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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 Genus ( plural genera ) is a taxonomic rank used in the biological classification of living and fossil organisms as well as viruses. In the hierarchy of biological classification, genus comes above species and below family. In binomial nom ...
name ''Apis'' is
Latin Latin (, or , ) is a classical language belonging to the Italic languages, Italic branch of the Indo-European languages. Latin was originally a dialect spoken in the lower Tiber area (then known as Latium) around present-day Rome, but through ...
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 In modern parlance, a colony is a territory subject to a form of foreign rule. Though dominated by the foreign colonizers, colonies remain separate from the administration of the original country of the colonizers, the '' metropolitan state'' ...
with a single fertile female (or "
queen Queen or QUEEN may refer to: Monarchy * Queen regnant, a female monarch of a Kingdom ** List of queens regnant * Queen consort, the wife of a reigning king * Queen dowager, the widow of a king * Queen mother, a queen dowager who is the mother ...
"), 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 Pollination is the transfer of pollen from an Stamen, anther of a plant to the stigma (botany), stigma of a plant, later enabling fertilisation and the production of seeds, most often by an animal or by Anemophily, wind. Pollinating agents can ...
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 2019, the western honey bee is listed as "Extinct" on the
IUCN Red List The International Union for Conservation of Nature (IUCN) Red List of Threatened Species, also known as the IUCN Red List or Red Data Book, founded in 1964, is the world's most comprehensive inventory of the global conservation status of biolo ...
, while also acknowledging "Data Deficient". 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 A model organism (often shortened to model) 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 workin ...
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.


Distribution and habitat

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 eastern Asia.


Subspecies

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 31 recognized
subspecies In biological classification, subspecies is a rank below species, used for populations that live in different areas and vary in size, shape, or other physical characteristics ( morphology), but that can successfully interbreed. Not all specie ...
. 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. The subspecies are divided into four major branches, based on work by Ruttner and confirmed by
mitochondrial DNA Mitochondrial DNA (mtDNA or mDNA) is the DNA located in mitochondria, cellular organelles within eukaryotic cells that convert chemical energy from food into a form that cells can use, such as adenosine triphosphate (ATP). Mitochondrial D ...
analysis. African subspecies belong to branch A, northwestern European subspecies branch M, southwestern European subspecies branch C and Middle Eastern subspecies branch O.


Life cycle


Colony life cycle

Unlike most other bee species, western honey bees have
perennial A perennial plant or simply perennial is a plant that lives more than two years. The term ('' per-'' + '' -ennial'', "through the years") is often used to differentiate a plant from shorter-lived annuals and biennials. The term is also widel ...
colonies In modern parlance, a colony is a territory subject to a form of foreign rule. Though dominated by the foreign colonizers, colonies remain separate from the administration of the original country of the colonizers, the '' metropolitan state'' ...
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". 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 Nectar is a sugar-rich liquid produced by plants in glands called nectaries or nectarines, either within the flowers with which it attracts pollinating animals, or by extrafloral nectaries, which provide a nutrient source to animal mutualist ...
and
pollen Pollen is a powdery substance produced by seed plants. It consists of pollen grains (highly reduced microgametophytes), which produce male gametes (sperm cells). Pollen grains have a hard coat made of sporopollenin that protects the gametop ...
. In response to these favorable conditions, the hive creates one to two dozen new queens. Just as the
pupa A pupa ( la, pupa, "doll"; plural: ''pupae'') is the life stage of some insects undergoing transformation between immature and mature stages. Insects that go through a pupal stage are holometabolous: they go through four distinct stages in thei ...
l 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 all of her rivals, the new queen, the only fertile
female Female ( symbol: ♀) is the sex of an organism that produces the large non-motile ova (egg cells), the type of gamete (sex cell) that fuses with the male gamete during sexual reproduction. A female has larger gametes than a male. Fema ...
, 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, 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 Honey is a sweet and viscous substance made by several bees, the best-known of which are honey bees. Honey is made and stored to nourish bee colonies. Bees produce honey by gathering and then refining the sugary secretions of plants (primar ...
, 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.


