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==Sociality== ===Haplodiploid breeding system=== {{further|Haplodiploidy}} [[File:Wasp attack.jpg|thumb|left|upright=1.3<!--size for low image-->|Willing to die for their sisters: worker honey bees killed defending their hive against [[yellowjacket]]s, along with a dead yellowjacket. Such [[altruism|altruistic]] behaviour may be favoured by the [[haplodiploid]] [[Sex-determination system|sex determination]] system of bees.]] According to [[inclusive fitness]] theory, organisms can gain fitness not just through increasing their own reproductive output, but also that of close relatives. In evolutionary terms, individuals should help relatives when ''Cost < Relatedness * Benefit''. The requirements for eusociality are more easily fulfilled by [[haplodiploid]] species such as bees because of their unusual relatedness structure.<ref name=Hughes2008>{{Cite journal | last1=Hughes | first1=W. O. H. | last2=Oldroyd | first2=B. P. | last3=Beekman | first3=M. | last4=Ratnieks | first4=F. L. W. | title=Ancestral Monogamy Shows Kin Selection is Key to the Evolution of Eusociality | doi=10.1126/science.1156108 | journal=Science | volume=320 | issue=5880 | pages=1213β1216 | year=2008 | pmid= 18511689| bibcode=2008Sci...320.1213H | s2cid=20388889 }}</ref> In haplodiploid species, females develop from fertilized eggs and males from unfertilized eggs. Because a male is [[haploid]] (has only one copy of each gene), his daughters (which are [[diploid]], with two copies of each gene) share 100% of his genes and 50% of their mother's. Therefore, they share 75% of their genes with each other. This mechanism of sex determination gives rise to what [[W. D. Hamilton]] termed "supersisters", more closely related to their sisters than they would be to their own offspring.<ref name="Hamilton1964II">{{cite journal | last=Hamilton | first=W. D. | title=The Genetical Evolution of Social Behaviour II | journal=Journal of Theoretical Biology | date=20 March 1964 | volume=7 | issue=1 | pages=17β52 | doi=10.1016/0022-5193(64)90039-6 | pmid=5875340| bibcode=1964JThBi...7...17H }}</ref> Workers often do not reproduce, but they can pass on more of their genes by helping to raise their sisters (as queens) than they would by having their own offspring (each of which would only have 50% of their genes), assuming they would produce similar numbers. This unusual situation has been proposed as an explanation of the multiple (at least nine) evolutions of eusociality within [[Hymenoptera]].<ref name="Hughes" /><ref>{{cite book |last1=Gullan |first1=P. J. |last2=Cranston |first2=P. S. |title=The Insects: An Outline of Entomology |date=2014 |publisher=Wiley Blackwell |isbn=978-1-118-84615-5 |edition=5th |pages=328, 348β350}}</ref> Haplodiploidy is neither necessary nor sufficient for eusociality. Some eusocial species such as [[termites]] are not haplodiploid. Conversely, all bees are haplodiploid but not all are eusocial, and among eusocial species many queens mate with multiple males, creating half-sisters that share only 25% of each other's genes.<ref name="NTW">{{cite journal | last=Nowak | first=Martin | author2=Tarnita, Corina | author3=Wilson, E.O. | title=The evolution of eusociality | journal=Nature | year=2010 | volume=466 | pmid=20740005 | issue=7310 | pages=1057β1062 | doi=10.1038/nature09205 | pmc=3279739| bibcode=2010Natur.466.1057N }}</ref> But, monogamy (queens mating singly) is the ancestral state for all eusocial species so far investigated, so it is likely that haplodiploidy contributed to the evolution of eusociality in bees.<ref name="Hughes">{{cite journal | author1=Hughes, William O. H. | author2=Oldroyd, Benjamin P. | author3=Beekman, Madeleine | author4=Ratnieks, Francis L. W. | title=Ancestral Monogamy Shows Kin Selection Is Key to the Evolution of Eusociality | journal=[[Science (journal)|Science]] | volume=320 | issue=5880 | pages=1213β1216 | publisher=American Association for the Advancement of Science | date=May 2008 | doi=10.1126/science.1156108 | pmid=18511689| bibcode=2008Sci...320.1213H | s2cid=20388889 }}</ref> ===Eusociality=== [[File:Bee swarm on fallen tree03.jpg|thumb|upright|A [[Western honey bee]] swarm]] [[File:Wildbienen.jpg|thumb|[[Western honey bee]] nest in the [[Trunk (botany)|trunk]] of a [[Picea abies|spruce]]]] {{further|Eusociality}} Bees may be solitary or may live in various types of communities. [[Eusociality]] appears to have originated from at least three independent origins in halictid bees.