Members of the Colony
Every female honey bee egg has, for its first three days of existence, the potential to become either a queen or worker. Its fate, in this regard, is determined solely by the attention the larvae receives from the workers. Female eggs destined to be raised as queens are generally laid in bowl shaped cups (the precursors to the large vertically oriented queen cells) and bathed in nutrient-rich royal jelly. As larvae they are fed copiously (more royal jelly) and tended often.
Royal jelly is composed of approximately: 67% water, 12.5% crude protein (including small number of amino acids), 11% simple sugars (monosaccharides), 5% fatty acids, and 4.5% other—trace minerals, enzymes, and vitamins B5, B6 and C.
The larval queen’s diet, thus enriched, encourages maturation of her reproductive organs but limits her development in other respects. Successfully mated queens will be cared for by a retinue of workers who will forage for, feed and groom her thus she does not need and so does not develop pollen baskets, a tongue sufficient in length to collect nectar nor a distensible rectum.
Developing queens go through larval and pupal instars (the period between molts) more quickly and immerge a full five days sooner (on day sixteen) than workers (who immerge on day twenty-one). Upon eclosion (immersion) queens generally weigh twice as much as workers (about two hundred milligrams), and their first concern is locating sister queens. Racing through the colony the queen emits a distinctive sound know as piping or tooting. If she finds other queen cells she feverishly tears until she has rent a hole large enough to allow her stinger purchase. She repeats this until all the other pupal queens have been eliminated or until a sister emerges with whom she battles to the death.
If she is not killed in her cell or upon eclosion by a rival, the nubile queen spends the next twelve to twenty-four hours drying and hardening. During this time, assuming she is accepted by the colony, she is fed and licked by workers who spread her pheromones throughout the hive.
After six-seven days of eating, resting, and stretching, (during which time her reproductive organ reach full maturity) the workers sense she is ready to be mated and begin to harass (chase and nip) her. If weather permits, the virgin queen takes to the skies in search of a drone congregation area (DCA) there to mate with on average fifteen drones.
The above description of queen rearing assumes nature is allowed to take its course without mishap or interference. However, as this is not always the case, below are two types of situations wherein queens are made under specific circumstances. The final entry describes the way in which a strong colony reproduces and in so doing makes some of the most healthy and hale queens—swarm queens.
When a colony’s queen is in some way compromised (such as waning in valality or fecundity) workers often raise supersedure queens. Alerted to their queen’s diminishing procreative powers by weakening of her distinctive pheromones and/or a decrease in her egg laying the workers encourage her to oviposit in queen cups located on the face of the frame near the middle of the brood nest. Often these cups are repurposed worker cells and their wax darker in color as a result. Workers often raise three to five supersedure queens simultaneously. Sometimes these efforts are aborted prior to capping, but generally, once the cells are capped, the reigning queen is killed by workers balling her. Balling is a behavior wherein many workers surround the queen creating a tight sphere within which she overheats, suffocates, and dies. Rarely, but occasionally, the workers and the supersedure queens deign to coexist with their aged mother for a period.
Emergency queens are reared in crisis situations (such as when the current queen suddenly dies). In these instances, the workers, usually within four hours of their queen’s loss, select worker eggs less than three days old and quickly build queen cells (usually several at once) around them. The workers begin depositing royal jelly in the cells as soon as they molt into larvae. Because the workers must construct the cells wherever the appropriate eggs can be found, these emergency queen cells are often located within the preexisting brood nest on the face of the frame.
When a colony of honey bees decides it is time to reproduce, they start by making swarm queens: workers build numerous queen cups toward the bottom of one or several frames, often from the freshest and creamiest-looking wax and encourage the queen to lay in them over a period of days to weeks. The queen larvae are lavished with royal jelly, the cells elongated, and ultimately capped, resembling a peanut shell. The staggered ages and successive emergence of queens provide the colony with multiple opportunities to swarm when conditions are most favorable as well as insurance against queenlessness should one or more of the virgins be forfeit in some way.
Most colonies swarm as soon as the first queen cells are capped and the weather is conducive, taking with them the mother queen. Often, the swarm instinct having been activated, the colony will cast afterswarms, much smaller than the initial swarm, as sucessive virgins emerge over the next six to eight days.
Virgin queens, encouraged by their biology and goaded by their nestmates, must, if they are to become proper queens, take to the sometimes-savage skies in search of several swift trysts on the wings (several meaning an average of fifteen and swift meaning about two seconds from inception to completion). Depending on the success of her nuptial flight, a virgin may make one flight only or many (usually lasting around fifteen minutes) over several days.
Jurgen Tautz (The Buzz about Bees: Biology of a Superorganism, 2008), convincingly argues that queen mating flights are not lone endeavors but instead are aided by workers who mask her departure and return a cloud of circling bees. This diversionary behavior is said to look like orientation flights but has been found to be made up of older foragers who begin to orbit the hive as the queen prepares to depart, persist during her absence, and cease immediately upon her return. Additionally, Tautz suggests some older workers may actively guide the inexperienced virgin through the areal neighborhood, helping her get to DCA(s) and back as quickly and safely as possible.
Queens prefer DCAs two or more miles away, while drones typically join DCAs two miles or less from their home. DCAs are usually located above geographically identifiable landscape features such as a tall tree or ecotone (transition area where forest meets field) and are populated by thousands of male honey bees.
Upon locating a DCA, a journey aided by strong pheromonal signals, the queen is spotted, and the chase is on. The virgin flies fast and high. When she is caught or allows herself to be caught, insemination is a brief affair. Copulation complete, the drone tumbles through the air, expiring and the no-longer-virgin queen flies on toward home or to mate again. Successful mating will involve her taking in seventy to one hundred million spermatozoa, ninety percent of which she will discard, leaving five to seven million to live in her spermatheca.
Two to three days after returning from her final mating flight the queen begins to lay eggs. Typically, she lays for five to fifteen minutes, pauses for about the same amount of time and resumes. During times of intense colony building, she can maintain this rhythm perpetually, night and day, as she is surrounded by a revolving (each attendee ministering to her for about a minute and then leaving to circulate her pheromones) retinue, who anticipate and meet her every need. Queens can lay up to (and even exceed) fifteen hundred eggs a day. However, rarely do they maintain this pace and instead, in synchronicity with environmental conditions and hive resources, modulate the expanding and contracting population of their colony.
Queens choose the sex of the eggs they lay, basing their decision in part on the diameter of the cell they encounter (which they measure with their fore legs)—female eggs in smaller and male in larger cells. After laying the eggs, the queens’ maternal obligations are complete. Childcare is the responsibility of the workers. Her other essential function, which is largely a by-product of simply existing, is the production of pheromones (collectively known as queen substance) which act to maintain colony cohesion.