|Western honey bee|
Temporal range: Oligocene–Recent
The western honey bee or European honey bee (Apis mellifera) is the most common of the 7–12 species of honey bee worldwide.The genus name Apis is Latin for "bee", and mellifera is the Latin for "honey-bearing", referring to the species' production of honey.
A honey bee is a eusocial, flying insect within the genus Apis of the bee clade. They are known for construction of perennial, colonial nests from wax, for the large size of their colonies, and for their surplus production and storage of honey, distinguishing their hives as a prized foraging target of many animals, including honey badgers, bears and human hunter-gatherers. In the early 21st century, between seven and nine species of honey bee are recognized, with a total of 44 subspecies, though historically seven to eleven species are recognized. The best known honey bee is the western honey bee which has been domesticated for honey production and crop pollination; modern humans also value the wax for candlemaking, soapmaking, lip balms, and other crafts. Honey bees represent only a small fraction of the roughly 20,000 known species of bees. Some other types of related bees produce and store honey and have been kept by humans for that purpose, including the stingless honey bees, but only members of the genus Apis are true honey bees. The study of bees, which includes the study of honey bees, is known as melittology.
A genus is a taxonomic rank used in the biological classification of living and fossil organisms, as well as viruses, in biology. In the hierarchy of biological classification, genus comes above species and below family. In binomial nomenclature, the genus name forms the first part of the binomial species name for each species within the genus.
Latin is a classical language belonging to the Italic branch of the Indo-European languages. The Latin alphabet is derived from the Etruscan and Greek alphabets and ultimately from the Phoenician alphabet.
Like all honey bees, the western honey bee is eusocial, creating colonies with a single fertile female (or "queen"), many normally non-reproductive females or "workers," and small proportion of fertile males or "drones." Individual colonies can house tens of thousands of bees. Colony activities are organized by complex communication between individuals, through both pheromones and the dance language.
Eusociality, the highest level of organization of sociality, is defined by the following characteristics: cooperative brood care, overlapping generations within a colony of adults, and a division of labor into reproductive and non-reproductive groups. The division of labor creates specialized behavioral groups within an animal society which are sometimes called castes. Eusociality is distinguished from all other social systems because individuals of at least one caste usually lose the ability to perform at least one behavior characteristic of individuals in another caste.
A beehive is an enclosed, man-made structure in which some honey bee species of the subgenus Apis live and raise their young. Though the word beehive is commonly used to describe the nest of any bee colony, scientific and professional literature distinguishes nest from hive. Nest is used to discuss colonies which house themselves in natural or artificial cavities or are hanging and exposed. Hive is used to describe an artificial, man-made structure to house a honey bee nest. Several species of Apis live in colonies, but for honey production the western honey bee and the eastern honey bee are the main species kept in hives.
Honey bees are sensitive to odors, tastes, and colors, including ultraviolet. They can demonstrate capabilities such as color discrimination through classical and operant conditioning and retain this information for several days at least; they communicate the location and nature of sources of food; they adjust their foraging to the times at which food is available; they may even form cognitive maps of their surroundings.
The western honey bee was one of the first domesticated insects, and it is the primary species maintained by beekeepers to this day for both its honey production and pollination activities. With human assistance, the western honey bee now occupies every continent except Antarctica. Because of its wide cultivation, this species is the single most important pollinator for agriculture globally [ citation needed ]. Honey bees are threatened by pests and diseases, especially the varroa mite and colony collapse disorder. As of 2019, the western honey bee is listed as Data Deficient on the IUCN Red List as numerous studies indicate that the species has undergone significant declines in Europe; however, it is not clear if they refer to population reduction of wild or managed colonies. Further research is required to enable differentiation between wild and non-wild colonies in order to determine the conservation status of the species in the wild.
Domestication is a sustained multi-generational relationship in which one group of organisms assumes a significant degree of influence over the reproduction and care of another group to secure a more predictable supply of resources from that second group.
Pollination is the transfer of pollen from a male part of a plant to a female part of a plant, later enabling fertilisation and the production of seeds, most often by an animal or by wind. Pollinating agents are animals such as insects, birds, and bats; water; wind; and even plants themselves, when self-pollination occurs within a closed flower. Pollination often occurs within a species. When pollination occurs between species it can produce hybrid offspring in nature and in plant breeding work.
A pollinator is an animal that moves pollen from the male anther of a flower to the female stigma of a flower. This helps to bring about fertilization of the ovules in the flower by the male gametes from the pollen grains.
Western honey bees are an important model organism in scientific studies, particularly in the fields of social evolution, learning, and memory; they are also used in studies of pesticide toxicity, to assess non-target impacts of commercial pesticides.
A model organism is a non-human species that is extensively studied to understand particular biological phenomena, with the expectation that discoveries made in the model organism will provide insight into the workings of other organisms. Model organisms are widely used to research human disease when human experimentation would be unfeasible or unethical. This strategy is made possible by the common descent of all living organisms, and the conservation of metabolic and developmental pathways and genetic material over the course of evolution.
Social evolution is a subdiscipline of evolutionary biology that is concerned with social behaviors that have fitness consequences for individuals other than the actor. It is also a subdiscipline of sociology that studies evolution of social systems.
