“supersedure.” The main factors that determine when a queen gets superseded are when she starts to run out of viable sperm with which to fertilize worker eggs, or by colony stress.
Practical application: Bees appear to “blame” their queen if the colony becomes seriously stressed by disease or parasites, which then triggers supersedure.
Worker bees often build “cell cups” on the face of a brood comb (Figure 5.9), but this does not necessarily mean that they are preparing to supersede or swarm. An “aging” queen will “willingly” lay an egg in a prepared cup, at which point the workers may turn it into a queen cell.
Practical application: The presence of cell cups does not indicate that a colony is about to supersede their queen. Even if supersedure larvae initially get fed, that doesn't mean that supersedure is inevitable, since the colony will often tear the cells down before emergence. Russian bees are noted for their continual starting of supersedure cells.
A mature supersedure cells looks similar to a peanut stuck to the side of the comb (Figure 5.10).
Practical application: beekeepers often ask whether they should destroy supersedure cells. I tell them that the bees likely have a better idea of the quality of their queen than they do. I see no reason to remove supersedure cells. 4
Figure 5.9 Preconstructed natural “queen cups.” It is not unusual to see these in a colony. Their presence does not necessarily mean that the queen is about to be replaced. Even when there is an egg or a larva with jelly in a queen cell, they are often destroyed by the bees during that new queen's development.
Figure 5.10 A typical supersedure cell. Supersedure cells are generally found singly, as opposed to swarm or emergency cells, of which there are generally several. They are also generally found near the center of the brood area, rather than along the edges, as are swarm cells, or scattered, as are emergency cells.
Figure 5.11 A dissected supersedure cell after emergence, showing that the egg had been laid in a preconstructed cell cup (as for a swarm cell), as opposed to the cell being postconstructed from a worker cell (as for an emergency cell). We are not yet clear as to the proportion of supersedure cells that are produced by these two different routes. There is also some evidence that workers may be able to transfer an egg or young larva from a worker cell to a queen cup (Punnett and Winston 1983).
Not all supersedure cells come from prepared cell cups. Some that I have dissected originated from worker cells (Figure 5.11), so the line between supersedure and emergency cells may be fuzzy.
Figure 5.12 Two laying queens on the same comb face (the lower one is a bit above the finger). On the other hand, I occasionally see a failing queen being “balled,” suggesting that her own workers are killing her.
Although sister queens will invariably fight to the death, an emerged daughter queen and her mother often peacefully coexist.5 It is not uncommon to observe mother and daughter laying eggs side by side for days or weeks, with the mother eventually disappearing from the hive (Figure 5.12).
Practical application: Although it is generally the case, never assume that there is only a single queen in a hive. The only way to be sure that all queens have been removed is to shake the bees through an excluder “sieve box” (Figures 5.13and 5.14).
“Balling” of the Queen
In the process of supersedure, the workers often show no mercy toward a no‐longer‐wanted queen. Bees kill a queen in the same way that they kill invading hornets, by biting her and forming a tight, heads‐in, walnut‐sized ball of bees around her (Figure 5.15). The bees then produce heat and direct it to their heads to “cook” the queen. Thus, even if one manages to “save” a queen being seriously balled, she rarely survives.
Practical application: There is great demand for “early mated” queens. But the weather does not necessarily cooperate, resulting in a percentage of early queens being poorly mated. And during shipping, temperature stress can cause loss of viability of the spermatozoa. Whatever the reason, many beekeepers report that newly‐introduced, or package bee queens, get superseded within the first month (this is not necessarily a problem).
Figure 5.13 Shaking bees through a sieve box to recover any queens. This is important when setting up a cell‐builder colony, as above, in order to mass‐produce emergency queens from selected mothers.
Figure 5.14 The thorax of a queen (center) or drone (lower right) is too broad to pass through a queen excluder.
Queen “Failure”
“Queen failure” is a nebulous and poorly‐defined term, nowadays given in surveys as a choice for the cause of colony mortality. As such, it is often listed as the most common cause of colony loss by beekeepers (BIP 2019). This is somewhat surprising, since “back in the day,” colonies tended to quietly and efficiently replace their queens via supersedure without any help from the beekeeper, and when I review older beekeeping textbooks, the term “queen failure” isn't mentioned.
