Category Archives: Queen failure

A June Gap

As far as the beekeeping season is concerned, we’ve had the starter and we’re now waiting for the main course. 

Like restaurants, the size of the ‘starter’ depends upon your location. If you live in an area with lots of oil seed rape (OSR) and other early nectar, the spring honey crop might account for the majority of your annual honey.

If you are in the west, or take your hives to the hills, you might have skipped the starter altogether hoping the heather is the all-you-can-eat buffet of the season.

Lockdown honey

In Fife they appear to be growing less OSR as the farmers have had problems with flea beetle since the neonicotinoid ban was introduced.

Nevertheless, my bees are in range of a couple of fields and – if the weather behaves – usually get a reasonable crop from it. My earlier plans to move hives directly onto the fields, saving the bees a few hundred yards of flying to and fro, was thwarted (like so much else this year) by the pandemic.

The timing of the spring honey harvest is variable, and quite important. You want it to be late enough that the bees have collected what they can and had a chance to ripen it properly so that the water content is below 20% 1.

However, you can’t leave it too late. Fast-granulating OSR honey sets hard in the frames and then cannot be extracted without melting. In addition, there’s often a dearth of nectar in the weeks after the OSR finishes and the bees can end up eating their stores, leaving the beekeeper with nothing 🙁

Judging all that from 150 miles away on the west coast where I’m currently based was a bit tricky. I had to timetable a return visit to also check on queen mating and the build up of all the colonies I’d used the nucleus method of swarm control on.

Ideally all in the same visit.

Blowin’ in the wind

I’d made up the nucs, added supers and last checked my colonies around the 17-19th of May. I finally returned on the 10th of June.

In the intervening period I’d been worried about one of my more exposed apiaries. I’d run out of ratchet straps to hold the hives together and was aware there had been some gales in late May.

Sure enough, when I got to the apiary, there was ample evidence of the gales …

How the mighty fall

The only unsecured hive was completely untouched and the bees were happily working away. However, one of the strapped hives had been toppled and was laying face (i.e. entrance) down. You can see the dent in the fence where it collided on its descent.

If she hadn’t already (and I expect she hadn’t based upon the date of the gales) I suspect the queen struggled to get out and mate from this hive 🙁

Nuked nucs

Two adjacent 8-frame nucs were also sitting lidless in the gentle rain. The lids and the large piece of timber they’d been held down with were on the ground. The perspex crownboards were shattered into dozens of pieces.

These bees were fine.

Both queens were laying and the bees were using the new top entrance (!) for entering and leaving the hive. They were a little subdued and the colonies were less well developed than the other nucs (see below). However, their survival for the best part of three weeks uncovered is a tribute to their resilience.

They were thoroughly confused how to get back into the hive after I replaced the lids 🙂

Slow queen mating

Other than extracting, the primary purpose of this visit was to check the queenright nucs from my swarm control weren’t running out of space, and to check on the progress of queen mating in the original colonies.

Queen mating always takes longer than you expect.

Or than I expect at least.

Poor weather hampered my inspection of all re-queening colonies but, of those I looked at, 50% had new laying queens and the others looked as though they would very soon.

By which I mean the colonies were calm and ‘behaved’ queenright, they were foraging well and the centre of the ‘broodnest’ (or what would be the centre if there was any brood) was being kept clear of nectar and had large patches of polished cells.

Overall it was a bit too soon to be sure everything was OK, but I expect it is.

However, it wasn’t too soon to check the nucs.

Overflowing nucs

In fact, it was almost too late …

With one exception the nucs were near to overflowing with bees and brood.

I favour the Thorne’s Everynuc which has an integral feeder at one end of the box. Once the bees start drawing comb in the feeder they’re running desperately short of space.

Most had started …

Here's one I prepared earlier

Here’s one I prepared earlier

I didn’t photograph any of the nucs, but the photo above (of an overly-full overwintered nuc) shows what I mean; the feeder is on the right.

The nucs had been made up with one frame of predominantly emerging brood, a few more nurse bees, two foundationless frames, a frame of drawn comb and a frame of stores.

They were now all packed with 5 frames of brood and would have started making swarm preparations within a few days if I hadn’t dealt with them.

Good laying pattern from queen in 5 frame nucleus

And the queens had laid beautiful solid sheets of brood (always reasonably easy if the comb is brand new).