<ref>{{cite journal |last1=Brady |first1=SeΓ‘n 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=1643β1649 |doi=10.1098/rspb.2006.3496 |issn=0962-8452 |pmc=1634925 |pmid=16769636}}</ref> The most advanced of these are species with [[Eusociality|eusocial]] colonies; these are characterized by cooperative brood care and a [[division of labour]] into reproductive and non-reproductive adults, plus overlapping generations.<ref>{{Cite book|title=The Insect Societies|last=Wilson|first=Edward O|publisher=Belknap Press of Harvard University Press|year=1971|location=Cambridge, Mass}}</ref> This division of labour creates specialized groups within eusocial societies which are called [[Eusociality#In insects|castes]]. In some species, groups of cohabiting females may be sisters, and if there is a division of labour within the group, they are considered [[semisocial]]. The group is called eusocial if, in addition, the group consists of a mother (the [[Queen bee|queen]]) and her daughters ([[Worker bee|workers]]). When the castes are purely behavioural alternatives, with no morphological differentiation other than size, the system is considered primitively eusocial, as in many [[paper wasp]]s; when the castes are morphologically discrete, the system is considered highly eusocial.<ref name=Michener1974>{{cite book|author=Michener, Charles Duncan |title=The Social Behavior of the Bees: A Comparative Study |url=https://books.google.com/books?id=aordrL_D-30C&pg=PA78 |year=1974 |publisher=Harvard University Press |isbn=978-0-674-81175-1 |pages=22β78}}</ref> True honey bees (genus ''[[Apis (genus)|Apis]]'', of which eight species are currently recognized) are highly eusocial, and are among the best known insects. Their colonies are established by [[Swarming (honey bee)|swarms]], consisting of a queen and several thousand workers. There are 29 subspecies of one of these species, ''[[Apis mellifera]]'', native to Europe, the Middle East, and Africa. [[Africanized bee]]s are a hybrid strain of ''A. mellifera'' that escaped from experiments involving crossing European and African subspecies; they are extremely defensive.<ref>{{cite web | last1=Sanford | first1=Malcolm T. | title=The Africanized Honey Bee in the Americas: A Biological Revolution with Human Cultural Implications | url=http://apisenterprises.com/papers_htm/Misc/AHB%20in%20the%20Americas.htm | publisher=Apis Enterprises | access-date=29 March 2015 | year=2006 | url-status=live | archive-url=http://archive.wikiwix.com/cache/20150329231615/http://apisenterprises.com/papers_htm/Misc/AHB%20in%20the%20Americas.htm | archive-date=29 March 2015}}</ref> [[Stingless bee]]s are also highly [[eusocial]]. They practice [[mass provisioning]], with complex nest architecture and perennial colonies also established via swarming.<ref name=":0" /><ref>{{cite journal | last1=Roubik | first1=D. W. | year=2006 | title=Stingless bee nesting biology | journal=Apidologie | volume=37 | issue=2 | pages=124β143 | doi=10.1051/apido:2006026| url=http://www.apidologie.org/articles/apido/pdf/2006/02/m6034sp.pdf |archive-url=https://ghostarchive.org/archive/20221009/http://www.apidologie.org/articles/apido/pdf/2006/02/m6034sp.pdf |archive-date=9 October 2022 |url-status=live | doi-access=free }}</ref> [[File:Bumblebee 05.JPG|thumb|left|A [[bumblebee]] carrying pollen in its [[pollen basket]]s (corbiculae)]] Many bumblebees are eusocial, similar to the eusocial [[Vespidae]] such as [[hornet]]s in that the queen initiates a nest on her own rather than by swarming. Bumblebee colonies typically have from 50 to 200 bees at peak population, which occurs in mid to late summer. Nest architecture is simple, limited by the size of the pre-existing nest cavity, and colonies rarely last more than a year.<ref name="BBCT-nest">{{cite web | title=Bumblebee nests | url=http://bumblebeeconservation.org/about-bees/habitats/bumblebee-nests/ | publisher=Bumblebee Conservation Trust | access-date=26 June 2015 | archive-url=https://web.archive.org/web/20170922230805/https://bumblebeeconservation.org/about-bees/habitats/bumblebee-nests/ | archive-date=22 September 2017}}</ref> In 2011, the [[International Union for Conservation of Nature]] set up the Bumblebee Specialist Group to review the threat status of all bumblebee species worldwide using the [[IUCN Red List]] criteria.