What is the name given to the largest bee in a hive?
TagMe TagMe
Bee.txt Beehive.txt
Bee Beehive
Bees are flying insects closely related to wasps and ants, known for their role in pollination and, in the case of the best-known bee species, the European honey bee, for producing honey and beeswax. Bees are a monophyletic lineage within the superfamily Apoidea, presently considered as a clade Anthophila. There are nearly 20,000 known species of bees in seven to nine recognized families,  though many are undescribed and the actual number is probably higher.  They are found on every continent except Antarctica, in every habitat on the planet that contains insect-pollinated flowering plants.

Some species including honey bees, bumblebees, and stingless bees live socially in colonies. Bees are adapted for feeding on nectar and pollen, the former primarily as an energy source and the latter primarily for protein and other nutrients. Most pollen is used as food for larvae. Bee pollination is important both ecologically and commercially; the decline in wild bees has increased the value of pollination by commercially managed hives of honey bees.

Bees range in size from tiny stingless bee species whose workers are less than 2 mm long, to Megachile pluto, the largest species of leafcutter bee, whose females can attain a length of 39 mm. The most common bees in the Northern Hemisphere are the Halictidae, or sweat bees, but they are small and often mistaken for wasps or flies. Vertebrate predators of bees include birds such as bee-eaters; insect predators include beewolves and dragonflies.

Human beekeeping or apiculture has been practised for millennia, since at least the times of Ancient Egypt and Ancient Greece. Apart from honey and pollination, honey bees produce beeswax, royal jelly and propolis. Bees have appeared in mythology and folklore, again since ancient times, and they feature in works of literature as varied as Virgil's Georgics, Beatrix Potter's The Tale of Mrs Tittlemouse, and W. B. Yeats's poem The Lake Isle of Innisfree. Bee larvae are included in the Javanese dish botok tawon, where they are eaten steamed with shredded coconut.

Evolution 

The ancestors of bees were 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. Until recently, the oldest non-compression bee fossil had been found in New Jersey amber, Cretotrigona prisca of Cretaceous age, a corbiculate bee.  A bee fossil from the early Cretaceous (~100 mya), Melittosphex burmensis, is considered "an extinct lineage of pollen-collecting Apoidea sister to the modern bees". Derived features of its morphology (apomorphies) place it clearly within the bees, but it retains two unmodified ancestral traits (plesiomorphies) of the legs (two mid-tibial spurs, and a slender hind basitarsus), showing its transitional status. By the Eocene (~45 mya) there was already considerable diversity among eusocial bee lineages.

The highly eusocial corbiculate Apidae appeared roughly 87 Mya, and the Allodapini (within the Apidae) around 53 Mya.
The Colletidae appear as fossils only from the late Oligocene (~25 Mya) to early Miocene. 
The Melittidae are known from Palaeomacropis eocenicus in the Early Eocene. 
The Megachilidae are known from trace fossils (characteristic leaf cuttings) from the Middle Eocene. 
The Andrenidae are known from the Eocene-Oligocene boundary, around 34 Mya, of the Florissant shale. 
The Halictidae first appear in the Early Eocene  with species    found in amber. The Stenotritidae are known from fossil brood cells of Pleistocene age. 

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  such as nectar and longer tubes, and bees developed longer tongues to extract the nectar. 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 bees have scopal hairs located on their hind legs or on the underside of their abdomens, some bees in the family Apidae possess pollen baskets on their hind legs while very few species lack these entirely and instead collect pollen in their crops.  This drove the adaptive radiation of the angiosperms, and, in turn, the bees themselves. Bees have not only coevolved with flowers but it is believed that some bees have coevolved with mites. Some bees provide tufts of hairs called acarinaria that appear to provide lodgings for mites; in return, it is believed that the mites eat fungi that attack pollen, so the relationship in this case may be mutualistc.  

Phylogeny

External

This cladogram is based on Debevic et al. 2012, which used molecular phylogeny to demonstrate that the bees arose from deep within the Crabronidae, which is therefore paraphyletic. The Heterogynaidae is also broken up. 

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.

Description

It is usually easy to recognise that a particular insect is a bee. They differ from closely related groups such as wasps by having branched or plume-like setae (bristles), combs on the forelimbs for cleaning their antennae, small anatomical differences in the limb structure and the venation of the hind wings, and in females, by having the seventh dorsal abdominal plate divided into two half-plates.

Behaviourally, one of the most obvious characteristics of bees is that they collect pollen to provide provisions for 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. The world's largest species of bee is thought to be the Indonesian resin bee Megachile pluto, whose females can attain a length of 39 mm.  The smallest species may be dwarf stingless bees in the tribe Meliponini whose workers are less than 2 mm in length. 

