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This article is about the group of flying insects. For other uses, seeBee (disambiguation).
"Anthophila" redirects here. For the moth genus, seeAnthophila (moth).

Bees
Temporal range:70–0MaLate Cretaceous–Present
The sugarbag bee,Tetragonula carbonaria
Scientific classificationEdit this classification
Domain: Eukaryota
Kingdom: Animalia
Phylum: Arthropoda
Class: Insecta
Order: Hymenoptera
(unranked): Unicalcarida
Suborder: Apocrita
Superfamily: Apoidea
Clade: Anthophila
Families
Synonyms

Apiformes(from Latin'apis')

Beesare wingedinsectsclosely related towaspsandants,known for their roles inpollinationand, in the case of the best-known bee species, thewestern honey bee,for producinghoney.Bees are amonophyleticlineage within the superfamilyApoidea.They are currently considered aclade,calledAnthophila.[1]There are over 20,000 known species of bees in seven recognizedbiological families.[2][3][4]Some species – includinghoney bees,bumblebees,andstingless bees– livesociallyin colonies while most species (>90%) – includingmason bees,carpenter bees,leafcutter bees,andsweat bees– are solitary.

Bees are found on every continent exceptAntarctica,in every habitat on the planet that contains insect-pollinatedflowering plants.The most common bees in theNorthern Hemisphereare theHalictidae,or sweat bees, but they are small and often mistaken for wasps or flies. Bees range in size from tiny stingless bee species, whose workers are less than 2 millimetres (0.08 in) long,[5]to the leafcutter beeMegachile pluto,the largest species of bee, whose females can attain a length of 39 millimetres (1.54 in).

Bees feed onnectarandpollen,the former primarily as an energy source and the latter primarily forproteinand other nutrients. Most pollen is used as food for theirlarvae.Vertebrate predators of bees includeprimatesand birds such asbee-eaters;insect predators includebeewolvesanddragonflies.

Beepollinationis important both ecologically andcommercially,and the decline in wild bees has increased the value of pollination by commercially managed hives of honey bees. The analysis of 353 wild bee and hoverfly species across Britain from 1980 to 2013 found the insects have been lost from a quarter of the places they inhabited in 1980.[6]

Humanbeekeepingor apiculture (meliponiculturefor stingless bees) has been practised for millennia, since at least the times ofAncient EgyptandAncient Greece.Bees have appeared in mythology and folklore, through all phases of art and literature from ancient times to the present day, although primarily focused in theNorthern Hemispherewhere beekeeping is far more common. InMesoamerica,theMayanshave practiced large-scale intensive meliponiculture since pre-Columbian times.[5]

Evolution

The immediate ancestors of bees werestinging waspsin the familyCrabronidae,which werepredatorsof 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 sameevolutionaryscenario may have occurred within thevespoidwasps, where thepollen waspsevolved from predatory ancestors.

Based on phylogenetic analysis, bees are thought to have originated during theEarly Cretaceous(about 124 million years ago) on the supercontinent ofWest Gondwana,just prior to its breakup intoSouth AmericaandAfrica.The supercontinent is thought to have been a largelyxericenvironment at this time; modern bee diversity hotspots are also in xeric and seasonal temperate environments, suggesting strongniche conservatismamong bees ever since their origins.[7]

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. TheMelittidae,Apidae,andMegachilidaehad 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 theColletidae,AndrenidaeandHalictidaein South America. The rapid radiation of the South American bee families is thought to have followed the concurrent radiation offlowering plantsin the same region. Later in the Cretaceous (80 million years ago), colletid bees colonizedAustraliafromSouth America(with an offshoot lineage evolving into theStenotritidae), and by the end of the Cretaceous, South American bees had also colonized North America.[7]The North American fossil taxonCretotrigonabelongs to a group that is no longer found in North America, suggesting that many bee lineages went extinct during theCretaceous-Paleogene extinction event.[7]

Following the K-Pg extinction, surviving bee lineages continued to spread into the Northern Hemisphere, colonizingEuropefrom Africa by thePaleocene,and then spreading east toAsia.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 theEocene(~45 mya) there was already considerable diversity among eusocial bee lineages.[8][a]A second extinction event among bees is thought to have occurred due to rapid climatic cooling around theEocene-Oligocene boundary,leading to the extinction of some bee lineages such as the tribeMelikertini.Over the Paleogene andNeogene,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.[7]

