Spotty brood ≠ failing queen

I thought I’d discuss real beekeeping this week, rather than struggle with the high finance of honey sales or grapple with the monetary or health consequences of leaving supers on the hive.

After all, the autumn equinox has been and gone and most of us won’t see bees for several months ūüôĀ

We need a reminder of what we’re missing.

Beekeeping provides lots of sensory pleasures – the smell of propolis on your fingers, the taste of honey when extracting, the sound of a full hive ‘humming’ as it dries stored nectar … and the sight of a frame packed, wall-to-wall, with sealed brood.

Brood frame with a good laying pattern

This is a sight welcomed by all beekeepers.

Nearly every cell within the laid up part of the frame is capped. All must therefore have been laid within ~12 days of each other (because that’s the length of time a worker cell is capped for).

However, the queen usually lays in concentric rings from the middle of the frame. Therefore, if you gently uncap a cell every inch or so from the centre of the frame outwards, you’ll see the oldest brood is in the centre and the most recently capped is at the periphery.

It’s even more reassuring if the age difference between the oldest and the youngest pupae is significantly less than 12 days. Hint … look at the eye development and colouration.

This shows that the queen was sufficiently fecund to lay up the entire frame in just a few days.

What are these lines of empty cells?

But sometimes, particularly on newly drawn comb, you’ll see lines of cells which the queen has studiously avoided laying up.

That'll do nicely

That’ll do nicely …

It’s pretty obvious that these are the supporting wires for the sheet of foundation. Until the frame has been used for a few brood cycles these cells are often avoided.

I don’t know why.

It doesn’t seem to be that the wire is exposed at the closed end of the cell. I suspect that either the workers don’t ‘prepare’ the cell properly for the queen – because they can detect something odd about the cell – or the queen can tell that there’s something awry.

However, after a few brood cycles it’s business as usual and the entire frame is used.

Good laying pattern ...

Good laying pattern …

All of these laid up frames contain a few apparently empty cells. There are perhaps four reasons why these exist:

  • Workers failed to prepare the cell properly for the queen to lay in
  • The queen simply failed to lay an egg in the cell
  • An egg was laid but it failed to hatch
  • The egg hatched but the larvae perished

Actually, there’s a fifth … the cell may have been missed (for whatever reason) but the queen laid in it later and so it now contains a developing larva, yet to be capped.

What are all these empty cells?

But sometimes a brood frame looks very different.

Worker brood 1 is present across the entire frame but there are a very large number of missed cells.

Patchy brood pattern

Patchy brood & QC’s …

Note: Ignore the queen cells on this frame! It was the only one I could find with a poor brood pattern.

This type of patchy or spotty brood pattern is often taken as a sign of a failing queen.

Perhaps she’s poorly mated and many of the eggs are unfertilised (but they should develop into drone brood)?

Maybe she or the brood are diseased, either reducing her fecundity or the survival and development of the larvae?

Sometimes spotty brood is taken as a sign of inbreeding or poor queen mating.

Whatever the cause, colonies producing frames like that shown above are clearly going to be less strong than those towards the top of the page 2.

So, if the queen is failing, it’s time to requeen the colony …


Perhaps, perhaps not …

Which brings me to an interesting paper published by Marla Spivak and colleagues published in Insects earlier this year 3.

This was a very simple and straightfoward study. There were three objectives, which were to:

  • Determine if brood pattern was a reliable indicator of queen quality
  • Identify colony-level measures associated with poor brood pattern colonies
  • Examine the change in brood pattern after queens were¬†exchanged into a colony with the opposite brood pattern (e.g. move a ‘failing queen’ into a colony with a good brood pattern)

If you are squeamish look away now.

Inevitably, measuring some of the variables relating to queen quality and mating success involve sacrificing the queen, dissecting her and counting ‘stuff’ … like viable sperm in the spermathecae.

Unpleasant, particularly for the queen(s) in question, but a necessary part of the study.

However, in the long run it might¬†save some queens, so it may have been a worthwhile sacrifice … so, on with the story.

Queen-level variables in ‘good’ and ‘poor’ queens

By queen level variables I mean things about the queen that could be measured – and that differ – between queens with a good laying pattern or a poor laying pattern.

