Beekeeping

At some point in history at least 10,000 years ago, humans began to attempt to maintain colonies of wild bees in artificial hives made from hollow logs, wooden boxes, pottery vessels, or woven straw baskets (known as skeps). Depictions of humans collecting honey from wild bees date to 10,000 years ago.[4] Beekeeping in pottery vessels began about 9,000 years ago in North Africa.[5] Traces of beeswax are found in potsherds throughout the Middle East beginning about 7000 BCE.[5] Domestication of bees is shown in Egyptian art from around 4,500 years ago.[6] Simple hives and smoke were used and honey was stored in jars, some of which were found in the tombs of pharaohs such as Tutankhamun. It was not until the 18th century that European understanding of the colonies and biology of bees allowed the construction of the movable comb hive so that honey could be harvested without destroying the entire colony.

Honeybees were kept in Egypt from antiquity.[7] On the walls of the sun temple of Nyuserre Ini from the Fifth Dynasty, before 2422 BCE, workers are depicted blowing smoke into hives as they are removing honeycombs.[8] Inscriptions detailing the production of honey are found on the tomb of Pabasa from the Twenty-sixth Dynasty (c. 650 BCE), depicting pouring honey in jars and cylindrical hives.[9]

An inscription records the introduction of honey bees into the land of Suhum in Mesopotamia, where they were previously unknown:

I am Shamash-resh-ușur, the governor of Suhu and the land of Mari. Bees that collect honey, which none of my ancestors had ever seen or brought into the land of Suhu, I brought down from the mountain of the men of Habha, and made them settle in the orchards of the town 'Gabbari-built-it'. They collect honey and wax, and I know how to melt the honey and wax – and the gardeners know too. Whoever comes in the future, may he ask the old men of the town, (who will say) thus: "They are the buildings of Shamash-resh-ușur, the governor of Suhu, who introduced honey bees into the land of Suhu."

— translated text from stele, (Dalley, 2002)[10]

The oldest archaeological finds directly relating to beekeeping have been discovered at Rehov, a Bronze and Iron Age archaeological site in the Jordan Valley, Israel.[11] Thirty intact hives, made of straw and unbaked clay, were discovered by archaeologist Amihai Mazar in the ruins of the city, dating from about 900 BCE. The hives were found in orderly rows, three high, in a manner that could have accommodated around 100 hives, held more than 1 million bees and had a potential annual yield of 500 kilograms of honey and 70 kilograms of beeswax, according to Mazar, and are evidence that an advanced honey industry existed in ancient Israel 3,000 years ago.[12][13][14]

The Beekeepers, 1568, by Pieter Bruegel the Elder

In ancient Greece (Crete and Mycenae), there existed a system of high-status apiculture, as can be concluded from the finds of hives, smoking pots, honey extractors and other beekeeping paraphernalia in Knossos. Beekeeping was considered a highly valued industry controlled by beekeeping overseers—owners of gold rings depicting apiculture scenes rather than religious ones as they have been reinterpreted recently, contra Sir Arthur Evans.[15] Aspects of the lives of bees and beekeeping are discussed at length by Aristotle. Beekeeping was also documented by the Roman writers Virgil, Gaius Julius Hyginus, Varro, and Columella.[16]

Beekeeping has also been practiced in ancient China since antiquity. In a book written by Fan Li (or Tao Zhu Gong) during the Spring and Autumn period there are sections describing the art of beekeeping, stressing the importance of the quality of the wooden box used and how this can affect the quality of the honey.[17] The Chinese word for honey (蜜 mì, reconstructed Old Chinese pronunciation *mjit) was borrowed from proto-Tocharian *ḿət(ə) (where *ḿ is palatalized; cf. Tocharian B mit), cognate with English mead.[18]