Fossils

The oldest non-compression bee fossil isCretotrigona prisca,acorbiculate beeofLate Cretaceousage (~70 mya) found inNew Jersey amber.[11]A fossil from the early Cretaceous (~100 mya),Melittosphex burmensis,was initially considered "an extinct lineage of pollen-collecting Apoideasisterto the modern bees ",[12]but subsequent research has rejected the claim thatMelittosphexis a bee, or even a member of the superfamilyApoideato which bees belong, instead treating the lineage asincertae sediswithin theAculeata.[13]

TheAllodapini(within the Apidae) appeared around 53 Mya.[14] The Colletidae appear as fossils only from the lateOligocene(~25 Mya) to earlyMiocene.[15] The Melittidae are known fromPalaeomacropis eocenicusin theEarly Eocene.[16] The Megachilidae are known from trace fossils (characteristic leaf cuttings) from theMiddle Eocene.[17] The Andrenidae are known from the Eocene-Oligocene boundary, around 34 Mya, of the Florissant shale.[18] The Halictidae first appear in the Early Eocene[19]with species[20][21]found in amber. The Stenotritidae are known from fossil brood cells ofPleistoceneage.[22]

Coevolution

Long-tongued bees and long-tubed flowerscoevolved,like thisAmegillaspecies (Apidae) onAcanthus ilicifolius.
Further information:Coevolution

The earliest animal-pollinated flowers were shallow, cup-shaped bloomspollinatedby insects such asbeetles,so thesyndrome of insect pollinationwas 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 ofcoevolution,flowers developed floral rewards[23]such asnectarand longer tubes, and bees developed longer tongues to extract the nectar.[24]Bees also developed structures known asscopal hairsandpollen basketsto 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 havepollen basketson their hind legs, while very few lack these and instead collect pollen in their crops.[3]The appearance of these structures drove theadaptive radiationof theangiosperms,and, in turn, bees themselves.[9]Beescoevolvednot only with flowers but it is believed that some species coevolved with mites. Some provide tufts of hairs calledacarinariathat 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 bemutualistic.[25][26]

Phylogeny

External

Thisphylogenetic treeis based on Debevicet al,2012, which used molecular phylogeny to demonstrate that the bees (Anthophila) arose from deep within theCrabronidae,which is thereforeparaphyletic.The placement of theHeterogynaidaeis uncertain.[27]The small familyMellinidaewas not included in this analysis.

Apoidea

Ampulicidae(Cockroach wasps)

Internal

This cladogram of the bee families is based on Hedtke et al., 2013, which places the former families Dasypodaidae and Meganomiidae as subfamilies inside the Melittidae.[28]English names, where available, are given in parentheses.

Anthophila(bees)

Melittidae(inc.Dasypodainae,Meganomiinae) at least 50 Mya

long-tongued bees

Apidae(inc. honeybees, cuckoo bees, carpenter bees) ≈87 Mya

Megachilidae(mason, leafcutter bees) ≈50 Mya

short-tongued bees

Andrenidae(mining bees) ≈34 Mya

Halictidae(sweat bees) ≈50 Mya

Colletidae(plasterer bees) ≈25 Mya

Stenotritidae(large Australian bees) ≈2 Mya

See also:Characteristics of common wasps and bees

Characteristics

The lappingmouthpartsof a honey bee, showing labium and maxillae

Bees differ from closely related groups such as wasps by having branched or plume-likesetae(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.[29]

Bees have the following characteristics:

  • A pair of largecompound eyeswhich cover much of the surface of the head. Between and above these are three small simple eyes (ocelli) which provide information on light intensity.
  • Theantennaeusually have 13 segments in males and 12 in females, and aregeniculate,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.
  • Themouthpartsare adapted for both chewing and sucking by having both a pair ofmandiblesand a longproboscisfor sucking up nectar.[30]
  • The thorax has three segments, each with a pair of robust legs, and a pair of membranous wings on the hind two segments. The front legs of corbiculate bees bear combs for cleaning the antennae, and in many species the hind legs bear pollen baskets, flattened sections with incurving hairs to secure the collected pollen. The wings are synchronised in flight, and the somewhat smaller hind wings connect to the forewings by a row of hooks along their margin which connect to a groove in the forewing.
  • The abdomen has nine segments, the hindermost three being modified into the sting.[30]
Head-on view of a malecarpenter bee,showing antennae, threeocelli,compound eyes,and mouthparts

