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Bees are winged
insect Insects (from Latin ') are pancrustacean hexapod invertebrates of the class Insecta. They are the largest group within the arthropod phylum. Insects have a chitinous exoskeleton, a three-part body ( head, thorax and abdomen), three pairs ...
s closely related to
wasp A wasp is any insect of the narrow-waisted suborder Apocrita of the order Hymenoptera which is neither a bee nor an ant; this excludes the broad-waisted sawflies (Symphyta), which look somewhat like wasps, but are in a separate suborder. Th ...
s and
ant Ants are eusocial insects of the family Formicidae and, along with the related wasps and bees, belong to the order Hymenoptera. Ants evolved from vespoid wasp ancestors in the Cretaceous period. More than 13,800 of an estimated total of 22 ...
s, known for their roles in
pollination Pollination is the transfer of pollen from an anther of a plant to the stigma of a plant, later enabling fertilisation and the production of seeds, most often by an animal or by wind. Pollinating agents can be animals such as insects, birds ...
and, in the case of the best-known bee species, the
western honey bee The western honey bee or European honey bee (''Apis mellifera'') is the most common of the 7–12 species of honey bees worldwide. The genus name ''Apis'' is Latin for "bee", and ''mellifera'' is the Latin for "honey-bearing" or "honey carrying", ...
, for producing
honey Honey is a sweet and viscous substance made by several bees, the best-known of which are honey bees. Honey is made and stored to nourish bee colonies. Bees produce honey by gathering and then refining the sugary secretions of plants (primar ...
. Bees are a
monophyletic In cladistics for a group of organisms, monophyly is the condition of being a clade—that is, a group of taxa composed only of a common ancestor (or more precisely an ancestral population) and all of its lineal descendants. Monophyletic gro ...
lineage within the superfamily
Apoidea The superfamily (zoology), superfamily Apoidea is a major group within the Hymenoptera, which includes two traditionally recognized lineages, the "sphecidae, sphecoid" wasps, and the bees. Molecular phylogeny demonstrates that the bees arose from ...
. They are presently considered a
clade A clade (), also known as a monophyletic group or natural group, is a group of organisms that are monophyletic – that is, composed of a common ancestor and all its lineal descendants – on a phylogenetic tree. Rather than the English term, ...
, called Anthophila. There are over 16,000 known species of bees in seven recognized biological families. Some speciesincluding
honey bee A honey bee (also spelled honeybee) is a eusocial flying insect within the genus ''Apis'' of the bee clade, all native to Afro-Eurasia. After bees spread naturally throughout Africa and Eurasia, humans became responsible for the current co ...
s,
bumblebee A bumblebee (or bumble bee, bumble-bee, or humble-bee) is any of over 250 species in the genus ''Bombus'', part of Apidae, one of the bee families. This genus is the only extant group in the tribe Bombini, though a few extinct related gener ...
s, and
stingless bee Stingless bees, sometimes called stingless honey bees or simply meliponines, are a large group of bees (about 550 described species), comprising the tribe Meliponini (or subtribe Meliponina according to other authors). They belong in the family A ...
slive
socially Social organisms, including human(s), live collectively in interacting populations. This interaction is considered social whether they are aware of it or not, and whether the exchange is voluntary or not. Etymology The word "social" derives from ...
in colonies while most species (>90%)including
mason bee Mason bee is a name now commonly used for species of bees in the genus ''Osmia'', of the family Megachilidae. Mason bees are named for their habit of using mud or other "masonry" products in constructing their nests, which are made in naturally ...
s,
carpenter bee Carpenter bees are species in the genus ''Xylocopa'' of the subfamily Xylocopinae. The genus includes some 500 bees in 31 subgenera. The common name "carpenter bee" derives from their nesting behavior; nearly all species burrow into hard plant m ...
s,
leafcutter bees The genus ''Megachile'' is a cosmopolitan group of solitary bees, often called leafcutter bees or leafcutting bees; it also includes the called resin bees and mortar bees. While other genera within the family Megachilidae may chew leaves or pet ...
, and sweat beesare solitary. Bees are found on every continent except
Antarctica Antarctica () is Earth's southernmost and least-populated continent. Situated almost entirely south of the Antarctic Circle and surrounded by the Southern Ocean, it contains the geographic South Pole. Antarctica is the fifth-largest contine ...
, in every habitat on the planet that contains insect-pollinated
flowering plant Flowering plants are plants that bear flowers and fruits, and form the clade Angiospermae (), commonly called angiosperms. The term "angiosperm" is derived from the Greek words ('container, vessel') and ('seed'), and refers to those plants th ...
s. The most common bees in the
Northern Hemisphere The Northern Hemisphere is the half of Earth that is north of the Equator. For other planets in the Solar System, north is defined as being in the same celestial hemisphere relative to the invariable plane of the solar system as Earth's Nort ...
are the
