Bee Buzz Box – October 2022 Honey Bee Colony Dynamics Part II – Queen Cell Signatures

Of the internal dark workings of the brood nest: Nothing to be seen here

Alan Wade

Ring out the old, ring in the new,
Ring, happy bells, across the snow:
The year is going, let him go;
Ring out the false, ring in the true.

In Memoriami
Alfred Lord Tennyson – 1809-1892

Last month we learnt about what to look for when inspecting the brood nest, the engine that drives the tractor that brings in the harvest. One of the legacies of trying to run bees with an extra queen is that I have had to pay far more attention to the relative performance of queens working in tandem. This is something we all do when comparing the performance of each of our hives – we are comparing queens – but it brings into sharp focus the need to only run hives with good queens.

What do we look for when we try to assess whether the queen in the hive is up to scratch. We have already had a shot at assessing how much brood she is producing taking into account the seasonal demands placed on her. What is less obvious is that bees may limp along quite happily because, though old and infirm, the queen may still lay the limited number of eggs required of her from late in the honey flow until mid spring. A good queen can do that at a canter.

Come mid spring and the equation changes. With an old queen (say 18 months or older) the colony will likely either swarm, never lay well enough for the colony to be productive or, as likely, be replaced. To my mind keeping such a queen is about as useful as appointing a politician to the New York trade commissioner office in the vain hope of making the taxpayer much richer.

Queen cell rules

You may get away with never requeening a colony. It may well survive – that is what wild hives do – but it is questionable if it is worth the effort of keeping bees if the sum total of your efforts is to get a measly two jars of honey out of your bees. Fortunately the bees realise this and more often than not will do the requeening for you that is, unbeknown to the beekeeper, simply supersede their queen. Butlerii presents a very clear picture of supersedure:

Queen supersedure, the process by which a colony of honeybees replaces its queen without swarming, is a [sic of] frequent occurrence … the process of queen supersedure differs radically from that of swarming by which colony reproduction is achieved, and in which one or more new queens are also reared and the old queen leaves the parent nest with a swarm if she is still alive.

The reason that sideline beekeepers often do so well in southeastern Australia is that there is a long time between the beginning of the buildup – by mid August in most districts – to when many major flows begin in around November. Such luxuries are not available in many parts of Europe, North America and northern Asia where critical early stimulus and very early colony establishment are essential.

However it is quite possible to lose the early momentum of building bees for the honey flow – for the lack of a good quality queen – where it is easy to miss a reasonable return. More critically bees may either swarm because the queen is old and cannot produce enough mandibular pheromone or fail to build. In any measure of productivity, the colony is a dud. And if the attempt of the bees to requeen itself fails the colony will die. As Tom Seeley so elegantly demonstrated of forest honey bee colonies in New York State:iii

…most (97%) survive summers, but only 23% of founder colonies and 84% of established colonies survive winters. Established colonies have a mean lifespan of 5–6 years and most (87%) have a queen turnover, probably by swarming, each summer.

Wild colonies or unmanaged backyard hives may survive for longer periods in more temperate climes but apiaries left untended will dwindle or at best languish unmanaged. That, in my experience, is what happened over several years of failing to attend bees in an out apiary when I were too busy trying to earn a more dishonest living.

Colony response to a queen lost to disease, predation or accident

All of this is a long way from reading what bees do when they take it upon themselves to replace their queen. We all know that if you squish a queen accidentally or divide a colony preparing to swarm that the now queenless bees will, within 48 h, attempt to start raising a new queen (Figure 1). We employ the emergency queen impulse in beekeeper initiated requeening practice speeding the process up by supplying a caged queen. Notice the flared modified worker cells with young queens fed more lavishly than worker larvae in adjacent cells. A fully developed emergency queen cell retains its origin in the cell matrix as you can observe by prising the cells from the comb. Most requeening schemes are predicated on removing the old queen first.