A bee has 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 for the bee on light intensity. The antennae usually have thirteen segments in males and twelve 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 and a long proboscis 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.

Sociality

Haplodiploid breeding system

According to inclusive fitness theory, organisms can gain fitness not just through increasing their own reproductive output, but also that of close relatives. In evolutionary terms, individuals should help relatives when Cost 
In haplodiploid species, females develop from fertilized eggs and males from unfertilized eggs. Because a male is haploid (has only one copy of each gene), his daughters (which are diploid, with two copies of each gene) share 100% of his genes and 50% of their mother's. Therefore, they share 75% of their genes with each other. This mechanism of sex determination gives rise to what W. D. Hamilton termed "supersisters", more closely related to their sisters than they would be to their own offspring.  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). This unusual situation has been proposed as an explanation of the multiple independent evolutions of eusociality (arising at least nine separate times) within the Hymenoptera. 
However, some eusocial species such as termites are not haplodiploid. Conversely, many bees are haplodiploid yet are not eusocial, and among eusocial species many queens mate with multiple males, creating half-sisters that share only 25% of their genes. Haplodiploidy is thus neither necessary nor sufficient for eusociality. 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. The most advanced of these are eusocial colonies found among the honey bees, bumblebees, and stingless bees; these are characterised by having cooperative brood care and a division of labour into reproductive and non-reproductive adults. Sociality, of several different types, is believed to have evolved separately many times within the bees. 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), with male drones at certain stages. When the castes are purely behavioural alternatives, the system is considered primitively eusocial as in many paper wasps; when the castes are morphologically discrete, the system is considered highly eusocial.

There are many more species of primitively eusocial than highly eusocial bees, but they have rarely been studied. 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. The orchid bees include some primitively eusocial species with similar biology. Some allodapine bees are 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. 

Bumblebees are eusocial, like the eusocial Vespidae such as hornets. The queen initiates a nest on her own. 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 world-wide using the IUCN Red List criteria. 

Stingless bees are highly eusocial. They practise mass provisioning, with complex nest architecture and perennial colonies. 

The true honey bees (genus Apis) are highly eusocial, and are among the best known of all insects. There are 29 subspecies of 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 unusually defensive. 

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.

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. A very few species of solitary bees 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, polyester bees, squash bees, dwarf carpenter bees, leafcutter bees, alkali bees and digger bees. 

Most solitary bees nest in the ground in a variety of soil textures and conditions while others create nests in hollow reeds or twigs, 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 either stingless or very unlikely to sting (only in self-defense, if ever).  

While solitary females each make individual nests, some species. such as the European mason bee Hoplitis anthocopoides, and the 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 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 there are multiple females using that same entrance on a regular basis.

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. 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. 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. In the case of solitary bees, each one is laid in a 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. In social species of bee there is progressive provisioning with the larva being fed 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.

The larvae are generally whitish grubs, roughly oval and bluntly-pointed at both ends. They have fifteen segments and spiracles in each segment for breathing. They have no legs but are able to move within the confines of the cell, helped by tubercles on their sides. They have short horns on the head, jaws for chewing their 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 their cocoons. The pupa can be seen through the semi-transparent cocoon and over the course of a few days, the insect undergoes metamorphosis into the form of the adult bee. When ready to emerge, it splits its skin dorsally and climbs out of the exuviae as a winged adult and breaks out of the cell.

Flight

In Antoine Magnan's 1934 book Le vol des insectes, he wrote 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".  This has led to a common misconception that bees "violate aerodynamic theory", but 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 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. 

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. Bees navigate using spatial memory with a "rich, map-like organization". 

Ecology

Floral relationships 

Most bees are polylectic (generalist) meaning they collect pollen from a range of flowering plants, however, 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 at least in part because their pollinator is also threatened. There is, however, a pronounced tendency for oligolectic bees to be associated with common, widespread plants which are visited by multiple pollinators. There are some forty oligoleges associated with the creosote bush in the arid parts of the United States southwest, for example. 

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,  all of which gain a measure of protection by superficially looking and behaving like bees.

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 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 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 apid subfamily Nomadinae.  Females of these bees 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.  The Arctic bee species, Bombus hyperboreus, in particular are an aggressive species that attack and enslave other bees of the same subgenus. However, unlike many other bee brood parasites, they have pollen baskets and often collect pollen. 

In the south of 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. 

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,  while the other species in the same genus attacks halictid bees. 