The ancient Maya domesticated a separate species of stingless bee, which they used for several purposes, including making balché, a mead-like alcoholic drink.[19] By 300 BCE they had achieved the highest levels of stingless beekeeping practices in the world.[20] The use of stingless bees is referred to as meliponiculture, named after bees of the tribe Meliponini—such as Melipona quadrifasciata in Brazil. This variation of bee keeping still occurs around the world today.[21] For instance, in Australia, the stingless bee Tetragonula carbonaria is kept for production of their honey.[22]

European natural philosophers began to study bee colonies scientifically in the 18th century. Preeminent among these scientific pioneers were Swammerdam, René Antoine Ferchault de Réaumur, Charles Bonnet, and François Huber. Swammerdam and Réaumur were among the first to use a microscope and dissection to understand the internal biology of honey bees. Réaumur was among the first to construct a glass walled observation hive to better observe activities within hives. He observed queens laying eggs in open cells, but still had no idea of how a queen was fertilized; nobody had ever witnessed the mating of a queen and drone and many theories held that queens were "self-fertile," while others believed that a vapor or "miasma" emanating from the drones fertilized queens without direct physical contact. Huber was the first to prove by observation and experiment that queens are physically inseminated by drones outside the confines of hives, usually a great distance away.[23]

Following Réaumur's design, Huber built improved glass-walled observation hives and sectional hives that could be opened like the leaves of a book. This allowed inspecting individual wax combs and greatly improved direct observation of hive activity. Although he went blind before he was twenty, Huber employed a secretary, François Burnens, to make daily observations, conduct careful experiments, and keep accurate notes over more than twenty years. Huber confirmed that a hive consists of one queen who is the mother of all the female workers and male drones in the colony. He was also the first to confirm that mating with drones takes place outside of hives and that queens are inseminated by a number of successive matings with male drones, high in the air at a great distance from their hive. Together, he and Burnens dissected bees under the microscope and were among the first to describe the ovaries and spermatheca, or sperm store, of queens as well as the penis of male drones. Huber is originally regarded as "the father of modern bee-science" and his "Nouvelles Observations sur Les Abeilles (or "New Observations on Bees")[24] revealed all the basic scientific truths for the biology and ecology of honeybees.[23]

Honey harvesting in its earliest times frequently resulted in the destruction of the whole colony as a result of the honey being taken. The wild hive was broken into, using smoke to quiet the bees. The honeycombs were pulled out and either immediately eaten whole or crushed up, along with the eggs, larvae, and honey they held. A sieve or basket was used to separate the liquid honey from the demolished brood nest. In mediaeval times in northern Europe, although skeps and other artificial containers were made to house bees, the precious honey and wax were still extracted only after killing the colony of bees.[25] It was impossible to replace old, dark-brown brood comb, in which larval bees are constricted by layers of shed pupal skins.[26]

The movable frames of modern hives are considered to be the descendants of the traditional basket top bar (movable comb) hives of Greece, which allowed the beekeeper to avoid killing the bees.[27] The oldest testimony on their use dates back to 1669 although it is probable that their use is more than 3000 years old.[28]

A beekeeper inspecting a hive frame from a Langstroth hive.

Intermediate stages in the transition from the old beekeeping to the new were recorded for example by Thomas Wildman in 1768, who described advances over the destructive old skep-based beekeeping so that the bees no longer had to be killed to harvest the honey.[29] Wildman for example fixed a parallel array of wooden bars across the top of a straw hive or skep about ten inches (about 25 cm) in diameter "so that there are in all seven bars of deal to which the bees fix their combs", foreshadowing more modern uses of movable-comb hives. He also described using such hives in a multi-storey configuration, foreshadowing the modern use of supers: he added (at a 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.[30]

Wildman's book acknowledged the advances in knowledge of bees previously made by Swammerdam, Maraldi, and de Réaumur—he included a lengthy translation of Réaumur'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. Another example of a hive design, was invented by Rev. John Thorley in 1744. The hive was placed in a bell jar that was screwed onto a wicker basket. The bees were free to move from the basket to the jar and the honey was produced and stored in the jar. The hive was designed to keep the bees from swarming as much as they would have in other hive designs.[31]