Halictidae Halictidae is the second-largest family of bees (clade Anthophila) with nearly 4,500 species. Halictid species are an extremely diverse group that can vary greatly in appearance. These bees occur all over the world and are found on every contine ...
, or sweat bees, but they are small and often mistaken for wasps or flies. Bees range in size from tiny
stingless bee Stingless bees, sometimes called stingless honey bees or simply meliponines, are a large group of bees (about 550 described species), comprising the tribe Meliponini (or subtribe Meliponina according to other authors). They belong in the family A ...
species, whose workers are less than long, to ''
Megachile pluto ''Megachile pluto'', also known as Wallace's giant bee or (), is a large resin bee found in Indonesia. With a wingspan of it is the largest known living bee species. It was believed to be extinct until several specimens were discovered in 1981 ...
'', the largest species of leafcutter bee, whose females can attain a length of . Bees feed on
nectar Nectar is a sugar-rich liquid produced by plants in glands called nectaries or nectarines, either within the flowers with which it attracts pollinating animals, or by extrafloral nectaries, which provide a nutrient source to animal mutualists ...
and
pollen Pollen is a powdery substance produced by seed plants. It consists of pollen grains (highly reduced microgametophytes), which produce male gametes (sperm cells). Pollen grains have a hard coat made of sporopollenin that protects the gametophyt ...
, the former primarily as an energy source and the latter primarily for
protein Proteins are large biomolecules and macromolecules that comprise one or more long chains of amino acid residues. Proteins perform a vast array of functions within organisms, including catalysing metabolic reactions, DNA replication, respo ...
and other nutrients. Most pollen is used as food for their
larva A larva (; plural larvae ) is a distinct juvenile form many animals undergo before metamorphosis into adults. Animals with indirect development such as insects, amphibians, or cnidarians typically have a larval phase of their life cycle. The ...
e. Vertebrate predators of bees include
primates Primates are a diverse order of mammals. They are divided into the strepsirrhines, which include the lemurs, galagos, and lorisids, and the haplorhines, which include the tarsiers and the simians (monkeys and apes, the latter including huma ...
and birds such as
bee-eater The bee-eaters are a group of non-passerine birds in the family Meropidae, containing three genera and thirty species. Most species are found in Africa and Asia, with a few in southern Europe, Australia, and New Guinea. They are characterised by ...
s; insect predators include beewolves and
dragonflies A dragonfly is a flying insect belonging to the infraorder Anisoptera below the order Odonata. About 3,000 extant species of true dragonfly are known. Most are tropical, with fewer species in temperate regions. Loss of wetland habitat threa ...
. Bee
pollination Pollination is the transfer of pollen from an anther of a plant to the stigma of a plant, later enabling fertilisation and the production of seeds, most often by an animal or by wind. Pollinating agents can be animals such as insects, birds ...
is important both ecologically and
commercially Commerce is the large-scale organized system of activities, functions, procedures and institutions directly and indirectly related to the exchange (buying and selling) of goods and services among two or more parties within local, regional, nation ...
, 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. Human
beekeeping Beekeeping (or apiculture) is the maintenance of bee colonies, commonly in man-made beehives. Honey bees in the genus '' Apis'' are the most-commonly-kept species but other honey-producing bees such as ''Melipona'' stingless bees are also kept. ...
or apiculture (meliponiculture for stingless bees) has been practised for millennia, since at least the times of Ancient Egypt and
Ancient Greece Ancient Greece ( el, Ἑλλάς, Hellás) was a northeastern Mediterranean civilization, existing from the Greek Dark Ages of the 12th–9th centuries BC to the end of classical antiquity ( AD 600), that comprised a loose collection of cult ...
. Bees have appeared in mythology and folklore, through all phases of art and literature from ancient times to the present day, although primarily focused in the
Northern Hemisphere The Northern Hemisphere is the half of Earth that is north of the Equator. For other planets in the Solar System, north is defined as being in the same celestial hemisphere relative to the invariable plane of the solar system as Earth's Nort ...
where
beekeeping Beekeeping (or apiculture) is the maintenance of bee colonies, commonly in man-made beehives. Honey bees in the genus '' Apis'' are the most-commonly-kept species but other honey-producing bees such as ''Melipona'' stingless bees are also kept. ...
is far more common. In
Mesoamerica Mesoamerica is a historical region and cultural area in southern North America and most of Central America. It extends from approximately central Mexico through Belize, Guatemala, El Salvador, Honduras, Nicaragua, and northern Costa Rica. W ...
, the
Mayans The Maya peoples () are an ethnolinguistic group of indigenous peoples of Mesoamerica. The ancient Maya civilization was formed by members of this group, and today's Maya are generally descended from people who lived within that historical reg ...
have practiced large-scale intensive meliponiculture since pre-Columbian times.