Nocturnal bees 

Four bee families (Andrenidae, Colletidae, Halictidae, and Apidae) contain some species that are crepuscular. Most are tropical or subtropical, but there are some which 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. 

Predators, parasites and pathogens

Vertebrate predators of bees include bee-eaters, shrikes and flycatchers, which make short sallies to catch insects in flight. Swifts and swallows fly almost continually, catching insects as they go. The 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; predatory bugs, and praying mantises,  some of which (the flower mantises of the tropics) wait motionless, aggressive mimics camouflaged as flowers.  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 robber flies and dragonflies.

Honey bees are affected by parasites including acarine and Varroa mites.  However, some bees are believed to have a mutualistic relationship with mites.

Bees and humans

In mythology and folklore

Three bee maidens with the power of divination and thus speaking truth are described in Homer's Hymn to Hermes, and the food of the gods is "identified as honey"; the bee maidens were originally associated with Apollo, and are probably not correctly identified 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 Mycenean times. Bees were associated, too, 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; 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 literature

Beatrix Potter's illustrated book The Tale of Mrs Tittlemouse (1910) features Babbity Bumble and her brood (pictured).

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. 

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 family of beekeepers, the Boatwrights.

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. 

The humorous 2007 animated 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. 

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.  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 the History of Animals Book 9 (a book not written by Aristotle himself). 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 άpοσειονται (aroseiontai, it waggles) and παρακολουθούσιν (parakolouthousin, they watch).

Beekeeping is described in detail by Virgil in his Eclogues; it is also mentioned in his Aeneid, and in Pliny's Natural History.

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, especially the domesticated European honey bee. 

Contract pollination has overtaken the role of honey production for beekeepers in many countries. From 1972 to 2006, feral honey bees declined dramatically in the US, and they are now almost absent.  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.  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, 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. 

However farmers have focused on alternative solutions in order to mitigate these problems. By raising native plants, they provide food for native bee pollinators like L. vierecki  and L. leucozonium,  leading to less reliance on honey bee populations.

As food

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 which have been dated to 15,000 BC.  Indigenous people in many countries eat insects, including consuming the larvae and pupae of bees, mostly stingless bees. They also gather "bee brood" (the larvae, pupae and surrounding cells) for consumption.  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.  

Honey bees are used commercially to produce honey.  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 allergic reactions.

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, and 22-docosanolide.  However, the secretions of these glands could also be used for nest construction. A beehive is an enclosed structure in which some honey bee species of the subgenus Apis live and raise their young. Natural beehives are naturally occurring structures occupied by honeybee colonies, such as hollowed-out trees, while domesticated honeybees live in man-made beehives, often in an apiary. These man-made structures are typically referred to as "beehives". Several species of Apis live in hives, but only the western honey bee (Apis mellifera) and the eastern honey bee (Apis cerana) are domesticated by humans. A natural beehive is comparable to a bird's nest built with a purpose to protect the dweller. 

The beehive's internal structure is a densely packed group of hexagonal cells made of beeswax, called a honeycomb. The bees use the cells to store food (honey and pollen) and to house the "brood" (eggs, larvae, and pupae).

Artificial beehives serve several purposes: production of honey, pollination of nearby crops, housing supply bees for apitherapy treatment, as safe housing for bees in an attempt to mitigate the effects of colony collapse disorder, and to keep bees as pets. Artificial hives are commonly transported so that bees can pollinate crops in other areas.   A number of patents have been issued for beehive designs.

Natural hives 

Honey bees use caves, rock cavities and hollow trees as natural nesting sites. Members of other subgenera have exposed aerial combs. The nest is composed of multiple honeycombs, parallel to each other, with a relatively uniform bee space. It usually has a single entrance. Western honey bees prefer nest cavities approximately 45 litres in volume and avoid those smaller than 10 or larger than 100 litres.[http://www.fao.org/docrep/X0083E/X0083E02.htm Honeybees of the genus Apis.] Food and Agriculture Organization of the United Nations Western honey bees show several  nest-site preferences: the height above ground is usually between 1 m and 5 m, entrance positions tend to face downward, Equatorial-facing entrances are favored, and nest sites over 300 m from the parent colony are preferred.  Bees usually occupy the nests for several years.

The bees often smooth the bark surrounding the hive entrance, and the cavity walls are coated with a thin layer of hardened plant resin (propolis). Honeycombs are attached to the walls along the cavity tops and sides, but small passageways are left along the comb edges.  The basic nest architecture for all honeybees is similar: honey is stored in the upper part of the comb; beneath it are rows of pollen-storage cells, worker-brood cells, and drone-brood cells, in that order. The peanut-shaped queen cells are normally built at the lower edge of the comb.