Individual bee life cycle

Like other insects that undergo
complete metamorphosis Holometabolism, also called complete metamorphosis, is a form of insect development which includes four life stages: egg, larva, pupa, and imago (or adult). Holometabolism is a synapomorphic trait of all insects in the superorder Endopterygota. ...
, the western honey bee has four distinct life stages: egg,
larva A larva (; plural larvae ) is a distinct juvenile form many animals undergo before metamorphosis into adults. Animals with indirect development such as insects, amphibians, or cnidarians typically have a larval phase of their life cycle. ...
,
pupa A pupa ( la, pupa, "doll"; plural: ''pupae'') is the life stage of some insects undergoing transformation between immature and mature stages. Insects that go through a pupal stage are holometabolous: they go through four distinct stages in thei ...
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 The ovipositor is a tube-like organ used by some animals, especially insects, for the laying of eggs. In insects, an ovipositor consists of a maximum of three pairs of appendages. The details and morphology of the ovipositor vary, but typical ...
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 Humans (''Homo sapiens'') are the most abundant and widespread species of primate, characterized by bipedalism and exceptional cognitive skills due to a large and complex brain. This has enabled the development of advanced tools, cultu ...
or other
mammal Mammals () are a group of vertebrate animals constituting the class Mammalia (), characterized by the presence of mammary glands which in females produce milk for feeding (nursing) their young, a neocortex (a region of the brain), fur ...
. The stinger and its venom sac, with musculature and a ganglion allowing them to continue delivering
venom Venom or zootoxin is a type of toxin produced by an animal that is actively delivered through a wound by means of a bite, sting, or similar action. The toxin is delivered through a specially evolved ''venom apparatus'', such as fangs or a st ...
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 honey bee subspecies (''A. m. mellifera'') previously used for beekeeping indicate that a queen can live up to eight years. Because a queen's store of
sperm Sperm is the male reproductive cell, or gamete, in anisogamous forms of sexual reproduction (forms in which there is a larger, female reproductive cell and a smaller, male one). Animals produce motile sperm with a tail known as a flagellum, ...
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. Except for the queen, all of a colony's workers are replaced about every four months.


Social caste

Behavioral and physiological differences between castes and subcastes arise from phenotypic plasticity, which relies on
gene expression Gene expression is the process by which information from a gene is used in the synthesis of a functional gene product that enables it to produce end products, protein or non-coding RNA, and ultimately affect a phenotype, as the final effect. T ...
rather than heritable genotypic differences.


Queens

The queen bee is a fertile female, who, unlike workers (which are also female), has a fully developed
reproductive system The reproductive system of an organism, also known as the genital system, is the biological system made up of all the anatomical organs involved in sexual reproduction. Many non-living substances such as fluids, hormones, and pheromones are ...
. 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
flavonoid Flavonoids (or bioflavonoids; from the Latin word ''flavus'', meaning yellow, their color in nature) are a class of polyphenolic secondary metabolites found in plants, and thus commonly consumed in the diets of humans. Chemically, flavonoids ...
, during the first three days. After that, workers are switched to a diet of mixed pollen and nectar (often called "bee bread"), while queens continue to receive royal jelly. In the absence of flavonoid and the presence of a high-protein diet, queen bees develop a healthy reproductive system, 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: # The hive is filled with honey, leaving little room for new eggs. This will trigger a swarm, where the old queen will take about half the worker bees to found a new colony and leave the new queen with the other half of the workers to continue the old one. # The old queen begins to fail, which is thought to be demonstrated by a decrease in queen pheromones throughout the hive. This is known as supersedure, and at its end, the old queen is usually killed. # The old queen dies suddenly, a situation known as emergency supersedure. The worker bees find several eggs (or larvae) of the appropriate age range and try to develop them into queens. 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 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) 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 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

Workers are females produced by the queen that develop from fertilized,
diploid Ploidy () is the number of complete sets of chromosomes in a cell, and hence the number of possible alleles for autosomal and pseudoautosomal genes. Sets of chromosomes refer to the number of maternal and paternal chromosome copies, respectiv ...
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

Drones are the colony's
male Male (symbol: ♂) is the sex of an organism that produces the gamete (sex cell) known as sperm, which fuses with the larger female gamete, or ovum, in the process of fertilization. A male organism cannot reproduce sexually without access to ...
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 Fertilisation or fertilization (see spelling differences), also known as generative fertilisation, syngamy and impregnation, is the fusion of gametes to give rise to a new individual organism or offspring and initiate its development. Pro ...
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 Ploidy () is the number of complete sets of chromosomes in a cell, and hence the number of possible alleles for autosomal and pseudoautosomal genes. Sets of chromosomes refer to the number of maternal and paternal chromosome copies, respective ...
(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 Thermoregulation is the ability of an organism to keep its body temperature within certain boundaries, even when the surrounding temperature is very different. A thermoconforming organism, by contrast, simply adopts the surrounding temperatur ...
. Drones are typically located near the center of hive clusters for unclear reasons. It is postulated that it is to maintain
sperm viability Semen quality is a measure of male fertility, a measure of the ability of sperm in semen to accomplish fertilization. Semen quality involves both sperm quantity and quality Cryptorchidism, hypospadias, testicular cancer and poor semen quality m ...
, 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 their ability to contribute declines.


