Bees Bees Are Winged Insects Closely Related To Wasps and Ants

Bee
Bees are winged insects closely related to wasps and ants,

known for their roles in pollination and, in the case of the best- Bees
known bee species, the western honey bee, for producing honey. Temporal range: Late Cretaceous –
Bees are a monophyletic lineage within the superfamily Present
Apoidea. They are presently considered a clade, called
Anthophila. There are over 16,000 known species of bees in
seven recognized biological families.[1][2] Some species –
including honey bees, bumblebees, and stingless bees – live
socially in colonies while most species (>90%) – including
mason bees, carpenter bees, leafcutter bees, and sweat bees – are
solitary.
Bees are found on every continent except Antarctica, in every

habitat on the planet that contains insect-pollinated flowering
plants. The most common bees in the Northern Hemisphere are The sugarbag bee, Tetragonula
the Halictidae, or sweat bees, but they are small and often carbonaria
mistaken for wasps or flies. Bees range in size from tiny stingless
Scientific classification
bee species, whose workers are less than 2 millimetres (0.08 in)
long,[3] to Megachile pluto, the largest species of leafcutter bee, Kingdom: Animalia
whose females can attain a length of 39 millimetres (1.54 in).
Phylum: Arthropoda
Bees feed on nectar and pollen, the former primarily as an Class: Insecta
energy source and the latter primarily for protein and other
Order: Hymenoptera
nutrients. Most pollen is used as food for their larvae. Vertebrate
predators of bees include primates and birds such as bee-eaters; (unranked): Unicalcarida
insect predators include beewolves and dragonflies.
Suborder: Apocrita
Bee pollination is important both ecologically and commercially, Superfamily: Apoidea
and the decline in wild bees has increased the value of
pollination by commercially managed hives of honey bees. The Clade: Anthophila
analysis of 353 wild bee and hoverfly species across Britain Families
from 1980 to 2013 found the insects have been lost from a
quarter of the places they inhabited in 1980.[4] Andrenidae
Apidae
Human beekeeping or apiculture (meliponiculture for stingless
bees) has been practised for millennia, since at least the times of Colletidae
Ancient Egypt and Ancient Greece. Bees have appeared in
Halictidae
mythology and folklore, through all phases of art and literature
from ancient times to the present day, although primarily focused Megachilidae
Melittidae
Stenotritidae
Synonyms
in the Northern Hemisphere where beekeeping is far more
common. In Mesoamerica, the Mayans have practiced large- Apiformes (from Latin 'apis')
scale intensive meliponiculture since pre-Columbian times.[3]
Evolution
The immediate ancestors of bees were stinging wasps in the family Crabronidae, which were predators of
other insects. The switch from insect prey to pollen may have resulted from the consumption of prey insects
which were flower visitors and were partially covered with pollen when they were fed to the wasp larvae.
This same evolutionary scenario may have occurred within the vespoid wasps, where the pollen wasps
evolved from predatory ancestors. The oldest non-compression bee fossil is found in New Jersey amber,
Cretotrigona prisca, a corbiculate bee of Cretaceous age (~65 mya).[5] A fossil from the early Cretaceous
(~100 mya), Melittosphex burmensis, was initially considered "an extinct lineage of pollen-collecting
Apoidea sister to the modern bees",[6] but subsequent research has rejected the claim that Melittosphex is a
bee, or even a member of the superfamily Apoidea to which bees belong, instead treating the lineage as
incertae sedis within the Aculeata.[7] By the Eocene (~45 mya) there was already considerable diversity
among eusocial bee lineages.[8][a]
The highly eusocial corbiculate Apidae appeared roughly 87 Mya, and the Allodapini (within the Apidae)
around 53 Mya.[11] The Colletidae appear as fossils only from the late Oligocene (~25 Mya) to early
Miocene.[12] The Melittidae are known from Palaeomacropis eocenicus in the Early Eocene.[13] The
Megachilidae are known from trace fossils (characteristic leaf cuttings) from the Middle Eocene.[14] The
Andrenidae are known from the Eocene-Oligocene boundary, around 34 Mya, of the Florissant shale.[15]
The Halictidae first appear in the Early Eocene[16] with species[17][18] found in amber. The Stenotritidae
are known from fossil brood cells of Pleistocene age.[19]
Coevolution
The earliest animal-pollinated flowers were shallow, cup-shaped