Evolution

The immediate ancestors of bees were stinging wasps in the family
Crabronidae The Crabronidae are a large paraphyletic group (nominally a family) of wasps, including nearly all of the species formerly comprising the now-defunct superfamily Sphecoidea. It collectively includes well over 200 genera, containing well over 9 ...
, which were
predators Predation is a biological interaction where one organism, the predator, kills and eats another organism, its prey. It is one of a family of common feeding behaviours that includes parasitism and micropredation (which usually do not kill the ...
of other insects. The switch from insect prey to pollen may have resulted from the consumption of prey insects which were flower visitors and were partially covered with pollen when they were fed to the wasp larvae. This same
evolution Evolution is change in the heritable characteristics of biological populations over successive generations. These characteristics are the expressions of genes, which are passed on from parent to offspring during reproduction. Variation ...
ary scenario may have occurred within the vespoid wasps, where the
pollen wasp Pollen wasps, the Masarinae, are unusual wasps that are typically treated as a subfamily of Vespidae, but have in the past sometimes been recognized as a separate family, "Masaridae", which also included the subfamilies Euparagiinae and Gayellina ...
s evolved from predatory ancestors. The oldest non-compression bee fossil is found in
New Jersey amber New Jersey Amber, sometimes called Raritan amber, is amber found in the Raritan and Magothy Formations of the Central Atlantic (Eastern) coast of the United States. It is dated to the Late Cretaceous, Turonian age, based on pollen analysis of th ...
, '' Cretotrigona prisca'', a corbiculate bee of Cretaceous age (~65 mya). A fossil from the
early Cretaceous The Early Cretaceous ( geochronological name) or the Lower Cretaceous (chronostratigraphic name), is the earlier or lower of the two major divisions of the Cretaceous. It is usually considered to stretch from 145  Ma to 100.5 Ma. Geology Pro ...
(~100 mya), ''
Melittosphex burmensis ''Melittosphex burmensis'' is an aculeate wasp that was formerly considered one of the two oldest-known species of bees. The species was described from an inclusion in Burmese amber in the year 2006 by George Poinar Jr., a zoologist at Oregon ...
'', was initially considered "an extinct lineage of pollen-collecting Apoidea
sister A sister is a woman or a girl who shares one or more parents with another individual; a female sibling. The male counterpart is a brother. Although the term typically refers to a familial relationship, it is sometimes used endearingly to refer to ...
to the modern bees", but subsequent research has rejected the claim that ''Melittosphex'' is a bee, or even a member of the superfamily
Apoidea The superfamily (zoology), superfamily Apoidea is a major group within the Hymenoptera, which includes two traditionally recognized lineages, the "sphecidae, sphecoid" wasps, and the bees. Molecular phylogeny demonstrates that the bees arose from ...
to which bees belong, instead treating the lineage as ''
incertae sedis ' () or ''problematica'' is a term used for a taxonomic group where its broader relationships are unknown or undefined. Alternatively, such groups are frequently referred to as "enigmatic taxa". In the system of open nomenclature, uncertainty ...
'' within the
Aculeata Aculeata is a subclade of Hymenoptera containing ants, bees, and stinging wasps. The name is a reference to the defining feature of the group, which is the modification of the ovipositor into a stinger. However, many members of the group cannot ...
. By the
Eocene The Eocene ( ) Epoch is a geological epoch (geology), epoch that lasted from about 56 to 33.9 million years ago (mya). It is the second epoch of the Paleogene Period (geology), Period in the modern Cenozoic Era (geology), Era. The name ''Eocene' ...
(~45 mya) there was already considerable diversity among eusocial bee lineages. The highly
eusocial Eusociality (from Greek εὖ ''eu'' "good" and social), the highest level of organization of sociality, is defined by the following characteristics: cooperative brood care (including care of offspring from other individuals), overlapping generat ...
corbiculate Apidae appeared roughly 87 Mya, and the
Allodapini The Allodapini is a tribe of bees in the subfamily Xylocopinae, family Apidae. They occur throughout sub-Saharan Africa, South East Asia, and Australasia. There is also a rare genus, Exoneuridia, that occurs in isolated regions of Turkey, Iraq, L ...
(within the Apidae) around 53 Mya. The
Colletidae The Colletidae are a family of bees, and are often referred to collectively as plasterer bees or polyester bees, due to the method of smoothing the walls of their nest cells with secretions applied with their mouthparts; these secretions dry int ...
appear as fossils only from the late
Oligocene The Oligocene ( ) is a geologic epoch of the Paleogene Period and extends from about 33.9 million to 23 million years before the present ( to ). As with other older geologic periods, the rock beds that define the epoch are well identified but the ...
(~25 Mya) to early
Miocene The Miocene ( ) is the first geological epoch of the Neogene Period and extends from about (Ma). The Miocene was named by Scottish geologist Charles Lyell; the name comes from the Greek words (', "less") and (', "new") and means "less recen ...
. The
Melittidae Melittidae is a small bee family, with over 200 described species in three subfamilies. The family has a limited distribution, with all described species restricted to Africa and the northern temperate zone. Fossil melittids have been found oc ...
are known from ''Palaeomacropis eocenicus'' in the
Early Eocene In the geologic timescale the Ypresian is the oldest age or lowest stratigraphic stage of the Eocene. It spans the time between , is preceded by the Thanetian Age (part of the Paleocene) and is followed by the Eocene Lutetian Age. The Ypresian i ...
. The
Megachilidae Megachilidae is a cosmopolitan family of mostly solitary bees. Both that their pollen-carrying structure (called a ''scopa'') is restricted to the ventral surface of the abdomen (rather than mostly or exclusively on the hind legs as in other b ...
are known from trace fossils (characteristic leaf cuttings) from the
Middle Eocene The Eocene ( ) Epoch is a geological epoch that lasted from about 56 to 33.9 million years ago (mya). It is the second epoch of the Paleogene Period in the modern Cenozoic Era. The name ''Eocene'' comes from the Ancient Greek (''ēṓs'', "dawn ...
. The
Andrenidae The Andrenidae (commonly known as mining bees) are a large, nearly cosmopolitan family of solitary, ground-nesting bees. Most of the family's diversity is located in temperate or arid areas (warm temperate xeric). It includes some enormous gene ...
are known from the Eocene-Oligocene boundary, around 34 Mya, of the Florissant shale. The
Halictidae Halictidae is the second-largest family of bees (clade Anthophila) with nearly 4,500 species. Halictid species are an extremely diverse group that can vary greatly in appearance. These bees occur all over the world and are found on every contine ...
first appear in the Early Eocene with species found in amber. The
Stenotritidae The Stenotritidae is the smallest of all formally recognised bee families, with only 21 species in two genera, all of them restricted to Australia. Historically, they were generally considered to belong in the family Colletidae, but the stenotrit ...
are known from fossil brood cells of
Pleistocene The Pleistocene ( , often referred to as the ''Ice age'') is the geological Epoch (geology), epoch that lasted from about 2,580,000 to 11,700 years ago, spanning the Earth's most recent period of repeated glaciations. Before a change was fina ...
age.


