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Queen bee
Queen bee
from Wikipedia
Queen (marked) surrounded by Africanized workers

A queen bee is typically an adult, mated female (gyne) that lives in a colony or hive of honey bees. With fully developed reproductive organs, the queen is usually the mother of most, if not all, of the bees in the beehive.[1] Queens are developed from larvae selected by worker bees and specially fed in order to become sexually mature. There is normally only one adult, mated queen in a hive, in which case the bees will usually follow and fiercely protect her.

The term "queen bee" can be more generally applied to any dominant reproductive female in a colony of a eusocial bee species other than honey bees. However, as in the Brazilian stingless bee (Schwarziana quadripunctata), a single nest may have multiple queens or even dwarf queens, ready to replace a dominant queen in case of a sudden death.[2]

Development

[edit]
Older queen larvae in queen cell lying on top of wax comb
A queen cup
Queen larvae floating on royal jelly in opened queen cups laid on top of wax comb

During the warm parts of the year, female "worker" bees leave the hive every day to collect nectar and pollen. While male bees serve no architectural or pollinating purpose, their primary function (if they are healthy enough) is to mate with a queen bee. If they are successful, they fall to the ground and die after copulation. Any fertilized egg has the potential to become a queen. Diet in the larval stage determines whether the bee will develop into a queen or a worker. Queens are fed only royal jelly, a protein-rich secretion from glands on the heads of young workers. Worker larvae are fed bee bread which is a mixture of nectar and pollen. All bee larvae are fed some royal jelly for the first few days after hatching but only queen larvae are fed the jelly exclusively. As a result of the difference in diet, the queen will develop into a sexually mature female, unlike the worker bees.[3]

Queens are raised in specially constructed queen cells. The fully constructed queen cells have a peanut-like shape and texture. Queen cells start out as queen cups, which are larger than the cells of normal brood comb and are oriented vertically instead of horizontally. Worker bees will only further build up the queen cup once the queen has laid an egg in a queen cup. In general, the old queen starts laying eggs into queen cups when conditions are right for swarming or supersedure. Swarm cells hang from the bottom of a frame while supersedure queens or emergency queens are generally raised in cells built out from the face of a frame.

As the young queen larva pupates with her head down, the workers cap the queen cell with beeswax. When ready to emerge, the virgin queen will chew a circular cut around the cap of her cell. Often the cap swings open when most of the cut is made, so as to appear like a hinged lid.

During swarming season, the old queen is likely to leave with the prime swarm before the first virgin queen emerges from a queen cell.

Virgin queen bee

[edit]
Metamorphosis of the queen bee
Egg hatches on day 3
Larva (several moltings) day 3 to day 8+12
Queen cell capped c. day 7+12
Pupa c. day 8 until emergence
Emergence c. day 15+12 – day 17
Nuptial flight(s) c. day 20 – 24
Egg laying c. day 23 and up

A virgin queen is a queen bee that has not mated with a drone. Virgins are intermediate in size between workers and mated, laying queens, and are much more active than the latter. They are hard to spot while inspecting a frame, because they run across the comb, climbing over worker bees if necessary, and may even take flight if sufficiently disturbed. Virgin queens can often be found clinging to the walls or corners of a hive during inspections.

Virgin queens appear to have little queen pheromone and often do not appear to be recognized as queens by the workers. A virgin queen in her first few hours after emergence can be placed into the entrance of any queenless hive or nuc and acceptance is usually very good, whereas a mated queen is usually recognized as a stranger and runs a high risk of being killed by the older workers.