Rule 1 Since emergency queens result from worker bees repurposing newly hatched worker larvae the cell will originate in an area that the old queen last laid eggs. The signatures are either of flared worker cells or of a fully developed queen cell both leaving their telltale origin at the base of a former worker cell.

Such queens raised under less than ideal conditions are themselves superseded.

Little known is the fact that colonies that lose their queen may swarm, that is abscond, rather than raise a new queen. While a frame of young brood is often added to a freshly hived swarm to help ‘hold the bees’, the alternative scenario where a queenless split has brood but no queen the situation is quite different. Mark Winstoniv argues that while absconding will disadvantage bee survival – they lose built comb, stores and the windfall of developing bees that might add to their numbers – the workers themselves have no queen pheromone to exert the cohesion required to maintain normal colony functioning. Sometimes colonies raise a new queen, the course normally followed when strong colonies are split. Other times they may swarm in response to queen loss. A further and common outcome is that, after being queenless for two to three weeks, laying workers will commence laying unfertilised eggs on worker cell walls and the colony will become almost impossible to requeen. The signature then becomes one of scattered domed drone brood in worker cells, old worker bees and numerous small drones.

Figure 1 Emergency queen development showing modified worker cells, 21 October 2017
Alan Wade

Colony response to a failing queen

We now move on to the scenario where the colony, by dint of failure of queen pheromone signalling, decides to replace its queen in a more orderly fashion. Here new queens are raised near the centre of the brood nest (Figure 2). From the outset they are optimally nurtured so mated well they can head very productive hives.

Rule 2 Supersedure queen cells are attached to the surface of the comb so they will come away freely from the comb face. They are also well formed and, with good nutrition and an area saturated with high calibre drones, they will be as good if not better than can be raised by a queen breeder.

Figure 2 Supersedure cells newly formed on the comb surface of in the presence of an old and previously particularly productive Carniolan queen. Jerrabomberra Wetlands Apiary 31 October 2016

Note eggs in adjacent worker cells, clearly visible here, that will be absent in colonies preparing to swarm and that they are formed on the comb surface and immediately point downwards. As the supersedure queen cells form (Figure 3) their attachment to the comb surface inside the brood nest becomes even more apparent.

Figure 3 Supersedure queen cell development attachment to comb face amongst sealed worker brood where the queen was clearly failing, 2 November 2016
Photo: Alan Wade

Simple queen replacement can occur at any time of the year but is most prevalent when colonies are under breeding stress (Figure 4), that is whenever the queen cannot lay fast enough for number of nurse bees available to raise brood. The presence of a single cell – generally no more than three – signals that the colony is not preparing to swarm. Note also that in this instance the developing queen cell has been formed over a large patch of sealed brood so the old queen, though pheromone deficient, was still laying well.

This is well illustrated by a lesser know facet of supersedure identified by John Hoggv:

Queen supersedure is never the cause of swarming. But supersedure may occur concurrently with the swarm whenever the queen’s failure was caused by being overtaxed while generating the bees for that swarm. Significantly, it is the virgin queen that is then selected to accompany the swarm. The failed queen is retained in the parent [hive] to be superseded in turn

Apparently, just as the survival instinct of the bees in control won’t allow a swarm to leave the hive without a replacement queen in the parent [hive], they won’t allow a failing queen to issue with the swarm.

Figure 4 Single supersedure cell development mid spring, 20 October 2016
Photo: Alan Wade

Colony response to reproductive swarming

We are now left with a very different scenario where the colony has a further inbuilt recipe for queen renewal. We call this swarming. The prime swarm almost always departs with the reigning colony queen so the parent colony ends up with a new and more fecund queen and a much invigorated hive, important as the parent colony needs to be rebuilt. It is the bees, not the beekeeper (unless he or she tries to crowd too many bees into a hive with too little space), that makes the call to swarm.