<ref>{{cite news | url=http://cmsdata.iucn.org/downloads/bumblebee_sg_proofed.pdf | title=Bumblebee Specialist Group: 2011 Update | publisher=[[IUCN]] | access-date=7 October 2012 | url-status=live | archive-url=https://web.archive.org/web/20121203034020/http://cmsdata.iucn.org/downloads/bumblebee_sg_proofed.pdf | archive-date=3 December 2012}}</ref> There are many more species of primitively eusocial than highly eusocial bees, but they have been studied less often. Most are in the family [[Halictidae]], or "sweat bees". Colonies are typically small, with a dozen or fewer workers, on average. Queens and workers differ only in size, if at all. Most species have a single season colony cycle, even in the tropics, and only mated females hibernate. A few species have long active seasons and attain colony sizes in the hundreds, such as ''[[Halictus hesperus]]''.<ref>{{cite journal | last1=Brooks | first1=R. W. | last2=Roubik | first2=D. W. | year=1983 | title=A Halictine bee with distinct castes: ''Halictus hesperus'' (Hymenoptera: Halictidae) and its bionomics in Central Panama | journal=Sociobiology | volume=7 | pages=263β282 }}</ref> Some species are eusocial in parts of their range and solitary in others,<ref>{{Cite journal|last1=Eickwort |first1=G. C. |last2=Eickwort |first2=J. M. |last3=Gordon|first3=J. |last4=Eickwort |first4=M. A. |last5=Wcislo |first5=W. T. |title=Solitary behavior in a high-altitude population of the social sweat bee Halictus rubicundus (Hymenoptera: Halictidae) |journal=Behavioral Ecology and Sociobiology |volume=38 |issue=4 |pages=227β233 |doi=10.1007/s002650050236 |year=1996|bibcode=1996BEcoS..38..227E |s2cid=12868253 }}</ref> or have a mix of eusocial and solitary nests in the same population.<ref>{{cite journal | last1=Yanega | first1=D. | year=1993 | title=Environmental effects on male production and social structure in ''Halictus rubicundus'' (Hymenoptera: Halictidae) | journal=Insectes Sociaux | volume=40 | pages=169β180 | doi=10.1007/bf01240705| s2cid=44934383 }}</ref> The [[orchid bee]]s (Apidae) include some primitively eusocial species with similar biology. Some [[Allodapini|allodapine]] bees (Apidae) form primitively eusocial colonies, with [[progressive provisioning]]: a larva's food is supplied gradually as it develops, as is the case in honey bees and some bumblebees.<ref>{{cite book |author=Michener, Charles Duncan |title=The Social Behavior of the Bees: A Comparative Study |url=https://books.google.com/books?id=aordrL_D-30C&pg=PA308 |year=1974 |publisher=Harvard University Press |isbn=978-0-674-81175-1 |page=308 |url-status=live |archive-url=https://web.archive.org/web/20161224104244/https://books.google.com/books?id=aordrL_D-30C&pg=PA308 |archive-date=24 December 2016}}</ref> ===Solitary and communal bees=== [[File:Megachile rotundata.JPG|thumb|A leafcutting bee, ''[[Megachile rotundata]]'', cutting circles from acacia leaves]] Most other bees, including familiar insects such as [[carpenter bee]]s, [[leafcutter bees]] and [[mason bees]] are solitary in the sense that every female is fertile, and typically inhabits a nest she constructs herself. There is no division of labor so these nests lack queens and ''worker'' bees for these species. Solitary bees typically produce neither honey nor [[beeswax]]. Bees collect pollen to feed their young, and have the necessary adaptations to do this. However, certain wasp species such as [[pollen wasp]]s have similar behaviours, and a few species of bee [[Scavenger|scavenge]] from carcases to feed their offspring.<ref name=Grimaldi/> Solitary bees are important pollinators; they gather pollen to provision their nests with food for their brood. Often it is mixed with nectar to form a paste-like consistency. Some solitary bees have advanced types of pollen-carrying structures on their bodies. Very few species of solitary bee are being cultured for commercial pollination. Most of these species belong to a distinct set of [[genus|genera]] which are commonly known by their nesting behavior or preferences, namely: carpenter bees, [[Halictidae|sweat bees]], mason bees, [[Colletes inaequalis|plasterer bees]], [[squash bee]]s, [[Ceratina|dwarf carpenter bees]], leafcutter bees, [[alkali bee]]s and [[Anthophorini|digger bees]].