Bees

Queen

Queen bees are longer than worker bees, and they and lack the hair color banding on the abdomenabdomen:
the segmented, posterior (third) part of the bee body that contains heart, honey, stomach, intestines, Malphigian tubules, reproductive organs, and sting
that is characteristic of workers. The queen’s head and thoraxthorax:
the middle region of the adult bee body, which lies in between the head and the abdomen; consists of three segments: pro-, meso-, and meta-thorax; thorax attachments include three pairs of legs and two pairs of wings
differ somewhat from a worker’s, but this difference is not readily apparent. We see the difference in the abdomenabdomen:
the segmented, posterior (third) part of the bee body that contains heart, honey, stomach, intestines, Malphigian tubules, reproductive organs, and sting
and in her behavior when we have the good fortune to spot her in the hive. Seeing a queen in the hive is mostly luck, enhanced by looking on appropriate frames where she might be laying eggs. However, we sometimes find her on honey frames, on the inside walls of the hive, or seeking to escape the light due to opening of the hive.

Adult queen; photo by Vera Kuttelvaserova, stock.adobe.com
Queen outside hive; photo by Robert Snyder
 

Our opportunity to spot a queen is reduced the longer the hive is opened and the more the colony disturbed. A strong retinue of a dozen or more bees tending their queen, if spotted, helps in finding her. Queens are light aversive. Thus, when hives are opened, a queen may abandon her normal behavior of egg laying and run on the comb. Some queens leave brood combs and go to a comb with honey stores or to internal hive surfaces. Each hive opening causes some disturbance, complicated further by use of smoke that may disrupt pheromonepheromone:
a chemical substance released externally by an individual (from an exocrine gland) which stimulates a response in a second individual of the same species
distribution. Africanized queens and workers exhibit pronounced comb running behavior when their colonies are examined.

The queen is the mother of all the workers and drones in the colony. A colony with a mated queen that is producing eggs is termed queenrightqueenright:
a colony with a healthy, worker egg-laying queen; the opposite of a queenless colony
. A queenless colony is one without a queen. Queen development takes 16 days on average. The queen, upon emergence, is a virgin queen. She gains strength and eventually performs orientation flights in preparation for her mating flight(s). Following mating (usually in the first week of her adult life, but before three weeks of adult life), she is termed the mated queen. Colonies normally have a single mated queen, but in a percentage of colonies there may be two queens; a mother-daughter queen in the same hive.

Capped vertical queen cell below horizontal capped worker cells; photo by Dewey M. Caron

A new queen develops in a vertical cell. The queen starts development of a replacement by laying a fertilized egg in a queen cup. Depending upon the conditions of queen rearing, the developing queen cells may be termed swarm queen cells, when the colony is in swarm behavior, or supersedure queen cells, when a colony is replacing the mated queen. (See more on brood page.)

If the queen is suddenly removed or the colony becomes queenless for any reason, the bees may rear one or more emergency queens. They modify the horizontal worker cell with a fertilized egg or a larva that is up to three days of larval age so the cell orientation is vertical. The large queen cells, eventually the size of a peanut, hang vertically from the margin or surface of the beeswax comb.

Queen size may vary considerably. Virgin (unmated) and recently mated queens have smaller and thinner abdomens. Queens preparing to swarm will lose weight and might appear thinner. Queens reared under emergency conditions may be smaller-bodied. Queens laying fewer eggs (in winter or in colonies with insufficient worker population, such as after a pesticide kill of adults, or following beekeeper splitting of a colony) will appear smaller. When the queen is laying an egg, her abdomenabdomen:
the segmented, posterior (third) part of the bee body that contains heart, honey, stomach, intestines, Malphigian tubules, reproductive organs, and sting
disappears into the cell and only her head and thoraxthorax:
the middle region of the adult bee body, which lies in between the head and the abdomen; consists of three segments: pro-, meso-, and meta-thorax; thorax attachments include three pairs of legs and two pairs of wings
are visible. Spotting a queen laying an egg is thus difficult (see YouTube video in Resources below to see a queen laying an egg).