blooms pollinated by insects such as beetles, so the syndrome of
insect pollination was well established before the first appearance
of bees. The novelty is that bees are specialized as pollination
agents, with behavioral and physical modifications that specifically
enhance pollination, and are the most efficient pollinating insects.
In a process of coevolution, flowers developed floral rewards[20]
such as nectar and longer tubes, and bees developed longer tongues
to extract the nectar.[21] Bees also developed structures known as
scopal hairs and pollen baskets to collect and carry pollen. The
location and type differ among and between groups of bees. Most
species have scopal hairs on their hind legs or on the underside of
their abdomens. Some species in the family Apidae have pollen
Long-tongued bees and long-tubed
baskets on their hind legs, while very few lack these and instead flowers coevolved, like this Amegilla
collect pollen in their crops.[2] The appearance of these structures cingulata (Apidae) on Acanthus
drove the adaptive radiation of the angiosperms, and, in turn, bees ilicifolius.
themselves.[9] Bees coevolved not only with flowers but it is
believed that some species coevolved with mites. Some provide tufts of hairs called acarinaria that appear to
provide lodgings for mites; in return, it is believed that mites eat fungi that attack pollen, so the relationship
in this case may be mutualistic.[22][23]
Phylogeny
External
This phylogenetic tree is based on Debevic et al, 2012, which used molecular phylogeny to demonstrate
that the bees (Anthophila) arose from deep within the Crabronidae, which is therefore paraphyletic. The
placement of the Heterogynaidae is uncertain.[24] The small subfamily Mellininae was not included in this
analysis.
⁠Ampulicidae (Cockroach wasps)

⁠
⁠
⁠Heterogynaidae (possible placement #1)
⁠Sphecidae (sensu stricto)

⁠
⁠
⁠
⁠Crabroninae (part of "Crabronidae")
⁠
⁠Bembicini
⁠
⁠Apoidea
⁠
⁠ ⁠Nyssonini, Astatinae
⁠
⁠ ⁠
⁠
⁠
⁠
⁠( rest of "Crabronidae") ⁠Heterogynaidae (possible placement #2)
⁠
⁠
⁠
⁠Pemphredoninae, Philanthinae
⁠
⁠
⁠
⁠Anthophila (bees)
⁠
⁠
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.[25] English names, where available,
are given in parentheses.
⁠Melittidae (inc. Dasypodainae, Meganomiinae) at least 50 Mya

⁠
Apidae (inc. honeybees, cuckoo bees, carpenter bees)

⁠
⁠≈87 Mya
⁠long-tongued bees
⁠
⁠Megachilidae (mason, leafcutter bees) ≈50 Mya

⁠
⁠Anthophila (bees) ⁠ ⁠Andrenidae (mining bees) ≈34 Mya

⁠ ⁠ ⁠
⁠
⁠
⁠Halictidae (sweat bees) ≈50 Mya
⁠
⁠short-tongued bees
⁠
⁠
⁠ ⁠Colletidae (plasterer bees) ≈25 Mya
⁠
⁠
⁠⁠
⁠Stenotritidae (large Australian bees) ≈2 Mya
Characteristics
Bees differ from closely related groups such as wasps by having branched
or plume-like setae (hairs), combs on the forelimbs for cleaning their
antennae, small anatomical differences in limb structure, and the venation
of the hind wings; and in females, by having the seventh dorsal abdominal
plate divided into two half-plates.[26]
Bees have the following characteristics:
A pair of large compound eyes which cover much of the surface

of the head. Between and above these are three small simple
eyes (ocelli) which provide information on light intensity.
The antennae usually have 13 segments in males and 12 in
females, and are geniculate, having an elbow joint part way
along. They house large numbers of sense organs that can
detect touch (mechanoreceptors), smell and taste; and small,
hairlike mechanoreceptors that can detect air movement so as The lapping mouthparts of a
to "hear" sounds. honey bee, showing labium
The mouthparts are adapted for both chewing and sucking by and maxillae
having both a pair of mandibles and a long proboscis for
sucking up nectar.[27]
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.[27]
The largest species of bee is thought to be Wallace's giant bee