Ancient artificial hives 

Bees were kept in man-made hives in Egypt in antiquity.  The walls of the Egyptian sun temple of Nyuserre Ini from the 5th Dynasty, dated earlier than 2422 BC, depict workers blowing smoke into hives as they remove honeycombs.  Inscriptions detailing the production of honey are found on the tomb of Pabasa from the 26th Dynasty (c. 650 BC), and describe honey stored in jars, and cylindrical hives. 

Archaeologist Amihai Mazar of the Hebrew University of Jerusalem cites 30 intact hives that were discovered in the ruins of the city of Rehov (2,000 residents in 900 BC, Israelites and Canaanites). This is evidence that an advanced honey industry existed in Israel, approximately 3,000 years ago. The beehives, made of straw and unbaked clay, were found in orderly rows, with a total of 100 hives, many broken. Ezra Marcus from the University of Haifa said the discovery provided a glimpse of ancient beekeeping seen in texts and ancient art from the Near East. An altar decorated with fertility figurines was found alongside the hives and may indicate religious practices associated with beekeeping. While beekeeping predates these ruins, this is the oldest apiary yet discovered.

Traditional artificial hives 

Traditional beehives simply provided an enclosure for the bee colony. Because no internal structures were provided for the bees, the bees created their own honeycomb within the hives.  The comb is often cross-attached and cannot be moved without destroying it. This is sometimes called a "fixed-frame" hive to differentiate it from the modern "movable-frame" hives. Harvest generally destroyed the hives, though there were some adaptations using extra top baskets which could be removed when the bees filled them with honey. These were gradually supplanted with box hives of varying dimensions, with or without frames, and finally replaced by newer modern equipment.

Honey from traditional hives was typically extracted by pressing – crushing the wax honeycomb to squeeze out the honey. Due to this harvesting, traditional beehives typically provided more beeswax, but far less honey, than a modern hive.

There are four basic styles of traditional beehive; mud hives, clay/tile hives, skeps and bee gums.

Mud and clay hives 

Mud hives are still used in Egypt. These are long cylinders made from a mixture of unbaked mud, straw, and dung. 

Clay tiles were the customary homes of domesticated bees in the eastern end of the Mediterranean. Long cylinders of baked clay were used in ancient Egypt, the Middle East and to some extent in Greece, Italy and Malta. They sometimes were used singly, but more often stacked in rows to provide some shade, at least for those not on top. Keepers would smoke one end to drive the bees to the other end while they harvested honey.

Skeps 

Skeps, which are baskets placed open-end-down, have been used for about 2000 years. Initially they were made from wicker plastered with mud and dung but from the Middle Ages they were made of straw. In northern and western Europe, skeps were made of coils of grass or straw. In its simplest form, there is a single entrance at the bottom of the skep. Again, there is no internal structure provided for the bees and the colony must produce its own honeycomb, which is attached to the inside of the skep.

Skeps have two disadvantages; beekeepers cannot inspect the comb for diseases and pests, and honey removal is not easy – often resulting in the destruction of the entire colony. To get the honey beekeepers either drove the bees out of the skep or, by the use of a bottom extension called an eke or a top extension called a cap, sought to create comb with just honey in it. Quite often the bees were just killed, sometimes using lighted sulfur, to allow the honeycomb to be removed. Skeps could also be squeezed in a vise to extract the honey. As of 1998, most US states prohibited the use of skeps because they can not be inspected for disease and parasites. 

Later skep designs included a smaller woven basket (cap) on top over a small hole in the main skep.  This cap acted as a crude super, allowing the harvesting of some honey with less destruction of brood and bees. In England such an extension piece consisting of a ring of about 4 or 5 coils of straw placed below a straw beehive to give extra room for brood rearing was called an eke, imp or nadir. An eke was used to give just a bit of extra room, or to "eke" some more space, a nadir is a larger extension used when a full story was needed beneath.

A person who made such woven beehives was called a "skepper", a surname that still exists in western countries. In England the thickness of the coil of straw was controlled using a ring of leather or piece of cows horn called a "girth" and the coils of straw could be sewn together using strips of briar. Likenesses of skeps can be found in paintings, carvings and old manuscripts. The skep is often used on signs as an indication of industry ("the busy bee").

In the late 18th century, more complex skeps appeared which had wooden tops with holes in them over which glass jars were placed. The comb was built in the glass jars which made it commercially attractive.

 

In the eastern United States, especially in the southeast, sections of hollow trees were used until the 20th century.  These were called "gums" because they often were from black gum (Nyssa sylvatica) trees. 