<ref>{{cite web |url=http://www.beesource.com/resources/usda/management-of-wild-bees/ |author1=Parker, Frank D. |author2=Torchio, Philip F. |title=Management of Wild Bees |publisher=Beesource Beekeeping Community |date=1 October 1980 |access-date=26 June 2015 |url-status=live |archive-url=https://web.archive.org/web/20150626235657/http://www.beesource.com/resources/usda/management-of-wild-bees/ |archive-date=26 June 2015}}</ref> [[File:Anthidium February 2008-1.jpg|thumb|A solitary bee, ''[[Anthidium florentinum]]'' (family [[Megachilidae]]), visiting ''[[Lantana]]'']] Most solitary bees are [[fossorial]], digging nests in the ground in a variety of soil textures and conditions, while others create nests in hollow [[Phragmites|reeds]] or twigs, or holes in [[wood]]. The female typically creates a compartment (a "cell") with an egg and some provisions for the resulting larva, then seals it off. A nest may consist of numerous cells. When the nest is in wood, usually the last (those closer to the entrance) contain eggs that will become males. The adult does not provide care for the brood once the egg is laid, and usually dies after making one or more nests. The males typically emerge first and are ready for mating when the females emerge. Solitary bees are very unlikely to sting (only in self-defense, if ever), and some (esp. in the family [[Andrenidae]]) are stingless.<ref>{{cite web |title=Solitary Bees (Hymenoptera) |url=http://www.royensoc.co.uk/insect_info/what/solitary_bees.htm |publisher=Royal Entomological Society |access-date=12 October 2015 |archive-url=https://web.archive.org/web/20170702005458/http://www.royensoc.co.uk/insect_info/what/solitary_bees.htm |archive-date=2 July 2017 |url-status=dead}}</ref><ref>{{cite web |title=Other bees |url=http://bumblebeeconservation.org/about-bees/identification/other-bees/ |publisher=Bumblebee Conservation Trust |access-date=12 October 2015 |url-status=dead |archive-url=https://web.archive.org/web/20150905113303/http://bumblebeeconservation.org/about-bees/identification/other-bees/ |archive-date=5 September 2015}}</ref> [[File:Osmia cornifrons.5.1.08.w.jpg|thumb|The [[mason bee]] ''[[Osmia cornifrons]]'' nests in a hole in dead wood. [[Insect hotel#Solitary bees and wasps|Bee "hotels"]] are often sold for this purpose.]] While solitary, females each make individual nests.<ref>{{Cite web|last=Westreich|first=Lila|title=Spring signals female bees to lay the next generation of pollinators|url=http://theconversation.com/spring-signals-female-bees-to-lay-the-next-generation-of-pollinators-134852|access-date=8 October 2020|website=The Conversation|date=May 2020 |language=en}}</ref> Some species, such as the European mason bee ''[[Hoplitis anthocopoides]]'',<ref name=Eickwort>{{cite journal |last1=Eickwort |first1=George C. |title=Gregarious Nesting of the Mason Bee Hoplitis anthocopoides and the Evolution of Parasitism and Sociality Among Megachilid Bees |journal=Evolution |date=1975 |volume=29 |issue=1 |pages=142β150 |doi=10.2307/2407147|jstor=2407147 |pmid=28563288 }}</ref> and the [[Amegilla dawsoni|Dawson's Burrowing bee]], ''Amegilla dawsoni,''<ref>{{Cite journal |title=The Nesting Behavior of Dawson's Burrowing Bee, Amegilla dawsoni (Hymenoptera: Anthophorini), and the Production of Offspring of Different Sizes |journal=Journal of Insect Behavior |date=1 May 1999 |issn=0892-7553 |pages=363β384 |volume=12 |issue=3 |doi=10.1023/A:1020843606530 |first=John |last=Alcock|bibcode=1999JIBeh..12..363A |s2cid=24832889 }}</ref> are gregarious, preferring to make nests near others of the same species, and giving the appearance of being social. Large groups of solitary bee nests are called ''aggregations'', to distinguish them from [[colony (biology)|colonies]]. In some species, multiple females share a common nest, but each makes and provisions her own cells independently. This type of group is called "communal" and is not uncommon. The primary advantage appears to be that a nest entrance is easier to defend from predators and parasites when multiple females use that same entrance regularly.<ref name=Eickwort/>
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