Honeybees have many adaptations for defense: Adults have orange and black striping that acts as warning coloration. Predators can learn to associate that pattern with a painful sting, and avoid them. Honeybees prefer to build their hives in protected cavities (small caves or tree hollows). They seal small openings with a mix of wax and resins called propolis, leaving only one small opening. Worker bees guard the entrance of the hive. They are able to recognize members of their colony by scent, and will attack any non-members that try to enter the hive. Workers and queens have a venomous sting at the end of the abdomen. Unlike queens, and unusual among stinging insects, the stings of Apis workers are heavily barbed and the sting and venom glands tear out of the abdomen, remaining embedded in the target. This causes the death of the worker, but may also cause a more painful sting, and discourage the predator from attacking other bees or the hive. A stinging worker releases an alarm pheromone which causes other workers to become agitated and more likely to sting, and signals the location of the first sting.
Honeybees are subject to many types of predators, some attacking the bees themselves, others consuming the wax and stored food in the hive. Some predators are specialists on bees, including honeybees.
Important invertebrate enemies of adult bees include crab spiders and orb-weaver spiders, wasps in the genus Philanthus (called “beewolves”), and many species of social wasps in the family Vespidae. Vespid wasp colonies are known to attack honeybee colonies en masse, and can wipe out a hive in one attack. Many vertebrate insectivores also eat adult honeybees. Toads (Bufo) that can reach the entrance of hive will sit and eat many workers, as will opossums (Didelphis). Birds are an important threat – the Meropidae (bee-eaters) in particular in Africa and southern Europe, but also flycatchers around the world (Tyrranidae and Muscicapidae). Apis mellifera in Africa are also subject to attack by honeyguides. These birds eat hive comb, consuming bees, wax, and stored honey. At least one species, the greater honeyguide (Indicator indicator) will guide mammal hive predators to hives, and then feed on the hive after the mammal has opened it up.
The main vertebrate predators of hives are mammals. Bears frequently attack the nests of social bees and wasps, as do many mustelids such as the tayra in the Neotropics and especially the honey badger of Africa and southern and western Asia. In the Western Hemisphere skunks, armadillos and anteaters also raid hives, as do pangolins (Manis) in Africa. Large primates, including baboons, chimpanzees (<>) and gorillas are reported to attack hives too. Smaller mammals such as mice (Mus) and rats (Rattus) will burrow into hives as well.
Some insects are predators in hives as well, including wax moth larvae (Galleria mellonella, Achroia grisella), and hive beetles (Hylostoma, Aethina), and some species of ants. In their native regions these tend not to be important enemies, but where honeybees have not co-evolved with these insects and have no defense, they can do great harm to hives.
See Ecosystem Roles section for information on honeybee parasites and pathogens.
Known Predators:
Anti-predator Adaptations: aposematic
Generally, Apis mellifera are red/brown with black bands and orange yellow rings on abdomen. They have hair on thorax and less hair on abdomen. They also have a pollen basket on their hind legs. Honeybee legs are mostly dark brown/black.
There are two castes of females, sterile workers are smaller (adults 10-15 mm long), fertile queens are larger (18-20 mm). Males, called drones, are 15-17 mm long at maturity. Though smaller, workers have longer wings than drones. Both castes of females have a stinger that is formed from modified ovipositor structures. In workers, the sting is barbed, and tears away from the body when used. In both castes, the stinger is supplied with venom from glands in the abdomen. Males have much larger eyes than females, probably to help locate flying queens during mating flights.
There are currently 26 recognized subspecies of Apis mellifera, with differences based on differences in morphology and molecular characteristics. The differences among the subspecies is usually discussed in terms of their agricultural output in particular environmental conditions. Some subspecies have the ability to tolerate warmer or colder climates. Subspecies may also vary in their defensive behavior, tongue length, wingspan, and coloration. Abdominal banding patterns also differ - some darker and some with more of a mix between darker and lighter banding patterns.
Honeybees are partially endothermic -- they can warm their bodies and the temperature in their hive by working their flight muscles.
Range length: 10 to 20 mm.
Other Physical Features: endothermic ; ectothermic ; heterothermic ; bilateral symmetry ; venomous
Sexual Dimorphism: female larger; sexes shaped differently
Apis mellifera queens usually live 2 to 3 years, but some have been known to last for 5 years. Workers typically only live for a few weeks, sometimes a few months if their hive becomes dormant in winter. Males live for 4-8 weeks at the most.
Typical lifespan
Status: wild: 2 to 3 years.
European honeybees prefer habitats that have an abundant supply of suitable flowering plants, such as meadows, open wooded areas, and gardens. They can survive in grasslands, deserts, and wetlands if there is sufficient water, food, and shelter. They need cavities (e.g. in hollow trees) to nest in.
