aculeataresearch.com/index.php/cuckoo-behavior/52-obligate-brood-parasitism|publisher=Aculeata Research Group|access-date=30 June 2015|url-status=live|archive-url=https://web.archive.org/web/20150707053542/http://www.aculeataresearch.com/index.php/cuckoo-behavior/52-obligate-brood-parasitism|archive-date=7 July 2015}}</ref> Females of these species lack pollen collecting structures (the [[scopa (biology)|scopa]]) and do not construct their own nests. They typically enter the nests of pollen collecting species, and lay their eggs in cells provisioned by the host bee. When the "cuckoo" bee larva hatches, it consumes the host larva's pollen ball, and often the host egg also.<ref>{{cite web |title=Brood Parasitism |url=http://www.amentsoc.org/insects/glossary/terms/brood-parasitism |publisher=Amateur Entomologists' Society |access-date=30 June 2015 |url-status=live |archive-url=https://web.archive.org/web/20150702165918/http://www.amentsoc.org/insects/glossary/terms/brood-parasitism |archive-date=2 July 2015}}</ref> In particular, the Arctic bee species, ''[[Bombus hyperboreus]]'' is an aggressive species that attacks and enslaves other bees of the same subgenus. However, unlike many other bee brood parasites, they have pollen baskets and often collect pollen.<ref>Gjershaug, Jan Ove (5 June 2009). "The social parasite bumblebee Bombus hyperboreus Schönherr, 1809 usurp nest of Bombus balteatus Dahlbom, 1832 (Hymenoptera, Apidae) in Norway" (PDF). ''Norwegian Journal of Entomology'' '''56'''(1): 28–31. Retrieved 26 September 2015.</ref> In Southern Africa, hives of African honeybees (''A. mellifera scutellata'') are being destroyed by parasitic workers of the Cape honeybee, ''A. m. capensis''. These lay [[diploid]] eggs ("[[thelytoky]]"), escaping normal [[worker policing]], leading to the colony's destruction; the parasites can then move to other hives.<ref>{{cite book |last1=Gullan |first1=P. J. |last2=Cranston |first2=P. S. |title=The Insects: An Outline of Entomology |date=2014 |publisher=Wiley Blackwell |isbn=978-1-118-84615-5 |edition=5th |page=347}}</ref> The [[cuckoo bee]]s in the ''[[List of bumblebee species|Bombus]]'' subgenus ''Psithyrus'' are closely related to, and resemble, their hosts in looks and size. This common pattern gave rise to the ecological principle "[[Emery's rule]]". Others parasitize bees in different families, like ''[[Townsendiella]]'', a [[nomadinae|nomadine]] [[apidae|apid]], two species of which are cleptoparasites of the [[Dasypodaidae|dasypodaid]] genus ''[[Hesperapis]]'',<ref>{{cite journal |author1=Rozen, Jerome George |author2=McGinley, Ronald J. |year=1991 |title=Biology and Larvae of the Cleptoparasitic Bee ''Townsendiella pulchra'' and Nesting Biology of its Host ''Hesperapis larreae'' (Hymenoptera, Apoidea) |journal=American Museum Novitates |issue=3005 |hdl=2246/5032 }}</ref> while the other species in the same genus attacks [[halictidae|halictid]] bees.<ref>{{cite book|author1=Moure, Jesus S.|author2=Hurd, Paul David |title=An Annotated Catalog of the Halictid Bees of the Western Hemisphere (Hymenoptera, Halictidae) |url=https://books.google.com/books?id=2iUlSfQt8vEC |year=1987 |publisher=Smithsonian Institution Press |pages=28–29}}</ref> === Nocturnal bees === Four bee families ([[Andrenidae]], [[Colletidae]], [[Halictidae]], and [[Apidae]]) contain some species that are [[crepuscular]]. Most are tropical or subtropical, but some live in arid regions at higher latitudes. These bees have greatly enlarged [[ocellus|ocelli]], which are extremely sensitive to light and dark, though incapable of forming images. Some have refracting superposition compound eyes: these combine the output of many elements of their compound eyes to provide enough light for each retinal photoreceptor. Their ability to fly by night enables them to avoid many predators, and to exploit flowers that produce nectar only or also at night.<ref>{{cite journal |last1=Warrant |first1=Eric J. |title=Seeing in the dark: vision and visual behaviour in nocturnal bees and wasps |journal=Journal of Experimental Biology |date=June 2008 |volume=211 |issue=11 |pages=1737–1746 |doi=10.1242/jeb.015396 |pmid=18490389|doi-access=free |bibcode=2008JExpB.211.1737W }}</ref> ===Predators, parasites and pathogens=== {{further|Diseases of the honey bee}} [[File:Pair of Merops apiaster feeding detail.jpg|thumb|right|The bee-eater, ''[[Merops apiaster]]'', specializes in feeding on bees; here a male catches a nuptial gift for his mate.]] Vertebrate predators of bees include [[bee-eater]]s, [[shrikes]] and [[Old World flycatcher|flycatcher]]s, which make short sallies to catch insects in flight.