The 19th century saw this revolution in beekeeping practice completed through the perfection of the movable comb hive by the American Lorenzo Lorraine Langstroth. Langstroth was the first person to make practical use of Huber's earlier discovery that there was a specific spatial measurement between the wax combs, later called the bee space, which bees do not block with wax, but keep as a free passage. Having determined this bee space (commonly given as between 6 and 9 millimetres or 1⁄4 and 3⁄8 inch),[32][33] though up to 15mm has been found in populations in Ethiopia.[34] Langstroth then designed a series of wooden frames within a rectangular hive box, carefully maintaining the correct space between successive frames, and found that the bees would build parallel honeycombs in the box without bonding them to each other or to the hive walls. This enables the beekeeper to slide any frame out of the hive for inspection, without harming the bees or the comb, protecting the eggs, larvae and pupae contained within the cells. It also meant that combs containing honey could be gently removed and the honey extracted without destroying the comb. The emptied honey combs could then be returned to the bees intact for refilling. Langstroth's book, The Hive and Honey-bee, published in 1853, described his rediscovery of the bee space and the development of his patent movable comb hive.

The invention and development of the movable-comb-hive fostered the growth of commercial honey production on a large scale in both Europe and the US (see also Beekeeping in the United States).

Langstroth's design for movable comb hives was seized upon by apiarists and inventors on both sides of the Atlantic and a wide range of moveable comb hives were designed and perfected in England, France, Germany and the United States.[35] Classic designs evolved in each country: Dadant hives and Langstroth hives are still dominant in the US; in France the De-Layens trough-hive became popular and in the UK a British National hive became standard as late as the 1930s although in Scotland the smaller Smith hive is still popular. In some Scandinavian countries and in Russia the traditional trough hive persisted until late in the 20th century and is still kept in some areas. However, the Langstroth and Dadant designs remain ubiquitous in the US and also in many parts of Europe, though Sweden, Denmark, Germany, France and Italy all have their own national hive designs. Regional variations of hive evolved to reflect the climate, floral productivity and the reproductive characteristics of the various subspecies of native honey bee in each bio-region.[35]

Honey-laden honeycomb in a wooden frame

The differences in hive dimensions are insignificant in comparison to the common factors in all these hives: they are all square or rectangular; they all use movable wooden frames; they all consist of a floor, brood-box, honey super, crown-board and roof. Hives have traditionally been constructed of cedar, pine, or cypress wood, but in recent years hives made from injection molded dense polystyrene have become increasingly important.[36]

Hives also use queen excluders between the brood-box and honey supers to keep the queen from laying eggs in cells next to those containing honey intended for consumption. Also, with the advent in the 20th century of mite pests, hive floors are often replaced for part of (or the whole) year with a wire mesh and removable tray.[36]

Western honey bee on a honeycomb

In 2015 the Flow Hive system was invented in Australia by Cedar Anderson and his father Stuart Anderson,[37] allowing honey to be extracted without cumbersome centrifuge equipment.

Beekeeping has seen improvements in the design and production of beehives, systems of management and husbandry, stock improvement by selective breeding, honey extraction and marketing, in the 19th century. Notable innovators of modern beekeeping include:

Petro Prokopovych used frames with channels in the side of the woodwork; these were packed side by side in boxes that were stacked one on top of the other. The bees traveled from frame to frame and box to box via the channels.[38] The channels were similar to the cutouts in the sides of modern wooden sections.[39]

Jan Dzierżon was the father of modern apiology and apiculture. All modern beehives are descendants of his design.[40]

François Huber made significant discoveries regarding the bee life-cycle and communication between bees. Despite being blind, Huber brought to light a large amount of information regarding the queen bee's mating habits and her contact with the rest of the hive. His work was published as New Observations on the Natural History of Bees.[41]

L. L. Langstroth revered as the "father of American apiculture"; no other individual has influenced modern beekeeping practice more than Lorenzo Lorraine Langstroth. His classic book The Hive and Honey-bee was published in 1853.[42]