Queen–worker conflict

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. This pits the worker against the queen and other workers, who try to maximize their
reproductive fitness Fitness (often denoted w or ω in population genetics models) is the quantitative representation of individual reproductive success. It is also equal to the average contribution to the gene pool of the next generation, made by the same individua ...
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. 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. 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. 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. According to the strategy of
kin selection Kin selection is the evolutionary strategy that favours the reproductive success of an organism's relatives, even when at a cost to the organism's own survival and reproduction. Kin altruism can look like altruistic behaviour whose evolution ...
, 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 ''Related'' is an American comedy-drama television series that aired on The WB from October 5, 2005, to March 20, 2006. It revolves around the lives of four close-knit sisters of Italian descent, raised in Brooklyn and living in Manhattan. The ...
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 Hymenoptera is a large order of insects, comprising the sawflies, wasps, bees, and ants. Over 150,000 living species of Hymenoptera have been described, in addition to over 2,000 extinct ones. Many of the species are parasitic. Females typic ...
.


Behavior


Thermoregulation

The western honey bee needs an internal body temperature of 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 . Western honey bees can forage over a 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 An evaporative cooler (also known as evaporative air conditioner, swamp cooler, swamp box, desert cooler and wet air cooler) is a device that cools air through the evaporation of water. Evaporative cooling differs from other air conditioning ...
from regurgitated honey-sac contents. Body temperatures differ, depending on caste and expected foraging rewards. The optimal air temperature for foraging is . 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 . Below bees are immobile, and above their activity slows. Western honey bees can tolerate temperatures up to for short periods.


Communication

Western honey bee behavior has been extensively studied. Karl von Frisch, who received the 1973
Nobel Prize in Physiology or Medicine The Nobel Prize in Physiology or Medicine is awarded yearly by the Nobel Assembly at the Karolinska Institute for outstanding discoveries in physiology or medicine. The Nobel Prize is not a single prize, but five separate prizes that, accordi ...
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. 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 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. Therefore, western honey bees seem to have evolved a better means of conveying information than their common ancestors with the dwarf honey bee. 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 Standard anatomical terms of location are used to unambiguously describe the anatomy of animals, including humans. The terms, typically derived from Latin or Greek roots, describe something in its standard anatomical position. This position prov ...
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

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 Defense or defence may refer to: Tactical, martial, and political acts or groups * Defense (military), forces primarily intended for warfare * Civil defense, the organizing of civilians to deal with emergencies or enemy attacks * Defense indus ...
, orientation, kin and
colony In modern parlance, a colony is a territory subject to a form of foreign rule. Though dominated by the foreign colonizers, colonies remain separate from the administration of the original country of the colonizers, the '' metropolitan state' ...
recognition, food production and integrating colony activities.Free, John B., ''Pheromones of social bees''. Ithaca, N.Y.: Comstock, 1987.Blum, M.S. 1992. Honey bee pheromones in The Hive and the Honey Bee, revised edition (Dadant and Sons: Hamilton, Illinois), pages 385–389.


Sociality

There is some degree of variability of sociality between individuals. Like a great many other social insects, ''A. mellifera'' engages in trophallaxis. 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. There is less inter-individual variation than found in humans however, possibly reflecting the higher genetic relatedness between hivemates.


Domestication

Humans have been collecting honey from western honey bees for thousands of years, with evidence in the form of
rock art In archaeology, rock art is human-made markings placed on natural surfaces, typically vertical stone surfaces. A high proportion of surviving historic and prehistoric rock art is found in caves or partly enclosed rock shelters; this type also m ...
found in France and Spain, dating to around 7,000 BCE. The western honey bee is one of the few
invertebrate Invertebrates are a paraphyletic group of animals that neither possess nor develop a vertebral column (commonly known as a ''backbone'' or ''spine''), derived from the notochord. This is a grouping including all animals apart from the chorda ...
animals to have been domesticated. Bees were likely first domesticated in ancient Egypt, where
tomb A tomb ( grc-gre, τύμβος ''tumbos'') is a repository for the remains of the dead. It is generally any structurally enclosed interment space or burial chamber, of varying sizes. Placing a corpse into a tomb can be called ''immureme ...
paintings depict beekeeping, before 2600BC. Europeans brought bees to North America in 1622.
Beekeeper A beekeeper is a person who keeps honey bees. Beekeepers are also called honey farmers, apiarists, or less commonly, apiculturists (both from the Latin '' apis'', bee; cf. apiary). The term beekeeper refers to a person who keeps honey bees i ...
s have selected western honey bees for several desirable features: * the ability of a colony to survive periods with little food * the ability of a colony to survive cold weather * resistance to disease * increased honey production * reduced aggressiveness * reduced tendency to swarm * reduced nest building * easy pacification with smoke 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. Authors do not agree on whether this degree of
artificial selection Selective breeding (also called artificial selection) is the process by which humans use animal breeding and plant breeding to selectively develop particular phenotypic traits (characteristics) by choosing which typically animal or plant ma ...
constitutes genuine domestication. In 1603,
John Guillim John Guillim (c. 1565 – 7 May 1621) of Minsterworth, Gloucestershire, was an antiquarian and officer of arms at the College of Arms in London. He is best remembered for his monumental work on heraldry, ''A Display of Heraldry'', first pu ...
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 western 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 western honey bees, at best, partial. 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". 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. 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".