The largest species of bee is thought to be Wallace's giant beeMegachile pluto,whose females can attain a length of 39 millimetres (1.54 in).[31]The smallest species may be dwarf stingless bees in the tribeMeliponiniwhose workers are less than 2 millimetres (0.08 in) in length.[32]

Sociality

Haplodiploid breeding system

Further information:Haplodiploidy
Willing to die for their sisters: worker honey bees killed defending their hive againstyellowjackets,along with a dead yellowjacket. Suchaltruisticbehaviour may be favoured by thehaplodiploidsex determinationsystem of bees.

According toinclusive fitnesstheory, 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 whenCost < Relatedness * Benefit.The requirements for eusociality are more easily fulfilled byhaplodiploidspecies such as bees because of their unusual relatedness structure.[33]

In haplodiploid species, females develop from fertilized eggs and males from unfertilized eggs. Because a male ishaploid(has only one copy of each gene), his daughters (which arediploid,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 whatW. D. Hamiltontermed "supersisters", more closely related to their sisters than they would be to their own offspring.[34]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 withinHymenoptera.[35][36]

Haplodiploidy is neither necessary nor sufficient for eusociality. Some eusocial species such astermitesare 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.[37]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.[35]

Eusociality

AWestern honey beeswarm
Western honey beenest in thetrunkof aspruce
Further information:Eusociality

Bees may be solitary or may live in various types of communities.Eusocialityappears to have originated from at least three independent origins in halictid bees.[38]The most advanced of these are species witheusocialcolonies; these are characterised by cooperative brood care and adivision of labourinto reproductive and non-reproductive adults, plus overlapping generations.[39]This division of labour creates specialized groups within eusocial societies which are calledcastes.In some species, groups of cohabiting females may be sisters, and if there is a division of labour within the group, they are consideredsemisocial.The group is called eusocial if, in addition, the group consists of a mother (thequeen) 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 manypaper wasps;when the castes are morphologically discrete, the system is considered highly eusocial.[24]

True honey bees (genusApis,of which eight species are currently recognized) are highly eusocial, and are among the best known insects. Their colonies are established byswarms,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 beesare a hybrid strain ofA. melliferathat escaped from experiments involving crossing European and African subspecies; they are extremely defensive.[40]

Stingless beesare also highlyeusocial.They practisemass provisioning,with complex nest architecture and perennial colonies also established via swarming.[5][41]

Abumblebeecarrying pollen in itspollen baskets(corbiculae)

Many bumblebees are eusocial, similar to the eusocialVespidaesuch ashornetsin 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.[42]In 2011, theInternational Union for Conservation of Natureset up the Bumblebee Specialist Group to review the threat status of all bumblebee species worldwide using theIUCN Red Listcriteria.[43]

There are many more species of primitively eusocial than highly eusocial bees, but they have been studied less often. Most are in the familyHalictidae,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 asHalictus hesperus.[44]Some species are eusocial in parts of their range and solitary in others,[45]or have a mix of eusocial and solitary nests in the same population.[46]Theorchid bees(Apidae) include some primitively eusocial species with similar biology. Someallodapinebees (Apidae) form primitively eusocial colonies, withprogressive provisioning:a larva's food is supplied gradually as it develops, as is the case in honey bees and some bumblebees.[47]

Solitary and communal bees

A leafcutting bee,Megachile rotundata,cutting circles from acacia leaves

Most other bees, including familiar insects such ascarpenter bees,leafcutter beesandmason beesare 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 andworkerbees for these species. Solitary bees typically produce neither honey norbeeswax. Bees collect pollen to feed their young, and have the necessary adaptations to do this. However, certain wasp species such aspollen waspshave similar behaviours, and a few species of beescavengefrom carcases to feed their offspring.[29]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 ofgenerawhich 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 beesanddigger bees.[48]

A solitary bee,Anthidium florentinum(familyMegachilidae), visitingLantana

Most solitary bees arefossorial,digging nests in the ground in a variety of soil textures and conditions, while others create nests in hollowreedsor twigs, or holes inwood.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 familyAndrenidae) are stingless.[49][50]

Themason beeOsmia cornifronsnests in a hole in dead wood.Bee "hotels"are often sold for this purpose.