Coevolution

The earliest animal-pollinated flowers were shallow, cup-shaped blooms
pollinated Pollination is the transfer of pollen from an anther of a plant to the stigma of a plant, later enabling fertilisation and the production of seeds, most often by an animal or by wind. Pollinating agents can be animals such as insects, birds, ...
by insects such as
beetle Beetles are insects that form the order Coleoptera (), in the superorder Endopterygota. Their front pair of wings are hardened into wing-cases, elytra, distinguishing them from most other insects. The Coleoptera, with about 400,000 describ ...
s, 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 In biology, coevolution occurs when two or more species reciprocally affect each other's evolution through the process of natural selection. The term sometimes is used for two traits in the same species affecting each other's evolution, as well ...
, flowers developed floral rewards such as
nectar Nectar is a sugar-rich liquid produced by plants in glands called nectaries or nectarines, either within the flowers with which it attracts pollinating animals, or by extrafloral nectaries, which provide a nutrient source to animal mutualists ...
and longer tubes, and bees developed longer tongues to extract the nectar. Bees also developed structures known as scopal hairs and
pollen baskets The pollen basket or corbicula (plural corbiculae) is part of the tibia on the hind legs of certain species of bees. They use the structure in harvesting pollen and carrying it to the nest or hive. Other species of bees have scopae instead. Et ...
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 The pollen basket or corbicula (plural corbiculae) is part of the tibia on the hind legs of certain species of bees. They use the structure in harvesting pollen and carrying it to the nest or hive. Other species of bees have scopae instead. Et ...
on their hind legs, while very few lack these and instead collect pollen in their crops. The appearance of these structures drove the
adaptive radiation In evolutionary biology, adaptive radiation is a process in which organisms diversify rapidly from an ancestral species into a multitude of new forms, particularly when a change in the environment makes new resources available, alters biotic int ...
of the
angiosperms Flowering plants are plants that bear flowers and fruits, and form the clade Angiospermae (), commonly called angiosperms. The term "angiosperm" is derived from the Greek words ('container, vessel') and ('seed'), and refers to those plants th ...
, and, in turn, bees themselves. Bees
coevolved In biology, coevolution occurs when two or more species reciprocally affect each other's evolution through the process of natural selection. The term sometimes is used for two traits in the same species affecting each other's evolution, as well ...
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.


Phylogeny


External

This
phylogenetic tree A phylogenetic tree (also phylogeny or evolutionary tree Felsenstein J. (2004). ''Inferring Phylogenies'' Sinauer Associates: Sunderland, MA.) is a branching diagram or a tree showing the evolutionary relationships among various biological spec ...
is based on Debevic ''et al'', 2012, which used molecular phylogeny to demonstrate that the bees (
Anthophila Bees are winged insects closely related to wasps and ants, known for their roles in pollination and, in the case of the best-known bee species, the western honey bee, for producing honey. Bees are a monophyletic lineage within the superfami ...
) arose from deep within the
Crabronidae The Crabronidae are a large paraphyletic group (nominally a family) of wasps, including nearly all of the species formerly comprising the now-defunct superfamily Sphecoidea. It collectively includes well over 200 genera, containing well over 9 ...
, which is therefore
paraphyletic In taxonomy (general), taxonomy, a group is paraphyletic if it consists of the group's most recent common ancestor, last common ancestor and most of its descendants, excluding a few Monophyly, monophyletic subgroups. The group is said to be pa ...
. The placement of the Heterogynaidae is uncertain. The small subfamily
Mellininae Mellininae is a very small subfamily of wasps, comprising only 17 described species in two genera. Occasionally this group has been treated as a separate family (Mellinidae), but has been more typically considered a subfamily within the all-inclu ...
was not included in this analysis.


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. English names, where available, are given in parentheses.


Characteristics

Bees differ from closely related groups such as wasps by having branched or plume-like
seta In biology, setae (singular seta ; from the Latin word for "bristle") are any of a number of different bristle- or hair-like structures on living organisms. Animal setae Protostomes Annelid setae are stiff bristles present on the body. Th ...
e (hairs), combs on the forelimbs for cleaning their antennae, small anatomical differences in limb structure, and the venation of the hind wings; and in females, by having the seventh dorsal abdominal plate divided into two half-plates. Bees have the following characteristics: * A pair of large
compound eyes A compound eye is a visual organ found in arthropods such as insects and crustaceans. It may consist of thousands of ommatidia, which are tiny independent photoreception units that consist of a cornea, lens, and photoreceptor cells which distin ...
which cover much of the surface of the head. Between and above these are three small simple eyes (
ocelli A simple eye (sometimes called a pigment pit) refers to a form of eye or an optical arrangement composed of a single lens and without an elaborate retina such as occurs in most vertebrates. In this sense "simple eye" is distinct from a multi-l ...
) 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 In anatomy, the mandible, lower jaw or jawbone is the largest, strongest and lowest bone in the human facial skeleton. It forms the lower jaw and holds the lower teeth in place. The mandible sits beneath the maxilla. It is the only movable bone ...
and a long
proboscis A proboscis () is an elongated appendage from the head of an animal, either a vertebrate or an invertebrate. In invertebrates, the term usually refers to tubular mouthparts used for feeding and sucking. In vertebrates, a proboscis is an elong ...
for sucking up nectar. * 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. The largest species of bee is thought to be Wallace's giant bee ''
Megachile pluto ''Megachile pluto'', also known as Wallace's giant bee or (), is a large resin bee found in Indonesia. With a wingspan of it is the largest known living bee species. It was believed to be extinct until several specimens were discovered in 1981 ...
'', whose females can attain a length of . The smallest species may be dwarf stingless bees in the tribe
Meliponini Stingless bees, sometimes called stingless honey bees or simply meliponines, are a large group of bees (about 550 described species), comprising the tribe Meliponini (or subtribe Meliponina according to other authors). They belong in the family A ...
whose workers are less than in length.