Beekeeping

The western honey bee is a colonial
insect Insects (from Latin ') are pancrustacean hexapod invertebrates of the class Insecta. They are the largest group within the arthropod phylum. Insects have a chitinous exoskeleton, a three-part body ( head, thorax and abdomen), three pa ...
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 Honey is a sweet and viscous substance made by several bees, the best-known of which are honey bees. Honey is made and stored to nourish bee colonies. Bees produce honey by gathering and then refining the sugary secretions of plants (primar ...
, 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 Pollen is a powdery substance produced by seed plants. It consists of pollen grains (highly reduced microgametophytes), which produce male gametes (sperm cells). Pollen grains have a hard coat made of sporopollenin that protects the gametop ...
which, after being processed to bee bread, supplies protein and fat for the
bee brood 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 ...
to grow. Centuries of
selective breeding Selective breeding (also called artificial selection) is the process by which humans use animal breeding and plant breeding to selectively develop particular phenotypic traits (characteristics) by choosing which typically animal or plant ...
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:
skep A beehive is an enclosed 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 ...
s, Langstroth hives,
top-bar hive A top-bar hive is a single-story frameless beehive in which the comb hangs from removable bars. The bars form a continuous roof over the comb, whereas the frames in most current hives allow space for bees to move up or down between boxes. Hives t ...
s, box hives, log gums, D. E. hives, 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 Dysentery (UK pronunciation: , US: ), historically known as the bloody flux, is a type of gastroenteritis that results in bloody diarrhea. Other symptoms may include fever, abdominal pain, and a feeling of incomplete defecation. Complications ...
and causes an excessive buildup of
carbon dioxide Carbon dioxide ( chemical formula ) is a chemical compound made up of molecules that each have one carbon atom covalently double bonded to two oxygen atoms. It is found in the gas state at room temperature. In the air, carbon dioxide is t ...
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.


Products


Honey bees

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.


Pollination

The western honey bee is an important
pollinator 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. Insects are the m ...
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. 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 California is a state in the Western United States, located along the Pacific Coast. With nearly 39.2million residents across a total area of approximately , it is the most populous U.S. state and the 3rd largest by area. It is also the m ...
,
almond The almond (''Prunus amygdalus'', syn. ''Prunus dulcis'') is a species of tree native to Iran and surrounding countries, including the Levant. The almond is also the name of the edible and widely cultivated seed of this tree. Within the genu ...
pollination occurs in February, early in the
growing season A season is a division of the year marked by changes in weather, ecology, and the amount of daylight. The growing season is that portion of the year in which local conditions (i.e. rainfall, temperature, daylight) permit normal plant growth. Whi ...
before local hives have built up their populations. Almond
orchard An orchard is an intentional plantation of trees or shrubs that is maintained for food production. Orchards comprise fruit- or nut-producing trees which are generally grown for commercial production. Orchards are also sometimes a feature of ...
s require two hives per acre, or 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 New York most commonly refers to: * New York City, the most populous city in the United States, located in the state of New York * New York (state), a state in the northeastern United States New York may also refer to: Film and television * '' ...
,
Michigan Michigan () is a state in the Great Lakes region of the upper Midwestern United States. With a population of nearly 10.12 million and an area of nearly , Michigan is the 10th-largest state by population, the 11th-largest by area, and t ...
, and
Washington Washington commonly refers to: * Washington (state), United States * Washington, D.C., the capital of the United States ** A metonym for the federal government of the United States ** Washington metropolitan area, the metropolitan area centered o ...
. 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. In 2000, Drs.
Roger Morse Roger A. Morse, Ph.D. (July 5, 1927 - May 12, 2000) was an American bee biologist who taught many beekeepers both the rudiments and the finer practices, through his research and publications. During his long career, three new parasites of the hone ...
and Nicholas Calderone of
Cornell University Cornell University is a private statutory land-grant research university based in Ithaca, New York. It is a member of the Ivy League. Founded in 1865 by Ezra Cornell and Andrew Dickson White, Cornell was founded with the intention to tea ...
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.