While solitary, females each make individual nests.[51]Some species, such as the European mason beeHoplitis anthocopoides,[52]and theDawson's Burrowing bee,Amegilla dawsoni,[53]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 calledaggregations,to distinguish them fromcolonies.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.[52]

Biology

Variousbeesvisit amorning gloryflower. ATumbling flower beetleremains in the flower with a bee visitor.

Life cycle

Further information: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 leglesslarva,apupationstage during which the insect undergoescomplete 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.[54]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 arehaplodiploid;the sex of a bee is determined by whether or not the egg is fertilised. After mating, a female stores the sperm, and determines which sex is required at the time each individual egg is laid, fertilised eggs producing female offspring and unfertilised eggs, males. Tropical bees may have several generations in a year and nodiapausestage.[55][56][57][58]

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.[59]

In most species, larvae are whitish grubs, roughly oval and bluntly-pointed at both ends. They have 15 segments andspiraclesin 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 theexuviaeand breaks out of the cell.[59]

Flight

Honeybee in flight carrying pollen inpollen basket
Further information:Insect flight

Antoine Magnan's 1934 bookLe vol des insectessays that he andAndré Sainte-Laguëhad applied the equations ofair resistancetoinsectsand 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".[60]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 byhelicopters.[61]In 1996 it was shown that vortices created by many insects' wings helped to provide lift.[62]High-speedcinematography[63]and robotic mock-up of a bee wing[64]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 smallarc,it flaps approximately 230 times per second, faster than afruitfly(200 times per second) which is 80 times smaller.[65]

Navigation, communication, and finding food

Karl von Frisch(1953) discovered that honey bee workers cannavigate,indicating the range and direction to food to other workers with awaggle dance.
Further information:Animal navigationandWaggle dance

The ethologistKarl von Frischstudiednavigationin the honey bee. He showed that honey bees communicate by thewaggle 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 thepolarizationpattern 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 darkbeehive.[66]Bees navigate usingspatial memorywith a "rich, map-like organization".[67]

Digestion

The gut of bees is relatively simple, but multiple metabolic strategies exist in the gutmicrobiota.[68]Pollinating bees consume nectar and pollen, which require different digestion strategies by somewhat specialized bacteria. While nectar is a liquid of mostlymonosaccharidesugars and so easily absorbed, pollen contains complexpolysaccharides:branchingpectinandhemicellulose.[69]Approximately five groups of bacteria are involved in digestion. Three groups specialize in simple sugars (Snodgrassellaand two groups ofLactobacillus), and two other groups in complex sugars (GilliamellaandBifidobacterium). Digestion of pectin and hemicellulose is dominated by bacterialcladesGilliamellaandBifidobacteriumrespectively. Bacteria that cannot digest polysaccharides obtain enzymes from their neighbors, and bacteria that lack certain amino acids do the same, creating multipleecological niches.[70]

Although most bee species arenectarivorousandpalynivorous,some are not. Particularly unusual arevulture beesin the genusTrigona,which consume carrion and wasp brood, turning meat into a honey-like substance.[71]Drinkingguttationdrops from leaves is also a source of energy and nutrients.[72]

Ecology

Floral relationships

Most bees are polylectic (generalist) meaning they collect pollen from a range of flowering plants, but some areoligoleges(specialists), in that they only gather pollen from one or a few species or genera of closely related plants.[73]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.[74]Male orchid bees in some species gather aromatic compounds fromorchids,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,[75]and even electromagnetic fields.[76]Once landed, a bee then uses nectar quality[75]and pollen taste[77]to determine whether to continue visiting similar flowers.

In rare cases, aplantspecies may only be effectively pollinated by a single bee species, and some plants areendangeredat 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, thecreosote bushin the arid parts of the United States southwest is associated with some 40 oligoleges.[78]

As mimics and models

The bee-flyBombylius major,aBatesian mimicof bees, taking nectar and pollinating a flower
Main articles:Mimicry,Batesian mimicry,andMüllerian mimicry
Bee orchidlures male bees to attempt to mate with the flower's lip, which resembles a bee perched on a pink flower.