Moses Quinby often termed "the father of commercial beekeeping in the United States," author of Mysteries of Bee-Keeping Explained. He invented the Bee smoker in 1873.[43][44]

Amos Root author of the A B C of Bee Culture, which has been continuously revised and remains in print. Root pioneered the manufacture of hives and the distribution of bee-packages in the United States.[45]

A. J. Cook author of The Bee-Keepers' Guide; or Manual of the Apiary, 1876.[46]

Dr. C.C. Miller was one of the first entrepreneurs actually to make a living from apiculture. By 1878 he made beekeeping his sole business activity. His book, Fifty Years Among the Bees, remains a classic, and his influence on bee management persists to this day.[47]

Honey Extractor

Franz Hruschka was an Austrian/Italian military officer who made one important invention that catalyzed the commercial honey industry. In 1865 he invented the simple machine for extracting honey from the comb by means of centrifugal force. His original idea was to support combs in a metal framework and then spin them around within a container to collect honey as it was thrown out by centrifugal force. This meant that honeycombs could be returned to a hive undamaged but empty, saving the bees a vast amount of work, time, and materials. This single invention significantly improved the efficiency of honey harvesting and catalyzed the modern honey industry.[48]

Walter T. Kelley was an American pioneer of modern beekeeping in the early and mid-20th century. He greatly improved upon beekeeping equipment and clothing and went on to manufacture these items as well as other equipment. His company sold via catalog worldwide, and his book, How to Keep Bees & Sell Honey, an introductory book of apiculture and marketing, allowed for a boom in beekeeping following World War II.[49]

In the U.K., practical beekeeping was led in the early 20th century by a few men, pre-eminently Brother Adam and his Buckfast bee and R.O.B. Manley, author of many titles, including Honey Production in the British Isles and inventor of the Manley frame, still universally popular in the U.K. Other notable British pioneers include William Herrod-Hempsall and Gale.[50][51]

Ahmed Zaky Abushady (1892–1955) was an Egyptian poet, medical doctor, bacteriologist, and bee scientist who was active in England and Egypt in the early part of the twentieth century. In 1919, Abushady patented a removable, standardized aluminum honeycomb. In 1919 he also founded The Apis Club in Benson, Oxfordshire, which was transitioned to the International Bee Research Association (IBRA). In Egypt in the 1930s, Abushady established The Bee Kingdom League and its organ, The Bee Kingdom.[52]

There are three types of vertical stackable hives: hanging or top-access frame, sliding or side-access frame, and top bar.

Hanging frame hives include Langstroth, the British National, Dadant, Layens, and Rose, differing primarily by size or number of frames. The Langstroth was the first successful top-opened hive with movable frames. Many other hive designs are based on the principle of bee space first described by Langstroth, and is a descendant of Jan Dzierzon's Polish hive designs. Langstroth hives are the most common size in the United States and much of the world; the British National is the most common size in the United Kingdom; Dadant and Modified Dadant hives are widely used in France and Italy, and Layens by some beekeepers, where their large size is an advantage. Square Dadant hives–often called 12 frame Dadant or Brother Adam hives–are used in large parts of Germany and other parts of Europe by commercial beekeepers.

Any hanging frame hive design can be built as a sliding frame design. The AZ Hive, the original sliding frame design, integrates hives using Langstroth-sized frames into a honey house so as to streamline the workflow of honey harvest by localization of labor, similar to cellular manufacturing. The honey house can be a portable trailer, allowing the beekeeper to haul the hives to a site and provide pollination services.

Top bar stackable hives simply use top bars instead of full frames. The most common type is the Warre hive, although any hive with hanging frames can be made into a top bar stackable hive by using only the top bar and not the whole frame. This may work less-well with larger frames, where crosscomb and attachment can occur more-readily.