Honey

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 Glucose is a simple sugar with the molecular formula . Glucose is overall the most abundant monosaccharide, a subcategory of carbohydrates. Glucose is mainly made by plants and most algae during photosynthesis from water and carbon dioxide, u ...
and
fructose Fructose, or fruit sugar, is a ketonic simple sugar found in many plants, where it is often bonded to glucose to form the disaccharide sucrose. It is one of the three dietary monosaccharides, along with glucose and galactose, that are absorb ...
. 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 ''Staphylococcus aureus'' is a Gram-positive spherically shaped bacterium, a member of the Bacillota, and is a usual member of the microbiota of the body, frequently found in the upper respiratory tract and on the skin. It is often posit ...
''. 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 Nectar is a sugar-rich liquid produced by plants in glands called nectaries or nectarines, either within the flowers with which it attracts pollinating animals, or by extrafloral nectaries, which provide a nutrient source to animal mutualist ...
, a clear liquid consisting of nearly 80 percent water and complex sugars. The collecting bees store the nectar in a second
stomach The stomach is a muscular, hollow organ in the gastrointestinal tract of humans and many other animals, including several invertebrates. The stomach has a dilated structure and functions as a vital organ in the digestive system. The stomach i ...
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.


Beeswax

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
candle A candle is an ignitable wick embedded in wax, or another flammable solid substance such as tallow, that provides light, and in some cases, a fragrance. A candle can also provide heat or a method of keeping time. A person who makes candle ...
s and
seals Seals may refer to: * Pinniped, a diverse group of semi-aquatic marine mammals, many of which are commonly called seals, particularly: ** Earless seal, or "true seal" ** Fur seal * Seal (emblem), a device to impress an emblem, used as a means of a ...
.


Bee bread

Bees collect
pollen Pollen is a powdery substance produced by seed plants. It consists of pollen grains (highly reduced microgametophytes), which produce male gametes (sperm cells). Pollen grains have a hard coat made of sporopollenin that protects the gametop ...
in a
pollen basket The pollen basket or corbicula (plural corbiculae) is part of the tibia on the hind legs of certain species of bees. They use the structure in harvesting pollen and carrying it to the nest or hive. Other species of bees have scopae instead. Ety ...
and carry it back to the hive where, after undergoing
fermentation Fermentation is a metabolic process that produces chemical changes in organic substrates through the action of enzymes. In biochemistry, it is narrowly defined as the extraction of energy from carbohydrates in the absence of oxygen. In food p ...
and turning into bee bread, it becomes a
protein Proteins are large biomolecules and macromolecules that comprise one or more long chains of amino acid residues. Proteins perform a vast array of functions within organisms, including catalysing metabolic reactions, DNA replication, res ...
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 Allergies, also known as allergic diseases, refer a number of conditions caused by the hypersensitivity of the immune system to typically harmless substances in the environment. These diseases include hay fever, food allergies, atopic derm ...
in susceptible individuals.


Bee brood

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

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

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.


