Like most topics in beekeeping, the supersession of queens has long been and still is a subject of controversy. Although there seems to be no particular mystery about the process of supersedure itself, by which a daughter-queen and her mother-queen can remain together in the hive for quite extended periods, the reasons why this might occur and the desirability of allowing it are strongly contested. I decided to look over some older books to find out quite what is so disputable in the commonplace, yet extraordinary, process of queen supersedure.
About seventy years ago, the controversy was already strongly argued between experts who regarded supersedure queens as inferior, and those who thought they were often superior to queens introduced by the beekeeper. For instance, both R.O.B. Manley (of the Manley frame) and E.B. Wedmore, who is still highly regarded today, consider that supersedure queens are less favourable for the beekeeper than the introduced queens, though for different reasons. Wedmore believed that a supersedure queen was likely to be produced in unfavourable circumstances and therefore less productive. (Successful Beekeeping, 1946) Manley, on the other hand, argued:
‘…It would seem at first sight that a strain of bees in which the supersedure instinct is strongly developed would be a very desirable one, but in my experience this is not so. I have not found that such stocks and their queens are up to average; but it is only right to say that there are other beekeepers who think differently. One must not forget that the establishment of an instinct of supersedure sufficiently strong to provide for a methodical renewal of queens by that process, would obviate the necessity for swarming as a method of queen replacement, and that any strain of hive-bees that ceases to swarm altogether is, in a state of nature, doomed to die out altogether, so that supersedure must be regarded, in so far as it is a desirable trait, as complementary and not alternative to swarming.’ (Beekeeping in Britain, 1948)
It is perfectly reasonable to argue that, without colony splitting to establish new colonies, the bees would ultimately risk dying out. Hooper argues this exactly in his Guide to Bees & Honey (rev. ed.1991).
However, at the same period we find another writer who claimed:
‘…This supersession seems to become more assured with heredity: some of my colonies have not swarmed or been requeened by me in five years, and have consistently given very high yields. Cut-down queen cells have often been found on their combs when they have not swarmed. This system is so simple that many bee-keepers will treat it with profound mistrust; but they should try it before condemning it. For three years I have had no swarms in four apiaries, and some hives under treatment have not swarmed for five or six years.’ (F.S. Stuart, Beekeeping Practice, 1947)
I have found that the same basic hypotheses and disagreements seem to have continued in the years since these books were written, with very little clear or new information about the scientific facts of supersedure. For example, I have been unable to establish that heredity (or as we now say, genetics) does really play a clear role in predisposing bees to supersede their queens. One persistent and logical opinion is that the colony uses supersedure as a means of compensating for an injured queen:
‘…It may even be true that the bees are slightly more likely to supersede a clipped queen, because she in some way seems older to them. Manley thinks this is so, if one of the wings is severely clipped, but not if both wings are slightly and equally clipped. If it is so, there is just a trifle more probability that a clipped queen will swarm than an unclipped queen, since the desire to supersede is one of the motives behind swarming. But there is hardly enough in it to influence your decision whether to clip your queens or not…’
(Kenneth C. Clark, Bee Keeping, Penguin Handbook 1951)
A couple of American beekeepers are even more insistent on this explanation:
‘Hybrid queens should never be clipped as this causes immediate supersedure in most cases! The bees think their queen is injured.’ (Ormond & Harry Aebi, Mastering the Art of Beekeeping, 1982, USA)
So it seems there are definite ‘for and against’ camps among expert beekeepers. Perhaps the most emphatically stated case for introducing specially selected queens comes from Brother Adam:
‘The same applies to queens which are being superseded – a sure indication that their life’s strength is at an end. As a matter of fact. I have never come across a supersedure queen whose performance equalled those raised from eggs derived from a breeder in her prime and whose laying abilities have been restricted in the way I have indicated. I am aware this sounds all very unorthodox and contrary to the commonly held views, but our comparative tests leave no doubt on this point. Indeed, for many years now we have replaced any supersedure queens found in the honey-producing colonies in the spring.’ (Brother Adam, The Bees at Buckfast Abbey 1950)
In the opposing camp, however, the natural beekeepers line up in favour of supersedure:
‘…Beekeepers have long known that colonies often supersede emergency queens with normal ones. The two types of queen may look the same but the bees can tell the difference. All commercial queens sold as queens are emergency queens, only they are usually raised in round cells from the outset after grafting the larva into a plastic cup. Alternatively they may restarted by confining the queen in a plastic cage mounted on a comb in the colony so that she is presented only with an array of plastic cell ‘plugs’ in which to lay. The plugs are then transferred to plastic holders that maintain them in the normal vertical orientation in which queen cells are constructed, and placed in a queenless colony. In order to introduce a queen resulting from this process, the colony needs to be brought to the point of ‘desperation’ to receive her…’ (David Heaf, The Bee-Friendly Beekeeper, 2010)
You will have seen by now that the underlying question for beekeepers concerns the excellence or otherwise of the new queen herself. On the one hand, some breeders definitely discourage the ‘natural’ and unmanaged process of supersedure and prefer to select queens from stocks which are quiet, productive, hygienic, and so on. On the other hand, a different philosophy of beekeeping encourages the idea that the bees know best and they will produce excellent queens by supersedure when they feel the need.