Many bees areaposematicallycoloured, typically orange and black, warning of their ability to defend themselves with a powerful sting. As such they are models forBatesian mimicryby non-stinging insects such asbee-flies,robber fliesandhoverflies,[79]all of which gain a measure of protection by superficially looking and behaving like bees.[79]

Bees are themselvesMüllerian mimicsof other aposematic insects with the same colour scheme, includingwasps,lycidand other beetles, and many butterflies and moths (Lepidoptera) which are themselves distasteful, often through acquiring bitter and poisonous chemicals from their plant food. All the Müllerian mimics, including bees, benefit from the reduced risk of predation that results from their easily recognised warning coloration.[80]

Bees are also mimicked by plants such as thebee orchidwhich 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.[81]

As brood parasites

Bombus vestalis,abrood parasiteof the bumblebeeBombus terrestris
Main articles:Brood parasiteandNest usurpation

Brood parasitesoccur in several bee families including theapidsubfamilyNomadinae.[82]Females of these species lack pollen collecting structures (thescopa) 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.[83]In particular, the Arctic bee species,Bombus hyperboreusis 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.[84]

In Southern Africa, hives of African honeybees (A. mellifera scutellata) are being destroyed by parasitic workers of the Cape honeybee,A. m. capensis.These laydiploideggs ( "thelytoky"), escaping normalworker policing,leading to the colony's destruction; the parasites can then move to other hives.[85]

Thecuckoo beesin theBombussubgenusPsithyrusare 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, likeTownsendiella,anomadineapid,two species of which are cleptoparasites of thedasypodaidgenusHesperapis,[86]while the other species in the same genus attackshalictidbees.[87]

Nocturnal bees

Four bee families (Andrenidae,Colletidae,Halictidae,andApidae) contain some species that arecrepuscular.Most are tropical or subtropical, but some live in arid regions at higher latitudes. These bees have greatly enlargedocelli,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.[88]

Predators, parasites and pathogens

Further information:Diseases of the honey bee
The bee-eater,Merops apiaster,specialises in feeding on bees; here a male catches a nuptial gift for his mate.

Vertebrate predators of bees includebee-eaters,shrikesandflycatchers,which make short sallies to catch insects in flight.[89]Swifts and swallows[89]fly almost continually, catching insects as they go. Thehoney buzzardattacks bees' nests and eats the larvae.[90]Thegreater honeyguideinteracts 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.[91]Among mammals, predators such as thebadgerdig up bumblebee nests and eat both the larvae and any stored food.[92]

The beewolfPhilanthus triangulumparalysing a bee with its sting

Specialist ambush predators of visitors to flowers includecrab spiders,which wait on flowering plants for pollinating insects;predatory bugs,andpraying mantises,[89]some of which (theflower mantisesof the tropics) wait motionless,aggressive mimicscamouflaged as flowers.[93]Beewolvesare large wasps that habitually attack bees;[89]theethologistNiko Tinbergenestimated that a single colony of the beewolfPhilanthus triangulummight kill several thousand honeybees in a day: all the prey he observed were honeybees.[94]Other predatory insects that sometimes catch bees includerobber fliesanddragonflies.[89]Honey bees are affected by parasites includingtrachealandVarroamites.[95]However, some bees are believed to have a mutualistic relationship with mites.[26]

Some mites of genusTarsonemusare 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.[96]

Symbiosis of Mycelium and Bees

Fungus properties

Recent studies have shown thatmyceliumprovideshoney beesandstingless beeswith vital nutrients. Specific fungi, such asZygosaccharomycessp,Candida sp., andMonascusruber, produce chemicals that fight againstbacteria,fungal infectionsfrom different species, andviruses.Recently these types of bees have been observed eatingmycelium,suggesting that honey bees have been “foragingmushroomsto collectantimicrobialmedicine to boost their collectiveimmunity”.[97]Without these vital nutrients, honey bee morbidity rates rise, and the possibility offungal infectionscan spike, leading to unhealthy bee hives and honey shortage.Fungal infectionscan also lead tocolony collapse disorder,so the ingestion ofmyceliumlowers the morbidity rate of honey bees by preventing thosefungal infectionsfrom happening.Colony collapse disorder(CCD) is when worker bees abandon thequeen beeand leave behind the brood and a few nurse bees. This however is not enough to sustain a hive as workers are required to construct and maintain the hive structure as well as produce honey.Colony collapse disordercan also happen whenvarroa mitesinfiltrate a hive. These mites will attack and eat bees inside a hive, making it impossible for them to continue to reproduce and makehoney.The presence ofvarroa mitesresults in a decrease in bee population, deformed bees, an inability to reproduce on the bees part, and overall weakening of the colony.Varroa mitesare only capable of reproducing inside of a honey bee colony, posing an even greater threat if they are able to infiltrate because it will destroy their home.Myceliumhas been shown togerminateinside ofvarroa mitesand grow from the inside out, killing the mites and protecting the bees. The extermination of mites bymyceliumis a better alternative topesticidesthat have shown to be toxic towards the bee colony.Myceliumalso plays a role in boostinganti-inflammatoryandantibacterial resistancein bees due to theecdysteroidsandZygosaccharomycesfound inmycelium,which are then fed tolarvae,boosting the next generations immunity and improving overall hive health.Zygosaccharomycesare “spoilage yeasts that have an extreme resistance to acids and preservatives” and can “tolerate high concentrations of sugars and salts”.[98]Honey bees depend on this source of steroids to allow them to develop properly during insectpupation.