Surprisingly, good and poor queens were essentially indistinguishable in terms of sperm counts, sperm viability, body size or weight.

Poor queens¬†i.e. those generating a spotty brood pattern, weren’t small queens, or poorly mated queens. They were also not more likely to have fewer than 3 million sperm in the spermathecae (a threshold for poorly mated queens in earlier studies).

Furthermore, the queens had no statistical differences in pathogen presence or load (i.e. amount), including viruses (DWV, Lake Sinai Virus, IAPV or BQCV), Nosema or trypanosomes (Crithidia). 

Hmmm … puzzling.

Colony-level variables

So if the queens did not differ, perhaps colonies with spotty brood patterns had other characteristics that distinguished them from colonies with good brood patterns?

Spivak and colleagues measured pathogen presence and amount in both the good-brood and poor-brood colonies.

Again, no statistical differences.

So what happens when queens laying poor-brood patterns are put into a good-brood pattern hive?

And vice versa …

Queen exchange studies

This was the most striking part of the study. The scientists exchanged queens between colonies with poor-brood and good-brood and then monitored the change in quality of the brood pattern 4.

Importantly, they monitored brood quality 21 days after queen exchange. I’ll return to this shortly.

Changes in sealed brood pattern after queen exchange

Queen¬†from good-brood colonies showed a slight decrease in brood pattern quality (but not so much that they’d be considered to now generate poor brood patterns).

However, surprisingly, queens from poor-brood colonies exhibited a¬†greater improvement in brood quality (+11.6% ¬Ī 9.9% more sealed cells) than the loss observed in the reverse exchange (-8.0% ¬Ī 10.9% fewer sealed cells).

These results indicate that the colony environment has a statistically significant impact on the sealed brood pattern.

Admittedly, a 10-20% increase (improvement) in the sealed brood pattern on the last frame photograph (above) might still not qualify as a ‘good brood pattern’ queen, but it would certainly be an improvement.

Matched and mismatched workers

Since exchanged queens were monitored just 21 days after moving them all the workers in the receiving hive were laid – and so genetically related to – the previous queen.

The authors acknowledge this and comment that it would be interesting to extend the period until surveying the hive to see if ‘matched’ workers reverted to the poor brood pattern (assuming that was what the queen originally laid).

This and a host of other questions remain unanswered and will undoubtedly form the basis of future studies.

The authors conclude that¬†“Brood pattern alone was an insufficient proxy of queen quality. In future studies, it is important to define the specific symptoms of queen failure being studied in order to address issues in queen health.”

Notwithstanding the improvements seen in some brood patterns I suspect they would be insufficient to justify not replacing an underperforming queen … when considering the issue as a practical beekeeper¬†i.e. there may be improvements but they were much less than could be achieved by replacing the queen from a known and reliable source.

But it might be worth thinking twice about this …

Insufficient storage space

In closing it’s worth noting that I’ve seen spotty or incomplete brood patterns when there’s a very strong nectar flow on and the colony is short of super storage space.

Under these conditions the bees start to backfill the brood box, taking up cells that the queen would lay in.

Usually this is resolved just by adding another super or two.

If there remains any doubt (about the queen) and you’ve provided more supers you can determine the quality of the laying pattern by putting a new frame of drawn comb into the brood nest.

The queen should lay this up in a day or two if she’s “firing on all cylinders”.

In which case, definitely keep her ūüôā



  1. Drone¬†cells spotted randomly across the frame are a pretty-sure sign of laying workers … I’m solely discussing worker brood here.
  2. And weak colonies are most likely to be robbed and less likely to produce a honey crop.
  3. Lee et al., 2019 Is the brood pattern within a honey bee colony a reliable indicator of queen quality? Insects 10:12 Рthe full text of this paper is available free.
  4. I’ve avoided discussion of this, but there are standard ways to quantify¬†how good a good brood pattern is, or¬†how bad a poor brood pattern is.

18 thoughts on “Spotty brood ≠ failing queen

  1. Janet Wilson

    An interesting topic! One other thing to consider is: is the queen in question a commercially raised queen? Here in N. America, 60% or more of package queens don’t survive their first season. This often leaves new beekeepers suddenly surprised by finding no eggs or new brood in their colonies, and flummoxed as to what is going on…did they swarm? Supercede? Is there a new queen in the works? How far along would she be??