Beekeepers often wear protective clothing to protect themselves from stings

Most beekeepers also wear some protective clothing. Novice beekeepers usually wear gloves and a hooded suit or hat and veil. Experienced beekeepers sometimes elect not to use gloves because they inhibit delicate manipulations. The face and neck are the most important areas to protect, so most beekeepers wear at least a veil.[57] Defensive bees are attracted to the breath, and a sting on the face can lead to much more pain and swelling than a sting elsewhere, while a sting on a bare hand can usually be quickly removed by fingernail scrape to reduce the amount of venom injected.

Traditionally beekeeping clothing was pale colored and this is still very common today. This is because of the natural color of cotton and cost of coloring was an expense not warranted for workwear, though some consider this is to provide better differentiation from the colony's natural predators (such as bears and skunks) which tend to be dark-colored. It is now known that bees see in ultraviolet and are also attracted to scent. So the type of fabric conditioner used has more impact than the color of the fabric.[58][59]

'Stings' retained in clothing fabric continue to pump out an alarm pheromone that attracts aggressive action and further stinging attacks. Washing suits regularly, and rinsing gloved hands in vinegar minimizes attraction.

Bee smoker with heat shield and hook

Most beekeepers use a smoker, which is a device designed to generate smoke from the incomplete combustion of various fuels. Although the exact mechanism is disputed, it is said that smoke calms bees. Some claim it initiates a feeding response in anticipation of possible hive abandonment due to fire.[60] It is also thought that smoke masks alarm pheromones released by guard bees or when bees are squashed in an inspection. The ensuing confusion creates an opportunity for the beekeeper to open the hive and work without triggering a defensive reaction.

Many types of fuel can be used in a smoker as long as it is natural and not contaminated with harmful substances. These fuels include hessian (or burlap), twine, pine needles, corrugated cardboard, and mostly rotten or punky wood. Indian beekeepers, especially in Kerala, often use coconut fibers as they are readily available, safe, and of negligible expense. Some beekeeping supply sources also sell commercial fuels like pulped paper and compressed cotton, or even aerosol cans of smoke. Other beekeepers use sumac as fuel because it ejects much smoke and lacks an odor.

Some beekeepers are using "liquid smoke" as a safer, more convenient alternative. It is a water-based solution that is sprayed onto the bees from a plastic spray bottle. A spray of clean water can also be used to encourage bees to move on.[61]

Torpor may also be induced by the introduction of chilled air into the hive – while chilled carbon dioxide may have harmful long-term effects.[62]

American hive tool

Most beekeepers use a Hive tool when working on their hives. There are two main types; the American hive tool; and the Australian hive tool often called a 'frame lifter'.

They are used to scrape off burr-comb from around the hive, especially on top of the frames. They are also used to separate the frames before lifting out of the hive.

Some beekeepers believe that the more stings a beekeeper receives, the less irritation each causes, and they consider it important for safety of the beekeeper to be stung a few times a season. Beekeepers have high levels of antibodies (mainly IgG) reacting to the major antigen of bee venom, phospholipase A2 (PLA).[63] Antibodies correlate with the frequency of bee stings.

The entry of venom into the body from bee-stings may also be hindered and reduced by protective clothing that allows the wearer to remove stings and venom sacs with a simple tug on the clothing. Although the stinger is barbed, a worker bee's stinger is less likely to become lodged into clothing than human skin.

Symptoms of a being stung include redness, swelling, and itching around the site of the sting. In mild cases, it will take about 2 hours for the pain and swelling to subside. In moderate cases, the red welt at the sting site will become slightly larger for 1–2 days before beginning to heal. A severe reaction, which is rare among beekeepers, results in anaphylactic shock.[64]

If a beekeeper is stung by a bee, there are many protective measures that should be taken in order to make sure the affected area does not become too irritated. The first cautionary step that should be taken following a bee sting is removing the stinger without squeezing the attached venom glands. A quick scrape with a fingernail is effective and intuitive. This step is effective in making sure that the venom injected does not spread, so the side effects of the sting will go away sooner. Washing the affected area with soap and water is also a good way to stop the spread of venom. The last step that needs to be taken is to apply ice or a cold compress to the stung area.[64]