Genome

Female bees are
diploid Ploidy () is the number of complete sets of chromosomes in a cell, and hence the number of possible alleles for autosomal and pseudoautosomal genes. Sets of chromosomes refer to the number of maternal and paternal chromosome copies, respectiv ...
and have 32
chromosomes A chromosome is a long DNA molecule with part or all of the genetic material of an organism. In most chromosomes the very long thin DNA fibers are coated with packaging proteins; in eukaryotic cells the most important of these proteins ar ...
, whereas males are
haploid Ploidy () is the number of complete sets of chromosomes in a cell, and hence the number of possible alleles for autosomal and pseudoautosomal genes. Sets of chromosomes refer to the number of maternal and paternal chromosome copies, respective ...
and have only 16. As of October 28, 2006, the
Honey Bee Genome Sequencing Consortium The Honey Bee Genome Sequencing Consortium is an international collaborative group of genomics scientists, scientific organisations and universities trying to decipher the genome sequences of the honey bee (''Apis mellifera''). It was formed in 200 ...
fully sequenced and analyzed the
genome In the fields of molecular biology and genetics, a genome is all the genetic information of an organism. It consists of nucleotide sequences of DNA (or RNA in RNA viruses). The nuclear genome includes protein-coding genes and non-coding ...
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. Scientists have found that
gene In biology, the word gene (from , ; "...Wilhelm Johannsen coined the word gene to describe the Mendelian units of heredity..." meaning ''generation'' or ''birth'' or ''gender'') can have several different meanings. The Mendelian gene is a b ...
s related to smell outnumber those for taste, and the European honey bee has fewer genes regulating
immunity Immunity may refer to: Medicine * Immunity (medical), resistance of an organism to infection or disease * ''Immunity'' (journal), a scientific journal published by Cell Press Biology * Immune system Engineering * Radiofrequence immunity de ...
than the fruit fly and the mosquito. The genome sequence also revealed that several groups of genes, particularly those related to
circadian rhythm A circadian rhythm (), or circadian cycle, is a natural, internal process that regulates the sleep–wake cycle and repeats roughly every 24 hours. It can refer to any process that originates within an organism (i.e., endogenous) and responds to ...
, 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 DNA methylation is a biological process by which methyl groups are added to the DNA molecule. Methylation can change the activity of a DNA segment without changing the sequence. When located in a gene promoter, DNA methylation typically acts ...
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 In biology, epigenetics is the study of stable phenotypic changes (known as ''marks'') that do not involve alterations in the DNA sequence. The Greek prefix '' epi-'' ( "over, outside of, around") in ''epigenetics'' implies features that are ...
to study gene expression and
regulation Regulation is the management of complex systems according to a set of rules and trends. In systems theory, these types of rules exist in various fields of biology and society, but the term has slightly different meanings according to context. ...
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 element A transposable element (TE, transposon, or jumping gene) is a nucleic acid sequence in DNA that can change its position within a genome, sometimes creating or reversing mutations and altering the cell's genetic identity and genome size. Transp ...
s, although they were present in the evolutionary past (remains and fossils have been found) and evolved more slowly than those in fly species. Since 2018 a new version of the honey bee genome is available on NCBI (Amel_HAv3.1, BioProject ID: PRJNA471592). 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 Hymenoptera is a large order of insects, comprising the sawflies, wasps, bees, and ants. Over 150,000 living species of Hymenoptera have been described, in addition to over 2,000 extinct ones. Many of the species are parasitic. Females typic ...
order compared to most other eukaryotic species except
fungi A fungus ( : fungi or funguses) is any member of the group of eukaryotic organisms that includes microorganisms such as yeasts and molds, as well as the more familiar mushrooms. These organisms are classified as a kingdom, separately fr ...
and
protozoa Protozoa (singular: protozoan or protozoon; alternative plural: protozoans) are a group of single-celled eukaryotes, either free-living or parasitic, that feed on organic matter such as other microorganisms or organic tissues and debris. Histo ...
. 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.


Hazards and survival


Parasites, diseases and pesticides

Western honey bee populations face threats to their survival increasing interests into other pollinator species, like the common eastern bumblebee.
North America North America is a continent in the Northern Hemisphere and almost entirely within the Western Hemisphere. It is bordered to the north by the Arctic Ocean, to the east by the Atlantic Ocean, to the southeast by South America and th ...
n 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. Some subspecies of ''Apis mellifera'' show naturally
varroa sensitive hygiene Varroa sensitive hygiene (VSH) is a behavioral trait of honey bees ''(Apis mellifera)'' in which bees detect and remove bee pupae that are infested by the parasitic mite ''Varroa destructor. V. destructor'' is considered to be the most dangerous p ...
, for example
Apis mellifera lamarckii Lamarck's honey bee or the Egyptian honey bee, ''Apis mellifera lamarckii'', is a subspecies of honey bee occurring in a narrow range along the Egyptian Nile Valley of Egypt and Sudan, named after Jean-Baptiste Lamarck and is considered the firs ...
and
Apis mellifera carnica The Carniolan honey bee (''Apis mellifera carnica'', Pollmann) is a subspecies of the western honey bee. The Carniolan honey bee is native to Slovenia, southern Austria, and parts of Croatia, Bosnia and Herzegovina, Montenegro, Serbia, Hungary, ...
. 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 Natural selection is the differential survival and reproduction of individuals due to differences in phenotype. It is a key mechanism of evolution, the change in the heritable traits characteristic of a population over generations. Cha ...
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 American foulbrood (AFB, ''Histolysis infectiosa perniciosa larvae apium'', ''Pestis americana larvae apium''), caused by the spore-forming bacterium '' Paenibacillus larvae'' (reclassified as one species without subspecies differentiation in 2 ...
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." 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. The 2020 study also found that heterozygosity may play a key role in buffering insecticide exposure.


Milkweed

In North America, various native
milkweed ''Asclepias'' is a genus of herbaceous, perennial, flowering plants known as milkweeds, named for their latex, a milky substance containing cardiac glycosides termed cardenolides, exuded where cells are damaged. Most species are toxic to hum ...
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 A pollinium (plural pollinia) is a coherent mass of pollen grains in a plant that are the product of only one anther, but are transferred, during pollination, as a single unit. This is regularly seen in plants such as orchids and many species of mil ...
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 A proboscis () is an elongated appendage from the head of an animal, either a vertebrate or an invertebrate. In invertebrates, the term usually refers to tubular mouthparts used for feeding and sucking. In vertebrates, a proboscis is an elong ...
, 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 The genus ''Halictus'' is a large assemblage of bee species in the family Halictidae. The genus is divided into 15 subgenera, some of dubious monophyly, containing over 200 species, primarily in the Northern Hemisphere (a few species occur in ...
'', '' Astata'', '' Lucilia'', ''
Trichius The bee beetles are scarab beetles of the subfamily Cetoniinae. They have hairy sides of the elytra like their relatives, and the upper sides of the elytra are usually yellow with prominent black blotches which form incomplete bands. This, and t ...
'', '' 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."