Pesticides vary in their effects on bees. Contact pesticides are usually sprayed on plants and can kill bees when they crawl over sprayed surfaces of plants or other areas around it. Systemic pesticides, on the other hand, are usually incorporated into the soil or onto seeds and move up into the stem, leaves, nectar, and pollen of plants.
The western honey bee can be found on every continent except Antarctica.The species is believed to have originated in Africa or Asia, and it spread naturally through Africa, the Middle East and Europe. Humans are responsible for its considerable additional range, introducing European subspecies into North America (early 1600s), South America, Australia, New Zealand, and East Asia.
In biogeography, a taxon is said to have a cosmopolitan distribution if its range extends across all or most of the world in appropriate habitats. Such a taxon is said to exhibit cosmopolitanism or cosmopolitism. The opposite extreme is endemism.
Western honey bees adapted to the local environments as they spread geographically.These adaptations include synchronizing colony cycles to the timing of local flower resources, forming a winter cluster in colder climates, migratory swarming in Africa, and enhanced foraging behavior in desert areas. All together, these variations resulted in 29 recognized subspecies, all of which are cross-fertile. The subspecies are divided into four major branches, based on work by Ruttner and confirmed by mitochondrial DNA analysis. African subspecies belong to branch A, northwestern European subspecies branch M, southwestern European subspecies branch C and Middle-Eastern subspecies branch O.
In beekeeping, a winter cluster is a well-defined cluster of honey bees that forms inside a beehive when the air temperature dips below 10 to 14 °C. Honey bees are one of but a few kinds of insects that survive the winter as a colony. As the outside air temperature decreases the winter cluster becomes tighter and more compact. The bees cling tightly together on the combs in the hive. The temperature within the winter cluster remains remarkably warm regardless of the outside air temperature. Larger clusters have a better chance for survival than smaller clusters. The winter cluster within the hive must move throughout the winter to reach the available honey stored in the combs.
Mitochondrial DNA is the DNA located in mitochondria, cellular organelles within eukaryotic cells that convert chemical energy from food into a form that cells can use, adenosine triphosphate (ATP). Mitochondrial DNA is only a small portion of the DNA in a eukaryotic cell; most of the DNA can be found in the cell nucleus and, in plants and algae, also in plastids such as chloroplasts.
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Unlike most other bee species, honey bees have perennial colonies which persist year after year. Because of this high degree of sociality and permanence, honey bee colonies can be considered superorganisms. This means that reproduction of the colony, rather than individual bees, is the biologically significant unit. Honey bee colonies reproduce through a process called "swarming".
In most climates, western honey bees swarm in the spring and early summer, when there is an abundance of blooming flowers from which to collect nectar and pollen. In response to these favorable conditions, the hive creates one to two dozen new queens. Just as the pupal stages of these "daughter queens" are nearly complete, the old queen and approximately two-thirds of the adult workers leave the colony in a swarm, traveling some distance to find a new location suitable for building a hive (e.g., a hollow tree trunk). In the old colony, the daughter queens often start "piping", just prior to emerging as adults,and, when the daughter queens eventually emerge, they fight each other until only one remains; the survivor then becomes the new queen. If one of the sisters emerges before the others, she may kill her siblings while they are still pupae, before they have a chance to emerge as adults.
Once she has dispatched her rivals, the new queen, the only fertile female, lays all the eggs for the old colony, which her mother has left. Virgin females are able to lay eggs, which develop into males (a trait shared with wasps, bees, and ants because of haplodiploidy). However, she requires a mate to produce female offspring, which comprise 90% or more of bees in the colony at any given time. Thus, the new queen goes on one or more nuptial flights, each time mating with 1–17 drones.Once she has finished mating, usually within two weeks of emerging, she remains in the hive, laying eggs.
Throughout the rest of the growing season, the colony produces many workers, who gather pollen and nectar as cold-season food; the average population of a healthy hive in midsummer may be as high as 40,000 to 80,000 bees. Nectar from flowers is processed by worker bees, who evaporate it until the moisture content is low enough to discourage mold, transforming it into honey, which can then be capped over with wax and stored almost indefinitely. In the temperate climates to which western honey bees are adapted, the bees gather in their hive and wait out the cold season, during which the queen may stop laying. During this time, activity is slow, and the colony consumes its stores of honey used for metabolic heat production in the cold season. In mid- through late winter, the queen starts laying again. This is probably triggered by day length. Depending on the subspecies, new queens (and swarms) may be produced every year, or less frequently, depending on local environmental conditions.
Like other insects that undergo complete metamorphosis, the western honey bee has four distinct life stages: egg, larva, pupa and adult. The complex social structure of honey bee hives means that all of these life stages occur simultaneously throughout much of the year. The queen deposits a single egg into each cell of a honeycomb prepared by worker bees. The egg hatches into a legless, eyeless larva fed by "nurse" bees (worker bees who maintain the interior of the colony). After about a week, the larva is sealed in its cell by the nurse bees and begins its pupal stage. After another week, it emerges as an adult bee. It is common for defined regions of the comb to be filled with young bees (also called "brood"), while others are filled with pollen and honey stores.