Megachile pluto, whose females can attain a length of 39
millimetres (1.54 in).[28] The smallest species may be dwarf
stingless bees in the tribe Meliponini whose workers are less than 2
millimetres (0.08 in) in length.[29]
Sociality
Haplodiploid breeding system

Head-on view of a male carpenter
According to inclusive fitness theory, organisms can gain fitness
bee, showing antennae, three ocelli,
not just through increasing their own reproductive output, but also
compound eyes, and mouthparts
that of close relatives. In evolutionary terms, individuals should
help relatives when Cost < Relatedness * Benefit. The
requirements for eusociality are more easily fulfilled by
haplodiploid species such as bees because of their
unusual relatedness structure.[30]
In haplodiploid species, females develop from fertilized

eggs and males from unfertilized eggs. Because a male
is haploid (has only one copy of each gene), his
daughters (which are diploid, with two copies of each
Willing to die for their sisters: worker honey bees gene) share 100% of his genes and 50% of their
killed defending their hive against yellowjackets, mother's. Therefore, they share 75% of their genes with
along with a dead yellowjacket. Such altruistic
each other. This mechanism of sex determination gives
behaviour may be favoured by the haplodiploid
rise to what W. D. Hamilton termed "supersisters", more
sex determination system of bees.
closely related to their sisters than they would be to their
own offspring.[31] Workers often do not reproduce, but
they can pass on more of their genes by helping to raise their sisters (as queens) than they would by having
their own offspring (each of which would only have 50% of their genes), assuming they would produce
similar numbers. This unusual situation has been proposed as an explanation of the multiple (at least nine)
evolutions of eusociality within Hymenoptera.[32][33]
Haplodiploidy is neither necessary nor sufficient for eusociality. Some eusocial species such as termites are
not haplodiploid. Conversely, all bees are haplodiploid but not all are eusocial, and among eusocial species
many queens mate with multiple males, creating half-sisters that share only 25% of each-other's genes.[34]
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.[32]
Eusociality
Bees may be solitary or may live in various types of communities.

Eusociality appears to have originated from at least three independent
origins in halictid bees.[35] The most advanced of these are species with
eusocial colonies; these are characterised by cooperative brood care and a
division of labour into reproductive and non-reproductive adults, plus
overlapping generations.[36] This division of labour creates specialized
groups within eusocial societies which are called castes. In some species,
groups of cohabiting females may be sisters, and if there is a division of
labour within the group, they are considered semisocial. The group is called
eusocial if, in addition, the group consists of a mother (the queen) and her
daughters (workers). When the castes are purely behavioural alternatives,
with no morphological differentiation other than size, the system is
considered primitively eusocial, as in many paper wasps; when the castes
are morphologically discrete, the system is considered highly eusocial.[21] A Western honey bee swarm
True honey bees (genus Apis, of which eight species are currently
recognized) are highly eusocial, and are among the best known insects. Their colonies are established by
swarms, consisting of a queen and several thousand workers. There are 29 subspecies of one of these
species, Apis mellifera, native to Europe, the Middle East, and Africa. Africanized bees are a hybrid strain
of A. mellifera that escaped from experiments involving crossing European and African subspecies; they
are extremely defensive.[37]
Stingless bees are also highly eusocial. They practise mass
provisioning, with complex nest architecture and perennial colonies
also established via swarming.[3][38]
Many bumblebees are