Sociality


Haplodiploid breeding system

According to
inclusive fitness In evolutionary biology, inclusive fitness is one of two metrics of evolutionary success as defined by W. D. Hamilton in 1964: * Personal fitness is the number of offspring that an individual begets (regardless of who rescues/rears/supports them ...
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 Haplodiploidy is a sex-determination system in which males develop from unfertilized eggs and are haploid, and females develop from fertilized eggs and are diploid. Haplodiploidy is sometimes called arrhenotoky. Haplodiploidy determines the se ...
species such as bees because of their unusual relatedness structure. In haplodiploid species, females develop from fertilized eggs and males from unfertilized eggs. Because a male is
haploid Ploidy () is the number of complete sets of chromosomes in a cell, and hence the number of possible alleles for autosomal and pseudoautosomal genes. Sets of chromosomes refer to the number of maternal and paternal chromosome copies, respectively ...
(has only one copy of each gene), his daughters (which are
diploid Ploidy () is the number of complete sets of chromosomes in a cell, and hence the number of possible alleles for autosomal and pseudoautosomal genes. Sets of chromosomes refer to the number of maternal and paternal chromosome copies, respectively ...
, 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 William Donald Hamilton (1 August 1936 – 7 March 2000) was a British evolutionary biologist, recognised as one of the most significant evolutionary theorists of the 20th century. Hamilton became known for his theoretical work expounding a ...
termed "supersisters", more closely related to their sisters than they would be to their own offspring. 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 9) evolutions of eusociality within
Hymenoptera Hymenoptera is a large order (biology), order of insects, comprising the sawfly, sawflies, wasps, bees, and ants. Over 150,000 living species of Hymenoptera have been described, in addition to over 2,000 extinct ones. Many of the species are Par ...
. Haplodiploidy is neither necessary nor sufficient for eusociality. Some eusocial species such as
termites Termites are small insects that live in colonies and have distinct castes (eusocial) and feed on wood or other dead plant matter. Termites comprise the infraorder Isoptera, or alternatively the epifamily Termitoidae, within the order Blattode ...
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. 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.


Eusociality

Bees may be solitary or may live in various types of communities.
Eusociality Eusociality (from Greek εὖ ''eu'' "good" and social), the highest level of organization of sociality, is defined by the following characteristics: cooperative brood care (including care of offspring from other individuals), overlapping genera ...
appears to have originated from at least three independent origins in halictid bees. The most advanced of these are species with
eusocial Eusociality (from Greek εὖ ''eu'' "good" and social), the highest level of organization of sociality, is defined by the following characteristics: cooperative brood care (including care of offspring from other individuals), overlapping generat ...
colonies; these are characterised by cooperative brood care and a
division of labour The division of labour is the separation of the tasks in any economic system or organisation so that participants may specialise (specialisation). Individuals, organizations, and nations are endowed with, or acquire specialised capabilities, and ...
into reproductive and non-reproductive adults, plus overlapping generations. This division of labour creates specialized groups within eusocial societies which are called
castes Caste is a form of social stratification characterised by endogamy, hereditary transmission of a style of life which often includes an occupation, ritual status in a hierarchy, and customary social interaction and exclusion based on cultura ...
. 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 Sociality is the degree to which individuals in an animal population tend to associate in social groups (gregariousness) and form cooperative societies. Sociality is a survival response to evolutionary pressures. For example, when a mother wasp ...
. The group is called eusocial if, in addition, the group consists of a mother (the
queen Queen or QUEEN may refer to: Monarchy * Queen regnant, a female monarch of a Kingdom ** List of queens regnant * Queen consort, the wife of a reigning king * Queen dowager, the widow of a king * Queen mother, a queen dowager who is the mother ...
) and her daughters (
workers The workforce or labour force is a concept referring to the pool of human beings either in employment or in unemployment. It is generally used to describe those working for a single company or industry, but can also apply to a geographic regio ...
). 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 Paper wasps are Vespidae, vespid wasps that gather fibers from dead wood and plant stems, which they mix with saliva, and use to construct nests made of gray or brown papery material. Some types of paper wasps are also sometimes called umbrell ...
s; when the castes are morphologically discrete, the system is considered highly eusocial. 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 The western honey bee or European honey bee (''Apis mellifera'') is the most common of the 7–12 species of honey bees worldwide. The genus name ''Apis'' is Latin for "bee", and ''mellifera'' is the Latin for "honey-bearing" or "honey carrying", ...
'', 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. Stingless bees are also highly eusocial. They practise mass provisioning, with complex nest architecture and perennial colonies also established via swarming. 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. 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. 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 Halictidae is the second-largest family of bees (clade Anthophila) with nearly 4,500 species. Halictid species are an extremely diverse group that can vary greatly in appearance. These bees occur all over the world and are found on every contine ...
, 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''. Some species are eusocial in parts of their range and solitary in others, or have a mix of eusocial and solitary nests in the same population. The orchid bees (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.


Solitary and communal bees

Most other bees, including familiar insects such as
carpenter bee Carpenter bees are species in the genus ''Xylocopa'' of the subfamily Xylocopinae. The genus includes some 500 bees in 31 subgenera. The common name "carpenter bee" derives from their nesting behavior; nearly all species burrow into hard plant m ...
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 Pollen wasps, the Masarinae, are unusual wasps that are typically treated as a subfamily of Vespidae, but have in the past sometimes been recognized as a separate family, "Masaridae", which also included the subfamilies Euparagiinae and Gayellina ...
s have similar behaviours, and a few species of bee Scavenger, 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 genus, genera which are commonly known by their nesting behavior or preferences, namely: carpenter bees, sweat bees, mason bees, Colletes inaequalis, plasterer bees, squash bees, Ceratina, dwarf carpenter bees, leafcutter bees, alkali bees and Anthophorini, digger bees. 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 The Andrenidae (commonly known as mining bees) are a large, nearly cosmopolitan family of solitary, ground-nesting bees. Most of the family's diversity is located in temperate or arid areas (warm temperate xeric). It includes some enormous gene ...
) are stingless. While solitary, females each make individual nests. Some species, such as the European mason bee ''Hoplitis anthocopoides'', and the Amegilla dawsoni, Dawson's Burrowing bee, ''Amegilla dawsoni,'' 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.