Habitat Regions: temperate ; tropical ; terrestrial
Terrestrial Biomes: desert or dune ; savanna or grassland ; chaparral ; forest
Wetlands: swamp
Other Habitat Features: urban ; suburban ; agricultural
Apis mellifera is native to Europe, western Asia, and Africa. Human introduction of Apis mellifera to other continents started in the 17th century, and now they are found all around the world, including east Asia, Australia and North and South America.
Biogeographic Regions: nearctic (Introduced ); palearctic (Native ); oriental (Introduced ); ethiopian (Native ); neotropical (Introduced ); australian (Introduced )
Other Geographic Terms: cosmopolitan
Apis mellifera feed on pollen and nectar collected from blooming flowers. They also eat honey (stored, concentrated nectar) and secretions produced by other members of their colony.
Workers forage for food (nectar and pollen) for the entire colony. They use their tongues to suck up nectar, and store it in the anterior section of the digestive tract, called the crop. They collect pollen by grooming it off the bodies and onto special structures on their hind legs called pollen baskets.
Returning foragers transfer the nectar they have collected to younger worker bees that in turn feed other members of the hive, or process it into honey for long-term storage. They add enzymes to the honey, and store it in open cells where the water can evaporate, concentrating the sugars.
Young workers eat pollen and nectar, and secrete food materials, called “royal jelly” and “worker jelly”, from glands in their heads. This material is fed to young larvae, and the amount and type they get determines if they will be queens or workers.
Honeybees forage during daylight hours, but are equally active on cloudy or sunny days. They will not fly in heavy rain or high winds, or if the temperature is too extreme (workers can't fly when they get below 10°C). During the warm, calm weather the honeybees collect the most pollen even if it is cloudy. If the light intensity changes rapidly, they immediately stop working and return to the hive. If it lightly rains, pollen collection stops, because moisture inhibits the bee’s ability to collect it. However, nectar collection is not inhibited by light rain. Wind also affects the rate of pollen collection.
Honeybee workers are opportunistic. They will steal from other hives if they can. Hive-robbing can be dangerous, but a weakened or damaged hive may be raided by workers from other hives, especially when nectar flows in flowers are not abundant. Honeybees will also collect “honeydew,” the sweet fluid excreted by sap-feeding insects like aphids.
Plant Foods: nectar; pollen; sap or other plant fluids
Foraging Behavior: stores or caches food
Primary Diet: herbivore (Nectarivore )
Honeybees are very important pollinators, and are the primary pollinator for many plants. Without honeybees, these plants have greatly reduced fertility. In North America and Australia, where there are no native bee species with large colonies, honeybees can have especially strong effects on native flowers, and on other pollinators such as solitary bee species. Honeybees ability to recruit fellow workers by “dancing” allows them to be more efficient than other pollinators at exploiting patches of flowers. This can create strong impacts on their competitors, especially solitary bees.
Like all social insects, honeybees are hosts to a variety of parasites, commensal organisms, and pathogenic microbes. Some of these can be serious problems for apiculture, and have been studied intensively. At least 18 types of viruses have been found to cause disease in bees, including Sacbrood disease. Several of them (but not sacbrood virus) are associated with parasitic mites. Bacteria infect bees, notably Bacillus larvae, agent of American Foulbrood disease, and Melissococcus pluton, agent of European Foulbrood. Fungi grow in bee hives, and Ascosphaera apis can cause Chalkbrood disease. One of the most common diseases in domesticated hives is Nosema disease, caused by a protozoan, Nosema apis. An amoeba, Malphigamoeba mellificae, also causes disease in honeybees.
In recent decades, two mite species have spread through domesticated and feral honeybee populations around the world. Acarapis woodi is a small mite species that lives in the tracheae of adult bees and feeds on bee hemolymph. It was first discovered in Europe, but its origin is unknown. Infestations of these mites weaken bees, and in cold climates, whole colonies may fail when the bees are confined in the hive during the winter. A much worse threat is Varroa destructor. This might evolved on an Asian honeybee, Apis cerana, but switched on to Apis mellifera colonies that were set up in east Asia. It has since spread all around the world, except Australia. Juvenile mites feed on bee larvae and pupae, and adult female mites feed and disperse on adult workers. This mite is known to spread several viruses as well. Infestations of V. destructor often wipe out colonies. Nearly all the feral, untended honeybee colonies in North American are believed to have been wiped out by mite infestations, along with a large proportion of domesticated colonies. Other mite species are known from honeybee colonies, but they are not considered harmful.
Another commensal or parasitic species is Braula coeca, the bee louse. Despite the common name, this is actually a wingless fly, that apparently feeds by intercepting food being transferred from one bee to another.
Beetles in the genera Hylostoma and Aethina are found in African honeybee nests, where they seem to do little harm. However, the "small hive beetle", Aethina tumida, has become a significant problem in European and North American hives. The larvae eat all the contents of comb: honey, pollen, and bee eggs and larvae.
Ecosystem Impact: pollinates; keystone species
Commensal/Parasitic Species:
Honeybees pollinate billions of US dollars worth of commercial agricultural crops around the world every year. They are important pollinators for economically important wild plant populations as well.