    In terms of this study, I wonder about two things:
    1. would we see poor brood patterns at all in properly raised queens we raise ourselves?
    2. if we exchange properly home raised queens would we see similar results?

    1. David Post author

      Hello Janet

      60% failure rate in packaged queens is pretty terrible. By the sound of things they start and then, very quickly, stop again. That certainly would perplex and frustrate a beginner.

      I’ve only rarely purchased queens. Many have failed, but nothing like 60% of them. My home grown queens are almost always more successful – certainly in their longevity and usually in their brood quality. However, some of my queens have had spotty brood (and I’m almost certain the frame of spotty brood in the post was from one of mine).

      I’ve never tried exchanging home grown underperforming queens with a better performing alternate. I might try next season, though – as I say in the post – I suspect the improvement would not be enough to raise the queen to an acceptable standard.

      But that’s one of the attractions of rearing your own queens … you can try a quick fix exchange or you have alternates ‘in the wings’ ready and waiting to take over if it doesn’t work.

      Best Wishes

  2. Amanda Millar (President, Brighton and Lewes Beekeepers)

    Interesting, thank you. Another reason for empty cells in the brood area is for the heater bees to warm the brood; up to10% of cells are left empty according to Jurgan Tautz, The Buzz about bees.

    1. David Post author

      Thanks Amanda
      I’d forgotten the heater bees …
      I can’t remember whether heater bees were there to warm brood when the ambient temperatures are low. The poor brood pattern frame I’ve shown was photographed in late June, so likely to be reasonably warm.

  3. Paul H

    How do you know the very spotty pattern is not simply because these cells are on the verge of hatching? I.e. maybe 50% of the sealed brood on this frame hatched in the last day, and the queen hasn’t been back to this frame to lay more yet.

    1. David Post author

      Usually the queen lays in concentric circles from the middle of the frame (the warmest bit presumably) outwards. If she was laying consistently and the brood were developing consistently entire blocks should emerge at the same time.

      Entire blocks emerge together

      In this photo the brood has emerged from the centre of the patch and the queen has already relaid the space.

      There are explanations for spotty brood that don’t involve the queen failing … like the heater bees mentioned above by Amanda, or the backfilling of the brood frames with nectar.

      However, the spotty frame in the body of the post was a failing queen back in summer 2014. The combination of really patchy brood (with the empty cells containing nothing, or nectar) and the multitude of queen cells were a pretty sure sign … and she was replaced.


  4. Janet Wilson

    The biggest reason to be hyper-aware of brood pattern integrity (wide sheets of brood, containing a seamless graduation of ages of brood, usually from the centre outwards) is to spot the symptoms of foulbrood as soon as they arise. Unfortunately, foulbrood is symptomless until larvae begin to die…they are removed, the cell is cleaned (both acts spread the infection) and the queen re-lays the cell, which now is surrounded by much older larvae. Over time, a surprisingly short time, you get a horribly spotty/shotty frame full of mixed age and dying brood. So a frame where brood is of mixed ages all over the frame, rather in concentrated areas of same/similar-age brood is a real cause for concern.

    1. David Post author

      I’d hope beekeepers would be alert for foulbroods at an earlier stage than the sort of spotty brood frame shown above. However, anything that helps identify these brood diseases – and then deal with them appropriately – is to be welcomed.

      Aside from help in spotting disease there’s also something very reassuring about pulling out frames and seeing sheet after sheet of uniform-looking brood.


  5. Kevin

    Excellent piece on releflecting real science against intuition! Perhaps we bekeepers should pause and think before squishing that “poor performing queen”. So long as there is enough time left (e.g. ultimately letting the colony decide – supercedure in Autumn for example), maybe it’s worth giving the resident queen a chance – what would happen in the wild? Unless you want buckets and buckets of honey . . . . . Some stats on ferral colonies suggest that sustainability is generally worse than with well-managed colonies, but perhaps that is also significantly due to unmanaged disease as well. So, if well-managed, most “well-managed” colonies ought to do better. Perhaps we beekeepers just have to make sure we try to manage our colonies optimally, sensitively and, perhaps also, cautiously.