Rule 3 Swarm queen cells are produced in abundance and are built from scratch. And like supersedure queen cells are not all produced at once: they are produced in sequence to accommodate the scenario where the replacement queen fails to mate or return to the hive. The other characterising feature of swarm cells is that they are located on the periphery of the brood nest, very commonly on or near the bottom bars of brood frames.

A common quick check technique for the presence of swarm cells is employed by commercial and savvy sideline beekeepers. They rely on examining the bottom bars of frames (Figure 5), especially in the upper box where double brood chambers are employed.

The alternative is to rely on snake oil salesmenvi claiming a cure all for swarming:

Now, brother and sister bee-keepers, it may cause you some surprise and it may awaken within you some skepticism when I make the bold broad statement that I can outline a method of treatment that will cure the swarming feature every time and all the time. It will work wherever bees can as it is founded upon an instinct of the bee which is infallible. The treatment will cure the swarming impulse if already acquired, or, if given before the bees think of swarming, will prevent swarm preparation. It is either a preventative measure or a curative measure, depending upon the time of giving treatment. It will and absolutely does prevent all swarming at all times and under all circumstances notwithstanding that old legend that ‘Bees Do Nothing Invariably.’

Figure 5 Swarm cells started at comb margin Jerrabomberra Wetlands Apiary, 12 November 2017
Alan Wade

Of all the colony and brood frame signatures that bees leave behind, swarming is the most obvious. The discovery that a colony that was building well is now suddenly depleted of bees is prima facie evidence that the colony has already swarmed. While emerging bees often quickly help fill the bee numbers gap, other evidence of swarming can be found in the presence of many opened queen cell caps or queen cells torn open at the side and stung by rival virgin queens. To confirm that swarming is in progress examine brood combs check to see if free roaming queens are present. We marked two such queens in a hive that had just swarmed a few years back and discovered an unmarked queen laying well a couple of weeks later.

Note that although there were few queen cells present in this instance (there are usually many more) that the cells are on the bottom bar and attached to the frame (often to the base of combs) and that while sealed worker brood is present the queen had long stopped laying. Workers from sealed brood would have helped the parent colony recover its population had the colony been allowed to swarm.

It is also worth noting that the occasional appearance of a queen cell is often unrelated to any intention of honey bees to replace their queen. Queen cell construction is promoted by worker tarsal pheromone and is often unassociated with swarming. Such queen cells are rarely used but I check them anyway and fellow beekeeper Dannielle Harden tells me that finding multiple eggs in such cells is a sure sign of the presence of laying workers.

Picking the difference

Are these queen cell signatures easy to recognise? That is hard to say as I have often had to look back at my notes and snapshots to assure myself that my diagnosis of queen replacenent was indeed correct. Picking the difference between different modes of queen replacement is not always straightforward but more often is in plain sight.

Since honey bee colonies are in most ways an expression of the colony queen, how a colony performs, disease and availability of flowers aside, can be sheeted home to the calibre of the queen. Your ability to read the bees and your willingness to maintain well bred and productive young queens in all hives at all times is one of the great secrets of beekeeping.

Ring out the old, ring in the new.


iTennyson, A.L. (1833-1850). In Memoriam, AAH.

iiButler, C.G. (1957). The process of queen supersedure in colonies of honeybees (Apis mellifera Linn.). Insectes Sociaux 4(3):211-223.
Butler, C.G. (1960). The significance of queen substance in swarming and supersedure in honey bee (Apis mellifera L.) colonies. Proceedings of the Royal Entomological Society of London A35:129-132.

iiiSeeley, T.D. (2017). Apidologie 48:743–754. Life-history traits of wild honey bee colonies living in forests around Ithaca, NY, USA.

ivWinston, M.L. (1991). The biology of the honey bee, pp.132-138.Harvard University Press.

viJones, H. (1909). A radical cure for the swarming habit of bees, 23pp.

vHogg, J.A. (2006). Colony level honey bee production: The anatomy of reproductive swarming. American Bee Journal 146(2):131-135.