Honeybee hives provide honey and wax, and pollen, propolis, and royal jelly that are sold for medicines and cosmetics.
Honeybees are important study organisms for research in the connections between nervous system structure and behavior.
Some research suggests honeybee venom may have medically useful applications in the treatment of auto-immune disease or inflammation.
Positive Impacts: food ; source of medicine or drug ; research and education; pollinates crops
Honeybee workers will sting humans and domesticated animals in defense of themselves or their hive. A single sting is painful but not dangerous unless the target is allergic to the venom, in which case it can be life threatening. Otherwise, it takes about 20 stings per kilogram of body weight to be life threatening.
Each subspecies of Apis mellifera has different behavioral patterns in regards to intruders near or around the hive. The African subspecies are particularly aggressive. One of them, Apis mellifera scutellata, was accidentally released in South America, and has spread north to the southern United States. This is the "killer bee." It is notable for having a much higher aggressive response to disturbance -- more workers attack than in other subspecies, and they pursue targets much longer than European bees do. The spread of these bees made beekeeping much more expensive and complicated, and the aggressive bees caused many deaths.
Negative Impacts: injures humans (bites or stings, venomous )
Honeybees build a hive out of wax secretions from their bodies, and queens lay their eggs in cells in the wax. The speed of subsequent development of the young is strongly affected by temperature, and is fastest at 33-36°C.
Honeybees are holometabolous insects, and have four stages in the life cycle: egg, larva, pupa, and adult.
A. mellifera eggs hatch in 28-144 hours, depending on their temperature. The larva that emerges is a small white grub. It stays in its wax cell, growing, and is fed and groomed by adult workers. The food that a female larva receives determines whether it will be a queen or worker. At 34°C, larvae feed and grow for 4-5 days, queens for 6 days, and males for 6-7 days. At the end of that period their cell is sealed by adult workers, and the larva molts, spins a silk cocoon, and transforms into the pupa stage. Pupae undergo a massive metamorphosis that takes about 7-8 days for queens, 12 days for workers, and 14-15 days for males. Once their final metamorphosis is complete, they chew their way out of the cell and begin their adult life. They will not grow or molt after emerging. Adult workers will live for 2-4 weeks in the summer, or as long as 11 months if they live through the winter. Males only survive for 4-8 weeks, and do not live through the winter. Queens live 2-5 years.
. The next stage is the larval stage where the larva is fed the royal jelly, pollen/nectar, and honey combination. Next the larva goes into the pupae stage where it caps itself into its cell to metamorphose into the mature stage.
Queens normally take 16 days to reach maturity, the worker bees take 21 days, and the drone takes 24 days to mature.
Development - Life Cycle: metamorphosis
While the species as a whole is still very numerous, there is concern in Europe that widespread commercialization of beekeeping is endangering locally-adapted populations and subspecies. This, combined with higher mortality of colonies due to Varroa mite and tracheal mite infestations, and the recent phenomenon of Colony Collapse Disorder in North America, has cause significant concern for the health of the population. Colony Collapse Disorder (CCD) is a condition of commercial beehives, where there are sudden massive waves of mortality among the workers. Beekeepers discover their hives simply empty of workers, with so few surviving that they cannot tend the queen and brood. This condition has occurred mainly in North America, and mainly in large commercial apiaries. No single cause has been identified yet.
US Federal List: no special status
CITES: no special status
State of Michigan List: no special status
Apis mellifera communication is based on chemical signals, and most of their communication and perception behaviors are centered around scent and taste. The members of the hive colony are bound chemically to each other. Each hive has a unique chemical signature that hivemates use to recognize each other and detect bees from other colonies.
Within the hive, bees are in constant chemical communication with each other. Workers feed and groom each other, as well as larvae, drones, and the queen. In the process they pass on pheromones, chemical signals that indicate information about the health of the queen and the state of the colony.
Chemicals not only help with detecting the right signature of hives but also with foraging. Honeybees use scent to locate flowers from a distance. When a successful forager returns to the hive, it passes the scent of the flowers to its nest mates, to help them find the same patch of flowers.
Bees also use chemicals to signal outside the hive. When a worker stings something, her stinger releases an alarm pheromone that causes other bees to become agitated, and helps them locate the enemy.
Thought it's always dark in the hive, vision is important to honeybees outside. They can see other animals, and recognize flowers. The eyes of Apis species can detect ultraviolet light wavelengths that are beyond the visible spectrum. This allows them to locate the sun on cloudy days, and see markings on flowers that are only visible in ultraviolet light. One portion of honeybee's eyes is sensitive to polarized light, and they use this to navigate.
Workers and queens can hear vibrations. New queens call to each other and workers when they first emerge. Workers hear the vibrations of the waggle dances made by returning foragers.