When a young virgin queen emerges from a queen cell, she will generally seek out virgin queen rivals and attempt to kill them. Virgin queens will quickly find and kill (by stinging) any other emerged virgin queen (or be dispatched themselves), as well as any unemerged queens. Queen cells that are opened on the side indicate that a virgin queen was likely killed by a rival virgin queen. When a colony remains in swarm mode after the prime swarm has left, the workers may prevent virgins from fighting and one or several virgins may go with after-swarms. Other virgins may stay behind with the remnant of the hive. Some virgins have been seen to escape the hive to avoid being killed and seek out another without a queen, such as in the eusocial bee Melipona scutellaris.[4] This can contain multiple virgin queens.[5] When the after-swarm settles into a new home, the virgins will then resume normal behavior and fight to the death until only one remains. If the prime swarm has a virgin queen and an old queen, the old queen will usually be allowed to live. The old queen continues laying. Within a couple of weeks she will die a natural death and the former virgin, now mated, will take her place.

Unlike the worker bees, the queen's stinger is not barbed and she is able to sting repeatedly without dying.

Capped queen cell opened to show queen pupa (with darkening eyes).

Piping

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Piping (listen) is a noise made by virgin and mated queen bees during certain times of the virgin queens' development. Fully developed virgin queens communicate through vibratory signals: "quacking" from virgin queens in their queen cells and "tooting" from queens free in the colony, collectively known as piping. A virgin queen may frequently pipe before she emerges from her cell and for a brief time afterwards. Mated queens may briefly pipe after being released in a hive.

Piping is most common when there is more than one queen in a hive. It is postulated that the piping is a form of battle cry announcing to competing queens and show the workers their willingness to fight. It may also be a signal to the worker bees which queen is the most worthwhile to support.

The adult queen pipes for a two-second pulse followed by a series of quarter-second toots.[6] The queens of African bees produce more vigorous and frequent bouts of piping.[7]

Reproduction cycle

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The surviving virgin queen will fly out on a sunny, warm day to a drone congregation area where she will mate with 12–15 drones. If the weather holds, she may return to the drone congregation area for several days until she is fully mated. Mating occurs in flight. The young queen stores up to 6 million sperm from multiple drones in her spermatheca. She will selectively release sperm for the remaining 2–7 years of her life.[8]

The young virgin queen has a limited time to mate. If she is unable to fly for several days because of bad weather and remains unmated, she will become a "drone layer." Drone-laying queens usually signal the death of the colony, because the workers have no fertilized (female) larvae from which to raise worker bees or a replacement queen.[9]

Though timing can vary, matings usually take place between the sixth and tenth day after the queen emerges. Egg laying usually begins 2 to 3 days after the queen returns to the beehive, but can start earlier than this.[10]

A special, rare case of reproduction is thelytoky: the reproduction of female workers or queens by laying worker bees by parthenogenesis. Thelytoky occurs in the Cape bee, Apis mellifera capensis, and has been found in other strains at very low frequency.[11]

Supersedure

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Capped swarm queen cells

As the queen ages, her pheromone output diminishes. A queen bee that becomes old, or is diseased or failing, is replaced by the workers in a procedure known as "supersedure".

Supersedure may be forced by a beekeeper, for example by clipping off one of the queen's middle or posterior legs. This makes her unable to properly place her eggs at the bottom of the brood cell; the workers detect this and then rear replacement queens. When a new queen becomes available, the workers kill the reigning queen by "balling" her, clustering tightly around her. Death through balling is accomplished by surrounding the queen and raising her body temperature, causing her to overheat and die. Balling is often a problem for beekeepers attempting to introduce a replacement queen.

If a queen suddenly dies, the workers will attempt to create an "emergency queen" by selecting several brood cells where a larva has just emerged which are then flooded with royal jelly. The worker bees then build larger queen cells over the normal-sized worker cells which protrude vertically from the face of the brood comb. Emergency queens are usually smaller and less prolific than normal queens.

Daily life

[edit]
Unmarked queen with attendants.