Housekeeping and more swarm prevention

The beauty of the nucleus method of swarm control is that you have the older queen ‘in reserve’ should the new queen not get mated, or be of poor quality.

The problem I was faced with was that the new queens weren’t all yet laying (and for those that were it was too soon to determine their quality), but the older queen was in a box they were rapidly outgrowing.

I therefore removed at least three frames of brood 2 from each nuc and used it to boost the re-queening colonies, replacing the brood-filled frames with fresh foundation 3.

The nucs will build up again strongly and the full colonies will benefit from a brood boost to make up for some of the bees lost during requeening. Some of the transferred frames had open brood. These produce pheromones that should hold back the development of laying workers.

Finally, if the requeening colonies actually lack a queen (the weather was poor and I didn’t search very hard in any of them) there should be a few larvae young enough on the transferred frames for them to draw a new queen cell if needed.

I marked the introduced frames so I can check them quickly on my next visit to the apiary.

This frame needs to be replaced … but could be used in a bait hive next year

The additional benefit of moving brood from the nucs to the full colonies is that it gave me an opportunity to remove some old, dark frames from the latter.

Shown above is one of the removed frames. As the colony is broodless 4 and there’s the usual reduction in available nectar in early/mid June, many of the frames in the brood box were largely empty and can easily be replaced with better quality comb.

Everyone’s a winner 😉

Drone laying queen

One of the nucs made in mid/late May had failed. The queen had developed into a drone layer.

Drone laying queen

The laying pattern was focused around the middle of frame indicating it had been laid by a queen. If it had been laying workers the drone brood would be scattered all over the frames.

There was no reasonable or efficient way to save this colony. The queen was removed and I then shook the bees out in front of a row of strong hives.

I was surprised I’d not seen problems with this queen when making up the nucs in May 5. I do know that all the colonies had worker brood because the nucs were all made containing one frame of emerging (worker) brood.

Perhaps the shock of being dumped into a new box stopped her laying fertilised eggs. Probably it was just a coincidence. We’ll ever know …


And, in between righting toppled hives, checking for queens, stopping nucs from swarming, moving a dozen hives/nucs, boosting requeening hives and replacing comb … I extracted a very good crop of spring honey.


Although I had fewer ‘production’ hives this season than previous years (to reduce my workload during the lockdown) I still managed to get a more than respectable spring harvest. In fact, it was my best spring since moving back to Scotland in 2015.

The crop wasn’t as large as I’d managed previously in Warwickshire, but the season here starts almost a month later.

A fat frame of spring honey

I start my supers with 10 or 11 frames, but once they are drawn I reduce to 9 frames. With a good nectar flow the bees draw out the comb very nicely.

The bees use less wax (many of my frames are also drawn on drone foundation, so even less wax than worker comb 6), it’s easier to uncap and I have fewer frames to extract.

Again … everyone’s a winner 😉

Not the June gap

Quite a few frames contained fresh nectar, so there was clearly a flow of something (other than rain, which seemed to predominate during my visit) going on. These frames are easy to identify as they drip nectar over the floor as you lift them out to uncap 🙁

In some years you find frames with a big central capped region – enough to usefully extract – but containing lots of drippy fresh nectar in the uncapped cells at the edges and shoulders. I’ve heard that some beekeepers do a low speed spin in the extractor to remove the nectar, then uncap and extract the ripe honey.

I generally don’t bother and instead just stick these back in the hive.

If there’s one task more tiresome than extracting it’s cleaning the extractor afterwards. To have to also clean the extractor during extracting (to avoid the high water content nectar from spoiling the honey) is asking too much!

Colonies can starve during a prolonged nectar dearth in June. All of mine were left with some stores in the brood box and with the returned wet supers. That, plus the clear evidence for some nectar being collected, means they should be OK.

National Honey monitoring Scheme

I have apiaries in different parts of Fife. The bees therefore forage in distinct areas and have access to a variety of different nectar sources.

It’s sometimes relatively easy to determine what they’ve been collecting nectar from – if the back of the thorax has a white(ish) stripe on it and it’s late summer they’re hammering the balsam, if they’ve got bags of yellow pollen and the bees are yellow and the fields all around are yellow it’s probably rape.

Mid-April in the apiary ...

Mid-April in a Warwickshire apiary …

But it might not be.

To be certain you need to analyse the pollen.