Sections of the hollow trees were set upright in "bee yards" or apiaries. Sometimes sticks or crossed sticks were placed under a board cover to give an attachment for the honeycomb. As with skeps, harvest of honey from these destroyed the colony. Often the human bee "robber" would kill the bees before even opening their nest. This was done by inserting a metal container of burning sulfur into the gum.

Natural tree hollows and artificially hollowed tree trunks were widely used in the past by beekeepers in Central Europe. For example, in Poland such a beehive was called a "barć" and was protected in various ways from unfavorable weather conditions (rain, frost) and predators (woodpeckers, bears). Harvest of honey from these did not destroy the colony, as only a protective piece of wood was removed from the opening and smoke was used to deter the bees for a short time.

Bee gums are still used by beekeepers today, for bee species whose honey output is less than that of the more productive honeybee. Unlike most manmade beehives (which are usable only with Apis mellifera and Apis cerana), the bee gum allows housing of many more bee species. The bee gum allows the bees themselves to make their hive (hive structure and cells in which to store their honey and pollen).

In some instances, bee gums are also still used, even with bee species that do produce large quantities of honey (i.e. Apis mellifera nigra). However, in these instances, part of the reason why bee gums are then used is that this allows the producers of the honey to distinguish themselves from other honey producers and to ask a higher price for the honey. An example where bee gums are still used is Mont-Lozère, France.     It should be noted however that the length of the bee gums used are rather on the short side; unlike regular bee gums, the bee gums are slightly modified artificially (hollowed out artificially and cut to a specific size).

Modern artificial hives 

The earliest recognizably modern designs of beehives arose in the 19th century, though they were perfected from intermediate stages of progress made in the 18th century.

Thus, intermediate stages in hive design were recorded for example by Thomas Wildman in 1768/1770, who described advances over the destructive old skep-based beekeeping so that the bees no longer had to be killed to harvest the honey.  Wildman, for example, fixed a parallel array of wooden bars across the top of a straw hive or skep (with a separate straw top to be fixed on later) "so that there are in all seven bars of deal" [in a 10 in hive] "to which the bees fix their combs".  He also described using such hives in a multi-story configuration, foreshadowing the modern use of supers:  he described adding (at the proper time) successive straw hives below, and eventually removing the ones above when free of brood and filled with honey, so that the bees could be separately preserved at the harvest for a following season.  Wildman also described  a further development, using hives with "sliding frames" for the bees to build their comb, foreshadowing more modern uses of movable-comb hives.  Wildman acknowledged the advances in knowledge of bees previously made by Swammerdam, Maraldi, and de Reaumur – he included a lengthy translation of Reaumur's account of the natural history of bees – and he also described the initiatives of others in designing hives for the preservation of bee-life when taking the harvest, citing in particular reports from Brittany dating from the 1750s, due to Comte de la Bourdonnaye.

In 1814, Petro Prokopovych, the founder of commercial beekeeping in Ukraine, invented one of the very first beehive frames. However, for easy operations in beehives the spaces between elements need to be correct. The correct distance between combs was described in 1845 by Jan Dzierżon as 1½ inches from the center of one top bar to the center of the next one. In 1848 Dzierzon introduced grooves into the hive's side walls replacing the strips of wood for moving top bars. The grooves were 8 x, the spacing later termed bee space. The Langstroth hive was the first successful top-opened hive with movable frames. Langstroth hive was however a direct descendant of Dzierzon's hive designs.

There are two basic types of modern hive in common use, the movable-frame "Langstroth hive" (including all the size variants) which has enclosed frames to hold the comb and the movable-comb "top-bar hive", which - as the name implies, has only a top-bar to support the comb. Movable frames and movable combs - as opposed to fixed combs - both allow the apiarist to inspect for diseases and parasites and also allow a beekeeper more easily to split the hive to make new colonies.

Bees occupy the new hive "correctly" only if it already contains frames with some honeycomb or wax plates. If only empty frames are present, bees often build honeycomb that does not follow them and cannot be later removed with the frame.

Hives optimized for Apis mellifera, Apis cerana 

Langstroth hives 

Named for their inventor, Rev. Lorenzo Langstroth, Langstroth hives are not the only hives of this style, but the most common in North America. Langstroth patented his design in 1852   originally for comb honey production; it has become the standard style hive for many of the world's beekeepers. This class of hives includes other styles, which differ mainly in the size and number of frames used. These include Smith, Segeberger Beute (German), D.E. hive, Frankenbeute (German), Normalmass (German), Langstroth hive, Modified Commercial, and Modified Dadant (developed in USA in 1920 from the Dadant-Blatt hive, a variant of the Dadant hive), plus regional variations such as the British Standard National Hive.