The primary function of a queen bee is to serve as the reproducer. A well-mated and well-fed queen of quality stock can lay about 1,500 eggs per day during the spring build-up—more than her own body weight in eggs every day. She is continuously surrounded by worker bees who meet her every need, giving her food and disposing of her waste. The attendant workers also collect and then distribute queen mandibular pheromone, a pheromone that inhibits the workers from starting queen cells.[12]

The queen bee is able to control the sex of the eggs she lays. The queen lays a fertilized (female) or unfertilized (male) egg according to the width of the cell. Drones are raised in cells that are significantly larger than the cells used for workers. The queen fertilizes the egg by selectively releasing sperm from her spermatheca as the egg passes through her oviduct.

Identification

[edit]
Color Year
ends in
white 1 or 6
yellow 2 or 7
red 3 or 8
green 4 or 9
blue 5 or 0
Marked queen

The queen bee's abdomen is longer than the worker bees surrounding her and also longer than a male bee's. Even so, in a hive of 60,000 to 80,000 honey bees, it is often difficult for beekeepers to find the queen with any speed; for this reason, many queens in non-feral colonies are marked with a light daub of paint on their thorax.[13] The paint usually does not harm the queen and makes her easier to find when necessary.

Although the color is sometimes randomly chosen, professional queen breeders use a color that identifies the year a queen hatched, which helps them to decide whether their queens are too old to maintain a strong hive and need to be replaced. The mnemonic taught to assist beekeepers in remembering the colour order is Will You Raise Good Bees (white, yellow, red, green, blue).[13][14]

Sometimes tiny convex disks marked with identification numbers (Opalithplättchen) are used when a beekeeper has many queens born in the same year— a method that can also be used to keep multiple bees in the same hive under observation for research purposes.[15]

Queen rearing

[edit]

Queen rearing is the process by which beekeepers raise queen bees from young fertilized worker bee larvae. The most commonly used method is known as the Doolittle method.[16] In the Doolittle method, the beekeeper grafts larvae, which are 24 hours or less of age, into a bar of queen cell cups. The queen cell cups are placed inside of a cell-building colony.[17] A cell-building colony is a strong, well-fed, queenless colony that feeds the larva royal jelly and develops the larvae into queen bees.[18]

After approximately 10 days, the queen cells are transferred from the cell building colony to small mating nuclei colonies, which are placed inside of mating yards. The queens emerge from their cells inside of the mating nuclei. After approximately 7–10 days, the virgin queens take their mating flights, mate with 10–20 drone bees, and return to their mating nuclei as mated queen bees.[17]

Queen rearing can be practiced on a small scale by hobbyist or sideline beekeepers raising a small number of queens for their own use, or can be practiced on a larger, commercial scale by companies that produce queen bees for sale to the public. As of 2017, the cost of a queen honeybee ranges from $25 to $32.[19]

Beekeepers can also utilize alternative methods of queen rearing. Examples are the Jenter kit, walk-away split, Cloake board, and artificial insemination.

See also

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References

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[edit]
Revisions and contributorsEdit on WikipediaRead on Wikipedia
from Grokipedia
The queen bee is the central reproductive female in a (Apis mellifera) colony, distinguished by her primary role in egg-laying and pheromonal regulation of colony unity and behavior. Unlike worker bees, which are sterile females focused on , , and hive maintenance, the queen specializes in producing up to 2,000 eggs per day during peak seasons, ensuring the colony's growth and genetic continuity. Physically larger than workers or drones, she measures approximately 18–20 mm in length and weighs 150–250 mg, with a notably elongated housing 200–400 ovarioles for prolific egg production and a for storing . Queens develop from fertilized eggs laid in specially constructed queen cells, where selected larvae are fed exclusively on —a nutrient-rich from worker bees' hypopharyngeal glands—starting from the third larval , which triggers epigenetic changes leading to their reproductive . This differential nutrition contrasts with worker larvae, which receive only for the first three days before switching to a mixture of , , and diluted , resulting in smaller, non-reproductive adults. Upon emergence after about 16 days, a virgin queen undertakes one or more nuptial flights at 4–6 days old, mid-air with 6–18 drones from multiple colonies at drone congregation areas up to 10 km away, collecting and storing enough sperm in her to fertilize eggs for the remainder of her 2–5 year lifespan. She then returns to the hive, destroys rival queens or cells, and begins laying eggs, with unfertilized ones developing into haploid drones. Beyond reproduction, the queen's mandibular (QMP), secreted from her head glands, acts as a chemical signal that inhibits worker development, promotes nurse behavior, suppresses swarming tendencies, and maintains overall cohesion, making her indispensable for hive stability and productivity. A failing or absent queen leads to rapid decline, often prompting workers to raise emergency queens from existing larvae or initiate supersedure to replace her. In , queen quality—assessed by traits like width, ovariole count, and production—is critical for health, honey yield, and resistance to stressors such as diseases and environmental changes.