eusocial, similar to the
eusocial Vespidae such as
hornets in that the queen
initiates a nest on her own Western honey bee nest in the trunk
rather than by swarming. of a spruce
Bumblebee colonies
typically have from 50 to
200 bees at peak population, which occurs in mid to late summer.
A bumblebee carrying pollen in its Nest architecture is simple, limited by the size of the pre-existing
pollen baskets (corbiculae) nest cavity, and colonies rarely last more than a year.[39] 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.[40]
There are many more species of primitively eusocial than highly eusocial bees, but they have been studied
less often. Most are in the family Halictidae, or "sweat bees". Colonies are typically small, with a dozen or
fewer workers, on average. Queens and workers differ only in size, if at all. Most species have a single
season colony cycle, even in the tropics, and only mated females hibernate. A few species have long active
seasons and attain colony sizes in the hundreds, such as Halictus hesperus.[41] Some species are eusocial in
parts of their range and solitary in others,[42] or have a mix of eusocial and solitary nests in the same
population.[43] The orchid bees (Apidae) include some primitively eusocial species with similar biology.
Some allodapine bees (Apidae) form primitively eusocial colonies, with progressive provisioning: a larva's
food is supplied gradually as it develops, as is the case in honey bees and some bumblebees.[44]
Solitary and communal bees
Most other bees, including familiar insects such as carpenter bees,

leafcutter bees and mason bees are solitary in the sense that every
female is fertile, and typically inhabits a nest she constructs herself.
There is no division of labor so these nests lack queens and worker
bees for these species. Solitary bees typically produce neither
honey nor beeswax. Bees collect pollen to feed their young, and
have the necessary adaptations to do this. However, certain wasp
species such as pollen wasps have similar behaviours, and a few
species of bee scavenge from carcases to feed their offspring.[26] A leafcutting bee, Megachile
Solitary bees are important pollinators; they gather pollen to rotundata, cutting circles from acacia
provision their nests with food for their brood. Often it is mixed leaves
with nectar to form a paste-like consistency. Some solitary bees
have advanced types of pollen-carrying structures on their bodies.
Very few species of solitary bee are being cultured for commercial pollination. Most of these species belong
to a distinct set of genera which are commonly known by their nesting behavior or preferences, namely:
carpenter bees, sweat bees, mason bees, plasterer bees, squash bees, dwarf carpenter bees, leafcutter bees,
alkali bees and digger bees.[45]
Most solitary bees are fossorial, digging nests in the ground in a
variety of soil textures and conditions, while others create nests in
hollow reeds or twigs, or holes in wood. The female typically
creates a compartment (a "cell") with an egg and some provisions
for the resulting larva, then seals it off. A nest may consist of
numerous cells. When the nest is in wood, usually the last (those
closer to the entrance) contain eggs that will become males. The
adult does not provide care for the brood once the egg is laid, and
usually dies after making one or more nests. The males typically
emerge first and are ready for mating when the females emerge.
Solitary bees are very unlikely to sting (only in self-defense, if A solitary bee, Anthidium florentinum
ever), and some (esp. in the family Andrenidae) are (family Megachilidae), visiting
stingless.[46][47] Lantana
While solitary, females each make individual nests.[48] Some

species, such as the European mason bee Hoplitis
anthocopoides,[49] and the Dawson's Burrowing bee, Amegilla
dawsoni,[50] are gregarious, preferring to make nests near others of
the same species, and giving the appearance of being social. Large
groups of solitary bee nests are called aggregations, to distinguish
them from colonies. In some species, multiple females share a
common nest, but each makes and provisions her own cells
independently. This type of group is called "communal" and is not
uncommon. The primary advantage appears to be that a nest The mason bee Osmia cornifrons
entrance is easier to defend from predators and parasites when nests in a hole in dead wood. Bee
multiple females use that same entrance regularly.[49] "hotels" are often sold for this
purpose.
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 pupation stage during which the insect undergoes complete
metamorphosis, followed by the emergence of a winged adult. The number of eggs laid by a female during
her lifetime can vary from eight or less in some solitary bees, to more than a million in highly social
species.[51] Most solitary bees and bumble bees in temperate climates overwinter as adults or pupae and
emerge in spring when increasing numbers of flowering plants come into bloom. The males usually emerge
first and search for females with which to mate. Like the other members of Hymenoptera bees are
haplodiploid; the sex of a bee is determined by whether or not the egg is 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.[52][53][54][55]
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.[56]
In most species, larvae are whitish grubs, roughly oval and bluntly-pointed at both ends. They have 15
segments and spiracles in each segment for breathing. They have no legs but move within the cell, helped
by tubercles on their sides. They have short horns on the head, jaws for chewing food and an appendage on
either side of the mouth tipped with a bristle. There is a gland under the mouth that secretes a viscous liquid
which solidifies into the silk they use to produce a cocoon. The cocoon is semi-transparent and the pupa
can be seen through it. Over the course of a few days, the larva undergoes metamorphosis into a winged
adult. When ready to emerge, the adult splits its skin dorsally and climbs out of the exuviae and breaks out
of the cell.[56]
Nest of common Carpenter bee nests in a Honeybees on brood