Worker bees secrete the wax used to build the hive, clean, maintain and guard it, raise the young and forage for nectar and pollen; the nature of the worker's role varies with age. For the first ten days of their lives, worker bees clean the hive and feed the larvae. After this, they begin building comb cells. On days 16 through 20, workers receive nectar and pollen from older workers and store it. After the 20th day, a worker leaves the hive and spends the remainder of its life as a forager. Although worker bees are usually infertile females, when some subspecies are stressed they may lay fertile eggs. Since workers are not fully sexually developed, they do not mate with drones and thus can only produce haploid (male) offspring.
Queens and workers have a modified ovipositor, a stinger, with which they defend the hive. Unlike bees of any other genus and the queens of their own species, the stinger of worker honey bees is barbed. Contrary to popular belief, a bee does not always die soon after stinging; this misconception is based on the fact that a bee will usually die after stinging a human or other mammal. The stinger and its venom sac, with musculature and a ganglion allowing them to continue delivering venom after they are detached, are designed to pull free of the body when they lodge. This apparatus (including barbs on the stinger) is thought to have evolved in response to predation by vertebrates, since the barbs do not function (and the stinger apparatus does not detach) unless the stinger is embedded in elastic material. The barbs do not always "catch", so a bee may occasionally pull its stinger free and fly off unharmed (or sting again).
Although the average lifespan of a queen in most subspecies is three to five years, reports from the German-European black bee subspecies previously used for beekeeping indicate that a queen can live up to eight years.Because a queen's store of sperm is depleted near the end of her life, she begins laying more unfertilized eggs; for this reason, beekeepers often replace queens every year or two.
The lifespan of workers varies considerably over the year in regions with long winters. Workers born in spring and summer will work hard, living only a few weeks, but those born in autumn will remain inside for several months as the colony clusters. On average during the year, about one percent of a colony's worker bees die naturally per day.Except for the queen, all of a colony's workers are replaced about every four months.
Behavioral and physiological differences between castes and subcastes arise from phenotypic plasticity, which relies on gene expression rather than heritable genotypic differences.
The queen bee is a fertile female, who, unlike workers (which are genetically also female), has a fully developed reproductive tract. She is larger than her workers, and has a characteristic rounder, longer abdomen. A female egg can become either a queen or a worker bee. Workers and queens are both fed royal jelly, which is high in protein and low in flavonoid, during the first three days of their larval stage. Workers are then switched to a diet of mixed pollen and nectar (often called "bee bread"), while queens will continue to receive royal jelly. In the absence of flavonoid and the presence of a high-protein diet, queen bees develop a healthy reproductive tract—a task necessary for maintaining a colony of tens of thousands of daughter-workers.
Periodically, the colony determines that a new queen is needed. There are three general causes:
Regardless of the trigger, workers develop the larvae into queens by continuing to feed them royal jelly.
Queens are not raised in the typical horizontal brood cells of the honeycomb. A queen cell is larger and oriented vertically. If workers sense that an old queen is weakening, they produce emergency cells (known as supersedure cells) made from cells with eggs or young larvae and which protrude from the comb. When the queen finishes her larval feeding and pupates, she moves into a head-downward position and later chews her way out of the cell. At pupation, workers cap (seal) the cell. The queen asserts control over the worker bees by releasing a complex suite of pheromones, known as queen scent.
After several days of orientation in and around the hive, the young queen flies to a drone congregation point – a site near a clearing and generally about 30 feet (9.1 m) above the ground – where drones from different hives congregate. They detect the presence of a queen in their congregation area by her smell, find her by sight and mate with her in midair; drones can be induced to mate with "dummy" queens with the queen pheromone. A queen will mate multiple times, and may leave to mate several days in a row (weather permitting) until her spermatheca is full.
The queen lays all the eggs in a healthy colony. The number and pace of egg-laying is controlled by weather, resource availability and specific racial characteristics. Queens generally begin to slow egg-laying in the early fall, and may stop during the winter. Egg-laying generally resumes in late winter when the days lengthen, peaking in the spring. At the height of the season, the queen may lay over 2,500 eggs per day (more than her body mass).
She fertilizes each egg (with stored sperm from the spermatheca) as it is laid in a worker-sized cell. Eggs laid in drone-sized (larger) cells are left unfertilized; these unfertilized eggs, with half as many genes as queen or worker eggs, develop into drones.
Workers are females produced by the queen that develop from fertilized, diploid eggs. Workers are essential for social structure and proper colony functioning. They carry out the main tasks of the colony, because the queen is occupied with only reproducing. These females will raise their sister workers and future queens that eventually leave the nest to start their own colony. They also forage and return to the nest with nectar and pollen to feed the young.
Drones are the colony's male bees. Since they do not have ovipositors, they do not have stingers. Drone honey bees do not forage for nectar or pollen. The primary purpose of a drone is to fertilize a new queen. Many drones will mate with a given queen in flight; each will die immediately after mating, since the process of insemination requires a lethally convulsive effort. Drone honey bees are haploid (single, unpaired chromosomes) in their genetic structure, and are descended only from their mother (the queen). In temperate regions drones are generally expelled from the hive before winter, dying of cold and starvation since they cannot forage, produce honey or care for themselves. There has been research into the role western honey bee drones play in thermoregulation within the hive. Given their larger size (1.5 times that of worker bees), drones may play a significant role. Drones are typically located near the center of hive clusters for unclear reasons. It is postulated that it is to maintain sperm viability, which may be compromised at cooler temperatures. Another possible explanation is that a more central location allows drones to contribute to warmth, since at temperatures below 25 °C (77 °F) their ability to contribute declines.