Apis species have a particularly notable form of communication called "dancing." Foragers that have located an abundant supply of food do a dance to communicate the location of the patch to other foragers. A "round dance" indicates food within about 300 meters of the hive, and only communicates the presence of the flowers, not the direction, though workers will also get the scent from the food the forager has brought back. The more complicated "waggle dance" indicates the direction and distance of food further away, using the location of the sun and the bee's memory of the distance it flew to return to the hive. Symbolic communication is quite unusual among invertebrates, and these honeybee "dances" have been intensively studied.
Communication Channels: visual ; tactile ; acoustic ; chemical
Other Communication Modes: pheromones ; scent marks ; vibrations
Perception Channels: visual ; ultraviolet; polarized light ; tactile ; acoustic ; chemical
The great majority of female A. mellifera in a hive are sterile workers. Only queens mate and lay eggs. Normally there is only a single reproductive queen in a hive.
During periods of suitably mild weather in spring and summer, males leave the hive and gather at "drone assembly areas" near the hive. Virgin queens will fly through these areas, attracting the males with pheromones. The males pursue, and attempt to mate with the queen in flight. Sometimes a "comet" forms, as a cluster of males forms around the female, with a string of other males trying to catch up. Each male who succeeds in mating drops away, and dies within a few hours or days. Males who do not mate will continue to loiter in the assembly areas until they mate or die trying. Queens will mate with up to 10 males in a single flight.
Queens may mate with males from their own hive, or from other hives in the area. The queen's mating behavior is centered around finding the best place to mate beforehand, by taking directional flights for a period of time, lasting no more than a couple of days. Afterward, she leaves the hive and flies to mate with drones in an assembly area. This normally starts to occur after their first week of birth. The queen does this up to four times. After this congregate of mating has occurred, she never mates again in her lifetime.
Mating System: polyandrous ; eusocial
Apis mellifera queens are the primary reproducers of the nest and all of the activities of the colony are centered around their reproductive behaviors and their survival. The queen is the only fertile female in the colony. She lays eggs nearly continuously throughout the year, sometimes pausing in late fall in cold climates. A particularly fertile queen may lay as many as 1,000 eggs/day, and 200,000 eggs in her lifetime. It takes a queen about 16 days to reach adulthood, and another week or more to begin laying eggs. Males take about 24 days to emerge as adults, and begin leaving the nest for assembly areas a few days after that.
Queen honeybees can control whether or not an egg they lay is fertilized. Unfertilized eggs develop as males and are haploid (have only one set of chromosomes). Fertilized eggs are diploid (two sets of chromosomes) and develop as workers or new queens, depending on how they are fed as larvae. Queens may increase the ratio of male to female eggs they lay if they are diseased or injured, or in response to problems in the colony.
Healthy, well-fed honeybee colonies reproduce by "swarming." The workers in the colony begin by producing numerous queen larvae. Shortly before the new queens emerge, the resident, egg-laying queen leaves the hive, taking up to half the workers with her. This "swarm" forms a temporary group in a tree nearby, while workers scout for a suitable location for a new hive. Once they find one, the swarm moves into the space, and begins building comb and starting the process of food collection and reproduction again.
Meanwhile at the old hive, the new queens emerge from their cells. If the population of workers is large enough, and there are few queens emerging, then the first one or two may leave with "afterswarms" of workers. After the swarming is completed, any remaining new queens try to sting and kill each other, continuing to fight until all but one is dead. After her competition is removed, the surviving queen begins to lay eggs.
Normally the pheromones secreted by a healthy queen prevent workers from reproducing, but if a colony remains queenless for long, some workers will begin laying eggs. These eggs are unfertilized, and so develop as males.
Breeding interval: Colonies typically swarm once or twice a year, usually at the beginning of the season that provides the most nectar.
Breeding season: Late spring until the winter months
Range eggs per season: 60,000 to 80,000.
Average gestation period: 3 days.
Range age at sexual or reproductive maturity (female): 15 to 17 days.
Average age at sexual or reproductive maturity (male): 24 days.
Key Reproductive Features: iteroparous ; seasonal breeding ; gonochoric/gonochoristic/dioecious (sexes separate); sexual ; induced ovulation ; fertilization (Internal ); oviparous ; sperm-storing
As in most eusocial insects, the offspring of fertile females (queens) are cared for other members of the colony. In honeybees, the caretakers are sterile females, daughters of the queen, called workers.
Workers build and maintain the comb where young bees are raised, gather food (nectar and pollen) feed and tend larvae, and defend the hive and its young from predators and parasites.
Young queens inherit their hive from their mothers. Often several new queens emerge after the old queen leaves with a swarm to found a new colony. The new queens fight for control of the hive, and only one survives the conflict.