There seem to be virtues on both sides of this controversy. It seems to me that most of the good beekeepers I know have reckoned on this and they tend to hedge their bets; so although they may use queen-rearing as such, they are also delighted to find a supersession queen working away in their hives. This is not uncommon after winter, for example; when they open the hive in springtime, they find their marked queen may still be there but an unmarked supersession queen is happily working beside her mother. The peacefulness of this arrangement has often been noted: it seems to run counter to the beekeeper’s expectation that the stronger queen will kill off any rival very quickly. Various suggestions have been put forward to explain the peaceable nature of supersedure: the diminution of the old queen’s pheromones, leading to her being tolerated as a harmless relic, perhaps? Or again, there may even be some question of patriline influence here, as seems to be the case with certain ant-societies where the recognition of kin relatedness gives rise to special treatment in the colony. Some research findings about honey bees have recently been put forward, showing that worker bees may preferentially feed their nearest relatives in the hive (full sisters with same father as well as mother). Until this aspect is teased out, the mystery remains.
However, maybe there is a clue in an article published in a Dutch journal in 2004: the researchers aimed to stimulate supersession artificially, in order to reduce colony losses overwinter. The team, led by Harmen P. Hendriksma, hypothesised that, if they could induce colonies to supersede early enough before the winter set in, they would enable them to survive the winter. They tried introducing queen cells containing 1-day-old larvae or capped queen cells into 50 needy hives, but although the larvae were fed at first, none was reared to maturity and not one of the colonies performed a supersedure. They concluded that ‘supersedure cannot be evoked by artificially by-passing the initial phase of the process.’ The inference is that the ‘initial phase’ is determined by the super-organism which is the colony itself. I think that this inference is consistent with a different study which found that ‘neither swarming nor supersedure can be predicted by prior queen replacement events in a colony’. (M.H. Allsopp & M.R. Hepburn, ‘Swarming, Supersedure and the Mating System of Apis mellifera capensis’, Journal of Apicultural Research, 1997). The interesting thing here may be that there could be much more to supersedure cells than we can yet understand.
At something of a loss to decide which side of the controversy I should stand, I thought it might help if I looked at related insects, and this confirmed that ants and wasps may also supersede their queens, as do some forms of stingless bees. I also found that some species of ant regularly support several queens working together in their nests, giving them a population advantage over less prolific types of colony. Yet another variation was found in a stingless bee (sweat bee) in USA where the single species could avail of two different strategies: it could either retain the queen throughout the season, leading to a ‘eusocial’ structure where all the workers are daughters of the queen and they raise their own sisters and brothers; or it could at need raise a supersedure queen, leading to a ‘parasocial’ structure where the workers were in fact sisters of the queen and therefore raise nieces and nephews. Honey bees are ‘eusocial’ and this confers a closer relationship among the members of the colony, with advantages which are beginning to be understood. However, perhaps the potential to supersede a queen when necessary is retained at a very profound evolutionary level. I also found an old, very interesting paper about bumblebees by a Russian entomologist, G.S Voveikov, called ‘Natural Requeening of Bumble Bee Colonies’ (1953). He shows that bumble bees can supersede their queens in a single season, and believes that ‘a two-fold supersedure, with two events at different times, appears very favourable for colony growth.’ He even proposes that the early season requirements of the colony can differ so much from the later ones that this change of queen is highly beneficial. [He also considers that this predisposes bumble bees to evolve cuckoo-bee species as well as true bumble bees.] For example, he argues that the overwinter queen may be exhausted by founding her new colony, and that a supersedure queen (probably a heavier bee) could lay with renewed vigour. I was reminded of Brother Adam’s contention that a well-bred queen, selected by him as the beekeeper, would be more vigorous and productive than a supersedure queen with uncertain breeding. How odd, that in bumble bees this might be considered exactly the reverse and supersession seen as an advantage to breeding. But then, bumble bees have no beekeepers!
Meanwhile, the best guideline, whichever side of the controversy you may prefer, would seem to be seasonal. There is consensus that at the early and the late part of the beekeeping season, you may well find a few supersedure cells in your hive. There will be far fewer than swarm cells at the height of the season, but you may see no other difference; some say that supersedure cells tend to be in the middle of the frame, but other writers deny this. In the early part of the season, the chances of a supersedure queen being properly mated are less good than at the height, and you may find that the bees will replace her during the season. In the late season, the bees may replace a failing or elderly queen in order to survive the winter. There seems to be some indication that there is no true ‘broodless’ period in our winters nowadays, and so perhaps late supersedures are a reversion to an evolutionary potential. This might be compared to the recent understanding of ‘hygienic’ behaviour in honey bees, which seems to be called forth under the extreme genetic ‘bottleneck’ effect of colony losses from varroa (as Professor Tom Seeley has argued). Perhaps one could say that this extreme environmental stress has brought out an evolutionary defence which has not been apparent until now. I wonder whether supercedure represents another underlying evolutionary trait, to complement the normal swarming strategy?
I will finish by quoting a clear and useful summary from Pam Gregory, Healthy Bees Are Happy Bees (2013)
‘…Really, it is the time of year that indicates whether the cells are swarm or supersedure cells….
There are two types of supersedure: perfect and imperfect. In a perfect supersedure the old queen does not die until the new queen in mated and laying and then she just seems to politely fade away. This is the only natural circumstance when two queens will be found in the hive together. In an imperfect supersedure the original queen dies before the new queen’s mating process is complete. This is then a risk for the colony’s survival because if the new queen is not well mated or does not return to the hive, the colony is unable to raise another queen. Lack of mating will not prevent a new queen from coming into lay, but of course, it will prevent her from laying anything other than drone eggs.
In my opinion, supersedure is rather a desirable trait to encourage, as the colony produces a new queen as well as collecting a honey crop, so beekeepers should always be aware of this possibility when thinking about replacing and old queen. Maybe the bees would do it better!’