When a fertile female worker produces drones, a conflict arises between her interests and those of the queen. The worker shares half her genes with the drone and one-quarter with her brothers, favouring her offspring over those of the queen. The queen shares half her genes with her sons and one-quarter with the sons of fertile female workers.This pits the worker against the queen and other workers, who try to maximize their reproductive fitness by rearing the offspring most related to them. This relationship leads to a phenomenon known as "worker policing". In these rare situations, other worker bees in the hive who are genetically more related to the queen's sons than those of the fertile workers will patrol the hive and remove worker-laid eggs. Another form of worker-based policing is aggression toward fertile females. Some studies have suggested a queen pheromone which may help workers distinguish worker- and queen-laid eggs, but others indicate egg viability as the key factor in eliciting the behavior. Worker policing is an example of forced altruism, where the benefits of worker reproduction are minimized and that of rearing the queen's offspring maximized.
In very rare instances workers subvert the policing mechanisms of the hive, laying eggs which are removed at a lower rate by other workers; this is known as anarchic syndrome. Anarchic workers can activate their ovaries at a higher rate and contribute a greater proportion of males to the hive. Although an increase in the number of drones would decrease the overall productivity of the hive, the reproductive fitness of the drones' mother would increase. Anarchic syndrome is an example of selection working in opposite directions at the individual and group levels for the stability of the hive.
Under ordinary circumstances the death (or removal) of a queen increases reproduction in workers, and a significant proportion of workers will have active ovaries in the absence of a queen. The workers of the hive produce a last batch of drones before the hive eventually collapses. Although during this period worker policing is usually absent, in certain groups of bees it continues.
According to the strategy of kin selection, worker policing is not favored if a queen does not mate multiple times. Workers would be related by three-quarters of their genes, and the difference in relationship between sons of the queen and those of the other workers would decrease. The benefit of policing is negated, and policing is less favored. Experiments confirming this hypothesis have shown a correlation between higher mating rates and increased rates of worker policing in many species of social hymenoptera.
The honey bee needs an internal body temperature of 35 °C (95 °F) to fly; this temperature is maintained in the nest to develop the brood, and is the optimal temperature for the creation of wax. The temperature on the periphery of the cluster varies with outside air temperature, and the winter cluster's internal temperature may be as low as 20–22 °C (68–72 °F).
Honey bees can forage over a 30 °C (86 °F) air-temperature range because of behavioral and physiological mechanisms for regulating the temperature of their flight muscles. From low to high air temperatures, the mechanisms are: shivering before flight, and stopping flight for additional shivering; passive body-temperature regulation based on work, and evaporative cooling from regurgitated honey-sac contents. Body temperatures differ, depending on caste and expected foraging rewards.
The optimal air temperature for foraging is 22–25 °C (72–77 °F). During flight, the bee's relatively large flight muscles create heat which must be dissipated. The honey bee uses evaporative cooling to release heat through its mouth. Under hot conditions, heat from the thorax is dissipated through the head; the bee regurgitates a droplet of warm internal fluid — a "honeycrop droplet" – which reduces the temperature of its head by 10 °C (18 °F).
Below 7–10 °C (45–50 °F) bees are immobile, and above 38 °C (100 °F) their activity slows. Honey bees can tolerate temperatures up to 50 °C (122 °F) for short periods.
Honey bee behavior has been extensively studied, since bees are widespread and familiar. Karl von Frisch, who received the 1973 Nobel Prize in Physiology or Medicine for his study of honey-bee communication, noticed that bees communicate with dance. Through these dances, bees communicate information regarding the distance, the situation, and the direction of a food source by the dances of the returning (honey bee) worker bee on the vertical comb of the hive. 50 metres (160 ft) of the hive, it provides insufficient information about direction. The waggle dance, which may be vertical or horizontal, provides more detail about the distance and direction of a food source. Foragers are also thought to rely on their olfactory sense to help locate a food source after they are directed by the dances. Unlike western honey bees, the dwarf honey bee does not change the precision of the waggle dance to indicate the type of site that is set as a new goal. Therefore, western honey bees are better at conveying information than its closely related species and this further supports the notion that former are more evolved than latter.Honey bees direct other bees to food sources with the round dance and the waggle dance. Although the round dance tells other foragers that food is within
Another means of communication is the shaking signal, also known as the jerking dance, vibration dance or vibration signal. Although the shaking signal is most common in worker communication, it also appears in reproductive swarming. A worker bee vibrates its body dorsoventrally while holding another bee with its front legs. Jacobus Biesmeijer, who examined shaking signals in a forager's life and the conditions leading to its performance, found that experienced foragers executed 92 percent of observed shaking signals and 64 percent of these signals were made after the discovery of a food source. About 71 percent of shaking signals occurred before the first five successful foraging flights of the day; other communication signals, such as the waggle dance, were performed more often after the first five successes. Biesmeijer demonstrated that most shakers are foragers and the shaking signal is most often executed by foraging bees on pre-foraging bees, concluding that it is a transfer message for several activities (or activity levels). Sometimes the signal increases activity, as when active bees shake inactive ones. At other times, such as the end of the day, the signal is an inhibitory mechanism. However, the shaking signal is preferentially directed towards inactive bees. All three forms of communication among honey bees are effective in foraging and task management.