Biology


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 larva (; plural larvae ) is a distinct juvenile form many animals undergo before metamorphosis into adults. Animals with indirect development such as insects, amphibians, or cnidarians typically have a larval phase of their life cycle. The ...
, a pupation stage during which the insect undergoes Holometabolism, complete metamorphosis, followed by the emergence of a winged adult. The number of eggs laid by a female during her lifetime can vary from eight or less in some solitary bees, to more than a million in highly social species. Most solitary bees and bumble bees in temperate climates overwinter as adults or pupae and emerge in spring when increasing numbers of flowering plants come into bloom. The males usually emerge first and search for females with which to mate. Like the other members of Hymenoptera bees are Haplodiploidy, haplodiploid; the sex of a bee is determined by whether or not the egg is fertilised. After mating, a female stores the sperm, and determines which sex is required at the time each individual egg is laid, fertilised eggs producing female offspring and unfertilised eggs, males. Tropical bees may have several generations in a year and no diapause stage. 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. In most species, larvae are whitish grubs, roughly oval and bluntly-pointed at both ends. They have 15 segments and Spiracle (arthropods), 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 Bombus pascuorum, common carder
bumblebee A bumblebee (or bumble bee, bumble-bee, or humble-bee) is any of over 250 species in the genus ''Bombus'', part of Apidae, one of the bee families. This genus is the only extant group in the tribe Bombini, though a few extinct related gener ...
, wax canopy removed to show winged worker (bee), workers and pupae in irregularly placed wax cells File:Carpenter Bee Galleries.jpeg, Carpenter bee nests in a cedar wood beam (sawn open) File:Bienen mit Brut 2.jpg, Honeybees on brood comb with eggs and larvae in cells


Flight

Antoine Magnan's 1934 book ''Le vol des insectes'', says that he and André Sainte-Laguë had applied the equations of air resistance to
insect Insects (from Latin ') are pancrustacean hexapod invertebrates of the class Insecta. They are the largest group within the arthropod phylum. Insects have a chitinous exoskeleton, a three-part body ( head, thorax and abdomen), three pairs ...
s and found that their flight could not be explained by fixed-wing calculations, but that "One shouldn't be surprised that the results of the calculations don't square with reality". 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. In 1996 it was shown that vortices created by many insects' wings helped to provide lift. High-speed cinematography and robotic mock-up of a bee wing Re: work of Michael Dickinson (biologist), 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 (geometry), arc, it flaps approximately 230 times per second, faster than a Drosophilidae, fruitfly (200 times per second) which is 80 times smaller.


Navigation, communication, and finding food

The ethologist Karl von Frisch studied animal navigation, navigation in the honey bee. He showed that honey bees communicate by the waggle dance, in which a worker indicates the location of a food source to other workers in the hive. He demonstrated that bees can recognize a desired compass direction in three different ways: by the sun, by the Polarization (waves), polarization pattern of the blue sky, and by the earth's magnetic field. He showed that the sun is the preferred or main compass; the other mechanisms are used under cloudy skies or inside a dark beehive. Bees navigate using spatial memory with a "rich, map-like organization".


Digestion

The gut of bees is relatively simple, but multiple metabolic strategies exist in the gut microbiota. 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. Approximately five groups of bacteria are involved in digestion. Three groups specialize in simple sugars (''Snodgrassella'' and two groups of ''Lactobacillus''), and two other groups in complex sugars (''Gilliamella'' and ''Bifidobacterium''). Digestion of pectin and hemicellulose is dominated by bacterial
clade A clade (), also known as a monophyletic group or natural group, is a group of organisms that are monophyletic – that is, composed of a common ancestor and all its lineal descendants – on a phylogenetic tree. Rather than the English term, ...
s ''Gilliamella'' and ''Bifidobacterium'' respectively. Bacteria that cannot digest polysaccharides obtain enzymes from their neighbors, and bacteria that lack certain amino acids do the same, creating multiple ecological niches. Although most bee species are Nectarivore, nectarivorous and Palynivore, 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.


Ecology


Floral relationships

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. Specialist pollinators also include bee species which gather floral oils instead of pollen, and male orchid bees, which gather aromatic compounds from orchids (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, and even electromagnetic fields. Once landed, a bee then uses nectar quality and pollen taste 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.


As mimics and models

Many bees are aposematically coloured, typically orange and black, warning of their ability to defend themselves with a powerful sting. As such they are models for Batesian mimicry by non-stinging insects such as bee-flies, Asilidae, robber flies and hoverflies, all of which gain a measure of protection by superficially looking and behaving like bees. Bees are themselves Müllerian mimicry, Müllerian mimics of other aposematic insects with the same colour scheme, including
wasp A wasp is any insect of the narrow-waisted suborder Apocrita of the order Hymenoptera which is neither a bee nor an ant; this excludes the broad-waisted sawflies (Symphyta), which look somewhat like wasps, but are in a separate suborder. Th ...
s, Lycidae, lycid and other beetles, and many butterflies and moths (Lepidoptera) which are themselves distasteful, often through acquiring bitter and poisonous chemicals from their plant food. All the Müllerian mimics, including bees, benefit from the reduced risk of predation that results from their easily recognised warning coloration. 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.