Biology and Anatomy

Physical Characteristics

The queen bee exhibits a distinctly larger body size than other castes in the , typically measuring 18 to 20 in length and weighing 150–250 mg, compared to worker bees at 10 to 15 and drones at 15 to 17 . This increased size is most evident in her elongated , which extends well beyond her folded wings—unlike in workers and drones, where the wings nearly reach the abdomen's tip—facilitating her role in egg production. Her is sleek and shiny, with a smooth that lacks the dense hair covering typical of workers, and it bears no baskets on the hind legs or glands, adaptations absent in the queen as they are irrelevant to her reproductive function. The queen's wings are proportionally reduced, spanning only about two-thirds of her when at rest, and her legs are slimmer and less robust relative to her overall body size, reflecting her limited involvement in or hive . Color variations occur among different breeds of honey bees; for instance, queens of the (Apis mellifera ligustica) often display a golden-yellow coloration, while those of the Carniolan bee (Apis mellifera carnica) tend to be darker, nearly black. In wild colonies, a queen's lifespan can extend up to 5 to 7 years, though her peak productivity in egg-laying generally wanes after 2 to 3 years, prompting colony succession.

Internal Anatomy

The queen bee's internal anatomy is specialized for high reproductive output and chemical signaling within the colony. Her ovaries are highly developed, each containing 150-180 ovarioles, compared to the 3-20 ovarioles found in worker bees, enabling the production of 1,500-2,000 eggs per day during peak season. These ovarioles facilitate the continuous maturation of oocytes, supported by the queen's larger that houses this extensive reproductive apparatus. The serves as a critical for , accommodating approximately 6 million viable spermatozoa obtained from with multiple drones, typically 10–20, during nuptial flights. This reservoir ensures lifelong fertility, as the queen does not remate after her initial period, utilizing stored to fertilize eggs selectively for producing female offspring. The spermathecal fluid provides a protective environment that maintains viability for years, preventing degradation and supporting over the queen's lifespan of up to 7 years. Mandibular glands in the queen's head produce queen mandibular (QMP), a blend of volatile fatty acids that acts as a primer to suppress worker development and promote for cohesion. QMP diffuses throughout the hive, inhibiting reproductive activity in workers and reinforcing the queen's dominance in . These glands are anatomically adapted for sustained , integrating with the queen's metabolic systems to regulate release. Accessory glands associated with the reproductive tract secrete coatings for eggs, providing and during oviposition, while the venom apparatus—derived from modified reproductive structures—produces a less potent compared to that in workers, primarily serving defensive rather than roles. The in queens retains epidermal characteristics from ancestral accessory glands but exhibits reduced protein content and activity. Metabolic adaptations include an expanded , the primary site for storage and intermediary , which accumulates higher reserves to fuel prolonged egg production and synthesis. This tissue undergoes dynamic changes, hypertrophying during reproductive phases to supply energy and vitellogenins essential for . The fat body's role extends to immune support, ensuring the queen's physiological resilience amid continuous reproductive demands.