carder bumblebee, cedar wood beam (sawn comb with eggs and
wax canopy removed open) larvae in cells
to show winged
workers and pupae in
irregularly placed wax
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
insects and found that their flight could not be explained by fixed-
wing calculations, but that "One shouldn't be surprised that the
results of the calculations don't square with reality".[57] 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, Honeybee in flight carrying pollen in
such as those used by helicopters.[58] In 1996 it was shown that pollen basket
vortices created by many insects' wings helped to provide lift.[59]
High-speed cinematography[60] and robotic mock-up of a bee
wing[61] showed that lift was generated by "the unconventional combination of short, choppy wing strokes,
a rapid rotation of the wing as it flops over and reverses direction, and a very fast wing-beat frequency".
Wing-beat frequency normally increases as size decreases, but as the bee's wing beat covers such a small
arc, it flaps approximately 230 times per second, faster than a fruitfly (200 times per second) which is 80
times smaller.[62]
Navigation, communication, and finding food
The ethologist Karl von Frisch studied navigation in the honey bee. He showed that honey bees
communicate by the waggle dance, in which a worker indicates the location of a food source to other
workers in the hive. He demonstrated that bees can recognize a desired compass direction in three different
ways: by the sun, by the polarization pattern of the blue sky, and by the
earth's magnetic field. He showed that the sun is the preferred or main
compass; the other mechanisms are used under cloudy skies or inside a
dark beehive.[63] Bees navigate using spatial memory with a "rich, map-
like organization".[64]
Digestion
The gut of bees is relatively simple, but multiple metabolic strategies exist Karl von Frisch (1953)
in the gut microbiota.[65] Pollinating bees consume nectar and pollen, discovered that honey bee
which require different digestion strategies by somewhat specialized workers can navigate,
bacteria. While nectar is a liquid of mostly monosaccharide sugars and so indicating the range and
easily absorbed, pollen contains complex polysaccharides: branching pectin direction to food to other
and hemicellulose.[66] Approximately five groups of bacteria are involved workers with a waggle
in digestion. Three groups specialize in simple sugars (Snodgrassella and dance.
two groups of Lactobacillus), and two other groups in complex sugars
(Gilliamella and Bifidobacterium). Digestion of pectin and hemicellulose is
dominated by bacterial clades Gilliamella and Bifidobacterium respectively. Bacteria that cannot digest
polysaccharides obtain enzymes from their neighbors, and bacteria that lack certain amino acids do the
same, creating multiple ecological niches.[67]
Although most bee species are nectarivorous and palynivorous, some are not. Particularly unusual are
vulture bees in the genus Trigona, which consume carrion and wasp brood, turning meat into a honey-like
substance.[68]
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.[69] 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,[70] and even electromagnetic fields.[71] Once landed, a bee then uses
nectar quality[70] and pollen taste[72] to determine whether to continue visiting similar flowers.
In rare cases, a plant species may only be effectively pollinated by a single bee species, and some plants are
endangered at least in part because their pollinator is also threatened. But, there is a pronounced tendency
for oligolectic bees to be associated with common, widespread plants visited by multiple pollinator species.
For example, the creosote bush in the arid parts of the United States southwest is associated with some 40
oligoleges.[73]
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, robber flies and hoverflies,[74] all of which gain a measure of protection by superficially
looking and behaving like bees.[74]
Bees are themselves Müllerian