Pheromones (substances involved in chemical communication) are essential to honey-bee survival. Honey bees rely on pheromones for nearly all behaviors, including mating, alarm, defense, orientation, kin and colony recognition, food production and integrating colony activities.
Humans have been collecting honey from bees for thousands of years, with evidence in the form of rock art found in France and Spain,dating to around 7000 BCE. The honey bee is one of the few invertebrate animals to have been domesticated. Bees were likely first domesticated in ancient Egypt, where tomb paintings depict bee-keeping. Europeans brought bees to North America in 1622.
Beekeepers have selected bees for several desirable features:
These modifications, along with artificial change of location, have improved bees from the point of view of the beekeeper, and simultaneously made them more dependent on beekeepers for their survival. In Europe, cold-weather survival was likely selected for, consciously or not, while in Africa, selection probably favoured the ability to survive heat, drought, and heavy rain.
Authors do not agree on whether this degree of artificial selection constitutes genuine domestication. In 1603, John Guillim wrote "The Bee I may well reckon a domestic insect, being so pliable to the benefit of the keeper."More recently, many biologists working on pollination take the domesticated status of honey bees for granted. For example, Rachael Winfree and colleagues write "We used crop pollination as a model system, and investigated whether the loss of a domesticated pollinator (the honey bee) could be compensated for by native, wild bee species." Similarly, Brian Dennis and William Kemp write: "Although the domestication of the honey bee is closely connected to the evolution of food-based socio-economic systems in many cultures throughout the world, in current economic terms, and in the U.S. alone, the estimated wholesale value of honey, more than $317 million dollars in 2013, pales in comparison to aggregate estimated annual value of pollination services, variously valued at $11–15 billion."
On the other hand, P. R. Oxley and B. P. Oldroyd (2010) consider the domestication of bees at best partial.Oldroyd observes that the lack of full domestication is somewhat surprising, given that people have kept bees for at least 7000 years. Instead, bee-keepers have found ways to manage bees using hives, while the bees remain "largely unchanged from their wild cousins".
Leslie Bailey and B. V. Ball, in their book Honey Bee Pathology, call honey bees "feral insects", in contrast to the silkworm (Bombyx mori) which they call "the only insect that has been domesticated", and refer to the "popular belief among many biologists as well as beekeepers that bees are domesticated". They argue that honey bees are able to survive without man's help, and in fact require to "be left at liberty" to survive. They argue further that even if bees could be raised away from the wild, they would still have to fly freely to gather nectar and pollinate plants. Therefore, they argue, bee-keeping is "the exploitation of colonies of a wild insect", with little more than the provision of a weatherproof cavity for them to nest in.Pilar de la Rua and colleagues likewise argue that honey bees are not fully domesticated, since "endemic subspecies-specific genetic footprints can still be identified in Europe and Africa", making conservation of wild-bee diversity important. They further argue that the difficulty of controlling drones for mating is a serious handicap and a sign that domestication is not complete, in particular as "extensive gene flow usually occurs between wild/feral and managed honeybee populations".
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The honey bee is a colonial insect which is housed, transported by and sometimes fed by beekeepers. Honey bees do not survive and reproduce individually, but as part of the colony (a superorganism).
Honey bees collect flower nectar and convert it to honey, which is stored in the hive. The nectar, transported in the bees' stomachs, is converted with the addition of digestive enzymes and storage in a honey cell for partial dehydration. Nectar and honey provide the energy for the bees' flight muscles and for heating the hive during the winter. Honey bees also collect pollen, which after being processed to bee bread supplies protein and fat for the bee brood to grow. Centuries of selective breeding by humans have created bees which produce far more honey than the colony needs, and beekeepers (also known as apiarists) harvest the surplus honey.
Beekeepers provide a place for the colony to live and store honey. There are seven basic types of beehive: skeps, Langstroth hives, top-bar hives, box hives, log gums, D. E. hive, and miller hives.All U.S. states require beekeepers to use movable frames to allow bee inspectors to check the brood for disease. This allows beekeepers to keep Langstroth, top-bar and D.E. hives without special permission, granted for purposes such as museum use. Modern hives also enable beekeepers to transport bees, moving from field to field as crops require pollinating (a source of income for beekeepers).
In cold climates, some beekeepers have kept colonies alive (with varying degrees of success) by moving them indoors for winter. While this can protect the colonies from extremes of temperature and make winter care and feeding more convenient for the beekeeper, it increases the risk of dysentery and causes an excessive buildup of carbon dioxide from the bees' respiration. Inside wintering has been refined by Canadian beekeepers, who use large barns solely for the wintering of bees; automated ventilation systems assist in carbon-dioxide dispersal.
Bees are one of the products of a beehive. They can be purchased as mated queens, in spring packages of a queen along with 2 to 5 pounds (0.91 to 2.27 kg) of bees, as nucleus colonies (which include frames of brood), or as full colonies. Commerce of bees dates back to as early as 1622, when the first colonies of bees were shipped from England to Virginia. Modern methods of producing queens and dividing colonies for increase date back to the late 1800s. Moses Quinby of New York is considered the first commercial beekeeper in the United States. Before Quinby and his experimentation with movable box hives, which allow honey to be extracted without first killing off the hive, bees and bee products were mainly an object of local trade. The expansion of better highways and increased use of motor vehicles after World War I, allowed commercial beekeepers to expand the size of their businesses.
