emergence (dependent upon weather), a queen will be mated and commence laying the first of the half a million eggs that she may produce over her lifetime. Each cell in the above photo contains a single egg.
Practical application: Many novice beekeepers, resistant to put on their reading glasses, are unable to identify eggs in the combs. To see eggs, face toward your shadow, which will allow sunlight to come from behind you to illuminate the bottoms of the cells. Don't confuse white reflections with eggs – eggs look like white grains of rice standing on end.
Be aware that an egg may carry virions on its shell (notably Deformed Wing Virus), but inside carry proteins that confer transgenerational immune priming to her offspring (Salmela et al. 2015).
Queen Performance
A queen's performance is mainly measured by how eggs she lays each day.
Practical application: A queen's potential performance is often throttled by the number of prepared, thermoregulated, empty brood cells available in the hive. A good queen cannot exhibit her full laying capacity until the cluster covers at least 10 deep combs, and even then she would be limited if there were appreciable amounts of honey or beebread in those combs. The exceptional queen can nearly completely fill 10 deep combs with brood.
Young queens are typically more “exuberant” in their egg‐laying than are older queens (newly‐mated queens may even lay multiple eggs in a cell if there is not adequate room in the cluster). Young queens as a rule outperform older queens, although many queens are highly productive in their second year (which then often leads to swarming).
The performance of a queen is based upon a few main factors (listed in approximate order of importance):
1 How well she was fed and cared for during her larval development, which determines her egg‐laying capacity, 3
2 How well she was mated – the number of viable spermatozoa in her spermatheca, and
3 Environmental factors, such as chilling and heating during shipment, or pathogen or pesticide exposure in the colony.
A queen's potential performance is largely fixed by how lavishly she is fed jelly during the approximately four days between the emergence of her larva from its egg until her cell is sealed (she continues to feed and grow after sealing). And then she must get properly mated.
Queen Performance vs. Colony Performance
The queen's contributions to colony “colony performance” are:
1 The number of eggs that she produces,
2 Her genetics, which then supply half the genetics to each of her daughters – the workers,
3 The genetics of the guys she mated with – the other half of the genetic equation, and
4 Some passing on of transgenerational epigenetic immunity.
After that, it's up to her offspring. Colony‐level performance is mostly the result of how well the genetically‐diverse patrilines of workers work as a “team.”
Practical application: In general, a well‐reared, well‐mated queen of average genetics will outperform a poorly reared, poorly mated queen of the best genetics. After that, it's largely a matter of chance of how the patrilines of daughters happen to function together as a “team” (Figure 5.8), “performance” being more the result of the workers than the queen.
Practical application: Unlike other livestock, honey bee queens are polyandrous, resulting in a colony of bees consisting of a mix of patrilines of half‐sisters, each fathered by one of the many drones with which their mother mated. Thus, there will always be genetic differences and diversity, even with colonies headed by sister queens. The performance of each colony as a whole is thus a matter of chance, similar to putting together a sports team of players (each patriline of worker bees) without knowing how well they will perform as a group (the colony). Bottom line: you might advise beekeepers to start in spring with twice as many colonies as they hope to take through the winter, and not waste their time on those that don't perform well. Some commercial beekeepers (the author included) simply cull any poor performers.
Figure 5.8 A histogram of colony performance (weight gain) of 35 colonies, started with sister queens mated and managed identically in one of the author's yards. Note the normal distribution of performance.
Practical application: On the other hand, many a poorly‐performing colony can completely turn around once it clears itself of disease, or enjoys better nutrition. If honey production is not the beekeeper's main goal, they may find joy in helping a struggling colony to get back on its feet.
Queen Longevity
Beekeepers often refer to a queen's “age” in years. But biologically, chronology has little to do with it – a queen's “age” is a function of the number of viable spermatozoa remaining in her spermatheca, or more specifically, her ability to fertilize each egg laid in a worker cell (Baer et al. 2016).
Practical application: Thus, in a colony in a commercial operation in California or Florida, supplementally fed for near year‐round broodrearing, a queen may begin to run out of spermatozoa in the late summer of her second year. On the other hand, a queen in a cold‐winter area, in which she rests for five months of the year, may be productive for several seasons.
Queen Succession
A colony of bees is theoretically immortal, but not so its queen. In the natural state, with colonies living in tree cavities, the queen is replaced at least once a year (due to swarming). And prior to varroa it was not uncommon for a relatively unmanaged hive to live for many years, also replacing its queen without help from the beekeeper.
Practical application: There are three ways in which a colony can replace its queen, each due to different circumstances:
Supersedure – when the workers build a supersedure cell to smoothly replace a failing queen.
Swarming – in which a colony in a crowded cavity builds swarm cells, and then divides by fission, with roughly half the workers flying off with their mother, leaving behind daughters in those cells to take her place.
Emergency – when workers respond to the sudden loss of a queen by converting a young larva in a worker cell into an emergency queen.
There is much overlap involved in the three methods above; the vet needs to be familiar with each.
Supersedure
A
