Honey bee
A honey bee queen, once she has mated, can lay up to 2,000 eggs in a single day. Around her may live tens of thousands of workers, and none of them survives alone. The honey bee is a eusocial flying insect of the genus Apis, which means its colony works as one tightly organized unit. Its family, Apidae, is the largest family of bees.
Roughly 20,000 species of bee are known, yet honey bees are the ones most familiar to people. They build perennial nests inside cavities, using hexagonal cells of wax that they secrete themselves. They also regurgitate digested carbohydrates and store them as honey. That store makes a hive a prized target for honey badgers, bears and human hunter-gatherers.
How did a bee native to Afro-Eurasia end up in the Americas and Australia? How does a colony decide which bees work, which mate and which are sent away? And is it true that humanity would starve without them? The answers begin with a surprisingly small family tree.
Eight living species of honey bee are recognized today, divided into 43 subspecies in total. Older classifications have counted anywhere from seven to 11 species. Honey bees are the only living members of the tribe Apini. Three clades hold them all. Micrapis contains the dwarf honey bees, Megapis the giant honey bees, and Apis proper the western honey bee and its close relatives.
South and Southeast Asia, including the Philippines, appears to be where honey bees began. Every living species except Apis mellifera is native there, and the two earliest-diverging lineages, Apis florea and Apis andreniformis, still live there. The oldest Apis fossils come from European deposits at the Eocene-Oligocene boundary, about 34 million years ago. That does not make Europe the homeland. Few fossil deposits are known from South Asia, and fewer still have been studied closely.
Apis florea and Apis andreniformis are the dwarf honey bees of southern and southeastern Asia. They hang very small, exposed nests in trees and shrubs. Their stings often cannot pierce human skin, so people can handle a hive or a swarm with little protection. Honey usually comes from A. florea, which is more widespread, because A. andreniformis is considerably more aggressive.
The giant honey bees are another matter. Apis dorsata, widespread across most of South and Southeast Asia, builds one or a few exposed combs on high tree limbs, on cliffs and sometimes on buildings. Human honey hunters periodically rob these colonies, and a provoked colony can sting a person to death. Apis laboriosa, the Himalayan giant honey bee, is the largest living honey bee. It looks much like its cousin, but behavioral adaptations let it nest in the open at high altitudes despite low temperatures.
Koschevnikov's honey bee, Apis koschevnikovi, is often called the red bee of Sabah. The name only half fits. The bee is pale reddish in Sabah State, Borneo, but a dark, coppery color on the Malay Peninsula and in Sumatra.
Of all eight species, only two have been truly domesticated: Apis mellifera and Apis cerana. A. mellifera has been cultivated since at least the building of the Egyptian pyramids, and it alone has been moved widely beyond its native range.
Eastern tropical Africa seems to be where Apis mellifera began. From there it spread into Europe and eastward into Asia as far as the Tian Shan range. Its split from its eastern relatives came only in the Late Miocene. One hypothesis blames desertification in the Middle East. Drying there thinned the food plants and the trees that offered nest sites, and gene flow between the African group and the tropical Asian group eventually stopped.
An Early Pleistocene radiation, shaped by climate and habitat shifts during the last ice age, probably produced the many subspecies. Human management has muddied the picture. People have hybridized and moved this bee for many millennia, which apparently sped up its evolution and scrambled the DNA evidence. Little of substance can now be said about how many of the subspecies relate to one another.
In 1622, European colonists brought the German honey bee, A. m. mellifera, to the Americas. The Italian honey bee, A. m. ligustica, and others came later. Before that, no Apis species had lived in the New World in human times, though indigenous peoples kept and traded other native bees. The western honey bee reached South America in the early 16th century, North America in the early 17th century and Australia in the early 19th century.
Escaped swarms, called wild but actually feral, ran ahead of the colonists as far as the Great Plains. The Rocky Mountains stopped them, because honey bees did not cross on their own. Mormon pioneers carried them to Utah in the late 1840s, and ships took them to California in the early 1850s.