The honey bee is an important pollinator of crops; this service accounts for much of the species' commercial value. In 2005, the estimated commercial value of honeybees was just under $200 billion.A large number of the crop species farmed worldwide depend on it, Although orchards and fields have increased in size, wild pollinators have dwindled. In a number of regions the pollination shortage is addressed by migratory beekeepers, who supply hives during a crop bloom and move them after the blooming period. Commercial beekeepers plan their movements and wintering locations according to anticipated pollination services. At higher latitudes it is difficult (or impossible) to overwinter sufficient bees, or to have them ready for early blooming plants. Much migration is seasonal, with hives wintering in warmer climates and moving to follow the bloom at higher latitudes. In California almond pollination occurs in February, early in the growing season before local hives have built up their populations.
Almond orchards require two hives per acre, or 2,000 m2 (22,000 sq ft) per hive, for maximum yield, and pollination is dependent on the importation of hives from warmer climates. Almond pollination (in February and March in the U.S.) is the largest managed pollination event in the world, requiring more than one-third of all managed honey bees in the country. Bees are also moved en masse for pollination of apples in New York, Michigan, and Washington. Despite honey bees' inefficiency as blueberry pollinators, large numbers are moved to Maine because they are the only pollinators who can be easily moved and concentrated for this and other monoculture crops. Bees and other insects maintain flower constancy by transferring pollen to other biologically specific plants; this prevents flower stigmas from being clogged with pollen from other species.
European honey bees are often described as being essential to all human food production, leading to claims that without their pollination, all of humanity would starve, or even die out.Einstein is sometimes misquoted as saying If bees disappeared off the face of the earth, man would only have four years left to live. Not only is this misquoted, but there is no science to support this prediction. In fact, many important crops need no insect pollination at all. The 10 most important crops, comprising 60% of all human food energy, all fall into this category: Plantains are sterile and propagated by cuttings, as are cassava. Potatoes, yams, and sweet potatoes are root vegetables propagated by tubers. Soybeans are self-pollinated. Rice, wheat, and corn are all wind-pollinated, because this is true of all grasses.
Similarly, no crops originating in the New World depend on the domesticated honey bee Apis mellifera for pollination at all. The insect is invasive to North America, having arrived with English colonists in the last few centuries. Thomas Jefferson mentioned this in his Notes on the State of Virginia:
|“||The honey-bee is not a native of our continent. Marcgrave indeed mentions a species of honey-bee in Brasil. But this has no sting, and is therefore different from the one we have, which resembles perfectly that of Europe. The Indians concur with us in the tradition that it was brought from Europe; but, when, and by whom, we know not. The bees have generally extended themselves into the country, a little in advance of the white settlers. The Indians therefore call them the white man's fly, and consider their approach as indicating the approach of the settlements of the whites.||”|
Tomatoes, peppers, squash, and all other New World crops evolved with native pollinators like squash bees, bumble bees, other native bees. The stingless bees mentioned by Jefferson are distant relatives of the honey bees, in the genus Melipona .
Honey is the complex substance made from nectar and sweet deposits from plants and trees, which are gathered, modified and stored in the comb by honey bees.Honey is a biological mixture of inverted sugars, primarily glucose and fructose. It has antibacterial and anti-fungal properties. Honey from the western honey bee, along with the bee Tetragonisca angustula , has specific antibacterial activity towards an infection causing bacteria, Staphylococcus aureus . Honey will not rot or ferment when stored under normal conditions, but it will crystallize over time. Although crystallized honey is acceptable for human use, bees can only use liquid honey and will remove and discard crystallized honey from the hive.
Bees produce honey by collecting nectar, a clear liquid consisting of nearly 80 percent water and complex sugars. The collecting bees store the nectar in a second stomach and return to the hive, where worker bees remove the nectar. The worker bees digest the raw nectar for about 30 minutes, using digestive enzymes to break down the complex sugars into simpler ones. Raw honey is then spread in empty honeycomb cells to dry, reducing its water content to less than 20 percent. When nectar is being processed, honey bees create a draft through the hive by fanning with their wings. When the honey has dried, the honeycomb cells are sealed (capped) with wax to preserve it.
Mature worker bees secrete beeswax from glands on their abdomen, using it to form the walls and caps of the comb.When honey is harvested, the wax can be collected for use in products like candles and seals.
Bees collect pollen in a pollen basket and carry it back to the hive, where after undergoing fermentation and turning into bee bread it becomes a protein source for brood-rearing.Excess pollen can be collected from the hive; although it is sometimes consumed as a dietary supplement by humans, bee pollen may cause an allergic reaction in susceptible individuals.
Bee brood, the eggs, larvae, or pupae of honey bees, is edible and highly nutritious. Bee brood contains the same amount of protein that beef or poultry does. Bee brood is often harvested as a byproduct when the farmer has excess bees and does not wish for more.