Langstroth hives make use of bee space so that frames are neither glued together nor filled with burr comb–comb joining adjacent frames.

Langstroth hives use standardized sizes of hive bodies (rectangular boxes without tops or bottoms placed one on top of another) and internal frames to ensure that parts are interchangeable and that the frames will remain relatively easy to remove, inspect, and replace without killing the bees. Langstroth hive bodies are rectangular in shape and can be made from a variety of materials that can be stacked to expand the usable space for the bees.  Inside the boxes, frames are hung parallel to each other.  The minimum size of the hive is dependent on outside air temperature and potential food sources in the winter months. The colder the winter, the larger the hive and food stores need to be. In regions with severe winter weather, a basketball-shaped cluster of bees typically survives in a "double-deep" box. In temperate and equatorial regions, a winter cluster will survive in a single box or in a "nuc" (short for nucleus colony).

Langstroth frames are thin rectangular structures made of wood or plastic and have a wax or plastic foundation on which the bees draw out the comb. The frames hold the beeswax honeycomb formed by the bees. Ten frames side by side will fill the hive body and leave the right amount of bee space between each frame and between the end frames and the hive body.

Langstroth frames are often reinforced with wire, making it possible to spin the honey out of the comb in a centrifuge. As a result, the empty frames and comb can be returned to the beehive for re-filling by the bees. Creating honeycomb involves a significant energy investment, conservatively estimated at 6.25 kilograms of honey needed to create 1 kilogram of comb in temperate climates.  Reusing comb can thus increase the productivity of a beekeeping enterprise. 

The modern Langstroth hive consists of the following parts:
* Hive stand: the upper hive components rest on this, providing a landing board for the bees and helping to protect the bottom board from rot and cold transfer.
* Bottom board: this has an entrance for the bees to get into the hive.
* Brood box: the lowest box of the hive and where the queen bee lays her eggs.
* Honey super: usually shorter than the brood box, but is the uppermost box(s), where honey is stored.
* Frames and foundation: wooden or plastic frames with wax or plastic sheets with honeycomb impression where bees build wax honey combs.
* Inner cover: provides separation from an overly hot or cold outer cover and can be used as a shelf for feeding or other purposes.
* Outer cover: provides weather protection for the hive.

Commercial hives 

Commercial hives have the same cross-sectional dimensions as a National hive (460mm x 460mm), but deeper brood box (267mm/10.5") and supers. The internal structure of the boxes is also simpler, resulting in wider frames (406mm/16") with shorter handles or lugs. The brood box is picked up using small hand holds cut into the external wall of the hive. Supers have this same feature, which some find difficult to use when the super is full of honey. Some beekeepers therefore use National supers on top of a Commercial brood box.

WBC hives 

The WBC, invented by and named after William Broughton Carr in 1890, is a double-walled hive with an external housing that splays out towards the bottom of each frame covering a standard box shape hive inside. The WBC is in many respects the 'classic' hive as represented in pictures and paintings, but despite the extra level of insulation for the bees offered by its double-walled design, many beekeepers avoid it, owing to the inconvenience of having to remove the external layer before the hive can be examined.

CDB hives 

An Irish hive. In 1894, the British Charles Nash Abbott (1830–1894), advisor to Ireland’s Department of Agriculture and Technical Instruction, oversaw the design of a new Congested Districts Board (CDB) hive, in Dublin.

Dartington Long Deep hives 

The Dartington Long Deep (DLD) hive takes 14 x 12 inch and can take up to 24 frames.  It is possible to have two colonies in the brood box as there is an entrance at either end.  It has half-size honey supers, which take 6 frames that are lighter than full supers and are correspondingly easier to lift.   The Dartington was originally developed by Robin Dartington so that he could keep bees on his London rooftop.

Beehaus 

The Beehaus is a proprietary design for a beehive launched in 2009.  It is based on similar principles to a Dartington. Kobe T Monzon was the first to consider this in the lower 48 states.

Long Box Hive 

The Long Box Hive is a single story hive utilizing fully enclosed frames (per the dimensions of Langstroth hives or deeper by variation,) but is worked horizontally in the manner of Kenya/Tanzanian Top-bar hives. This non-stacked style had higher popularity a century ago in the Southeast United States, but faded from use due to lack of portability. With the recent popularity of horizontal Top-bar hives, the Long Box Hive is gaining renewed but limited utilization. Alternative names "New Idea Hive", "Single Story Hive", "Poppleton Hive", or simply "Long Hive". 