Development and Emergence

Larval Development

The development of a queen bee begins with the laying of a fertilized egg by the reigning queen into a specialized queen cell constructed by worker bees, where the resulting larva will receive an exclusive diet of royal jelly after hatching. These cells are larger than worker cells and can be either swarm cells, constructed vertically along the lower edges or face of the comb in preparation for colony reproduction, or emergency cells, converted from existing worker cells when the colony becomes queenless. The egg, standing upright, hatches after approximately three days into a small, C-shaped larva. From hatching, the queen-destined larva receives an exclusive diet of secreted by nurse bees, delivered through thousands of feeding visits that support rapid growth to over 1,500 times its initial size. In contrast, worker larvae are fed only for the first three days before switching to a mixture of and (worker jelly). This sustained nutrition, containing compounds like (E)-10-hydroxy-2-decenoic acid, acts as a , promoting epigenetic modifications such as increased at specific sites (e.g., H3K9 and H3K27), which enhance chromatin accessibility and trigger caste-specific favoring queen traits like larger body size and reproductive capacity. Around day 8 or 9 after egg laying, worker bees cap the queen cell with beeswax, at which point the mature larva spins a silken cocoon inside. The larva then pupates, undergoing metamorphosis over 7-8 days in a stable colony-maintained temperature of 34-35°C, which is essential for proper development. Queen bees develop from diploid female zygotes, just as workers do, with caste differentiation driven primarily by environmental factors like nutrition rather than genetics; however, the age of the larva when transferred to a queen cell (as in grafting) significantly affects queen quality, as younger larvae (e.g., under 24 hours old) yield heavier, more viable adults due to longer exposure to optimal feeding conditions.

Virgin Queen Phase

The virgin queen emerges from her queen cell after a total development period of approximately 16 days from the laying of the , chewing through the wax cap at the tip of the cell to exit. At emergence, she typically weighs between 180 and 200 mg, with variations depending on factors such as larval nutrition and . This lightweight, pale adult queen, with her soft still hardening, immediately interacts with the while other potential queens may still be developing in nearby cells. Upon emergence, the virgin queen begins communicating through acoustic signals known as , which serve to announce her presence and challenge rivals. She emits "toots," consisting of a long introductory pulse followed by shorter pulses at frequencies of 400-500 Hz, often from outside her cell to signal dominance. Rival still within their cells respond with "quacks," a series of shorter pulses at lower frequencies of 200-350 Hz, escalating tensions that can lead to direct confrontations if multiple emerge. These vocalizations, produced by vibrating the thoracic muscles against the cell walls or hive structures, help coordinate rival interactions and are most intense in the first few days post-emergence. Rival elimination is a critical during this phase, ensuring only one queen survives to lead the . The first-emerged virgin queen often seeks out and stings unemerged rivals through their cells, targeting those closest to emergence first to neutralize immediate threats; her smooth allows repeated use without fatality to herself. If multiple queens emerge simultaneously, they engage in physical fights involving , stinging, and , with the dominant queen typically prevailing through superior size or aggression, resulting in the death of subordinates. Workers may facilitate this by isolating cells or supporting the victor, though intervention is minimal. Throughout this period, the virgin queen depends on nurse worker bees for sustenance, receiving exclusive feedings of —a protein-rich glandular secretion that supports her rapid maturation and energy needs. Within hours of emergence, she initiates production of queen mandibular pheromones, such as 9-octadecenoic acid (9-ODA) and 10-hydroxy-2-decenoic acid (10-HDA), which begin influencing worker behavior and colony cohesion even in her unmated state. The virgin queen remains hive-bound for 5-10 days post-emergence, using this time to gain strength, fully harden her , and develop her flight muscles while avoiding premature exposure. This rest period's duration can vary based on health, nutritional status, and environmental factors like , with optimal conditions promoting quicker readiness. During this phase, she hides in protected areas of the hive, fed and groomed by workers, as the prepares for her eventual role.