mimics of other aposematic insects
with the same colour scheme,
including wasps, lycid and other
beetles, and many butterflies and
moths (Lepidoptera) which are
themselves distasteful, often
through acquiring bitter and
poisonous chemicals from their
The bee-fly Bombylius major, a plant food. All the Müllerian
Batesian mimic of bees, taking mimics, including bees, benefit Bee orchid lures male bees
nectar and pollinating a flower from the reduced risk of predation to attempt to mate with the
that results from their easily flower's lip, which resembles
recognised warning coloration.[75] a bee perched on a pink
flower.
Bees are also mimicked by plants such as the bee orchid which imitates
both the appearance and the scent of a female bee; male bees attempt to
mate (pseudocopulation) with the furry lip of the flower, thus pollinating it.[76]
As brood parasites
Brood parasites occur in several bee families including the apid

subfamily Nomadinae.[77] Females of these species lack pollen
collecting structures (the scopa) and do not construct their own
nests. They typically enter the nests of pollen collecting species,
and lay their eggs in cells provisioned by the host bee. When the
"cuckoo" bee larva hatches, it consumes the host larva's pollen ball,
and often the host egg also.[78] 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.[79]
Bombus vestalis, a brood parasite of
the bumblebee Bombus terrestris
In Southern Africa, hives of African honeybees (A. mellifera
scutellata) are being destroyed by parasitic workers of the Cape
honeybee, A. m. capensis. These lay diploid eggs ("thelytoky"),
escaping normal worker policing, leading to the colony's destruction; the parasites can then move to other
hives.[80]
The cuckoo bees in the Bombus subgenus Psithyrus are closely related to, and resemble, their hosts in
looks and size. This common pattern gave rise to the ecological principle "Emery's rule". Others parasitize
bees in different families, like Townsendiella, a nomadine apid, two species of which are cleptoparasites of
the dasypodaid genus Hesperapis,[81] while the other species in the same genus attacks halictid bees.[82]
Nocturnal bees
Four bee families (Andrenidae, Colletidae, Halictidae, and Apidae) contain some species that are
crepuscular. Most are tropical or subtropical, but some live in arid regions at higher latitudes. These bees
have greatly enlarged 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.[83]
Predators, parasites and pathogens
Vertebrate predators of bees include bee-eaters, shrikes and

flycatchers, which make short sallies to catch insects in flight.[84]
Swifts and swallows[84] fly almost continually, catching insects as
they go. The honey buzzard attacks bees' nests and eats the
larvae.[85] 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.[86] Among mammals, predators such as the badger dig up
bumblebee nests and eat both the larvae and any stored food.[87]
Specialist ambush predators

of visitors to flowers The bee-eater, Merops apiaster,
include crab spiders, which specialises in feeding on bees; here
wait on flowering plants for a male catches a nuptial gift for his
pollinating insects; mate.
predatory bugs, and
praying mantises,[84] some
of which (the flower mantises of the tropics) wait motionless,
aggressive mimics camouflaged as flowers.[88] Beewolves are large
The beewolf Philanthus triangulum wasps that habitually attack bees;[84] the ethologist Niko Tinbergen
paralysing a bee with its sting 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.[89] Other predatory insects that
sometimes catch bees include robber flies and dragonflies.[84] Honey bees are affected by parasites
including tracheal and Varroa mites.[90] However, some bees are believed to have a mutualistic relationship
with mites.[23]
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.[91]
Relationship with humans
In mythology and folklore
Homer's Hymn to Hermes describes three bee-maidens with the power of divination and thus speaking
truth, and identifies the food of the gods as honey. Sources associated the bee maidens with Apollo and,
until the 1980s, scholars followed Gottfried Hermann (1806) in incorrectly identifying the bee-maidens
with the Thriae.[92] Honey, according to a Greek myth, was
discovered by a nymph called Melissa ("Bee"); and honey was
offered to the Greek gods from Mycenean times. Bees were also
associated with the Delphic oracle and the prophetess was
sometimes called a bee.[93]
The image of a community of honey bees has been used from

