This series of queen introduction articles grew from time with observation hives, watching the fluctuation in the number of bees balling on the screen of the queen cages, and later on, even watching the queens and attendant bees inside the cages.
Figure 1 shows a queen inside her introduction cage –– in the moment of laying an egg. Her sting chamber is open and the egg has become visible. I had just installed the cage and queen. A laying queen, suddenly removed from her brood nest, cannot immediately stop laying eggs. Rather, she releases several eggs before fully stopping (see Figure 2). Now let’s return to the apiary.
To further explore the unknowns in queen introduction, we turn back to my first queen introduction work with attendant bees in the apiary. Before I began those queen introduction experiments, counting the number of ballers, described in the previous articles, I conducted a very small preliminary test run. Proceeding carefully, I needed to ensure my procedure of counting the bees balling the queen cage in the apiary worked properly, as modified from the observation hives in the bee house.
For this test run, I only had two queens purchased directly from a queen producer at the Eastern Apicultural Society (EAS). I followed the dequeening and setup procedures of the previous articles except that I began the test run in the afternoon and took the first count on the following morning. For both queen introductions, I removed the attendant bees. The results were both expected and surprising.
Figure 3 shows the time series of the baller counts from both colonies, designated Hive 128NA and Hive 60NA, which began on August 8, 2000. (I am retaining the NA notation for “no attendants” to be consistent with the previous article in this series.) Hive 128NA, shown in orange, has a fairly typical transition to zero balling (B = 0), indicating a provisional queen acceptance without attendant bees. The baller count initially began from the low twenties. Queen acceptance occurred by the third day, a little longer compared to some of the quicker transitions absent attendant bees that we have observed earlier. (The previous baller count was 2 at 64.5 hrs or 2.7 days, not quite queen acceptance.)
If the other colony in the test run, Hive 60NA, had shown a similar decrease in the number of ballers to queen acceptance, I would have been merely satisfied. See how the graph for Hive 60NA, color-coded in black, contrasts dramatically with the one for Hive 128NA, keeping in mind there are no attendant bee effects here.
For all the unusual balling behavior to come, Hive 60NA began with only five ballers, much less than the 20 ballers initially counted on the queen cage in Hive 128NA. Instead of the baller counts showing an immediate decrease to zero or showing a modest increase before turning down, Hive 60NA showed fairly large oscillations in its baller counts from about the mid teens (17, 16, 16) down to single digits (5, 7, 1). During this time, I knew something stalled the acceptance of the queen in Hive 60NA. What was it?
For prolonged balling during routine requeening, the typical cause is a second queen in the hive. In general, however, be sure the environment and “hive” conditions support a second-queen conclusion. For example, the test run occurred in the summer during a dearth or marginal nectar flow. That breaks any connection to reproductive after-swarming, another possible time, though brief, with multiple free-roaming queens in the hive. Of course, the conclusion assumes no attendant bees since we know they alone can cause long periods of balling. Another possibility, which I currently think is more unlikely, but also more difficult to detect, is the colony should not show any symptoms of being taken over by a usurpation swarm. So far in my experience, usurpation has been occurring in the summer during dearth conditions. Dead bees in front of the hive and bees fighting inside of the hive would indicate a very recent swarm invasion. If a queen were being introduced (no attendant bees) during this time, my educated guess is that the bees would ball the cage relentlessly.
But wait –– basic beekeeping teaches a bee colony has only one queen. That is true –– most of the time. As one learns more bee behavior, important exceptions can occur to the basic beginning “facts.” Sometimes a colony has two queens, coexisting as a mother-daughter pair, during the supersedure of an older queen. The pair can remain together without fighting, for several weeks, something with luck a beekeeper can watch.
I have taken an old queen being superseded by her large colony in an apiary, and started an observation hive colony with her in my bee house. Strangely, curiously, the small observation hive colony will sometimes supersede the old queen again. The old queen, still laying, seems defective to the bees –– independent of colony size. (The new mother-daughter pair coexists on only a one-comb brood nest.) Even when I check these dual-queen observation hives most every day for the queen pair, invariably the day comes. The old queen has vanished without a trace, leaving me wondering what caused her abrupt elimination.
My counting procedure of the bees balling the queen cage had the top-bar colony opened for a bare minimum of time, usually less than four or five minutes. I just needed to remove a few combs from the front of the top-bar hive, entering the front part of the brood nest until getting to the comb with the attached queen cage. I mostly ignored the comb contents –– including the brood.
Motivated by suspicions of a second queen in Hive 60NA, I began briefly scanning the brood combs as I removed them, when working my way to the comb with the queen caged, into about the front third of the brood nest. Then, at 97.5 hrs, a bit above four days, I found –– a mere three worker eggs scattered among the older brood.
Carefully explaining the origin of the eggs, even a few of them, was critical. Beginning beekeeping literature typically reports three days for worker bee eggs to hatch. Saying the eggs found at four days could not come from the removed queen because worker bee eggs hatch in three days is not the correct reasoning, although it sounds correct. Reporting three days for the hatch time is another oversimplification, although useful as an introduction to beekeeping, not to overwhelm beginners with too much information. Again as one’s understanding of honey bee biology progresses, those beginner bee facts need reexamination to understand more complexity of bee life (like only one queen in the hive).
In advanced literature, it is well known that worker bee hatch times vary. Turning back to a little more than a century ago, to 1915, we find Nelson’s herculean monograph, “The Embryology of the Honey Bee,” an intense 282-page description, including detailed sketches of the embryonic transition from egg to larva. All that complicated development occurs during a reported time of 74-76 hrs (3.01-3.17 days).1 Far more recently, we have Collins’ (2004) scientific research and her fascinating pictures of hatching bee eggs. She reported egg hatch variations from two sources. First, eggs from individual queens varied in their hatch times. Secondly, upon averaging the hatch time of eggs from different queens, those queen averages were not the same number. But rather, the hatch time averages of different queens themselves varied, which was the hatch time variation due to the queens.2 So for example, if a queen lays 100 eggs, the hatch times of those 100 eggs will vary. But that is not all of the hatch time variation. If, say, for 10 queen bees, we averaged the hatch times of each of their 100 eggs (randomly sampled) –– some of those 10 averages would be so different from the other averages that it could not be explained by some random occurrence. (They were statistically significant as Collins found.) Think of it as: Some queens had slightly faster-hatching eggs and other queens had slightly slower-hatching eggs.
A proper biological argument takes into account this natural variation in the hatch time, something like this: Even from the ….