    1. David Post author

      Hello Kevin

      I think the problem with letting nature take its course is that it often doesn’t go the way we’d like …

      It’s worth remembering that we generally manage a higher density of colonies than would occur naturally. With a failing (or just temporarily underperforming?!) queen there’s a risk that the colony will be so weakened that it will be the target for robbing by adjacent colonies if there’s a nectar dearth.

      Notwithstanding this recent paper I think I’d still intervene seeing a frame as poor as the one shown in the body of the post … though I might be tempted to pop the queen in a nuc to see if things improved … just on the off chance.


  6. Yuri

    A more experienced beekeeper suggested that I give my hives full sun after I told him I was worried about them overheating in Northern California. After I did that the queens are laying wall to wall and no more chalk brood. Just throwing it out there as an observation.

    1. David Post author

      Hi Yuri

      There are several studies on the influence of temperature and development of chalkbrood. Under lab conditions you can get almost 100% mummification by chilling at particular stages of larval development. Presumably your colonies were responding to warming in the earlier or later parts of the day.

      I try and place hives in a location that gets early morning sun, but that is lightly shaded at midday … though this is much less of an issue here in Scotland than when I lived further south.


    2. Chris

      Very true. My mentor told me to keep my hives in the sun here in zone 5 early in my beekeeping years and it has worked for the most part. Occasionally we get a wet spring and I sometimes see a little chalkbrood but very seldom.

  7. Barry Crabtree

    Hi David

    I’ve just come across this problem in a couple of my colonies & we put it down a poorly mated queen – poorly mated in that she’s mated with some of her brothers, and although the eggs she is laying are fertilized they are drones & are detected as abnormal and eaten by the workers.

    I’m sure you’re aware of this!

    Anyway. If the sperm from the various drones was not mixed thoroughly in the queen’s spermatheca, then you could get periods of variability in the viability of the laid eggs which may account for the difference. Alternatively, just the fact that the queens have past ‘shot’ comb to lay in, where random cells may be filled with nectar/pollen may account for the +/- 10% difference.

    Anyway, I’ll be replacing my queens as soon as the season lets me! My notes showed that at the end of last season these colonies were spotty….

    1. David Post author

      Hi Barry

      Surely if the eggs are fertilized she will be laying worker brood? Drone brood is unfertilized. Drones are haploid, not diploid. A drone laying queen has usually run out of sperm or is otherwise poorly mated.

      You’re absolutely right that spotty brood can result from ‘shot’ comb. Spotty drone brood is usually a sign of laying workers who don’t adhere to the typical concentrated rings or patches that the queen does.

      Whatever the cause I hope there are sufficient workers to support the new queens if/when they arrive. Going by the temperatures at the moment it’s going to be some time until UK mated queens are available.


  8. Barry Crabtree

    Hi David

    Honey bee mating biology is not my strong point, but here goes….

    So I’m talking about *diploid* drones. They are fertilized, but because the queen mated with some of her brothers the two sex alleles are the same and the offspring become diploid drones rather than workers. From what I’ve read nurse bees detect this and cannibalise them, hence the gaps rather than seeing pepperpot brood with a drone-laying queen. So the queen’s not failing as such, she’s just got poor material to work with!

    I think it will be a couple of weeks before we see plenty of flying drones in my part of the world (Suffolk), so hopefully I’ll have raised some replacement queens by the beginning of May.

    Cheers. Barry.

    1. David Post author

      Hi Barry

      My mistake … Jerzy Woyke determined the fate of diploid drones back in the 60’s.

      I’d be surprised if queens were poorly mated in Suffolk. It’s a county that’s awash with beekeepers and bees! Perhaps the queen got mated late in the season or in poor weather and the drone congregation areas (DCA) were underpopulated. I’ve written elsewhere that drones and queens from the same hive exhibit behaviour that should prevent them meeting as they tend to fly different distances to the DCA.

      I need to improve my understanding of mating biology as here on the west coast there are very few bees … Beebase reports only one apiary within 10 km of mine. I’ll be lucky if my queens get mated at all.

      Perhaps I should keep Cape honey bees Apis mellifera capensis and take advantage of thelytokous parthenogenesis.


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