Predators

Insect predators of western honey bees include the Asian giant hornet and other
wasp A wasp is any insect of the narrow-waisted suborder Apocrita of the order Hymenoptera which is neither a bee nor an ant; this excludes the broad-waisted sawflies (Symphyta), which look somewhat like wasps, but are in a separate suborder ...
s,
robber flies The Asilidae are the robber fly family, also called assassin flies. They are powerfully built, bristly flies with a short, stout proboscis enclosing the sharp, sucking hypopharynx. The name "robber flies" reflects their notoriously aggressive pr ...
, dragonflies such as the
green darner The green darner or common green darner (''Anax junius''), after its resemblance to a darning needle, is a species of dragonfly in the family Aeshnidae. One of the most common and abundant species throughout North America, it also ranges south t ...
, some mantises, water striders and the
European beewolf The European beewolf (''Philanthus triangulum''), also known as the bee-killer wasp or the bee-eating philanthus (from the now obsolete synonym ''Philanthus apivorus''), is a solitary wasp that lives in the Western Palearctic and Afrotropics. Alt ...
. Arachnid predators of western honey bees include fishing spiders, lynx spiders, goldenrod spiders 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
amphibian Amphibians are four-limbed and ectothermic vertebrates of the class Amphibia. All living amphibians belong to the group Lissamphibia. They inhabit a wide variety of habitats, with most species living within terrestrial, fossorial, arbo ...
s including the American toad, the
American bullfrog The American bullfrog (''Lithobates catesbeianus''), often simply known as the bullfrog in Canada and the United States, is a large true frog native to eastern North America. It typically inhabits large permanent water bodies such as swamps, p ...
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,
hummingbird Hummingbirds are birds native to the Americas and comprise the Family (biology), biological family Trochilidae. With about 361 species and 113 genus, genera, they occur from Alaska to Tierra del Fuego, but the vast majority of the species are ...
s, 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. Mammals that sometimes take western honey bees include
giant armadillo The giant armadillo (''Priodontes maximus''), colloquially ''tatu-canastra'', ''tatou'', ''ocarro'' or ''tatú carreta'', is the largest living species of armadillo (although their extinct relatives, the glyptodonts, were much larger). It live ...
s, both
brown bear The brown bear (''Ursus arctos'') is a large bear species found across Eurasia and North America. In North America, the populations of brown bears are called grizzly bears, while the subspecies that inhabits the Kodiak Islands of Alaska is k ...
s and black bears,
opossum Opossums () are members of the marsupial order Didelphimorphia () endemic to the Americas. The largest order of marsupials in the Western Hemisphere, it comprises 93 species in 18 genera. Opossums originated in South America and entered No ...
s, raccoons,
skunk Skunks are mammals in the family Mephitidae. They are known for their ability to spray a liquid with a strong, unpleasant scent from their anal glands. Different species of skunk vary in appearance from black-and-white to brown, cream or gin ...
s, the
North American least shrew The North American least shrew (''Cryptotis parvus'') is one of the smallest mammals, growing to be only up to 3 inches long. It has a long pointed snout and a tail never more than twice the length of its hind foot. The dense fur coat is ei ...
and the honey badger.


