What Queen Quality Actually Means, and How to Measure It

When you look at reasons beekeepers give in surveys for colony deaths, ‘queen issues’ feature prominently. We also hear ‘old timers’ saying that queens aren’t what they used to be, but don’t old timers always think things were better in the past? I wonder when I will become an old timer? Probably after I’ve kept bees for 20 years, which will be in six years time. Anyway, even I, in my quiet corner of obscurity, have noticed that in some years my queens don’t perform as hoped. I don’t just mean in terms of productivity, but earlier than that; things like early supersedure and being poorly mated, or disappearing from a mating box. The point of this article is to look at what queen quality actually means, and how to measure it.

What Queen Quality Is Not

The job of the queen is to lay eggs for her colony so that it can, in the words of Mr Spock, “live long and prosper.” Behavioural traits expressed by the colony, such as defensiveness and swarming tendency, are influenced by the queen, but those things are also very much down to the drones that she mated with. Those calm, productive colonies that my friend Paul Horton uses – which seem to regard swarming as beneath them – came about because the breeder had complete control of the drones. I don’t have such control, and nor do most beekeepers.

So, even though we beekeepers tend to blame the queen for everything, I don’t think it’s that simple. There are often multiple patrilines in the colony, and it is the combination of queen DNA + drone DNA + environment that determines many behaviours. A defensive colony may be so because of nutritional shortage, poor weather, or being regularly bothered by livestock (as well as who their Mum and Dad are). Or it may be because one patriline from particularly nasty drones dominates¹.

Quality in the strict sense means reproductive competency. Will this queen produce enough fertilised eggs, for long enough, to sustain a productive colony?

Markers of a Quality Queen

Body weight at emergence

Under normal developmental conditions, queen body size at emergence is a reasonable proxy for reproductive potential. Larger queens tend to have larger spermathecae and store more sperm. Collins and Pettis (2004)² demonstrated this directly, finding that in normally reared queens, body weight correlated with both spermatheca volume and sperm count. I think the consensus seems to be that 200 mg+ is a heavy queen, and therefore likely to be a ‘quality queen’. My breeder queen for this year, from Andy Sweet, weighed 221 mg. More importantly, she got through the winter and is heading a strong and healthy colony.

The important caveat is that this relationship between heavy queens and high quality can be disrupted. The study found that miticide (coumaphos) exposure during development broke the weight-to-spermatheca correlation entirely, meaning a queen can look physically normal (or big) while being reproductively compromised in ways that will only become apparent months later.

Other studies have shown a similar effect by tau-fluvalinate (Apistan) but the evidence for amitraz (Apivar) is more mixed. The synthetic miticides linger in wax for years, which casts a shadow on their use when other effective solutions exist. Having said that, I do use amitraz as part of integrated pest management, but maybe I should reserve that for emergencies only. I’ll have to think about that, especially as I had queen issues last season. Many more of my queens than normal failed to mate, or were quickly superseded.

Tarpy et al. (2011)³ went further, showing that queens produced from older larvae were smaller and mated with significantly fewer males, with downstream consequences for the genetic diversity of their colonies. So, the real problem was that older larvae were used to make queens, but it showed up as smaller-sized queens, meaning that ‘queen body weight at emergence’ is a good indicator of quality. This also ties in with my previous post about nutrition, in which I referred to grafting from larvae that are in ‘lakes’ of brood food, rather than those that look a little ‘dry’.

Sperm Count and Viability

A naturally mated queen typically starts off with 4–7 million sperm stored in her spermatheca. Three million sperm or fewer would be what most researchers consider to be ‘poorly mated’, often citing a 1962 paper by J.Woyke, which I can’t find. Not all sperm will be viable, which is what makes mating with multiple drones such a clever idea; you get genetic diversity plus reduce the risks of holding too much dodgy sperm from the odd male with low fertility. A good source for this sperm count and viability stuff is Delaney et al⁴.

Beekeepers tend to know when a colony is headed by a good queen or a poor one. I think you can tell a lot by the brood pattern. This was covered by a study⁵ that compared beekeepers’ rating of queens to the viability of the sperm in their spermatheca. It found: “Queens removed from colonies rated in good health averaged high viability (ca. 85%) while those rated as failing or in poor health had significantly lower viability (ca. 50%). Thus, low sperm viability was indicative of, or linked to, colony performance.”