Propolis is a resinous mixture collected by honey bees from tree buds, sap flows or other botanical sources, which is used as a sealant for unwanted open spaces in the hive.Although propolis is alleged to have health benefits (tincture of Propolis is marketed as a cold and flu remedy), it may cause severe allergic reactions in some individuals. Propolis is also used in wood finishes, and gives a Stradivarius violin its unique red color.
Royal jelly is a honey-bee secretion used to nourish the larvae and queen.It is marketed for its alleged but unsupported claims of health benefits. On the other hand, it may cause severe allergic reactions in some individuals.
As of October 28, 2006, the Honey Bee Genome Sequencing Consortium fully sequenced and analyzed the genome of Apis mellifera, the western honey bee. Since 2007, attention has been devoted to colony collapse disorder, a decline in European honey bee colonies in a number of regions.
The European honey bee is the third insect, after the fruit fly and the mosquito, to have its genome mapped. According to scientists who analyzed its genetic code, the honey bee originated in Africa and spread to Europe in two ancient migrations.Scientists have found that genes related to smell outnumber those for taste, and the European honey bee has fewer genes regulating immunity than the fruit fly and the mosquito. The genome sequence also revealed that several groups of genes, particularly those related to circadian rhythm, resembled those of vertebrates more than other insects. Another significant finding from the honey bee genome study, was that the honey bee was the first insect to be discovered with a functional DNA methylation system since functional key enzymes (DNA methyltransferase-1 and -3) were identified in the genome. DNA methylation is one of the important mechanisms in epigenetics to study gene expression and regulation without changing the DNA sequence, but modifications on DNA activity. DNA methylation later was identified to play an important role in gene regulation and gene splicing. The genome is unusual in having few transposable elements, although they were present in the evolutionary past (remains and fossils have been found) and evolved more slowly than those in fly-species.
European honey bee populations face threats to their survival increasing interests into other pollinator species, like the common eastern bumblebee. [ citation needed ] as bee pests and diseases (including American foulbrood and tracheal mites) are becoming resistant to medications.North American and European populations were severely depleted by Varroa mite infestations during the early 1990s, and U.S. beekeepers were further affected by colony collapse disorder in 2006 and 2007. Improved cultural practices and chemical treatments against Varroa mites saved most commercial operations; new bee breeds are beginning to reduce beekeeper dependence on acaricides. Feral bee populations were greatly reduced during this period; they are slowly recovering, primarily in mild climates, due to natural selection for varroa resistance and repopulation by resistant breeds. Insecticides, particularly when used in excess of label directions, have also depleted bee populations
Africanized bees have spread across the southern United States, where they pose a slight danger to humans (making beekeeping—particularly hobby beekeeping—difficult). As an invasive species, feral honey bees have become a significant environmental problem in non-native areas. Imported bees may displace native bees and birds, and may also promote the reproduction of invasive plants ignored by native pollinators. Unlike native bees, they do not properly extract or transfer pollen from plants with pore anthers (anthers which only release pollen through tiny apical pores); this requires buzz pollination, a behavior rarely exhibited by honey bees. Honey bees reduce fruiting in Melastoma affine , a plant with pore anthers, by robbing its stigmas of previously deposited pollen.
Insect predators of honeybees include the Asian giant hornet and other wasps, robber flies, dragonflies such as the green darner, the European beewolf, some praying mantises and the water strider.
Arachnid predators of honeybees include fishing spiders, lynx spiders, goldenrod spidersand the St Andrew's Cross spider.
Reptile and amphibian predators of honeybees include the American toad, anoles, the American bullfrog and the wood frog.
Specialist bird predators include the bee-eaters; other birds that may take bees include grackles, hummingbirds, summer tanagers, and tyrant flycatchers. Most birds that eat bees do so opportunistically, however, summer tanagers will sit on a limb and catch dozens of bees from the hive entrance.
Mammals that sometimes take bees include bears, least shrews, opossums, raccoons, honey badgers and skunks.
Apart from Apis mellifera, there are six other species in the genus Apis. These are Apis andreniformis, Apis florea, Apis dorsata, Apis cerana, Apis koschevnikovi, and Apis nigrocincta.These other species all originated in South and Southeast Asia. Only Apis mellifera is thought to have originated in Europe, Asia, and Africa.
The Africanized bee, also known as the Africanised honey bee, and known colloquially as the "killer bee", is a hybrid of the western honey bee species, produced originally by cross-breeding of the East African lowland honey bee (A. m. scutellata) with various European honey bees such as the Italian honey bee A. m. ligustica and the Iberian honey bee A. m. iberiensis.
Beekeeping is the maintenance of bee colonies, commonly in man-made hives, by humans. Most such bees are honey bees in the genus Apis, but other honey-producing bees such as Melipona stingless bees are also kept. A beekeeper keeps bees in order to collect their honey and other products that the hive produce, to pollinate crops, or to produce bees for sale to other beekeepers. A location where bees are kept is called an apiary or "bee yard."
The honey bee life cycle, here referring exclusively to the domesticated Western honey bee, depends greatly on their social structure.
In beekeeping, bee brood or brood refers to the eggs, larvae and pupae of honeybees. The brood of Western honey bees develops within a bee hive. In man-made, removable frame hives, such as Langstroth hives, each frame which is mainly occupied by brood is called a brood frame. Brood frames usually have some pollen and nectar or honey in the upper corners of the frame. The rest of the brood frame cells may be empty or occupied by brood in various developmental stages. During the brood raising season, the bees may reuse the cells from which brood has emerged for additional brood or convert it to honey or pollen storage. Bees show remarkable flexibility in adapting cells to a use best suited for the hive's survival.