As brood parasites

Brood parasites occur in several bee families including the apidae, apid subfamily Nomadinae. 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. 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. 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 Ploidy () is the number of complete sets of chromosomes in a cell, and hence the number of possible alleles for autosomal and pseudoautosomal genes. Sets of chromosomes refer to the number of maternal and paternal chromosome copies, respectively ...
eggs ("thelytoky"), escaping normal worker policing, leading to the colony's destruction; the parasites can then move to other hives. The cuckoo bees 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'', while the other species in the same genus attacks halictidae, halictid bees.


Nocturnal bees

Four bee families (
Andrenidae The Andrenidae (commonly known as mining bees) are a large, nearly cosmopolitan family of solitary, ground-nesting bees. Most of the family's diversity is located in temperate or arid areas (warm temperate xeric). It includes some enormous gene ...
,
Colletidae The Colletidae are a family of bees, and are often referred to collectively as plasterer bees or polyester bees, due to the method of smoothing the walls of their nest cells with secretions applied with their mouthparts; these secretions dry int ...
,
Halictidae Halictidae is the second-largest family of bees (clade Anthophila) with nearly 4,500 species. Halictid species are an extremely diverse group that can vary greatly in appearance. These bees occur all over the world and are found on every contine ...
, 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.


Predators, parasites and pathogens

Vertebrate predators of bees include
bee-eater The bee-eaters are a group of non-passerine birds in the family Meropidae, containing three genera and thirty species. Most species are found in Africa and Asia, with a few in southern Europe, Australia, and New Guinea. They are characterised by ...
s, shrikes and Old World flycatcher, flycatchers, which make short sallies to catch insects in flight. Swifts and swallows fly almost continually, catching insects as they go. The Pernis (bird), honey buzzard attacks bees' nests and eats the larvae. 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. Among mammals, predators such as the badger dig up bumblebee nests and eat both the larvae and any stored food. Specialist ambush predators of visitors to flowers include crab spiders, which wait on flowering plants for pollinating insects; Hemiptera, predatory bugs, and praying mantises, some of which (the flower mantises of the tropics) wait motionless, aggressive mimicry, aggressive mimics camouflaged as flowers. Beewolf, 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. Other predatory insects that sometimes catch bees include Asilidae, robber flies and
dragonflies A dragonfly is a flying insect belonging to the infraorder Anisoptera below the order Odonata. About 3,000 extant species of true dragonfly are known. Most are tropical, with fewer species in temperate regions. Loss of wetland habitat threa ...
. Honey bees are affected by parasites including Acarapis woodi, tracheal and ''Varroa'' mites. 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.


Relationship with humans


In mythology and folklore

Homer's ''Homeric Hymns, Hymn to Hermes'' describes three bee-maidens with the power of divination and thus speaking truth, and identifies the food of the gods as honey. Sources associated the bee maidens with Apollo and, until the 1980s, scholars followed Gottfried Hermann (1806) in incorrectly identifying the bee-maidens with the Thriae. Honey, according to a Greek myth, was discovered by a nymph called Melissa ("Bee"); and honey was offered to the Greek gods from Helladic period, Mycenean times. Bees were also associated with the Delphic oracle and the prophetess was sometimes called a bee. The image of a community of honey bees has been used from ancient to modern times, in Aristotle and Plato; in Virgil and Seneca the Younger, Seneca; in Erasmus and Shakespeare; Tolstoy, and by political and social theorists such as Bernard Mandeville and Karl Marx as a model for human society. In English folklore, bees would be told of important events in the household, in a custom known as "Telling the bees".


In art and literature

Some of the oldest examples of bees in art are Cave painting, rock paintings in Spain which have been dated to 15,000 BC. W. B. Yeats's poem ''The Lake Isle of Innisfree'' (1888) contains the couplet "Nine bean rows will I have there, a hive for the honey bee, / And live alone in the bee loud glade." At the time he was living in Bedford Park, London, Bedford Park in the West of London. Beatrix Potter's illustrated book ''The Tale of Mrs Tittlemouse'' (1910) features Babbity Bumble and her brood ''(pictured)''. Kit Williams' Treasure hunt (game), treasure hunt book ''The Bee on the Comb'' (1984) uses bees and beekeeping as part of its story and puzzle. Sue Monk Kidd's ''The Secret Life of Bees (novel), The Secret Life of Bees'' (2004), and the The Secret Life of Bees (film), 2009 film starring Dakota Fanning, tells the story of a girl who escapes her abusive home and finds her way to live with a family of beekeepers, the Boatwrights. The 2007 animated comedy film ''Bee Movie'' used Jerry Seinfeld's first script and was his first work for children; he starred as a bee named Barry B. Benson, alongside Renée Zellweger. Critics found its premise awkward and its delivery tame. Dave Goulson's ''A Sting in the Tale'' (2014) describes his efforts to save bumblebees in Britain, as well as much about their biology. The playwright Laline Paull's fantasy ''The Bees'' (2015) tells the tale of a hive bee named Flora 717 from hatching onwards.