Tunnel entrance with baffle

The bees maintain the internal temperature of their hive at about 35 °C (95 °F).[65] Their ability to do this is known as social homeostasis and was first described by Gates.[66]

During hot weather, the bees cool the hive by circulating cool air from the entrance up through the hive and out again;[67] and if necessary, by placing water, which they fetch, throughout the hive to create evaporative cooling.[68]

Hive with a second skin of polystyrene

In cold weather the packing/insulation of the bee hive is essential.[69] The extra insulation reduces the amount of honey the bees consume and makes it easier for them to maintain the hive's ideal temperature. This need for insulation has encouraged the use of double walled hives with an outer wall of timber, or polystyrene as in the photograph; and even hives constructed from a ceramic.[70]

There has been considerable debate about the best location for hives. Virgil thought they should be located near clear springs, ponds or shallow brooks. Wildman thought they should face to the south or west. One thing all writers agreed on is that hives should be sheltered from strong winds. In hot climates, they were often placed under the shade of trees in summer.[71]

Researchers found that domestic honey bees placed in national parks in the USA competed with native bee species for resources. A further review of the literature concluded that large concentrations of beehives, in continents where they were not native, such as North and South America, could compete against the native bees, however this was not as strongly observed in areas where domestic bees are native such as Europe and Africa, where the different bee species have adapted over millennia to have a narrower overlapping of forage preferences.[72]

The natural beekeeping movement believes that bee hives are weakened by modern beekeeping and agricultural practices, such as crop spraying, hive movement, frequent hive inspections, artificial insemination of queens, routine medication, and sugar water feeding.[73]

Practitioners of "natural beekeeping" tend to use variations of the top-bar hive, which is a simple design that retains the concept of having a movable comb without the use of frames or a foundation. The horizontal top-bar hive, as championed by Marty Hardison, Michael Bush, Philip Chandler, Dennis Murrell and others, can be seen as a modernization of hollow log hives, with the addition of wooden bars of specific width from which bees hang their combs. Its widespread adoption in recent years can be attributed to the publication in 2007 of The Barefoot Beekeeper[74] by Philip Chandler, which challenged many aspects of modern beekeeping and offered the horizontal top-bar hive as a viable alternative to the ubiquitous Langstroth-style movable-frame hive.

A vertical top-bar hive is the Warré hive, based on a design by the French priest Abbé Émile Warré (1867–1951) and popularized by Dr. David Heaf in his English translation of Warré's book L'Apiculture pour Tous as Beekeeping For All.[75]

Honey bee in Toronto

Related to natural beekeeping, urban beekeeping is an attempt to revert to a less industrialized way of obtaining honey by utilizing small-scale colonies that pollinate urban gardens.

Some have found that "city bees" are actually healthier than "rural bees" because there are fewer pesticides and greater biodiversity in the urban gardens.[76] Urban bees may fail to find forage, however, and homeowners can use their landscapes to help feed local bee populations by planting flowers that provide nectar and pollen. An environment of year-round, uninterrupted bloom creates an ideal environment for colony reproduction.[77]

Modern beekeepers have experimented with raising bees indoors, in a controlled environment, or indoor observation hives. This may be done for reasons of space and monitoring or in the off-season. In the off-season, large commercial beekeepers may move colonies to "wintering" warehouses with fixed temperature, light, and humidity. This helps the bees remain healthy but relatively dormant. These relatively dormant or "wintered" bees survive on stored honey, and new bees are not born.[78]

Experiments in raising bees for longer durations indoors have looked into more precise and varying environment controls. In 2015, MIT's Synthetic Apiary project simulated springtime inside a closed environment for several hives throughout a winter. They provided food sources and simulated long days and saw activity and reproduction levels comparable to the levels seen outdoors in warm weather. They concluded that such an indoor apiary could be sustained year-round if needed.[79][80]