Top-bar hives 

The top-bar or Kenya-hives were developed as a lower-cost alternative to the standard Langstroth hives and equipment. They are becoming very popular in the US due to their alignment with the organic, treatment-free philosophies of many new beekeeping devotees in the United States. They are also popular, owing to their simplicity and low cost, in developing countries. Top-bar hives have movable comb and make use of the concept of bee space.

The top-bar hive is so named because the bees draw their comb from  a top bar, suspended across the top of a cavity, and not inside a full rectangular frame with sides and a bottom bar.  The beekeeper does not provide foundation wax (or provides only a small starter piece of foundation) for the bees to build from.  The bees build the comb so it hangs down from the top bar.  This is in keeping with the way bees build wax in a natural cavity.  There is some belief that the use of natural wax in a top-bar hive supports the bees' natural systems in ways that improve their health.

The hive body of a common style of top-bar hive is often shaped as an inverted trapezoid* in order to reduce the tendency of bees to attach the comb to the hive-body walls, though this reasoning has become less popular recently. It may be more likely that the trapezoid shape helps to improve the ratio of the weight of the comb to the amount of attachment at the bar and helps to lessen the likelihood of heavy combs detaching from the top bar when being handled or harvested.  Unlike the Langstroth design, *this style of top-bar hive is expanded horizontally, not vertically. The top-bar design is a single, much longer box, with the bars hanging in parallel. This common style is sometimes referred to as a horizontal Top Bar hive, or hTBH. Most horizontal top bar hives in the UK are based on the tandem-follower system designed by Phil Chandler, and free plans are available at http://www.biobees.com

Some top bar hives do stack vertically, much like the Langstroth, National, or other frame hives, although vertical designs use bars with gaps between them, unlike the adjacent bars of the horizontal design. Vertical top bar hives have been largely unknown in the United States until about 2010. This is largely due to the work by Dr David Heaf, who, along with wife Patricia, translated a book by Abbé Emile Warré (~1852) from French and implemented some hives in 2007. These are commonly referred to as Warré Hives, but the abbé named them, "The People's Hive," for their intended economy and simplicity of construction. Since then, the book has been translated into about 6 other languages as of 2014, and is a free ebook download on some sites.

Unlike the Langstroth hive, the honey is usually extracted by crushing and straining rather than centrifuging because a top bar does not have wire reinforcement. There is a laterally oriented extractor available, made by Swienty, but these are not overly popular in the US. Because the bees have to rebuild their comb after honey is harvested, a top-bar hive yields a beeswax harvest in addition to honey.

However, like the Langstroth hive, the bees will store most of their honey separately from the areas where they are raising the brood. For this reason, bees are not killed when harvesting from a top-bar hive, unlike the old practice of harvesting from a skep by driving bees out or killing them over a sulphur pit.

Warré hives 

The Warré hive was invented by the abbot Émile Warré, and is also called "ruche populaire" (fr) or "The People's Hive" (en), the Warré hive is a modular and storied design similar to a Langstroth hive. The hive body is made of boxes stacked vertically; however, it uses Top Bars for comb support instead of full frames, as a general rule. The popularity of this hive is growing among 'sustainable-practice' beekeepers.

The Warre hive differs from other stacked hive systems in one fundamental aspect: when the bees need more space as the colony expands, the new box is "nadired". i.e. positioned underneath the existing box(es). This serves the purpose of warmth retention within the brood nest of the hive, considered vital to colony health. The 1948 book "Beekeeping for All" by Abbé Warré has been translated from French in 2010 by Dr. David and Patricia Heaf and is available as a free download at; http://www.users.callnetuk.com/~heaf/beekeeping_for_all.pdf More information on the Abbé and the hive is available at http://warre.biobees.com/bfa.htm

Perone hives 

The Perone or Automatic Hive was designed by Oscar Perone, and aims at managing bees in a way more closely analogous to their natural behavior. It is favoured by proponents of 'Natural Beekeeping', who seek to allow more natural bee behaviour and minimise intervention into the hive.

Perone Hives are very large 2m-high vertical top bar hives that remain the same size all year, split into a bee area underneath, and a bee keepers area above (Mark 1) or side by side (Mark 2). The total hive volume is large, around 280 litres, which it is proposed allows the bees to develop into a 'super-colony' differing in behaviour to colonies in smaller hives. They are managed so that under normal circumstances the bee area is never opened, and the bee keepers area is opened only once per year when any honey is harvested. These hives are managed around bees using their own honey supplies in winter, not replacing these with sugar syrup.