The term "queen bee" is typically used to refer to an adult, mated female (gyne) that lives in a honey bee colony or hive; she is usually the mother of most, if not all, of the bees in the beehive. Queens developed from larvae selected by worker bees and specially fed in order to become sexually mature. There is normally only one adult, mated queen in a hive, in which case the bees will usually follow and fiercely protect her.
A worker bee is any female (eusocial) bee that lacks the full reproductive capacity of the colony's queen bee; under most circumstances, this is correlated to an increase in certain non-reproductive activities relative to a queen, as well. Worker bees occur in many bee species other than honey bees, but this is by far the most familiar colloquial use of the term.
Nosema apis is a microsporidian, a small, unicellular parasite recently reclassified as a fungus that mainly affects honey bees. It causes nosemosis, also called nosema, which is the most common and widespread disease of adult honey bee diseases. The dormant stage of N. apis is a long-lived spore which is resistant to temperature extremes and dehydration, and cannot be killed by freezing the contaminated comb. Nosemosis is a listed disease with the Office International des Epizooties (OIE).
For bees, their forage or food supply consists of nectar and pollen from blooming plants within flight range. The forage sources for honey bees are an important consideration for beekeepers. In order to determine where to locate hives for maximum honey production and brood one must consider the off-season. If there are no honey flows the bees may have to be fed. Bees that are used for commercial pollination are usually fed in the holding yards. Forage is also significant for pollination management with other bee species. Nectar contains sugars that are the primary source of energy for the bees' wing muscles and for heat for honey bee colonies for winter. Pollen provides the protein and trace minerals that are mostly fed to the brood in order to replace bees lost in the normal course of life cycle and colony activity.
Hive management in beekeeping refers to intervention techniques that a beekeeper may perform to ensure hive survival and to maximize hive production. Hive management techniques vary widely depending on the objectives.
Stingless bees, sometimes called stingless honey bees or simply meliponines, are a large group of bees, comprising the tribe Meliponini. They belong in the family Apidae, and are closely related to common honey bees, carpenter bees, orchid bees, and bumblebees. Meliponines have stingers, but they are highly reduced and cannot be used for defense, though these bees exhibit other defensive behaviors and mechanisms. Meliponines are not the only type of "stingless" bee; all male bees and many female bees of several other families, such as Andrenidae, also cannot sting.
The dwarf honey bee, Apis florea, is one of two species of small, wild honey bees of southern and southeastern Asia. It has a much wider distribution than its sister species, Apis andreniformis. First identified in the late 18th century, Apis florea is unique for its morphology, foraging behavior and defensive mechanisms like making a piping noise. Apis florea have open nests and small colonies, which makes them more susceptible to predation than cavity nesters with large numbers of defensive workers. These honey bees are important pollinators and therefore commodified in countries like Cambodia.
The Cape honey bee or Cape bee is a southern South African subspecies of the Western honey bee. They play a major role in South African agriculture and the economy of the Western Cape by pollinating crops and producing honey in the Western Cape region of South Africa.
Apis andreniformis, or the black dwarf honey bee, is a relatively rare species of honey bee whose native habitat is the tropical and subtropical regions of Southeast Asia.
The East African lowland honey bee is a subspecies of the western honey bee. It is native to central and eastern Africa, though at the southern extreme it is replaced by the Cape honey bee. This subspecies has been determined to constitute one part of the ancestry of the Africanized bees spreading through America.
Apis cerana, the eastern honey bee, is a species of honey bee found in southern and southeastern Asia, including China, Pakistan, India, Korea, Japan, Malaysia, Nepal, Bangladesh, Papua New Guinea, the Solomon Islands and Australia. This species is the sister species of Apis koschevnikovi and both are in the same subgenus as the western (European) honey bee, Apis mellifera. A. cerana is known to live sympatrically along with Apis koschevnikovi within the same geographic location. Apis cerana colonies are known for building nests consisting of multiple combs in cavities containing a small entrance, presumably for defense against invasion by individuals of another nest. The diet of this honey bee species consists mostly of pollen and nectar, or honey. Moreover, Apis cerana is known for its highly social behavior, reflective of its classification as a type of honey bee.
Tetragonula carbonaria is a stingless bee, endemic to the north-east coast of Australia. Its common name is sugarbag bee. They are also occasionally referred to as bush bees. The bee is known to pollinate orchid species, such as Dendrobium lichenastrum, D. toressae, and D. speciosum. It has been identified as an insect that collects pollen from the cycad Cycas media. They are also known for their small body size, reduced wing venation, and highly developed social structure comparable to honey bees.
Melipona beecheii is a species of eusocial, stingless bees of the order Hymenoptera and the genus Melipona. It is native to Central America from the Yucatán Peninsula in the north to Costa Rica in the south. M. beecheii was cultivated in the Yucatán Peninsula starting in the pre-Columbian era by the ancient Maya civilization. The Mayan name for M. beecheii is "xunan kab," which translates roughly to "regal lady bee." M. beecheii once served as the subject of various Mayan religious ceremonies.
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