Reproduction and Mating

Mating Flights

A newly emerged queen honey bee typically undertakes her nuptial flights 5 to 10 days after from the pupal stage, during warm afternoons when temperatures exceed 20°C, with flight activity peaking between 13:00 and 16:00 hours. These flights occur 1 to 5 times over successive days, allowing the virgin queen to leave the hive and travel several kilometers—often 2 to 6 km—to reach drone congregation areas (DCAs), elevated sites where thousands of drones from multiple colonies gather for . During these preparatory orientation flights in the virgin queen phase, the queen familiarizes herself with the surroundings before committing to full nuptial excursions. In mid-air at the DCA, the queen engages in polyandrous , copulating with 10 to 20 drones on average during her flights, though numbers can range up to 44 in exceptional cases. Each lasts mere seconds, during which the drone everts its endophallus to transfer before dying shortly thereafter, while the queen stores the in her for lifelong use in fertilizing eggs. This rapid process enables the queen to mate multiple times in a single flight, accumulating 5 to 7 million viable cells overall. The exhibited by the queen promotes within the colony through multiple patrilines—distinct paternal lineages among worker offspring—which enhances overall fitness by mitigating and bolstering resistance to diseases and parasites. Studies show that colonies headed by multiply mated queens exhibit improved , , and reduced loads compared to those with singly mated queens, as the varied allow for a broader range of immune responses and behavioral adaptations. Upon returning to the hive after her final mating flight, the queen is noticeably heavier due to the stored and initiates physiological changes, including a ramp-up in queen mandibular pheromone (QMP) production from her mandibular glands to signal her mated status and inhibit worker . This pheromonal shift helps stabilize colony organization but comes with significant risks: approximately 10 to 20% of queens fail to return due to predation by birds or , adverse weather, or disorientation, potentially triggering supersedure by a rival virgin queen.

Egg-Laying Process

The queen bee's enables prolific egg production through her paired ovaries, each containing 150–200 ovarioles that facilitate , the process by which immature oocytes develop into mature s. During each oviposition cycle, one matures per ovariole and is transported to the common for laying, allowing the queen to deposit eggs sequentially at a high rate after , when sperm is stored in the for selective use. The eggs themselves are barrel-shaped, measuring approximately 1.5 mm in length and 0.35 mm in width, and stand upright on their ends within cells shortly after deposition. Fertilization occurs selectively as the passes through the median oviduct toward the , where the queen controls the release of a precise volume of spermathecal fluid—typically containing just 2 cells—via the Bresslau pump, a structure equipped with a at the junction of the and its duct. This mechanism allows the queen to decide whether to fertilize the , drawing from the millions of stored post-mating; fertilized eggs develop into diploid females (workers or ), while unfertilized ones become haploid males (drones). The process is highly efficient, with usage remaining consistent until stores deplete in older , ensuring sustained throughout her lifespan. In laying, the queen actively inspects and selects empty comb cells, preferentially depositing fertilized eggs in smaller worker cells (approximately 5.4 mm in diameter) to produce offspring, while laying unfertilized eggs in larger drone cells (about 6.5 mm in diameter) only when colony needs demand increased drone production. During peak season, she can achieve a laying rate of up to 2,000 eggs per day, matching colony growth demands, but this rate adjusts dynamically based on environmental and . Egg-laying is tightly regulated by nutritional status, colony pheromones, and space availability, with worker-produced s and feedback from queen mandibular (QMP) influencing ovarian activity to prevent overproduction. Laying ceases or significantly reduces if the colony becomes overcrowded or faces , as reduced worker feeding limits the queen's access to essential proteins and lipids needed for . Seasonally, production peaks in spring and summer but tapers in fall, often entering a diapause-like state by late autumn or winter, where laying halts entirely due to cooler temperatures, limited , and shorter days, resuming only when conditions improve.