The limitations for direct inspection of this type of hive and its management raise issues around disease monitoring, and they are criticised as sources of varroa for other nearby hives, as the large colony will have many foragers and many standard techniques for managing varroa populations cannot be practiced. However the hive will also have more bees available for defence, and many standard treatments for varroa also cause damage to the bees.

Flow Hives 

The Flow Hive  is a prototype easy-harvest hive describing a design for supers filled with a split plastic deep comb. The Flow Hive became the subject of a crowdfunding campaign, collecting just over US $12 million as of 2015 Apr 20.   A patent was applied for in 2011. 

The bees use their wax to seal the splits, build out the comb and fill the cells with honey. According to the 2011 patent, once the comb is filled with honey and capped, turning a crank splits the hexagonal cells in the artificial comb vertically, allowing the honey to flow down the resulting zig-zag channels into a sealed trough and out of the hive. According to the inventors, this allows harvesting the honey with minimal disruption to the bees who are standing on the still capped comb. When drained, the crank is used to reset the comb. The bees then chew open the capping, reseal the splits and refill the cells.

Hives optimized for meliponines 

Hives have been designed for some meliponines such as  Melipona beecheii. Examples of such hives are the Nogueira-Neto hive and the UTOB hive. 

Hives optimized for bumblebees 

Some designs have been optimized for housing bumblebees. See [http://www.bumblebee.org/nestboxes.htm here].

Symbolism 

The beehive is a commonly used symbol dating at least to Roman times.  In medieval heraldry it was considered a symbol of industry.

In modern times, it is used in Freemasonry. In masonic lectures it is explained as a symbol of industry and co-operation,  and as cautioning against intellectual laziness, warning that "he that will so demean himself as not to be endeavoring to add to the common stock of knowledge and understanding, may be deemed a drone in the hive of nature, a useless member of society, and unworthy of our protection as Masons." 

The beehive is also used with similar meaning by The Church of Jesus Christ of Latter-day Saints, or Mormons. From Mormon usage it has become one of the State symbols of Utah.  (See deseret.)

In Wellington, New Zealand, the round building used for Parliamentary offices is known as the "Beehive". The official website of the New Zealand Government is [http://www.beehive.govt.nz/ www.beehive.govt.nz].

Beehive Brand matches made by Bryant and May popular in New Zealand have a logo based on the traditional skep beehive design.

Relocation 

A common misconception of non-beekeepers is that smoke will cause bees to move their hive. Smoke is used by beekeepers to hide the alarm pheromone emitted by bees during inspections or harvest.  One-way traps can be used to reduce the size of a colony, but is a limited and time-consuming solution.  Beekeepers and companies can be hired to do "cut-outs" (remove bees from a home).  Once honey, pollen, and wax are built up in a hive, only physical removal is an option.  Some homeowners resort to using spray-foam, but the bees will wait and live off reserves until the foam is weakened, then will burrow through it.  An important fact in understanding a hive is that fewer than 10% of the bees will ever leave at once, other than when swarming.  A fertilized queen will lay up to 2000 eggs a day. If bees are killed and the hive products are left, heat and time will loosen the wax and honey causing further problems in the future.

Destruction 

By other natural organisms 

Black bears destroy hives in their quest for honey. 

Elephants sometimes also destroy hives erected by humans as a defense for their crops against elephants. These hives are hung on a single metal wire that encircles the crop field of some farms in African elephant territory. The installation is called a BeeHive Fence and was conceived by Lucy King.  

By humans 

Humans often destroy hives of honey-producing bee species in the interest of obtaining honey and other bee products.

Humans may at times also determine that a beehive must be destroyed in the interest of public safety or in the interest of preventing the spread of bee diseases. The U.S. state of Florida destroyed the hives of Africanized honey bees in 1999.  The state of Alaska has issued regulations governing the treatment of diseased beehives via burning followed by burial, fumigation using ethylene oxide or other approved gases, sterilization by treatment with lye, or by scorching.  In New Zealand, the treatment of hives infected with the disease American foulbrood with antibiotics is prohibited, and beekeepers are required by law to destroy such colonies and hives with fire.

Method of destruction 

Spraying the hive with a solution of soap and water may be effective, since soap dissolves the bees' waxy exterior that protects them from drowning. However, the procedure should be undertaken with caution, as it angers the bees.
Quene Queene Queen Queen (disambiguation) Queen (automobile) Queen (album) Queen (song)
quene queen disambiguation queen album queene queen song queen queen automobile
queen
WikipediaEntity
Queen
 What is the name given to the largest biome? 

 The largest barrier is Dr Dr Dr to is a fascinating topic to a sea relevant context or or mind to chat E is reliable of the context of the