Parental Investment: pre-fertilization (Protecting: Female); pre-hatching/birth (Provisioning: Female, Protecting: Female); pre-weaning/fledging (Provisioning: Female, Protecting: Female); pre-independence (Provisioning: Female, Protecting: Female); inherits maternal/paternal territory; maternal position in the dominance hierarchy affects status of young
Chemical Structure
Melittin is the main component of apitoxin (Apis mellifera venom), accounting for approximately 50% of its dry weight (Terra et al., 2006). The water-soluble, 26 amino acid-long polypeptide chain, weighing 2,840 Da, is largely composed of hydrophobic residues, with the exception of the cationic and hydrophilic carboxy-terminal sequence (Vogel et al., 1986). It is this amphiphilic nature that gives melittin its characteristic detergent-like properties (Maulet et al., 1980).
Using a range of techniques, including X-ray crystallography, NMR spectroscopy, and molecular dynamics simulations, melittin was found to adopt a variety of conformations, depending on factors including the pH and the type of aqueous medium. For instance, when dissolved in water, the hydrophilic residues 22-26 were shown to form a non-helical segment, whereas the remaining hydrophobic residues of melittin were reported to form a bent helix, composed of two smaller α-helical segments of residues 1-11 and 12-21. The concave side of the bent helix was found to be hydrophobic, while the convex side was shown to be hydrophilic (Vogel, et al., 1986). Additionally, melittin was found to be tetrameric at high pH, a random coil at pH 7.0, and monomeric in plasma (Terra, et al., 2006).
Mode of Action
In the bloodstream, melittin is able to rapidly bind to erythrocytes (red blood cells), inducing the release of haemoglobin and other cellular contents into the extracellular medium. Once melittin has penetrated the erythrocyte, it causes micellisation of phosphatidylcholine bilayers, ultimately leading to haemolysis and cell death (Dempsey, et al., 1990).
Apart from its ability to disrupt lipid bilayers, melittin can also inhibit transmembrane proteins, including Na+/K+-ATPase, leading to a rise in sodium concentration within cells (Yang, et al., 2001). The increase in sodium induces an increase in the concentration of intracellular calcium, which results in the increased contraction of cardiac and smooth muscle.
Potential Therapeutic Use
Melittin is currently one of the most extensively used peptides in the research on lipid-peptide and peptide-peptide interactions (Wessman, et al., 2010). The presence of a single tryptophan residue at position 19 allows for a facilitated interpretation of fluorescence data via the tryptophan fluorescence technique, whereby intrinsic fluorescence emissions can be measured via the excitation of tryptophan residues (Raghuraman, et al., 2004).
More recently, the peptide has been shown to possess a variety of therapeutic uses. For instance, melittin is currently being analysed as a potential treatment and preventative for HIV. In a study currently being conducted at Washington University School of Medicine in St. Louis, a melittin-nanoparticle complex was shown to effectively destroy the AIDS-causing virus by forming pores in its protective viral envelope, required for viral reproduction (Evangelou Strait, 2013).
Another use of melittin is in the treatment of cancer. A promising study, once again conducted by researchers at Washington University School of Medicine in St. Louis, involves the attaching of melittin to a different nanoparticle. The novel melittin-nanoparticle complex, named the “nanobee”, selectively targets tumour cells, thus avoiding healthy cells. Once attached to a tumour cell, melittin is able to break down the tumour by forming pores in the cell membrane (Loftus, 2009).
The western honey bee or European honey bee (Apis mellifera) is the most common of the 7–12 species of honey bees worldwide.[3][4] The genus name Apis is Latin for "bee", and mellifera is the Latin for "honey-bearing" or "honey carrying", referring to the species' production of honey.[5]
Like all honey bee species, the western honey bee is eusocial, creating colonies with a single fertile female (or "queen"), many normally non-reproductive females or "workers", and a 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.
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. Western honey bees are threatened by pests and diseases, especially the Varroa mite and colony collapse disorder. There are indications that the species is rare, if not extinct in the wild in Europe and as of 2014, the western honey bee was assessed as "Data Deficient" on the IUCN Red List. 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, meaning self sustaining, without treatments or management.[6]
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, especially via pollen, to assess non-target impacts of commercial pesticides.
The western honey bee can be found on every continent except Antarctica.[7] The species is believed to have originated in Africa[8] or Asia,[9] and it spread naturally through Africa, the Middle East and Europe.[7] Humans are responsible for its considerable additional range, introducing European subspecies into North America (early 1600s),[10] South America, Australia, New Zealand, and eastern Asia.[11]
Western honey bees adapted to the local environments as they spread geographically.[8] 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 31 recognized subspecies.[3]
Previously it was believed that the various subspecies were all cross-fertile, but in 2013 it was found that the A. m. mellifera queens do not mate with non-A. m. mellifera drones.[12]
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.
Unlike most other bee species, western honey bees have perennial colonies which persist year after year. Because of this high degree of sociality and permanence, western honey bee colonies can be considered superorganisms. This means that reproduction of the colony, rather than individual bees, is the biologically significant unit. Western honey bee colonies reproduce through a process called "swarming".[13]
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,[14] 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 all of 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 found in bees, wasps, 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.[15] Once she has finished mating, usually within two weeks of emerging, she remains in the hive, playing the primary role of 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 and a number of characteristics inside the hive.