A swarm about to land

New wax combs between basement joists

All colonies are totally dependent on their queen, who is the only egg-layer. Although queens have a 3–4 year adult lifespan, diminished longevity of queens (less than 1 year) is commonly and increasingly observed.[81] She can choose whether or not to fertilize an egg as she lays it; if she does so, it develops into a female worker bee; if she lays an unfertilized egg it becomes a male drone. She decides which type of egg to lay depending on the size of the open brood cell she encounters on the comb. In a small worker cell, she lays a fertilized egg; if she finds a larger drone cell, she lays an unfertilized drone egg.[82]

All the time that the queen is fertile and laying eggs she produces a variety of pheromones, which control the behavior of the bees in the hive. These are commonly called queen substance, but there are various pheromones with different functions. As the queen ages, she begins to run out of stored sperm, and her pheromones begin to fail.[83]

Inevitably, the queen begins to falter, and the bees decide to replace her by creating a new queen from one of her worker eggs. They may do this because she has been damaged (lost a leg or an antenna), because she has run out of sperm and cannot lay fertilized eggs (has become a "drone laying queen"), or because her pheromones have dwindled to where they cannot control all the bees in the hive. At this juncture, the bees produce one or more queen cells by modifying existing worker cells that contain a normal female egg. They then pursue one of two ways to replace the queen: one is to supersedure, that is, replacing or superseding the queen without swarming, or, two, swarm cell production, that is, dividing the hive into two colonies through swarming.[84]

Supersedure is a valued behavioral trait by some beekeepers. A hive that supersedes its old queen does not lose any stock. Instead it creates a new queen and the old one fades away or is killed when the new queen emerges. In these hives, the bees produce just one or two queen cells, characteristically in the center of the face of a broodcomb.[85]

Swarm cell production involves creating many queen cells, typically a dozen or more. These are large, peanut-shaped protrusions requiring space, for which reason they are often located around the edges of a broodcomb, commonly at the sides and the bottom.[85]

Once either process has begun, the old queen leaves the hive with the hatching of the first queen cells. She leaves accompanied by a large number of bees, predominantly young bees (wax-secretors), who form the basis of the new hive. Scouts are sent out from the swarm to find suitable hollow trees or rock crevices. As soon as one is found, the entire swarm moves in. Within a matter of hours, they build new wax brood combs, using honey stores that the young bees have filled themselves with before leaving the old hive. Only young bees can secrete wax from special abdominal segments, and this is why swarms tend to contain more young bees. Often a number of virgin queens accompany the first swarm (the "prime swarm"), and the old queen is replaced as soon as a daughter queen mates and begins laying. Otherwise, she is quickly superseded in the new home.[85]

Different sub-species of Apis mellifera exhibit differing swarming characteristics. In general the more northerly black races are said to swarm less and supersede more, whereas the more southerly yellow and grey varieties are said to swarm more frequently. The truth is complicated because of the prevalence of cross-breeding and hybridization of the sub-species.[85]

A swarm attached to a branch

George S. Demuth describes the main factors that increase the swarming tendency of bees.[86] They are:

• The genetics of bees; that is, how strong is the swarming instinct
• Congestion of the brood nest
• Insufficient empty combs for ripening nectar and storing honey
• Inadequate ventilation
• Having an old queen
• Warming weather conditions.