Bee broods

Thesymbioticrelationship between bees andmyceliumis found primarily in Brazilian stingless bees and Malaysian stingless bees - or more commonly honey bees.Bee broodsare thelarvaeof honeybees. They can typically be found inside of a bee hive, and in man made hives especially, the honeybees can be found developing at different stages (eggs, larvae, andpupae) inside ahexagonal shape.Bee larvae are incapable of producingsteroidsat birth, so they ingestmyceliumto receive vital nutrients they cannot create on their own such asecdysteroidsand Zygosaccharomyces sp.[99]Once the honey bee eggs hatch, a white microbial film starts to grow on the boundary between the brood cell and the larval food supply, and is then ingested by thelarvaeto complete their development.[99]

Gut microbiotaplay an immense role in the health of the entire bee colony. Three studies were recently conducted and each introduced a new organism to the beesgut microbiota.The bees were fed agedpollen,the assembly of thegut microbiotawas disturbed, and antibiotic tetracycline entered their diet. All three studies showed that the honey bees' ability to survive decreased drastically and they became more likely to contractparasitesandfungal infections.[100]The introduction of certainmyceliumto the honey beesgut microbiotahas the opposite effect to what took place in these three studies, highlighting the importance of what bees ingest and the impacts it has on their survivability during both the development and adult stages.

Bee-fungus symbiosis

As mentioned above, honey bees cannot producesteroidsthemselves, they must be ingested through their diet, specifically in the early development process. Larvae eat the fungus and theecdysteroidsandZygosaccharomycesproduced by themyceliumbenefit thelarvae.Ecdysteroidsare naturally occurringsteroidsfound inmyceliumand they help enhance performance and reproduction, boosting honey production and keeping the hive population running at a stable rate. “Zygosaccharomycessp. is essential for S. depilis larvae”.[99]Thesesterolsthus have a high impact on the survival rate of honey bees. Their ingestion determines whether the honey bees will be able to protect themselves againstfungal infections,viruses,and whether or not they will have sufficient strength to increase honey production and the ability topollinatea larger area and more frequently.

Knowledge of howmyceliumboosts honey bees immunity could be pivotal to the increase of a honey bee's lifespan and boost reproduction by helping implement new policies to prevent the use of harmfulpesticides.[99]

Impact of pesticides

Pesticideshave been diminishing the bee population recently due to a lack of regulations regarding what can and cannot be sprayed on produce to protect it from being damaged during growth. When honey bees collectpollenandnectarfor nutrition and to makehoney,they are also ingesting harmful chemicals. Such chemicals take a toll on the honey bees' already sensitivegut microbiomeand lead to a higher morbidity rate in honey bees. "Thesemicrobescan suffer with toxicpesticidesapplied in agriculture, causing dangerous changes in the colony fitness and perturbing bees' health.” (Yordanova, M. et al., 2022)[101]

With the research provided on the positive impact ofmyceliumon bees, the relationship betweenmyceliumand honey bees issymbioticin that the survival of bees and the mycelium's ability to help boost bee pollination, boosts the ability of thefungito grow because bee pollination improves air and soil quality, thus boosting plant life. Allowing for a higher survivability rate for both bees andmyceliumif they are able to perform their environmental roles properly without the interruption of harmful government approvedpesticides.The recent studies done on thesymbioticrelationship betweenmyceliumand honey bees will prove to be vital in the argument towards lessening the types of chemicals legally allowed to be sprayed on produce. The use ofpesticideson lawns and for other agricultural uses destroy the livelihood ofmyceliumby killing the soil it grows in, inhibiting bees from ingesting the necessary nutrientsmyceliumprovides to survive.