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 western 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 10 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 called a stinger, with which they defend the hive. Unlike those of bees of any other genus and of the queens of their own species, the stinger of worker western 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).[13]
Although the average lifespan of a queen in most subspecies is three to five years, reports from the German honey bee subspecies (A. m. mellifera) previously used for beekeeping indicate that a queen can live up to eight years.[16] 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 work hard, and live only a few weeks, but those born in autumn remain inside for several months as the colony clusters. On average during the year, about 1% of a colony's worker bees die naturally per day.[17] 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.[18][19]
The queen bee is a fertile female, who, unlike workers (which are also female), has a fully developed reproductive system. 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 queen larva are both fed royal jelly, which is high in protein and low in flavonoids, during the first three days. After that, larval prospective workers are switched to a diet of mixed pollen and nectar (often called "bee bread"), while prospective queens continue to receive royal jelly. In the absence of flavonoids and the presence of a high-protein diet, female bees grow into queens by developing the vigorous reproductive system[20] necessary to maintain a colony of tens of thousands of daughter workers.
Periodically, the colony determines that a new queen is needed. There are three general causes:
Emergency supersedure can generally be recognized because new queen cells are built out from comb cells, instead of hanging from the bottom of a frame. Regardless of the trigger, workers develop existing larvae into queens by continuing to feed them royal jelly, rather than switching them to bee bread, and by extending the selected larvae's cells to house the developing larger-bodied queens.
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) 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 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 mate with a given queen in flight; each dies 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. Given their larger size (1.5 times that of worker bees), inside the hive it is believed that drones may play a significant role in thermoregulation. 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.[21]
When a fertile female worker produces drones, a conflict arises between her interests and those of the queen. The worker shares one-half of her genes with the drone and one-quarter with her brothers, favouring her offspring over those of the queen. The queen shares one-half of her genes with her sons and one-quarter with the sons of fertile female workers.[22] 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 called "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, patrol the hive and remove worker-laid eggs.
Another form of worker policing is aggression toward fertile females.[23] Some studies suggest a queen pheromone which may help workers distinguish worker-laid and queen-laid eggs, but others indicate egg viability as the key factor in eliciting the behavior.[24][25]
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 faster than other workers remove them; 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 decreases the overall productivity of the hive, it increases the reproductive fitness of the drones' mother. Anarchic syndrome is an example of selection working in opposite directions at the individual and group levels for the stability of the hive.[26]
Under ordinary circumstances, if the queen dies or is removed, reproduction in workers increases because a significant proportion of workers then have activated ovaries. The workers produce a last batch of drones before the hive collapses. Although during this period worker policing is usually absent, in certain groups of bees it continues.[27]
According to the strategy of kin selection, worker policing is not favored if a queen mates just once. In that case, workers are related by three-quarters of their genes, and the sons of workers are related more than usual to sons of the queen. Then the benefit of policing is negated. Experiments confirming this hypothesis have shown a correlation between higher mating rates and increased rates of worker policing in many species of social hymenoptera.[28]
The western 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).
Western 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.[29]
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).[30]
Below 7–10 °C (45–50 °F) bees are immobile, and above 38 °C (100 °F) their activity slows. Western honey bees can tolerate temperatures up to 50 °C (122 °F) for short periods.
Western honey bee behavior has been extensively studied. 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.[31] 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 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.
Western honey bees also change the precision of the waggle dance to indicate the type of site that is set as a new goal. Their close relatives, dwarf honey bees, do not.[32] Therefore, western honey bees seem to have evolved a better means of conveying information than their common ancestors with the dwarf honey bee.[33]
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% of observed shaking signals and 64% of these signals were made after the discovery of a food source. About 71% 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. Western honey bees rely on pheromones for nearly all behaviors, including mating, alarm, defense, orientation, kin and colony recognition, food production and integrating colony activities.[34][35]
There is some degree of variability of sociality between individuals.[36][37] Like a great many other social insects, A. mellifera engages in trophallaxis.[36][37] When the duration of trophallaxis pairings was measured, it was found that like human social interactions, there are durable long-term trends for each individual bee.[36][37] There is less inter-individual variation than found in humans however, possibly reflecting the higher genetic relatedness between hivemates.[36]
Humans have been collecting honey from western honey bees for thousands of years, with evidence in the form of rock art found in France and Spain,[38] dating to around 7,000 BCE.[39] The western 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 beekeeping, before 2600BC.[40] Europeans brought bees to North America in 1622.[41][42]
Beekeepers have selected western honey bees for several desirable features:[41]
These modifications, along with artificial change of location, have improved western honey 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.[41]
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."[43] More recently, many biologists working on pollination take the domesticated status of western honey bees for granted.[44][45] 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."[46] 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."[47]
On the other hand, P. R. Oxley and B. P. Oldroyd (2010) consider the domestication of western honey bees, at best, partial.[48] Oldroyd observes that the lack of full domestication is somewhat surprising, given that people have kept bees for at least 7,000 years. Instead, beekeepers have found ways to manage bees using hives, while the bees remain "largely unchanged from their wild cousins".[49]
Leslie Bailey and B. V. Ball, in their book Honey Bee Pathology, call western honey bees "feral insects", in contrast to the domestic silkmoth (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 western honey bees are able to survive without human help, and in fact require to "be left at liberty" to survive. Further, they argue 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, beekeeping is "the exploitation of colonies of a wild insect", with little more than the provision of a weatherproof cavity for them to nest in.[50] Likewise, Pilar de la Rua and colleagues argue that western honey bees are not fully domesticated, because "endemic subspecies-specific genetic footprints can still be identified in Europe and Africa", making conservation of wild bee diversity important. Further, they 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".[51]
The western 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).