ancient to modern times, in Aristotle and Plato; in Virgil and Gold plaques embossed with winged
Seneca; in Erasmus and Shakespeare; Tolstoy, and by political and bee goddesses. Camiros, Rhodes.
social theorists such as Bernard Mandeville and Karl Marx as a 7th century BC.
model for human society.[94] In English folklore, bees would be
told of important events in the household, in a custom known as
"Telling the bees".[95]
In art and literature
Some of the oldest examples of bees in art are rock paintings in Spain
which have been dated to 15,000 BC.[96]
W. B. Yeats's poem The Lake Isle of Innisfree (1888) contains the couplet
"Nine bean rows will I have there, a hive for the honey bee, / And live
alone in the bee loud glade." At the time he was living in Bedford Park in
the West of London.[97] Beatrix Potter's illustrated book The Tale of Mrs
Tittlemouse (1910) features Babbity Bumble and her brood (pictured). Kit
Williams' treasure hunt book The Bee on the Comb (1984) uses bees and
beekeeping as part of its story and puzzle. Sue Monk Kidd's The Secret Life
of Bees (2004), and the 2009 film starring Dakota Fanning, tells the story of
a girl who escapes her abusive home and finds her way to live with a Beatrix Potter's illustration
family of beekeepers, the Boatwrights. of Babbity Bumble in The
Tale of Mrs Tittlemouse,
The 2007 animated comedy film Bee Movie used Jerry Seinfeld's first script 1910
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.[98] 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.[99]
Beekeeping
Humans have kept honey bee colonies, commonly in hives, for

millennia. Beekeepers collect honey, beeswax, propolis, pollen, and
royal jelly from hives; bees are also kept to 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.[100] Simple hives and smoke
were used;[101][102] jars of honey were found in the tombs of A commercial beekeeper at work
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.[103][104] Among Classical
Era authors, beekeeping with the use of smoke is described in
Aristotle's History of Animals Book 9.[105] The account mentions
that bees die after stinging; that workers remove corpses from the
hive, and guard it; castes including workers and non-working
drones, but "kings" rather than queens; predators including toads
and bee-eaters; and the waggle dance, with the "irresistible
suggestion" of άροσειονται ("aroseiontai", it waggles) and
παρακολουθούσιν ("parakolouthousin", they watch).[106][b] Western honey bee on a honeycomb
Beekeeping is described in detail by Virgil in his Georgics; it is also