As an environmental threat

As an
invasive species An invasive species otherwise known as an alien is an introduced organism that becomes overpopulated and harms its new environment. Although most introduced species are neutral or beneficial with respect to other species, invasive species adv ...
, 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. Honey bees are not native to
the Americas The Americas, which are sometimes collectively called America, are a landmass comprising the totality of North and South America. The Americas make up most of the land in Earth's Western Hemisphere and comprise the New World. Along with th ...
, arriving with colonists in North America in the 18th century.
Thomas Jefferson Thomas Jefferson (April 13, 1743 – July 4, 1826) was an American statesman, diplomat, lawyer, architect, philosopher, and Founding Fathers of the United States, Founding Father who served as the third president of the United States from 18 ...
mentioned this in his ''
Notes on the State of Virginia ''Notes on the State of Virginia'' (1785) is a book written by the American statesman, philosopher, and planter Thomas Jefferson. He completed the first version in 1781 and updated and enlarged the book in 1782 and 1783. It originated in Jeffers ...
'': Honey bees have become an invasive species in the US, outcompeting native pollinators when resources are tight. 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. 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. In the
United Kingdom The United Kingdom of Great Britain and Northern Ireland, commonly known as the United Kingdom (UK) or Britain, is a country in Europe, off the north-western coast of the continental mainland. It comprises England, Scotland, Wales and ...
, honey bees are known to compete with native
bumblebee A bumblebee (or bumble bee, bumble-bee, or humble-bee) is any of over 250 species in the genus ''Bombus'', part of Apidae, one of the bee families. This genus is the only Extant taxon, extant group in the tribe Bombini, though a few extinct r ...
s 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. Most flowering plants depend on specialized pollinators to efficiently fertilize them.
Cucurbits The Cucurbitaceae, also called cucurbits or the gourd family, are a plant family consisting of about 965 species in around 95 genera, of which the most important to humans are: *''Cucurbita'' – squash, pumpkin, zucchini, some gourds *'' Lagena ...
, 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. 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 ''Melastoma affine'', also known by the common names blue tongue or native lassiandra, is a shrub of the family Melastomataceae. Distributed in tropical and sub-tropical forests of India, South-east Asia and Australia, it is a plant of rainfo ...
'', a plant with pore anthers, by robbing its stigmas of previously deposited pollen.


Close relatives

Apart from ''Apis mellifera'', there are six other species in the genus ''Apis''. These are ''
Apis andreniformis ''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. ''A. andreniformis'' was the fifth honey bee species to be described ...
'', '' Apis cerana'', '' Apis dorsata'', ''
Apis florea The dwarf honey bee (or red 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 ...
'', ''
Apis koschevnikovi ''Apis koschevnikovi'', Koschevnikov's honey bee, is a species of honey bee which inhabits Malaysian and Indonesian Borneo, where it lives sympatrically with other honey bee species such as '' Apis cerana'' (specifically '' A. c. nuluensis''). ...
'', and ''
Apis nigrocincta ''Apis nigrocincta'' is a species of honey bee that inhabits the Philippine island of Mindanao as well as the Indonesian islands of Sangihe and Sulawesi. The species is known to have queens with the highest mating frequencies of any species of ...
''. These species all originated in southern and southeastern Asia. Only ''Apis mellifera'' is thought to have originated in Europe, Asia, and Africa.


See also

* Apitherapy *
Bee bearding Bee bearding is the practice of wearing several thousand honey bees on the face, usually as a sideshow-type demonstration at agricultural shows. Hive bees are attracted into position by a queen in a small cage worn under the chin. History Thoug ...
*
Beeline A bee line is an idiom for the shortest route or a straight line between two points (see "as the crow flies"): bee line, bee-line, or beeline may also refer to: Brands and enterprises * Beeline (brand), a telecommunications brand by VimpelCom L ...
*
Characteristics of common wasps and bees While observers can easily confuse common wasps and bees at a distance or without close observation, there are many different characteristics of large bee Bees are winged insects closely related to wasps and ants, known for their roles in p ...
* Worker policing


References


Bibliography

* A. I. Root's The ABC and XYZ of Beekeeping
Molecular confirmation of a fourth lineage in honeybees from the Near East
Apidologie 31 (2000) 167–180, accessed Oct 2005 *Biesmeijer, Jacobus. "The Occurrence and Context of the Shaking Signal in Honey Bees (''Apis mellifera'') Exploiting Natural Food Sources". ''
Ethology Ethology is the scientific study of animal behaviour, usually with a focus on behaviour under natural conditions, and viewing behaviour as an evolutionarily adaptive trait. Behaviourism as a term also describes the scientific and objecti ...
''. 2003. * *Lindauer, Martin. "Communication among social bees". Harvard University Press 1971. * *Schneider, S. S., P. K. Visscher, Camazine, S. "Vibration Signal Behavior of Waggle-dancers in Swarms of the Honey Bee, ''Apis mellifera'' (Hymenoptera: Apidae). ''Ethology''. 1998.


External links


FAO: Beekeeping explained
* ttp://entnemdept.ufl.edu/creatures/MISC/BEES/euro_honey_bee.htm IFAS: ''Apis mellifera''
Sound recordings of ''Apis mellifera'' at BioAcoustica
* {{DEFAULTSORT:Western Honey Bee Apis (genus) Beekeeping Hymenoptera of Asia Hymenoptera of Africa Hymenoptera of Europe Insects of the Middle East Insects of North Africa Insects described in 1758 Taxa named by Carl Linnaeus Symbols of Georgia (U.S. state) Symbols of South Dakota Symbols of Utah Symbols of Vermont Symbols of Wisconsin Symbols of West Virginia Vermont culture Articles containing video clips Hymenoptera of New Zealand Symbols of Kentucky Symbols of Mississippi Taxobox binomials not recognized by IUCN