Queens have two basic functions. Although critical to hive population the queen is not the bee hive “ruler” in the sense of human connotations for this term. Her functions are to lay eggs to populate her family with workers and drones, once conditions improve and the colony can grow in population. Under favorable conditions she may also lay the eggs that will lead to colony reproduction (i.e., swarming) or her replacement (supersedure). Her second function is to produce pheromones that will serve to unify the family.

Queen bees lay fertilized eggs that will develop into females or she may withhold release of sperm as she lays an unfertilized egg. Unfertilized eggs yield the male drones. These unfertilized eggs are laid in comb cells that are larger in diameter; the queen uses her front legs like a pair of calipers to measure cell size to determine the appropriate egg to lay. Queens normally lay one egg in a cell but on occasion cells might have multiple eggs. Usually only a small number of cells will have multiple eggs since younger aged (cleaning) worker bees police the brood area to remove (cannibalize) the “extra” eggs. If the queen places unfertilized eggs in smaller worker cells, or fertilized eggs in the larger drone cells, the workers remove the incorrectly placed eggs.

Queens deposit one egg per cell but occasionally may put more than one egg in a cell (arrow); photo by Robert Snyder

Young, vigorous queens just starting their egg laying may put more than one egg in a cell. Multiple eggs happen in colonies without enough cleaned, polished cells for the queen to lay eggs in; this can occur in spring when there is an imbalance of more older-aged bees compared to fewer numbers of newly emerging younger bees. Colonies running out of space in the brood area, such as instances when there is a pollen- or honey-bound colony, can lead to too few empty cells for queen egg laying and the queen laying more than one egg in a cell. Rarely, the queen may put an egg in a cell with bee bread. When colonies become queenless, worker bee ovaries will develop and some workers then lay unfertilized eggs. This condition, termed laying workers, usually leads to numerous eggs in a cell. Laying worker eggs in cells will look very different from cells in which the queen might lay an extra egg. If drone brood is being reared in worker cells it is a different situation. The queen is running out of stored sperm or the sperm is not alive. This condition is called a drone laying queen or drone layer.

A difficult-to-diagnose condition is queens that perform poorly in their egg laying or pheromonepheromone:
a chemical substance released externally by an individual (from an exocrine gland) which stimulates a response in a second individual of the same species
production. Such a condition can result from inbreeding in a queen rearing operation. It can also mean a queen whose worker offspring are not productive or are susceptible to pests and diseases. Such queens are likely to have spotty brood patterns with eggs, larvae, and cappedcapping:
the covering that bees add over comb cells containing fully ripened honey or to cap brood that has reached the pupal stage; bee bread cells are not capped
cells scattered on the comb and brood patterns that are mixed with various interspersed life stages.

There can also be considerable variation in queen coloration. While variation of body color is normal, some variations might be due to breeding for specific coloration, such as golden (yellow) Italian queens, or from their racial mixture, as with, for example, darker Cape bees, or Carniolan bees.. Some beekeepers prefer the lighter color Italian queens over the darker Carniolan or even darker Caucasian bees, as they are easier to spot on darker colored brood comb.

Light-colored Italian queen; photo by Robert Snyder
Darkly colored queen; photo by Robert Snyder
 
Queen marked with number disc (25); note clipped queen wing on right side; this is not an injury but an alternative marking technique; photo by Robert Snyder

Marking queens

Beekeepers mark queens with a paint mark on the thoraxthorax:
the middle region of the adult bee body, which lies in between the head and the abdomen; consists of three segments: pro-, meso-, and meta-thorax; thorax attachments include three pairs of legs and two pairs of wings
or a numbered disc glued to the top of the thoraxthorax:
the middle region of the adult bee body, which lies in between the head and the abdomen; consists of three segments: pro-, meso-, and meta-thorax; thorax attachments include three pairs of legs and two pairs of wings
. The marking can make it easier to spot a queen. Queens may have part of a wing cut as a mark of year reared or may be missing a wing or part of a leg, due to injury or from cutting by a beekeeper. Cutting a queen’s leg is a technique to promote supersedure. Wing cutting does not appear to harm a queen. Wing cutting might also be done as an (ineffective) swarm control technique. Without intact wings, the queen will not be able to fly to the swarm bivouacbivouac:
a temporary cluster of swarming honey bees intermediate between leaving home and finding a new nest cavity
site. In such instances, the worker bees of the swarm will return to their hive and then often await the emergence of one or more virgin queens and subsequently successfully swarm with a virgin or several virgin queens.