mentioned in his Aeneid, and in Pliny's Natural History.[106]
As commercial pollinators
Bees play an important role in pollinating flowering plants, and are the major type of pollinator in many
ecosystems that contain flowering plants. It is estimated that one third of the human food supply depends on
pollination by insects, birds and bats, most of which is accomplished by bees, whether wild or
domesticated.[107][108] 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.[109]
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.[110][111] The number of colonies kept by beekeepers declined slightly, through
urbanization, systematic pesticide use, tracheal and Varroa mites, and the closure of beekeeping businesses.
In 2006 and 2007 the rate of attrition increased, and was described as colony collapse disorder.[112] In 2010
invertebrate iridescent virus and the fungus Nosema ceranae were shown to be in every killed colony, and
deadly in combination.[113][114][115][116] Winter losses increased to about 1/3.[117][118] Varroa mites were
thought to be responsible for about half the losses.[119]
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.[120][121] From 2013 the European Union restricted some pesticides to stop bee populations
from declining further.[122] In 2014 the Intergovernmental Panel on Climate Change report warned that
bees faced increased risk of extinction because of global warming.[123] 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.[124]
Farmers have focused on alternative solutions to mitigate these problems. By raising native plants, they
provide food for native bee pollinators like Lasioglossum vierecki[125] and L. leucozonium,[126] leading to
less reliance on honey bee populations.
Squash bees (Apidae) are Bee covered in pollen
important pollinators of
squashes and cucumbers.
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 rock paintings in
Spain dated to 15,000 BC.[96]
Honey bees are used commercially to produce honey.[127] They also produce some substances used as
dietary supplements with possible health benefits, pollen,[128] propolis,[129] and royal jelly,[130] though all
of these can also cause allergic reactions.
As food
Bees are considered edible insects. People in some countries eat insects, including the larvae and pupae of
bees, mostly stingless species. They also gather larvae, pupae and surrounding cells, known as bee brood,
for consumption.[131] In the Indonesian dish botok tawon from Central and East Java, bee larvae are eaten
as a companion to rice, after being mixed with shredded coconut, wrapped in banana leaves, and
steamed.[132][133]
Bee brood (pupae and larvae) although low in calcium, has been found to be high in protein and
carbohydrate, and a useful source of phosphorus, magnesium, potassium, and trace minerals iron, zinc,
copper, and selenium. In addition, while bee brood was high in fat, it contained no fat soluble vitamins
(such as A, D, and E) but it was a good source of most of the water-soluble B vitamins including choline as
well as vitamin C. The fat was composed mostly of saturated and monounsaturated fatty acids with 2.0%
being polyunsaturated fatty acids.[134][135]
Bee larvae as food in the Fried whole bees served in a
Javanese dish botok tawon 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).[136] The claim that apitherapy treats cancer,
which some proponents of apitherapy make, remains unsupported by evidence-based medicine.[137][138]
Stings
The painful stings of bees are mostly associated with the poison gland and the Dufour's gland which are
abdominal exocrine glands containing various chemicals. In Lasioglossum leucozonium, the Dufour's
Gland mostly contains octadecanolide as well as some eicosanolide. There is also evidence of n-triscosane,
n-heptacosane,[139] and 22-docosanolide.[140] However, the secretions of these glands could also be used
for nest construction.[139]
See also
Australian native bees
Fear of bees (apiphobia)
Superorganism
World Bee Day
Explanatory notes
a. Triassic nests in a petrified forest in Arizona, implying that bees evolved much earlier,[9] are
now thought to be beetle borings.[10]
b. 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".[105]
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External links
"Bees" (https://eol.org/pages/677). Encyclopedia of Life.
"Apoidea" (https://www.discoverlife.org/20/q?search=Apoidea) at All Living Things –
images, identification guides, and maps of bees
Bee Genera of the World (https://web.archive.org/web/20190411210512/http://cache.ucr.ed
u/~heraty/beepage.html)
Anthophila (Apoidea) – Bees (https://bugguide.net/node/view/8267) – North American
species of bees at BugGuide
Native Bees of North America (https://bugguide.net/node/view/475348) at BugGuide
"Bee declines driven by combined stress from parasites, pesticides, and lack of flowers" (htt
ps://www.science.org/doi/10.1126/science.1255957) – Science
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Bees Bees Are Winged Insects Closely Related To Wasps and Ants

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Bee

Bees are winged insects closely related to wasps and ants,

Bees are found on every continent except Antarctica, in every

The earliest animal-pollinated flowers were shallow, cup-shaped

⁠Ampulicidae (Cockroach wasps)

⁠Sphecidae (sensu stricto)

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

Apidae (inc. honeybees, cuckoo bees, carpenter bees)

⁠Megachilidae (mason, leafcutter bees) ≈50 Mya

⁠Anthophila (bees) ⁠ ⁠Andrenidae (mining bees) ≈34 Mya

Bees have the following characteristics:

A pair of large compound eyes which cover much of the surface

The largest species of bee is thought to be Wallace's giant bee

Haplodiploid breeding system

In haplodiploid species, females develop from fertilized

Bees may be solitary or may live in various types of communities.

Many bumblebees are

Solitary and communal bees

Most other bees, including familiar insects such as carpenter bees,

While solitary, females each make individual nests.[48] Some

Nest of common Carpenter bee nests in a Honeybees on brood

Navigation, communication, and finding food

As mimics and models

Bees are themselves Müllerian

Brood parasites occur in several bee families including the apid

Predators, parasites and pathogens

Vertebrate predators of bees include bee-eaters, shrikes and

Specialist ambush predators

Relationship with humans

In mythology and folklore

The image of a community of honey bees has been used from

In art and literature

Humans have kept honey bee colonies, commonly in hives, for

Depictions of humans collecting honey from wild bees date to

Beekeeping is described in detail by Virgil in his Georgics; it is also

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