Relationship with humans

In mythology and folklore

Main article:Bees in mythology
Gold plaques embossed with winged bee goddesses.Camiros,Rhodes.7th century BC.

Homer'sHymn to Hermesdescribes three bee-maidens with the power ofdivinationand thus speaking truth, and identifies the food of the gods as honey. Sources associated the bee maidens withApolloand, until the 1980s, scholars followed Gottfried Hermann (1806) in incorrectly identifying the bee-maidens with theThriae.[102]Honey, according to a Greek myth, was discovered by a nymph calledMelissa( "Bee" ); and honey was offered to the Greek gods fromMycenean times.Bees were also associated with theDelphic oracleand the prophetess was sometimes called a bee.[103]

The image of a community of honey bees has been used from ancient to modern times, inAristotleandPlato;inVirgilandSeneca;inErasmusandShakespeare;Tolstoy,and by political and social theorists such asBernard MandevilleandKarl Marxas a model for humansociety.[104]In English folklore, bees would be told of important events in the household, in a custom known as "Telling the bees".[105]

In art and literature

Beatrix Potter's illustration of Babbity Bumble inThe Tale of Mrs Tittlemouse,1910

Some of the oldest examples of bees in art arerock paintingsinSpainwhich have been dated to 15,000 BC.[106]

W. B. Yeats's poemThe 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 inBedford Parkin the West of London.[107]Beatrix Potter's illustrated bookThe Tale of Mrs Tittlemouse(1910) features Babbity Bumble and her brood(pictured).Kit Williams'treasure huntbookThe Bee on the Comb(1984) uses bees and beekeeping as part of its story and puzzle.Sue Monk Kidd'sThe Secret Life of Bees(2004), and the2009 filmstarringDakota Fanning,tells the story of a girl who escapes her abusive home and finds her way to live with a family of beekeepers, the Boatwrights.

The 2007 animated comedy filmBee MovieusedJerry Seinfeld's first script and was his first work for children; he starred as a bee named Barry B. Benson, alongsideRenée Zellweger.Critics found its premise awkward and its delivery tame.[108]Dave Goulson'sA Sting in the Tale(2014) describes his efforts to save bumblebees in Britain, as well as much about their biology. The playwrightLaline Paull'sfantasyThe Bees(2015) tells the tale of a hive bee named Flora 717 from hatching onwards.[109]

Beekeeping

Main article:Beekeeping
A commercial beekeeper at work
Western honey bee on a honeycomb

Humans have kept honey bee colonies, commonly inhives,for millennia.Beekeeperscollecthoney,beeswax,propolis,pollen,androyal jellyfrom hives; bees are also kept topollinatecrops and to produce bees for sale to other beekeepers.

Depictions of humans collecting honey from wild bees date to 15,000 years ago; efforts to domesticate them are shown in Egyptian art around 4,500 years ago.[110]Simple hives and smoke were used;[111][112]jars of honey were found in the tombs ofpharaohssuch asTutankhamun.

Among Classical Era authors, beekeeping with the use of smoke is described in Aristotle'sHistory of AnimalsBook 9.[113]The account mentions that bees die after stinging; that workers remove corpses from the hive, and guard it; castes including workers and non-workingdrones,but "kings" rather than queens; predators including toads and bee-eaters; and thewaggle dance,with the "irresistible suggestion" ofάροσειονται( "aroseiontai",it waggles) andπαρακολουθούσιν( "parakolouthousin",they watch).[114][b]

Beekeeping is described in detail byVirgilin hisGeorgics;it is also mentioned in hisAeneid,and inPliny'sNatural History.[114]

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.[115][116]

As commercial pollinators

See also:List of crop plants pollinated by bees,Pollinator decline,andPesticide toxicity to bees

Role of bees

Bees play an important role inpollinatingflowering plants,and are the major type ofpollinatorin manyecosystemsthat contain flowering plants. It is estimated that one third of the human food supply depends onpollinationby insects, birds and bats, most of which is accomplished by bees, whether wild or domesticated.[117][118]