Beekeeping

Humans have kept honey bee colonies, commonly in beehive, hives, for millennia. Beekeepers collect
honey Honey is a sweet and viscous substance made by several bees, the best-known of which are honey bees. Honey is made and stored to nourish bee colonies. Bees produce honey by gathering and then refining the sugary secretions of plants (primar ...
, beeswax, propolis,
pollen Pollen is a powdery substance produced by seed plants. It consists of pollen grains (highly reduced microgametophytes), which produce male gametes (sperm cells). Pollen grains have a hard coat made of sporopollenin that protects the gametophyt ...
, and royal jelly from hives; bees are also kept to pollination, pollinate crops and to produce bees for sale to other beekeepers. Depictions of humans collecting honey from wild bees date to 15,000 years ago; efforts to domesticate them are shown in Egyptian art around 4,500 years ago. Simple hives and smoke were used; jars of honey were found in the tombs of pharaohs such as Tutankhamun. 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. Among Classical Era authors, beekeeping with the use of smoke is described in Aristotle's ''History of Animals'' Book 9. The account mentions that bees die after stinging; that workers remove corpses from the hive, and guard it; castes including workers and non-working drone (bee), drones, but "kings" rather than queens; predators including toads and bee-eaters; and the waggle dance, with the "irresistible suggestion" of άροσειονται ("aroseiontai", it waggles) and παρακολουθούσιν ("parakolouthousin", they watch). Beekeeping is described in detail by Virgil in his ''Georgics''; it is also mentioned in his ''Aeneid'', and in Pliny's ''Natural History (Pliny), Natural History''.


As commercial pollinators

Bees play an important role in pollination, pollinating
flowering plant Flowering plants are plants that bear flowers and fruits, and form the clade Angiospermae (), commonly called angiosperms. The term "angiosperm" is derived from the Greek words ('container, vessel') and ('seed'), and refers to those plants th ...
s, 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 Pollination is the transfer of pollen from an anther of a plant to the stigma of a plant, later enabling fertilisation and the production of seeds, most often by an animal or by wind. Pollinating agents can be animals such as insects, birds ...
by insects, birds and bats, most of which is accomplished by bees, whether wild or domesticated. 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. pollination management, 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. The number of colonies kept by beekeepers declined slightly, through urbanization, systematic pesticide use, Acarapis woodi, tracheal and ''Varroa'' mites, and the closure of beekeeping businesses. In 2006 and 2007 the rate of attrition increased, and was described as colony collapse disorder. In 2010 invertebrate iridescent virus and the fungus ''Nosema ceranae'' were shown to be in every killed colony, and deadly in combination. Winter losses increased to about 1/3. ''Varroa'' mites were thought to be responsible for about half the losses. 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. From 2013 the European Union restricted some pesticides to stop bee populations from declining further. In 2014 the Intergovernmental Panel on Climate Change report warned that bees faced increased risk of extinction because of global warming. 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. Farmers have focused on alternative solutions to mitigate these problems. By raising native plants, they provide food for native bee pollinators like ''Lasioglossum vierecki''Kuehn, F. Coordinator. (2015). Farming for native bees. World Wide Web electronic publication. Retrieved from . (Accessed: 22 September 2015). and ''Lasioglossum leucozonium, L. leucozonium'',Adamson, Nancy Lee
An Assessment of Non-Apis Bees as Fruit and Vegetable Crop Pollinators in Southwest Virginia
. 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 Cucurbita, squashes and cucumbers. File:A bee covered with pollen.jpg, Bee covered in pollen


As food producers

Honey is a natural product produced by bees and stored for their own use, but its sweetness has always appealed to humans. Before domestication of bees was even attempted, humans were raiding their nests for their honey. Smoke was often used to subdue the bees and such activities are depicted in Cave painting, rock paintings in Spain dated to 15,000 BC. Honey bees are used commercially to produce
honey Honey is a sweet and viscous substance made by several bees, the best-known of which are honey bees. Honey is made and stored to nourish bee colonies. Bees produce honey by gathering and then refining the sugary secretions of plants (primar ...
. They also produce some substances used as dietary supplements with possible health benefits, pollen, propolis, and royal jelly, though all of these can also cause allergy, allergic reactions.


As food

Bees are considered edible insects. People in some countries entomophagy, eat insects, including the larvae and pupae of bees, mostly stingless species. They also gather larvae, pupae and surrounding cells, known as bee brood, for consumption. In the Indonesian dish ''botok, botok tawon'' from Central and East Java, bee larvae are eaten as a companion to rice, after being mixed with shredded coconut, wrapped in banana leaves, and steamed. (This particular Botok recipe uses anchovies, not bees) Bee brood (pupae and larvae) although low in calcium, has been found to be high in
protein Proteins are large biomolecules and macromolecules that comprise one or more long chains of amino acid residues. Proteins perform a vast array of functions within organisms, including catalysing metabolic reactions, DNA replication, respo ...
and carbohydrate, and a useful source of phosphorus, magnesium, potassium, and trace minerals iron, zinc, copper, and selenium. In addition, while bee brood was high in fat, it contained no fat soluble vitamins (such as A, D, and E) but it was a good source of most of the water-soluble B-vitamins including choline as well as vitamin C. The fat was composed mostly of Saturated fat, saturated and monounsaturated fatty acids with 2.0% being polyunsaturated fatty acids. File:Botoktawon.jpg, Bee larvae as food in the Javanese dish ''botok, botok tawon'' File:Fried bees dish.jpg, Fried whole bees served in a Ukrainian restaurant


As alternative medicine

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


Stings

The painful Bee sting, stings of bees are mostly associated with the poison gland and the Dufour's gland which are abdominal exocrine glands containing various chemicals. In ''Lasioglossum leucozonium'', the Dufour's Gland mostly contains octadecanolide as well as some eicosanolide. There is also evidence of n-triscosane, n-heptacosane, and 22-docosanolide. However, the secretions of these glands could also be used for nest construction.


See also

* Australian native bees * Superorganism * World Bee Day


Explanatory notes


References


External links

*
"Apoidea"
at All Living Thingsimages, identification guides, and maps of bees


Anthophila (Apoidea) – Bees
orth American species of bees at BugGuide
Native Bees of North America
at BugGuide
"Bee declines driven by combined stress from parasites, pesticides, and lack of flowers"
'Science (journal), Science'' {{Authority control Bees, Extant Early Cretaceous first appearances