Role in the Colony

Daily Behaviors

The queen honey bee exhibits a continuous routine of movement within the hive, roaming through the brood cluster to inspect cells and distribute pheromones, covering an average daily distance of 46 to 60 meters. This patrolling behavior allows her to balance exploration of the hive structure with targeted stops for egg-laying and interactions, with short pauses often triggered by worker contacts that facilitate and feeding. Workers actively attend to the queen during these movements, antennating and grooming her to maintain her cleanliness and spread pheromones, while she in turn is fed regurgitated multiple times between oviposition bouts, typically after laying 2 to 26 eggs per cycle. Central to her daily activities is the dispersal of queen mandibular (QMP), produced at approximately 500 micrograms per day, which she trails through physical contact during walking and self-grooming. This volatile blend suppresses ovarian development in workers and attracts a of 10 to 20 attendants who lick, feed, and groom her, further amplifying pheromone distribution via worker-to-worker contacts throughout the colony. The queen generally avoids interactions with drones post-mating, focusing her social engagements on workers who form a protective "" around her, prioritizing her safety even at the cost of their own lives during hive disturbances. Her activity alternates between egg-laying, rest, and maintenance, with no distinct ; she remains active around the clock, resting 37 to 61 percent of the time under or constant conditions. Peak behaviors occur in spring and summer when demands are high, supporting up to 2,000 eggs laid daily, whereas winter confines her to minimal movement within the insulating cluster, with reduced feeding and output until brood rearing resumes. In response to threats or disturbances, such as hive manipulations, workers intensify retinue formation to shield the queen through grooming and clustering, mitigating stress and exposure.

Supersedure and Succession

Supersedure refers to the natural replacement of a queen by workers in a , ensuring continuity without the disruption of swarming or complete abandonment. This is triggered primarily by declining levels of queen mandibular (QMP), which decreases due to the queen's old age, disease, or preparation for swarming. Workers detect these changes through declining QMP levels, signaling her diminished reproductive capacity. Diseases such as viral infections vectored by mites can cause queen failure by affecting function and production, including components like methyl oleate. In emergency supersedure, which occurs when the queen is failing but the remains intact, workers select young worker larvae less than three days old—still in their totipotent phase—for rearing as new . These larvae, typically from existing brood, are transferred to specially constructed vertical queen cells and fed copious to promote queen development, a process that builds on the basic larval rearing mechanisms outlined in queen . Workers prioritize larvae in good nutritional condition, as non-deprived ones yield significantly higher rates of successful pupation into queens compared to deprived ones. Swarm supersedure, in contrast, integrates with reproductive swarming, where the existing queen departs with a portion of the , leaving behind developing queen cells in the original hive for a new queen to emerge and lead the remnant group. This ensures the colony's division while replacing the outgoing queen. Upon emergence, if multiple virgin queens develop, they engage in piping challenges—audible signals produced by emerging queens to assess rivals—often leading to combat where the dominant queen eliminates others using her sting. Success rates for supersedure are influenced by factors like larval quality, environmental conditions, and queen viability post-emergence, with estimated at 80-90% in free-living ; and fighting typically resolve multiple to one. Evolutionarily, supersedure maintains genetic fitness by timely replacement of suboptimal , enhancing survival and productivity, unlike absconding, where the entire relocates due to severe threats without producing new . This adaptive mechanism underscores the 's collective , balancing stability with renewal.