Mated queens alternate egg-laying behavior with a resting period. A retinue of worker bees often forms around the queen to groom, feed, and remove her waste during the resting phrase. The workers are also picking up the essential pheromones secreted by the queen. Depending upon the season, a queen may produce eggs approaching her body weight each day. During winter clusterwinter cluster:
a closely packed colony of bees forming in winter to conserve heat
a queen lays few to no eggs.

Queens confined in cages

Mated queens confined in cages (rarely virgin queens) are a common method used when starting a bee colony. Queens are caged for shipping from queen producer to beekeeper to requeenrequeen:
to replace a queen; old queens are often removed and replaced by a ripe queen cell or by a mated queen via an introduction cage
a colony, to make a nucnucleus:
also called a "nuc;" a smaller colony of bees usually with three to five frames. Nucs are splits (divides) made from larger colonies. Nucs are purchased as a method of starting a new colony or as a resource to use to bolster weaker colonies. See more information about nucs <a href="index.cfm?pageID=3093#standard">here</a> and <a href="index.cfm?pageID=3417#nuc">here</a>. 
, or to make a division/split queenrightqueenright:
a colony with a healthy, worker egg-laying queen; the opposite of a queenless colony
, for example when the beekeeper splits a larger colony to reduce chances it will swarm. Queens are caged in package bees shipped to beekeepers starting a new colony.

The queen may be released immediately from a cage (direct release), but it is preferred to allow the worker bees to release the queen (indirect release). Direct release when the queen cage is opened and the queen allowed to run onto a comb is risky if the bees have not had a chance to become used to her pheromonepheromone:
a chemical substance released externally by an individual (from an exocrine gland) which stimulates a response in a second individual of the same species
signature. If the queen has been confined in a cage among package bees it may be OK to direct release when hiving the package. However, indirect release is the norm. With the indirect release technique the caged queen is placed into a hive within the cage; she is released when worker bees chew a passage through the sugar candy plug (or masking tape) of the queen cage. The workers have time to become used to the new queen as her release is delayed two or three days. There is considerable lore around how to place the cage to ensure introduction is successful. Some advocate moving queens from their shipping cage to (homemade) introducing wire cages as they provide more queen and worker interaction before her release (see photos).

Queen cage; bees calm, so queen is ready for release; photo by Penn State University

Before release, beekeepers can assess if the worker bees are ready to accept the queen by observing the behavior of the worker bees on the cage. If the workers move freely, it is okay to release the queen, but if workers are moving rapidly and/or there is evidence of workers biting the cage, the queen should remain caged for later release. If released too soon, the bees may ball and kill the queen.

Beekeepers may cage a queen to create a brood break for varroa mite control. Some prefer to make a cage around a frame with queen excluderqueen excluder:
a metal or plastic device that allows workers to pass through opening slots or wires, but is spaced to keep the queen and drones from passing through.
material to hold the queen during this period. Others use the familiar shipping cage to sequester the queen. An alternative is to use a push-in cage (plastic or wire) to confine the queen, usually without worker attendants, over a small portion of drawn brood cells. Homemade cages of wire may also be used.

There are three common cages used to ship queens - the Benton (3-hole) wooden cage, the California queen cage (a single cavity, normally to confine a queen without attendants), and the JZ BZ plastic cage. Caged queens in Benton cages often have attendant worker, bees but the smaller California cage usually does not include worker attendants. When shipping a large numbers of queens, the queens may be individually caged and put into a battery box; the shipping (battery box) package is stocked with younger-aged attendant bees. 

California queen shipping cage; queen without attendants. Sugar candy tube replaces cork for bees to release the queen; photo by The BeeMD photo collection
Caged queens with worker attendants in three-hole Porter shipping cage; photo by Jen Larsen
Queen in JZBZ queen cage; photo by The BeeMD photo collection
 

Benton and California queen cages have the exit plugged with a queen candy; in the California cage, a tube of candy is added to plug the single cage opening (after removal of a cork plug). In the Benton cage, corks plug both the opening at the sugar candy end and at the opposite end. The cork at the candy end must be removed so the bees can remove the sugar candy plug. In some instances workers have been known to chew the cork to release a caged queen. Lacking corks, the exit holes can be covered with tape.