Africanized honey bees, nicknamed killer bees, began with an accident in Brazil. They are hybrids of European stock and the African subspecies A. m. scutellata, and they escaped from quarantine. They are often more aggressive than European bees and store less surplus honey. In exchange, they resist disease better and forage better. They have spread to North America and count as a pest in some regions, but they overwinter poorly, so colder northern areas rarely see them.
The original breeding experiment that brought African bees to Brazil continues, though not as planned. New hybrid strains combine tolerance of tropical conditions with good yields, and Brazilian beekeepers like them.
Honey bees divide into three castes: drones, workers and queens. Drones are male, while workers and queens are female. The queen decides which she lays. Using a sperm-storing organ called the spermatheca, she can fertilize an egg or leave it unfertilized, usually depending on the cell she is laying into. Unfertilized eggs become drones, which are haploid, with a single set of chromosomes. Fertilized eggs become diploid females.
Development times differ by caste. A queen takes 16 days, a worker 21 days and a drone 24 days. What separates a queen from a worker is diet. Workers feed a single female larva nothing but royal jelly, in an oversized cell. The result is a larger bee with working ovaries and a spermatheca. Her sting has no barbs, and she lacks the glands that make beeswax.
Work in a colony of as many as 60,000 workers follows a schedule set by age. After eating her way out of her capped cell, a worker cleans it. She then feeds brood, receives nectar, cleans the hive, stands guard and finally forages. Some take on special jobs such as undertaking, which means carrying dead nestmates out of the hive.
Drones number as many as 500 per hive and exist mainly to reproduce. They appear from summer through autumn. Their large eyes help them spot queens during mating flights. They have no stinger and do not defend the hive. When winter turns the hive's focus to warmth and food, the drones are expelled.
A virgin queen flies to a drone congregation area to mate, and she never mates with drones from her own colony. Queens ready to mate make between 1 and 6 nuptial flights over consecutive days. The Philippine honey bee holds the documented record, with queens mating with 42-69 drones. Each drone dies in or soon after the act. Because it all happens in free flight, the process is hard to observe. When workers raise new queens, the young queens fight one another until one remains.
Genetics sets up a quiet conflict inside the hive. A laying worker shares half her genes with her own son but only one-quarter with her brothers, the queen's sons. The queen is half related to her sons and one-quarter related to the sons of workers. Other workers, who are more closely related to the queen's sons, patrol the hive and remove worker-laid eggs. This is called worker policing.
Policing weakens if a queen mates only once, because the genetic gap between her sons and other workers' sons shrinks. Experiments confirm that higher mating rates go with more policing in many social hymenoptera. In very rare cases, workers subvert the system. This is anarchic syndrome: their eggs are removed at a lower rate and more drones result. The hive becomes less productive, but the laying worker's reproductive fitness rises.
When a queen dies or is removed, a significant share of workers activate their ovaries. The hive produces a final batch of drones before it collapses.
Every living species of Apis performs some form of the waggle dance, though the details vary from species to species. Through it, workers report the distance and direction of a target, usually flowers or water. When the food is very close, a less specific round dance serves instead. Tremble dances do something different, recruiting receiver bees to collect nectar from foragers that have just returned.
The two smallest species, Apis andreniformis and A. florea, dance differently from the rest. They dance on the upper surface of a comb that is horizontal rather than vertical. Their dancers orient the dance to the actual compass direction of the resource. Chemicals and odors carry much of the rest of the colony's conversation, as is common in insects.
Most bees start a colony with a solitary queen, but honey bees send out groups called swarms. A swarm is a mated queen with a large contingent of workers. Scouts choose the nest site beforehand, and the same dance language is used to report its location and quality. On arrival, the swarm immediately builds new wax comb and begins raising a new worker brood.