Beekeeping Practices

Queen Identification

Beekeepers locate the queen bee in a hive through careful observation of her distinct physical traits, which set her apart from the more numerous worker bees. She possesses an elongated that extends noticeably beyond her wings, a larger and more tapered body overall, a shiny and hairless often darker in color, and no pollen baskets (corbiculae) on her hind legs, unlike workers who return from with visible loads. These features are most reliably spotted during spring inspections when activity peaks and brood frames are lighter, or in the evening when reduced bee movement allows for steadier viewing of frames. Additionally, the queen is typically attended by a small of 5–10 worker bees that follow her closely, forming a visible circle or escort as she moves across the . To facilitate repeated identification during hive management, beekeepers often mark the queen by applying a small dot of non-toxic, quick-drying to her using a specialized pen, such as a Posca PC-7M. This practice follows an international tied to the year of her emergence: white or gray for years ending in 1 or 6, yellow for 2 or 7, red for 3 or 8, green for 4 or 9, and blue for 5 or 0. , where one or both forewings are trimmed with small scissors, may also be performed after marking to prevent swarming by limiting her flight capability, though this does not hinder her egg-laying or function. Various tools assist in safely locating, capturing, and confirming the queen without excessive disturbance to the . Puffing from a disperses workers temporarily, exposing the queen on the frame; a or handheld catcher then allows gentle immobilization for marking or removal. Queen excluders—metal or plastic grids with openings sized to permit worker passage but block the larger queen—can confine her to specific hive sections for easier monitoring during inspections. Identifying the queen presents challenges, particularly her tendency to blend into the dense cluster of bees or against the dark wax of brood combs, where workers may obscure her position. In rare instances of supersedure failure, multiple virgin or mated queens may coexist briefly before fighting, further complicating visual confirmation until only one remains dominant. The ability to reliably identify the queen advanced significantly in the mid-19th century with the invention of the movable-frame hive by Rev. Lorenzo Lorraine Langstroth in 1852, which allowed beekeepers to inspect individual frames without destroying the comb structure, transforming hive management practices.

Artificial Rearing

Artificial rearing of queen bees enables beekeepers to produce and introduce new queens to enhance colony , replace aging or failing queens, and expand apiaries, paralleling aspects of larval development but under controlled conditions. This process begins with selecting high-quality larvae from superior colonies and involves several stages to ensure viability and mating success. The technique is a foundational method in artificial queen rearing, where 1-2 day-old larvae are carefully transferred from worker cells to artificial queen cups using specialized tools like grafting needles. These cups, often made of wax or plastic, are primed with to provide initial nutrition and prevent , mimicking the jelly-rich environment that promotes queen development. The grafted frame is then placed in a cell builder colony for incubation, where nurse bees feed the larvae extensively with over 5-6 days until queen cells are capped. Cell builders are strong, queenless colonies specifically prepared to nurture grafted larvae, consisting of abundant nurse bees that secrete to provision the developing queens. These colonies are typically set up with frames of emerging brood to maximize nurse bee populations, along with ample food stores like sugar syrup and patties to support secretion. Cell builder colonies can rear 100 or more queens during spring nectar flows when nurse bees' hypopharyngeal glands are highly productive, but in fall, rearing more than 15–20 queens is challenging due to resource limitations and reduced gland activity. Once queen cells mature and virgins emerge after about 16 days, they are placed into mating nuclei—small, isolated hives stocked with worker bees, brood, and stores, positioned near drone congregation areas. These nuclei allow the virgin queens to undertake mating flights, collecting semen from multiple drones (typically 10-20), with beekeepers monitoring for successful returns by observing colony activity and egg-laying after 7-10 days. Success depends on weather, drone availability, and nucleus strength, often achieving mated queens in 2-3 weeks. Introducing mated queens to recipient colonies requires gradual acceptance to minimize rejection, commonly using protective cages with candy plugs that workers must chew through over 2-4 days. The queen is confined in a bent-wire or push-in cage between brood frames, allowing pheromones to familiarize the colony while preventing attacks; timing introductions to queenless hives in the evening boosts acceptance. Success rates range from 70-90% with proper preparation, such as reducing the colony to nurse bees and ensuring no laying workers are present. Modern advances in artificial rearing include instrumental insemination, pioneered in the 1920s and refined by Harry H. Laidlaw Jr. in the mid-20th century, which involves collecting semen from selected drones and directly injecting it into the queen's oviducts using a specialized . This technique enables precise genetic control, such as selecting for varroa mite resistance by inseminating queens with sperm from hygienic or suppressed-mite-reproduction drone lines, achieving heritable gains in survival rates over generations. Recent developments include artificial rearing methods, allowing controlled queen production in settings without full involvement. Additionally, genetic selection programs leverage instrumental insemination to promote hybrid vigor through controlled crosses of diverse , enhancing traits like tolerance and without relying on natural .

References

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