<ref name="ChittkaThomson2001"/> Swifts and swallows<ref name="ChittkaThomson2001"/> fly almost continually, catching insects as they go. The [[Pernis (bird)|honey buzzard]] attacks bees' nests and eats the larvae.<ref>{{cite web | url=https://www.theguardian.com/world/2013/sep/26/hornet-attacks-kill-18-china | work=The Guardian | title=Hornet attacks kill dozens in China | date=26 September 2013 | access-date=18 June 2015 | url-status=live | archive-url=https://web.archive.org/web/20150906191229/http://www.theguardian.com/world/2013/sep/26/hornet-attacks-kill-18-china | archive-date=6 September 2015}}</ref> The [[greater honeyguide]] interacts with humans by guiding them to the nests of wild bees. The humans break open the nests and take the honey and the bird feeds on the larvae and the wax.<ref>{{cite journal |author=Friedmann, Herbert |year=1955 |title=The Honey-Guides |journal=Bulletin of the United States National Museum |doi=10.5479/si.03629236.208.1 |pages=1–292 |issue=208|hdl=10088/10101 }}</ref> Among mammals, predators such as the [[badger]] dig up bumblebee nests and eat both the larvae and any stored food.<ref>{{cite web |title=What predators do bumblebees have? |url=http://bumblebeeconservation.org/about-bees/faqs/bumblebee-predators/ |publisher=Bumblebee Conservation Trust |access-date=29 June 2015 |url-status=dead |archive-url=https://web.archive.org/web/20150629025444/http://bumblebeeconservation.org/about-bees/faqs/bumblebee-predators/ |archive-date=29 June 2015}}</ref> [[File:Wasp and bee August 2008-2.jpg|thumb|left|The beewolf ''[[Philanthus triangulum]]'' paralysing a bee with its sting]] Specialist ambush predators of visitors to flowers include [[crab spiders]], which wait on flowering plants for pollinating insects; [[Hemiptera|predatory bugs]], and [[praying mantis]]es,<ref name="ChittkaThomson2001">{{cite book |last1=Chittka |first1=Lars |last2=Thomson |first2=James D. |title=Cognitive Ecology of Pollination: Animal Behaviour and Floral Evolution |url=https://books.google.com/books?id=g2Km4B6n-mQC&pg=PA215 |date=28 May 2001 |publisher=Cambridge University Press |isbn=978-1-139-43004-3 |pages=215–216 |url-status=live |archive-url=https://web.archive.org/web/20161224104844/https://books.google.com/books?id=g2Km4B6n-mQC&pg=PA215 |archive-date=24 December 2016}}</ref> some of which (the [[flower mantis]]es of the tropics) wait motionless, [[aggressive mimicry|aggressive mimics]] camouflaged as flowers.<ref>{{cite web |last1=Choi |first1=Charles Q. |title=Found! First Known Predator To Lure Prey By Mimicking Flowers |url=http://www.livescience.com/41605-predator-lures-prey-by-mimicking-flowers.html |publisher=LiveScience |access-date=2 July 2015 |date=30 November 2013 |quote=the color of the orchid mantis was indistinguishable from 13 species of wild flowers in the areas the predator lived. ... The orchid mantis is unique in that the mantis itself is the attractive stimulus. |url-status=live |archive-url=https://web.archive.org/web/20150630181114/http://www.livescience.com/41605-predator-lures-prey-by-mimicking-flowers.html |archive-date=30 June 2015}}</ref> [[Beewolf|Beewolves]] are large wasps that habitually attack bees;<ref name="ChittkaThomson2001"/> the [[ethologist]] [[Niko Tinbergen]] estimated that a single colony of the beewolf ''[[Philanthus triangulum]]'' might kill several thousand honeybees in a day: all the prey he observed were honeybees.<ref>{{cite book | author=Tinbergen, Niko | title=Curious Naturalists | publisher=Methuen | year=1958 | page=21}}</ref> Other predatory insects that sometimes catch bees include [[Asilidae|robber flies]] and [[dragonfly|dragonflies]].<ref name="ChittkaThomson2001"/> Honey bees are affected by parasites including [[Acarapis woodi|tracheal]] and ''[[Varroa]]'' [[mite]]s.<ref>{{cite web |title=Honey Bee Disorders: Honey Bee Parasites |url=http://www.ent.uga.edu/bees/disorders/honey-bee-parasites.html |publisher=University of Georgia |access-date=29 June 2015 |url-status=live |archive-url=https://web.archive.org/web/20150701225054/http://www.ent.uga.edu/bees/disorders/honey-bee-parasites.html |archive-date=1 July 2015}}</ref> However, some bees are believed to have a mutualistic relationship with mites.<ref name="Pavel B 2007"/> Some mites of genus ''[[Tarsonemus]]'' are associated with bees. They live in bee nests and ride on adult bees for dispersal. They are presumed to feed on fungi, nest materials or pollen. However, the impact they have on bees remains uncertain.<ref>{{Cite web |title=Tarsonemus {{!}} Bee Mite ID |url=http://idtools.org/id/mites/beemites/factsheet.php?name=15293 |access-date=25 August 2022 |website=idtools.org}}</ref>
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