No other living bee genus founds nests this way. Several groups of vespid wasps do swarm, sometimes with multiple queens. Stingless bees start nests with large numbers of workers, but they finish the nest before a queen is escorted there, so theirs is not a true swarm.
Carniolan honey bees, Apis mellifera carnica, use their antennae asymmetrically in social interactions, with a strong preference for the right antenna. There has also been speculation about honey bee consciousness. Bees lack the cerebral cortex, but they have a tiny structure that resembles the human midbrain. If it works the same way, they may have a small, simple awareness of their own bodies.
Below about 10 C, honey bees in cold climates stop flying and crowd into a winter cluster at the center of the hive. Workers huddle around the queen and shiver to hold the core between 27 C at the start of winter, when no brood is present, and 34 C once the queen resumes laying. Bees rotate from the outside to the inside so that none gets too cold. The colder the weather, the tighter the cluster. Stored honey is the fuel. In summer, the western honey bee and Apis cerana cool the nest by fanning and by evaporating collected water.
Workers alone carry barbs on the sting, and those barbs set honey bees apart from all other bees. The barbs may have evolved in response to predation by vertebrates, since they only work when the sting is lodged in fleshy tissue. A torn-away stinger keeps delivering venom, because it has its own musculature and ganglion. It also keeps releasing alarm pheromone, which draws more defenders. The worker dies. The venom, called apitoxin, has melittin as its most abundant component. Laboratory and clinical research is probing it for rheumatoid arthritis and allergy immunotherapy, but as of 2018 no clinical use is approved.
The Japanese honey bee, Apis cerana japonica, answers predatory wasps such as the Asian giant hornet by surrounding the intruder in a mass of workers. The bees vibrate their muscles vigorously to raise the temperature to a lethal level. This is called balling. Heat alone was long thought to do the killing, but experiments show that rising carbon dioxide inside the ball is needed too. Beekeepers use the same phrase, balling the queen, when bees kill a queen they see as an intruder or as defective.
Species with open combs, such as Apis dorsata, warn predators with a wave that ripples across the dense layer of bees on the comb surface. Bees momentarily arch their bodies and flick their wings. Cavity dwellers such as Apis cerana, Apis mellifera and Apis nigrocincta guard their entrances and check incoming traffic. Another defense is body shaking, a violent, pendulum-like swaying of the abdomen.
A 2020 study of Apis cerana in Vietnam found something stranger. These bees smear feces and even human urine around their hives to defend against raids by hornets of the species Vespa soror. European and North American bees do not do this.
Albert Einstein is sometimes misquoted as saying, "If bees disappeared off the face of the earth, man would only have four years left to live." Einstein did not say this, and no science supports the prediction. The ten most important crops, which supply 60% of all human food energy, need no insect pollination. Plantains and cassava are propagated by cuttings, while potatoes, yams and sweet potatoes grow from tubers. Soybeans pollinate themselves. Rice, wheat, sorghum and maize are wind-pollinated.
No crop that originated in the New World depends on the western honey bee, which arrived with colonists. Tomatoes, peppers and squash evolved with native pollinators such as squash bees and bumble bees. Tomatoes are not pollinated effectively by honey bees, so bumblebees of the genus Bombus are used for them in several regions.
Still, apples, blueberries and cherries are 90 percent dependent on honey bee pollination. The scale of commercial pollination services is commonly measured in the billions of dollars, credited with adding about 9% to the value of crops worldwide. The USDA says honey bees pollinate more than 130 types of fruits and vegetables. Modern hives let beekeepers move bees from field to field and charge for the service, which favors large commercial operations. In Japan, where A. mellifera is vulnerable to local hornets and disease, the Japanese honey bee A. cerana japonica is used instead.
Honey bees are generalist visitors, and that generality has a cost. Without specialized adaptations for particular flowers, they often pollinate inefficiently, and they are the most commonly documented pollen thieves. For plants with specialized pollinators, experiments show that more honey bee visits can reduce pollination. Where honey bees are invasive, they compete with native pollinators and can push them out.