In the United States

Contract pollinationhas overtaken the role of honey production forbeekeepersin many countries. After the introduction ofVarroa mites,feralhoney bees declined dramatically in the US, though their numbers have since recovered.[119][120]The number of colonies kept by beekeepers declined slightly, throughurbanization,systematic pesticide use,trachealandVarroamites, and the closure of beekeeping businesses. In 2006 and 2007 the rate of attrition increased, and was described ascolony collapse disorder.[121]In 2010 invertebrate iridescent virus and the fungusNosema ceranaewere shown to be in every killed colony, and deadly in combination.[122][123][124][125]Winter losses increased to about 1/3.[126][127]Varroamites were thought to be responsible for about half the losses.[128]

In the European Union

Apart from colony collapse disorder, losses outside the US have been attributed to causes including pesticide seed dressings, usingneonicotinoidssuch asclothianidin,imidaclopridandthiamethoxam.[129][130]From 2013 theEuropean Unionrestricted some pesticides to stop bee populations from declining further.[131]In 2014 theIntergovernmental Panel on Climate Changereport warned that bees faced increased risk of extinction because ofglobal warming.[132]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.[133]

Raising native plants

Farmers have focused on alternative solutions to mitigate these problems. By raising native plants, they provide food for native bee pollinators likeLasioglossum vierecki[134]andL. leucozonium,[135]leading to less reliance on honey bee populations.

As food producers

Honey is a natural product produced by bees and stored for their own use, but its sweetness has always appealed to humans. Before domestication of bees was even attempted, humans were raiding their nests for their honey. Smoke was often used to subdue the bees and such activities are depicted inrock paintingsin Spain dated to 15,000 BC.[106]

Honey bees are used commercially to producehoney.[136]They also produce some substances used as dietary supplements with possible health benefits, pollen,[137]propolis,[138]androyal jelly,[139]though all of these can also causeallergic reactions.

As food

Bees are considerededible insects.People in some countrieseat insects,including the larvae and pupae of bees, mostly stingless species. They also gather larvae, pupae and surrounding cells, known asbee brood,for consumption.[140]In theIndonesiandishbotok tawonfrom Central and EastJava,bee larvae are eaten as a companion torice,after being mixed with shreddedcoconut,wrapped inbananaleaves, and steamed.[141][142]

Bee brood (pupae and larvae) although low incalcium,has been found to be high inproteinandcarbohydrate,and a useful source ofphosphorus,magnesium,potassium,and trace mineralsiron,zinc,copper,andselenium.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-solubleB vitaminsincludingcholineas well as vitamin C. The fat was composed mostly ofsaturatedandmonounsaturated fatty acidswith 2.0% beingpolyunsaturated fatty acids.[143][144]

  • Bee larvae as food in the Javanese dish botok tawon
    Bee larvae as food in theJavanesedishbotok tawon
  • Fried whole bees served in a Ukrainian restaurant
    Fried whole bees served in a Ukrainian restaurant

As alternative medicine

Apitherapyis a branch ofalternative medicinethat uses honey bee products, includingraw honey,royal jelly,pollen,propolis,beeswaxandapitoxin(Beevenom).[145]The claim that apitherapy treats cancer, which some proponents of apitherapy make, remains unsupported byevidence-based medicine.[146][147]

Stings

The painfulstings of beesare mostly associated with the poison gland and theDufour's glandwhich are abdominal exocrine glands containing various chemicals. InLasioglossum leucozonium,the Dufour's Gland mostly containsoctadecanolideas well as someeicosanolide.There is also evidence of n-triscosane, n-heptacosane,[148]and 22-docosanolide.[149]However, the secretions of these glands could also be used for nest construction.[148]

Bees population decline

Further information:Pollinator decline

Over the last half century, there has been a general decline in the species richness of wild bees and other pollinators, probably attributable to stress from increased parasites and disease, the use of pesticides, and a general decrease in the number of wild flowers. Climate change probably exacerbates the problem.[150]This is a major cause of concern, as it can cause biodiversity loss and ecosystem degradation as well as increase climate change.[151]

See also

Explanatory notes

  1. ^Triassicnests in a petrified forest in Arizona, implying that bees evolved much earlier,[9]are now thought to be beetle borings.[10]
  2. ^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".[113]

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External links

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