My ‘back of an envelope’ calculation on queen egg laying suggests that a queen lays about 225,000 eggs per year (lots in May/June, very little in winter). Apparently when fertilising eggs the queen is remarkably efficient, using only two sperm per egg⁶. Even with 3 million sperm at 70% viability, that gives a useful ‘laying life’ of 4.67 years. However, what seems to happen is that the sperm viability declines over time, so the higher it is at the start, the better. Queen failure would be well before 4.67 years because of declining sperm viability rather than running out of sperm. We know that, in addition to time, pesticides, and high temperatures can also mess with sperm health.

Plenty of Daddies

A queen that mates with more drones produces a more genetically diverse colony, and there is solid evidence this is important for colony resilience, disease resistance, and hygienic behaviour. Polyandry is not an accident of bee biology; it is good old evolution at work. For a queen rearer, this means conditions during the mating period matter enormously: drone availability and flight weather influence how many matings occur. I covered this in another post.

Ovary Development and Pheromones

A queen with well-developed ovaries and a full spermatheca produces a stronger mandibular pheromone signal, which suppresses worker ovary development and maintains colony coherence. This is a good thing. A poorly mated or virus-compromised queen produces a weaker signal, which can trigger premature supersedure attempts.

On the matter of pheromones, may I refer you to another really interesting post looking at when newly mated queens are caught, and how well they are accepted by their new colony. The Australian work by Rhodes and Sommerville showed clearly that queens caught at 21 to 28 days post-emergence had substantially better long-term survival than those caught earlier. A queen caught at 14 days is still mid-development in ways that are invisible to the beekeeper, but not to the workers she is introduced to.

Are Queens Really Getting Worse?

In a word, yes. But that is a grand generalisation. There are some great queens out there, even in 2026. It seems to me that the turning point may have been the arrival of varroa mites. People who remember the pre-varroa days go all misty eyed and dreamy when recalling those times.

There is an unfortunate Catch-22⁷ involving drones and varroa mites, or more pertinently, the deformed wing virus (DWV) associated with mite infestation. Being riddled with viruses tends to bugger up your drones – lower sperm viability, poorer ability to fly strongly – things which make your queens ‘poorly mated’ and therefore low quality. However, the use of synthetic miticides, which build up in the wax in the combs, is also associated with lower sperm viability. I make drone combs using foundation-less frames so that at least they are on clean wax (hopefully), and I use organic acids as my primary weapon against varroa. Furthermore, queens that spent their larval days in beeswax contaminated with coumaphos and fluvalinate had fewer live sperm.

An article in Biotechniques refers to a paper⁸ which states: “we observed that the median lifespan of caged honey bees has been declining in the US since the 1970s, from an average of 34.3 days to 17.7 days (48% reduction). In response to this, we again turned to historical record and found a relationship between this trend and a decline in the average amount of honey produced per colony per year in the US over the last 5 decades.”

So there you have it – the old timers are right. Nowadays, the practice of replacing queens after they have had one full laying season makes sense for commercial honey producers because most queens do not perform so well when given more time.

Update: see the comment by David Evans – perhaps the old timers aren’t quite so right after all!

What Can Beekeepers Do?

Beekeepers cannot count sperm or weigh spermathecae, but there are things they can observe and act on.

At the grafting stage: Only graft from larvae with plentiful brood food — lakes, not puddles. Reject dry larvae. This is nutritional quality control at source. There are plenty of challenges with producing quality queens, and starting off with poorly nourished larvae, or old larvae, is doomed to failure from the start.

At the emergence stage: Weigh your virgin queens if you have a precision scale. This sounds exotic, but a letter scale accurate to 0.01g will do it. Discard obvious runts. Queens below around 180 mg⁹ are more likely to have a smaller spermatheca and be poorly mated. If you would rather not be all nerdy and weigh queens, you can just look at them. With experience, you will know which ones look small.

After mating: The speed and pattern of early laying is informative. A queen who takes more than three weeks from emergence to start laying, or whose initial brood pattern is scattered, might have problems. But location, and therefore weather, makes a big difference. If the weather goes bad, they have to wait before going on a mating flight. A solid brood pattern in the first frame is a good indicator of adequate insemination.