Caged queens may suffer from being sticky if cage candy softens too much or sugar water feeders leak onto cages. Queen health is negatively affected by temperature extremes during shipment. Queens (or queen attendants) may harbor varroa mites. In a few instances a queen cage may include small hive beetle adults or wax moth caterpillars or fly maggots.

Introducing cages

There are numerous designs of cages for queen introduction into a colony that will influence queen acceptance/rejection. The shipping cage may be used with the escape hole filled with queen candy that must be consumed to release the queen. The candy plug helps ensure a slower release so worker bees become accustomed to the distinctive queen odors for improved queen acceptance. Alternatively, a push-in cage of wire or plastic can be used to confine the queen over some beeswax comb cells. Push-in cages can include (or exclude) worker bee attendants; when constructed of queen excluderqueen excluder:
a metal or plastic device that allows workers to pass through opening slots or wires, but is spaced to keep the queen and drones from passing through.
material worker bees may come and go. Push-in cages can be purchased or homemade.

Selection of cages: homemade wire introduction cage; a wire push-in cage (below); plastic and metal queen shipping cages; California shipping queen cage (top); photo by Dewey M. Caron

Specialty cages to confine queens (for example when seeking to create a brood break for varroa mite control) may be purchased or homemade., Such cages usually enclose a single frame and may exclude workers or be made with queen excluderqueen excluder:
a metal or plastic device that allows workers to pass through opening slots or wires, but is spaced to keep the queen and drones from passing through.
material to allow attendant bees to enter/exit. Cages with workers allow for normal egg laying by a queen; the frame can be removed when filled by queen egg laying and substituted with another empty one.

For protection, queen cells close to emergence of virgin queens may be enclosed in a queen cell protector cage (plastic hair rollers sometimes substituted). This can protect queen cells from being destroyed by another queen see chewed hole in queen cell).

Resources

Bee-Health. 2019. Biology of Individual Honey Bees. Bee Health. Accessed 2023. https://bee-health.extension.org/biology-of-individual-honey-bees/?msclkid=c3cb798ccfcc11ecb5773b5bfd6f1e2d

Conrad R. 2015. Introducing Her Royal Highness. Bee Culture. Accessed 2023. https://www.beeculture.com/introducing-her-royal-higness/

“Caring for Caged Queens”. YouTube, uploaded by University of Guelph Honey Bee Research Centre, 20 March 2019. https://www.youtube.com/watch?v=E2aRpTkoDeM

Mangum W. 2020. Queen Introduction: Part 3 Seeing into the Forbidden Time When the Queen will Live or Die. American Bee Journal 160(8): 867-872. https://bluetoad.com/publication/?i=666377&p=43&view=issueViewer https://americanbeejournal.com/queen-introduction-part-2-the-eft-bees/

Anderson C. 2023. Care of a Caged Queen Bee. Carolina HoneybeesCarolina Honeybees:
Carolina Honeybees. Accessed October 2022. https://carolinahoneybees.com/
. Accessed 2023. https://carolinahoneybees.com/care-caged-queen-bee/

“How the bees act when they reject the queen new beekeepers don’t miss this”. YouTube, uploaded by Barnyard Bees, 6 September 2017. https://www.youtube.com/watch?v=qXVVOn5uZCY

“Balling the Queen”. YouTube, uploaded by The Honey Company, 28 July 2022. https://www.youtube.com/watch?v=y25HIUQX9hk

“Installing Queens in Various Queen Cages into a Hive”. YouTube, uploaded by Betterbee, 1 June 2021. https://www.youtube.com/watch?v=uO2biHgsfmQ

Cushman D. 2022. Queen Cages for Travelling and Introduction of Queen Honey Bees. David A. Cushman. Accessed 2023. http://www.dave-cushman.net/bee/queencages.html

“Honey Bee Queen Laying Eggs” YouTube, uploaded by AskaBiologist, 20 August 2020, https://www.youtube.com/watch?v=1NO7sXltrbE