In Colorado in 1891 and 1896, colony collapses were called "May Disease." Limited events resembling modern colony collapse were documented as early as 1869. Fungus, mites and starvation have all been blamed. Starting in the first decade of the 21st century, die-offs of 30-70% of hives struck North America. The pattern was named colony collapse disorder and was at first unexplained. It seems to come from a combination of factors, possibly including neonicotinoid pesticides or Israeli acute paralysis virus.
A survey by the University of Maryland and Auburn University, published in 2023, found United States colony numbers relatively stable. Even so, 48% of colonies were lost in the year that ended on the 1st of April 2023, against a 12-year average annual mortality of 39.6%. Beekeepers told the surveyors that a 21% winter loss is acceptable. More than three-fifths said their 2022-2023 losses were higher than that.
Varroa mites are arguably the biggest threat to honey bees in the United States. They lay eggs on the pupa, and the hatching mites eat it, causing deformities and spreading disease. Many people believed the mites drank bee blood. A 2018 study in PNAS showed they feed on the fat body tissue of live bees. Another parasite, Acarapis woodi, lives in the breathing tubes of adult bees. The resulting infestation is called acarine, and it has been known as "Isle of Wight disease."
Napoleon I of France used the bee as a symbol of government. The Merovingians took honey bees as royal emblems, signifying immortality and resurrection. Political theorists from Aristotle and Plato to Virgil used a community of honey bees as a model of human society.
The Barberini family, from which Pope Urban VIII came, bore three bees on its coat of arms. Those bees appear throughout St. Peter's Basilica and other Rome monuments as signs of the pope's authority and industry.
Ancient Egyptian mythology held that honey bees were born from the tears of the sun god Ra. Their divine origin made them a stand-in for the Pharaoh. Paired with the sedge, which stood for Upper Egypt, the bee symbolized Lower Egypt. In Joseph and Asenath, a work composed by Egyptian Jews who may have been tied to the Leontopolis temple, bee and honey imagery appears when Asenath converts and is visited by an angel. If that link holds, the bees likely stand for Levite priests, and the imagery signals the legitimacy of a Jewish temple in Egypt.
The Hindu Atharva Veda and the ancient Greeks both linked lips anointed with honey to eloquence and even prescience. The priestess at Delphi was known as the Delphic Bee. The Quran has a chapter titled "The Bee," which compares the industry and adaptability of honey bees to the industry of people. It records the line: "Make your homes in the mountains, the trees, and in what people construct."
The U.S. state of Utah is called the Beehive State. Its state emblem is the beehive, its state insect is the honey bee, and a beehive appears on both the state flag and the state seal.
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Common questions
What is a honey bee?
A honey bee is a eusocial flying insect of the genus Apis, in Apidae, the largest bee family. It builds perennial wax nests in cavities and stores regurgitated carbohydrates as honey.
How many species of honey bee are there?
Eight living species of honey bee are recognized, with a total of 43 subspecies. Historically, anywhere from seven to 11 species have been counted.
Which honey bee species have been domesticated?
Only Apis mellifera, the western honey bee, and Apis cerana, the eastern honey bee, have been truly domesticated. A. mellifera has been cultivated since at least the building of the Egyptian pyramids.
How do honey bees communicate where food is?
Honey bees use the waggle dance to convey the distance and direction of flowers or water. Bees use a round dance when the resource is very close, and scouts also use dance to report nest sites during swarming.
Do honey bees keep humans alive through pollination?
No. The ten most important crops, supplying 60% of human food energy, need no insect pollination. Einstein did not say that humanity would last only four years without bees.
What is colony collapse disorder in honey bees?
Colony collapse disorder describes abnormally high honey bee die-offs of 30-70% of hives in North America, beginning in the first decade of the 21st century. It seems to come from a combination of factors, possibly including neonicotinoid pesticides or Israeli acute paralysis virus.
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