Sperm viability You cannot measure sperm viability directly, but the fertilisation rate in the brood is a reasonable proxy. A queen producing more than a few percent drone cells in worker comb is showing signs of sperm depletion or poor mating. A good brood pattern is a comforting sight, and seems to be a sign that many things are working as they should.

The Drone Side of the Equation

As I have said many times, the drones are crucial. The bees love them too, so I often have a frame of drone comb in my colonies. I know it sounds silly, but I think the bees are happier when the sun is shining and drones are at large in the colony, as long as forage is available.

I like to have ‘drone donor’ colonies in and around my mating apiary, so that my virgin queens have a chance of mating with drones of my choosing. With open mating, it’s not foolproof, but worth it nonetheless. It’s important for the drones to be from a different queen-line than the virgins that you want them to mate with. I know drone congregation areas are a thing, and a fascinating one at that, but I know of commercial queen producers who see matings happen at the mating apiary, just a few metres above ground. Ray Olivarez in California told me about that, but he does have a LOT of virgins and drones in the same area at once. Perhaps drones get attracted to mating apiaries if there are hundreds of virgins present?

As previously stated, to give the drones a good chance of being super-sexy they need to be on clean wax and, ideally, any miticides used are organic acids (formic or oxalic) and essential oils (thymol) rather than the synthetic types.

Finally, to get good quality queens, you need to be producing virgins at a time when drones are plentiful and sexually mature. For my area, depending on what type of spring we get, that’s May and June.

Conclusion

Queen quality is all about having good-sized queens with large egg laying equipment (ovaries and spermatheca) and she must be mated with multiple drones of good quality (viable sperm). As beekeepers, nothing is certain, but we can shift the odds in our favour by applying the advice above.

References

1. Drone patriline determines which workers defend — shown clearly in European × Africanized crosses: DeGrandi-Hoffman et al. (1998). Nest defense behavior in colonies from crosses between Africanized and European honey bees. Journal of Insect Behavior 11: 37–45.
   https://doi.org/10.1023/A:1020862432087
2. Body weight correlates with spermatheca size and sperm count — and miticide disrupts this: Collins, A.M. and Pettis, J.S. (2013). Correlation of queen size and spermathecal contents and effects of miticide exposure during development. Apidologie 44: 351–357.
   https://doi.org/10.1007/s13592-012-0186-1
3. Smaller queens (from older larvae) mate with fewer males: Tarpy, D.R., Keller, J.J., Caren, J.R. and Delaney, D.A. (2011). Experimentally induced variation in the physical reproductive potential and mating success in honey bee queens. Insectes Sociaux 58: 569–574.
   https://doi.org/10.1007/s00040-011-0180-z
4. Delaney, D.A., Keller, J.J., Caren, J.R. et al. The physical, insemination, and reproductive quality of honey bee queens (Apis mellifera L.). Apidologie 42, 1–13 (2011).
   https://doi.org/10.1051/apido/2010027
5. Pettis JS, Rice N, Joselow K, vanEngelsdorp D, Chaimanee V (2016) Colony Failure Linked to Low Sperm Viability in Honey Bee (Apis mellifera) Queens and an Exploration of Potential Causative Factors. PLoS ONE 11(2): e0147220. https://doi.org/10.1371/journal.pone.0147220
6. Baer, B., Collins, J., Maalaps, K. and den Boer, S.P.A. (2016), Sperm use economy of honeybee (Apis mellifera) queens. Ecol Evol, 6: 2877-2885. https://doi.org/10.1002/ece3.2075
7. A “Catch-22” is “a problem for which the only solution is denied by a circumstance inherent in the problem or by a rule”. For example, losing something is typically a conventional problem; to solve it, one looks for the lost item until one finds it. But if the thing lost is one’s glasses, one cannot see to look for them – a Catch-22. The term “Catch-22” is also used more broadly to mean a tricky problem or a no-win or absurd situation. Source: Wikipedia.
8. Nearman, A., vanEngelsdorp, D. Water provisioning increases caged worker bee lifespan and caged worker bees are living half as long as observed 50 years ago. Sci Rep 12, 18660 (2022).
   https://doi.org/10.1038/s41598-022-21401-2
9. De SOUZA, D.A., HUANG, M.H. & TARPY, D.R. Experimental improvement of honey bee (Apis mellifera) queen quality through nutritional and hormonal supplementation. Apidologie 50, 14–27 (2019).
   https://doi.org/10.1007/s13592-018-0614-y