paths switch, with larvae chosen by the nurses to be a queen being fed to excess, with much of the jelly not being consumed until after her cell is “sealed.”
Although the common conception is that queen larvae are fed a special “royal jelly,” whereas worker larvae are fed “worker jelly” and pollen, this explanation is controversial, especially considering how logistically difficult it would be for the nurse bees which are constantly moving.1 The most parsimonious (but debatable) explanation is that the nurses adjust the proportions of the three components of jelly dependent upon the age and caste of the larva being fed. Those components are (i) a protein‐rich secretion from the hypopharyngeal glands, (ii) a lipid‐rich secretion from the mandibular glands, and (iii) the amount of nectar added. Queen larvae receive not only a far greater amount of jelly, but higher proportions of lipids and sugars (Winston 1987; Wang et al. 2016). Queen larvae thus grow more rapidly than do worker larvae, to a greater size, and emerge at an earlier age.
Figure 5.1 The queen functions not only as the ovary of the honey bee superorganism, but also as the pheromonal “heart” of the hive, critical for colony cohesiveness. In a rapidly‐growing colony, roughly a thousand of her daughters die each day from natural aging and mortality. Thus, in order to maintain colony growth, the queen must not only lay an adequate number of eggs to replace those lost workers, but also enough for population increase, as many as 1500 a day.
Figure 5.2 Whenever a “good” queen pauses on the comb, an ad hoc group of adjacent nurse bees will turn to face her, offering her food, and antennating her to pick up her pheromones (Collison 2017). This ring of attendants is called a “retinue.” Since the advent of varroa, some are suggesting that queens these days don't seem to attract retinues the way they used to. It is possible that this may correlate with what appear to be reports of greater rates of queen failure.
Figure 5.3 A well‐fed queen is an egg‐laying machine, capable of producing an egg per minute, 24 hours a day (and up to double that rate in bursts). If she cannot locate an empty cell, she will just drop the egg (as this queen is doing), which will be consumed and recycled by a nurse bee.
Queen Diet
Similar to workers and drones, upon emergence, a young queen may consume nectar or honey, but unlike them, she does not consume pollen as a protein source,2 instead depending upon receiving a diet of jelly begged from nurse bees for her entire life. A queen can lay more than her body weight in eggs in a day, and thus requires an exceptionally nutritious diet. The jelly diet is so perfect that a queen's feces look like droplets of water. Due to this diet, queens carry a different endosymbiotic microbial community structure than do workers (Anderson et al. 2018), and typically live to a much longer age (Figure 5.3).
Queen Mating
A queen is not really a queen until she has successfully mated, and is barely recognized by the workers until then. A few to several days after emergence, a “virgin queen” (Figure 5.4) flies out to mate with about 15 drones (often more) in what is known as a “drone congregation area,” during a one to two day period when the temperature is above 70 °F (Figure 5.5). She returns to the hive exhibiting the torn endophallus of the last drone with which she mated, which must be removed by the workers (Figure 5.6). The workers then begin paying attention to her (Figure 5.2) (Richard et al. 2007), and she shifts her pheromonal output to signal that she is adequately mated. A few days later (typically 10–14 days after emergence, weather permitting), the queen begins to lay eggs (Figure 5.7).
Practical application: It's important to know that if a virgin is constrained from mating by weather, that the chance of her ever successfully being mated decreases greatly after three weeks.
Figure 5.4 To the untrained eye, virgin queens are difficult to spot. Look for her longer legs, slightly more angular “hips,” and rapid movement atypical of the rest of the bees on the comb.
Figure 5.5 The leading edge of a “drone comet” chasing a “virgin” queen. At the top left you can see a drone starting to mount the queen. Slightly lower is what appears to be the previously‐successful drone paralyzed and falling to his death after his explosive ejaculation.
After mating, the queen homogenizes the received semen, and discards roughly 95% of it, holding the remaining mixed spermatazoa in a clear sac called the spermatheca, in which the spermatozoa can remain viable for years.
Practical application: Temperature extremes, or certain insecticides and beekeeper‐applied miticides, may diminish the viability of the spermatozoa, causing early failure of the queen. The seminal fluid received may confer some immunity to pathogens (as well as pathogen exposure), and affect the spermatozoa of other drones.
Figure 5.6 A freshly‐returned no‐longer‐a‐virgin exhibiting “mating sign” – the torn endophallus of the last drone to successfully mate with her. This will soon be removed by the workers.