Western 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. Western 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 western honey 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. hives, and miller hives.[13] 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.
Honey bees are one of the products of a beehive. They can be purchased as mated queens, in spring packages of a queen along with two to five pounds (0.91 to 2.27 kg) of honey bees, as nucleus colonies (which include frames of brood), or as full colonies. Commerce of western honey 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. Honey was extracted by killing off the hive, and bees and bee products were mainly an object of local trade. The first commercial beekeeper in the United States is considered Moses Quinby of New York, who experimented with movable box hives, which allow extraction without killing the hive. The improvements in roads and motor vehicles after World War I allowed commercial beekeepers to expand the size of their businesses.[52]
The western 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 western honey bees was just under $200 billion worldwide.[53] A large number of the crop species farmed worldwide depend on it.[54] 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,[55] 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;[56] this prevents flower stigmas from being clogged with pollen from other species.[57] In 2000, Drs. Roger Morse and Nicholas Calderone of Cornell University attempted to quantify the effects of the western honey bee on only US food crops. Their calculations came up with a figure of US$14.6 billion in food crop value.[58]
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.[59] 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.[60] 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 any 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.[61] 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.[62] 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 beekeeper has excess bees and does not wish to have any 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.[63] 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.[64] Propolis is also used in wood finishes, and gives a Stradivarius violin its unique red color.[65]
Royal jelly is a honey bee secretion used to nourish the larvae and queen.[66] It is marketed for its alleged but unsupported claims of health benefits.[67][68] On the other hand, it may cause severe allergic reactions in some individuals.[69]
Female bees are diploid and have 32 chromosomes, whereas males are haploid and have only 16.
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 western honey bee colonies in a number of regions.
The western honey bee is the third insect, after the fruit fly and the mosquito, whose genome has been mapped. According to scientists who analyzed its genetic code, the honey bee originated in Africa and spread to Europe in two ancient migrations.[8] 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.[70] 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 western 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.[71] DNA methylation later was identified to play an important role in gene regulation and gene splicing.[72] 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.[70]
Since 2018 a new version of the honey bee genome is available on NCBI (Amel_HAv3.1, BioProject ID: PRJNA471592).[73] This assembly contains full chromosome length scaffolds, which means that the sequence data for each chromosome is contiguous, and not split between multiple pieces called scaffolds. The existence of a highly contiguous reference genome for a species enables more detailed investigations of evolutionary processes that affect the genome as well as more accurate estimations of for example differentiation between populations and diversity within populations.
An important process that shapes the honey bee genome is meiotic recombination, the rate of which is strongly elevated in honey bees and other social insects of the Hymenoptera order compared to most other eukaryotic species except fungi and protozoa.[74] The reason for elevated recombination rates in social Hymenoptera is not fully understood, but one theory is that it is related to their social behaviour. The increased genetic diversity resulting from high recombination rates could make the workers less vulnerable to parasites and facilitate their specialisations to different tasks in the colony.[74]
Western honey bee populations face threats to their survival increasing interests into other pollinator species, like the common eastern bumblebee.[75] 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.[76] Some subspecies of Apis mellifera show naturally varroa sensitive hygiene, for example Apis mellifera lamarckii[77] and Apis mellifera carnica.[78] 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. Although it is generally believed that insecticides have also depleted bee populations, particularly when used in excess of label directions, as bee pests and diseases (including American foulbrood and tracheal mites) are becoming resistant to medications, research in this regard has not been conclusive. A 2012 study of the effect of neonicotinoid-based insecticides showed "no effects observed in field studies at field-realistic dosages."[79] A new study in 2020 found that neonicotinoid insecticides affected the developmental stability of honey bees, particularly haploid males were more susceptible to neonicotinoids than diploid females.[80] The 2020 study also found that heterozygosity may play a key role in buffering insecticide exposure.[80]
In North America, various native milkweed species may be found with dead western honey bees stuck to their flowers. The non-native western honey bees are attracted to the flowers but are not adapted to their pollination mechanisms. The milkweed pollinium is collected when the tarsus (foot) of an insect falls into one of the flower's stigmatic slits as it obtains nectar from the flower's hood. If the insect is unable to remove its tarsus from the stigmatic slit it is likely to die due to predation or starvation/exhaustion. If the insect is able to escape with damaged or missing tarsi it may also be likely to die from its injuries. Western honey bees which escape with their tarsi intact may have their nectar gathering ability obstructed by parts of the pollinia being stuck to the bee's proboscis, resulting in starvation. The pollinia may also stick to the bee's tarsal claws, causing a lack of climbing ability and honey gathering which may result in expulsion from the colony leading to death. Native butterflies, moths, flies, beetles, bees and wasps are common milkweed visitors which are often able to escape without issue, though some species of Megachile, Halictus, Astata, Lucilia, Trichius, Pamphila and Scepsis have been found dead on the flowers. After removing over 140 dead bees from a patch of A. sullivantii, entomologist Charles Robertson quipped "... it seems that the flowers are better adapted to kill hive-bees than to produce fruit through their aid."[81]
Insect predators of western honey bees include the Asian giant hornet and other wasps, robber flies, dragonflies such as the green darner, some mantises, water striders and the European beewolf.