Demuth attributed some of his comments to Snelgrove.[87]

Some beekeepers may monitor their colonies carefully in spring and watch for the appearance of queen cells, which are a dramatic signal that the colony is determined to swarm.[85]

This swarm looks for shelter. A beekeeper may capture it and introduce it into a new hive, helping meet this need. Otherwise, it reverts to a feral state, in which case it finds shelter in a hollow tree, excavation, abandoned chimney, or even behind shutters.[85]

A small after-swarm has less chance of survival and may threaten the original hive's survival if the number of individuals left is unsustainable. When a hive swarms despite the beekeeper's preventative efforts, a good management practice is to give the reduced hive a couple frames of open brood with eggs. This helps replenish the hive more quickly and gives a second opportunity to raise a queen if there is a mating failure.[85]

Each sub-species of honey bee has its own swarming characteristics. Italian bees are very prolific and inclined to swarm; Northern European black bees have a strong tendency to supersede their old queen without swarming. These differences are the result of differing evolutionary pressures in the regions where each sub-species evolved.[85]

When a colony accidentally loses its queen, it is said to be "queenless".[88] The workers realize that the queen is absent after as little as an hour, as her pheromones fade in the hive. Instinctively, the workers select cells containing eggs aged less than three days and enlarge these cells dramatically to form "emergency queen cells". These appear similar to large peanut-like structures about an inch long that hang from the center or side of the brood combs. The developing larva in a queen cell is fed differently from an ordinary worker-bee; in addition to the normal honey and pollen, she receives a great deal of royal jelly, a special food secreted by young "nurse bees" from the hypopharyngeal gland.[89] This special food dramatically alters the growth and development of the larva so that, after metamorphosis and pupation, it emerges from the cell as a queen bee. The queen is the only bee in a colony which has fully developed ovaries, and she secretes a pheromone which suppresses the normal development of ovaries in all her workers.[90]

Beekeepers use the ability of the bees to produce new queens to increase their colonies in a procedure called splitting a colony.[91] To do this, they remove several brood combs from a healthy hive, taking care to leave the old queen behind. These combs must contain eggs or larvae less than three days old and be covered by young nurse bees, which care for the brood and keep it warm. These brood combs and attendant nurse bees are then placed into a small "nucleus hive" with other combs containing honey and pollen. As soon as the nurse bees find themselves in this new hive and realize they have no queen, they set about constructing emergency queen cells using the eggs or larvae they have in the combs with them.[85]

The common agents of disease that affect adult honey bees include fungi, bacteria, protozoa, viruses, parasites, and poisons. The gross symptoms displayed by affected adult bees are very similar, whatever the cause, making it difficult for the apiarist to ascertain the causes of problems without microscopic identification of microorganisms or chemical analysis of poisons.[92] Since 2006, colony losses from colony collapse disorder have been increasing across the world although the causes of the syndrome are, as yet, unknown.[93][94] In the US, commercial beekeepers have been increasing the number of hives to deal with higher rates of attrition.[95]

Nosema apis is a microsporidian which causes the most common and widespread disease of the adult honey bee, nosemosis, also called nosema.[96]

Galleria mellonella and Achroia grisella "wax moth" larvae that hatch, tunnel through, and destroy comb that contains bee larvae and their honey stores. The tunnels they create are lined with silk, which entangles and starves emerging bees. Destruction of honeycombs also results in honey leaking and being wasted. A healthy hive can manage wax moths, but weak colonies, unoccupied hives, and stored frames can be decimated.[97]

Small hive beetle (Aethina tumida) is native to Africa but has now spread to most continents. It is a serious pest among honey bees unadapted to it.[98]

Varroa destructor, the Varroa mite, is an established pest of two species of honey bee through many parts of the world, and is blamed by many researchers as a leading cause of CCD.[99]

Tropilaelaps mites, of which there are four species, are native to Apis dorsata, Apis laboriosa, and Apis breviligula, but spread to Apis mellifera after they were introduced to Asia.[100]

Acarapis woodi, the tracheal mite, infests the trachea of honey bees.[101]

Most predators prefer not to eat honeybees due to their unpleasant sting, but they still have some predators. These include large animals such as skunks or bears, which are after the honey and brood in the nest as well as the adult bees themselves.[102] Some birds will also eat bees (for example, bee-eaters, which are named for their bee-centric diet), as do some robber flies, such as Mallophora ruficauda, which is a pest of apiculture in South America due to its habit of eating workers while they are foraging in meadows.[103]