Arachnid predators of western honey bees include fishing spiders, lynx spiders, goldenrod spiders[82] and St. Andrew's cross spiders.
Reptile and amphibian predators of western honey bees include the black girdled lizard, anoles, and other lizards, and various anuran amphibians including the American toad, the American bullfrog and the wood frog.
Specialist bird predators of western honey bees include the bee-eaters; other birds that may take western honey bees include grackles, hummingbirds, tyrant flycatchers and the summer tanager. 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 as they emerge.[83]
Mammals that sometimes take western honey bees include giant armadillos, [84]both brown bears and black bears,[85] opossums, raccoons, skunks, the North American least shrew and the honey badger.
As an invasive species, feral western honey bees are a significant environmental problem in non-native areas. Imported bees may displace native bees and birds, and may also promote reproduction of invasive plants ignored by native pollinators.[86]
Honey bees are not native to the Americas, arriving with colonists in North America in the 18th century. 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.[87]
Honey bees have become an invasive species in the US, outcompeting native pollinators when resources are tight.[88] With an increased number of honey bees in a specific area due to beekeeping, domesticated bees and native wild bees often have to compete for the limited habitat and food sources available.[89] Western honey bees may become defensive in response to the seasonal arrival of competition from other colonies, particularly Africanized bees which may be on the offence and defence year round due to their tropical origin.[90] In the United Kingdom, honey bees are known to compete with native bumblebees such as Bombus hortorum, because they forage at the same sites. To resolve the issue and maximize both their total consumption during foraging, bumblebees forage early in the morning, while honey bees forage during the afternoon.[91]
Most flowering plants depend on specialized pollinators to efficiently fertilize them. Cucurbits, for example, are pollinated by squash bees that specifically visit the early-blooming male flowers before sunrise, when honey bees are inactive, and then return to pollinate the female flowers later in the day. Such symbiotic relationships also mean that the specialized pollinator will be covered mainly in its host's specific pollen.
The very generalized nature of the honey bee's nectar-gathering activities, potentially visiting dozens of different species in a single day, means that a flower visited by a honey bee will often get very little pollen from its own species. This diminished pollination can reduce the plant's ability to produce seeds, especially when the honey bees are squeezing out the native pollinators for a species, a problem occurring all over the United States because of honey bees and other invasive species.[92][93]
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.[94]
Apart from Apis mellifera, there are six other species in the genus Apis. These are Apis andreniformis, Apis cerana, Apis dorsata, Apis florea, Apis koschevnikovi, and Apis nigrocincta.[95] These species all originated in southern and southeastern Asia. Only Apis mellifera is thought to have originated in Europe, Asia, and Africa.[96]
.jpg)
.jpg)
{{cite journal}}: CS1 maint: multiple names: authors list (link)old-form url {{cite journal}}: CS1 maint: multiple names: authors list (link) {{cite journal}}: CS1 maint: multiple names: authors list (link) {{cite book}}: CS1 maint: multiple names: authors list (link) {{cite journal}}: CS1 maint: multiple names: authors list (link) The western honey bee or European honey bee (Apis mellifera) is the most common of the 7–12 species of honey bees worldwide. The genus name Apis is Latin for "bee", and mellifera is the Latin for "honey-bearing" or "honey carrying", referring to the species' production of honey.
Like all honey bee species, the western honey bee is eusocial, creating colonies with a single fertile female (or "queen"), many normally non-reproductive females or "workers", and a 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.
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. Western honey bees are threatened by pests and diseases, especially the Varroa mite and colony collapse disorder. There are indications that the species is rare, if not extinct in the wild in Europe and as of 2014, the western honey bee was assessed as "Data Deficient" on the IUCN Red List. 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, meaning self sustaining, without treatments or management.
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, especially via pollen, to assess non-target impacts of commercial pesticides.