The old skool way of doing this is by microscopy. Honey – at least the top quality honey produced by local amateur beekeepers 7 – contains lots of pollen. Broadly speaking, the relative proportions of the different pollens – which can usually be distinguished microscopically – tells you the plants the nectar was collected from.

The cutting edge way to achieve the same thing in a fraction of the time (albeit at great expense) is to use so-called next generation sequencing to catalogue all the pollen present in the sample.

Pollen contains nucleic acid and the sequence of the nucleotides in the nucleic acid are uniquely characteristics of particular plant species. You can easily get both qualitative and quantitative data.

And this is exactly what the National Honey Monitoring Scheme is doing.

They use the data to monitor long-term changes in the condition and health of the countryside” but they provide the beekeeper’s involved with the information of pollen types and proportions in their honey.

National Honey Monitoring Scheme samples

Samples must be taken directly from capped comb. It’s a messy business. Fortunately the labelling on the sample bottles is waterproof so everything can be thoroughly rinsed before popping them into the post for future analysis.

I have samples analysed already from last year and will have spring and summer samples from a different apiary this season. I’ll write in the future about what the results look like, together with a more in-depth explanation of the technology used.

When I last checked you could still register to take part and have your own honey analysed.


Under (re)construction

Lockdown means there have been more visitors than ever to this site, with numbers up at least 75% over this time last year.

This, coupled with the need to upgrade some of the underlying software that keeps this site together, means I’m in the middle of moving to a bigger, faster, better (more expensive 🙁 ) server. I’m beginning to regret the bloat of wordpress over the lean and mean Hugo or Jekyll-type templating systems (and if this means nothing to you then I’m in good company) and may yet switch.

In the meantime, bear with me … there may be some broken links littering a few pages. If it looks and works really badly, clear your browser cache, re-check things and please send me an email using the link at the bottom of the right hand column.

Thank you


Queenright … or not?

A brief follow-up to the (ridiculously long) post last week about leaving queen cells in the colony after a) it swarms, or b) implementing swarm control 1.

How long does it take for the new queen to emerge, mate and start laying? 

And what if she doesn’t?

How did we get here?

We are approaching the peak of the beekeeping season. Colonies have built up strongly and should now be topped by comfortingly heavy supers of spring honey. 

Mind your back 😯 

The box you inspected in early April and found three frames of brood in is now bursting at the seams with bees and brood. Everything is getting busier and bigger. You may have already run out of supers or – lucky you – are frantically extracting to free-up supers to return to the colonies.

Depending upon your location you may already have discovered that your swarm prevention efforts, whilst temporarily effective, were soon treated with disdain as the colonies started to build queen cells.

Sealed queen cell ...

Sealed queen cell …

You are now using some form of swarm control and the colony now contains a mature queen cell.

Or they swarmed … leaving a mature queen cell 🙁

Queenless colonies

Is a colony with a charged, capped, queen cell queenless? 

A philosophical question 🙂

I guess the answer is technically no, but practically yes.

There’s clearly a queen in the hive, but she’s really a potential queen. To be useful to the colony (and the beekeeper) she has to emerge, mature, mate and start laying.

It’s at that stage that the colony can be described as queenright.

All of this takes time and all of which significantly changes the tempo of the season.

Colonies that are requeening should generally not be disturbed and the change from full-on to full-off can feel strange.

Doubly so, because the lack of reassuring inspections can make the wait seem interminable. 

It’s tempting to have a quick peek … after all, what harm could it do?

Tick tock

The development of a queen takes 16 days from egg to eclosed virgin. The first three days as an egg, then six days as a larva before a further week as a developing pupa. The rapid development is due to the very rich diet that queens are fed in the first couple of days. This triggers a host of changes in gene expression 2 which dramatically alters the morphology, behaviour and longevity of the queen from the genetically identical worker.

After a virgin queen emerges she needs to mature sexually which takes 5-6 days. During this period they don’t look or behave much like queens. They tend to be quite small and, if disturbed, rush about the frame skittishly. They are also a lot more willing to fly than a mature laying queen – you have been warned! 3

Where have all my young girls gone?

What a beauty

Virgin queens are not lavished with attention by a retinue of workers, all of which often makes them more difficult to find in the hive.

The queen goes on one or more mating flights which usually take place on warm, calm, sunny early afternoons.

She then returns to the hive and, 2-3 days later, starts laying eggs. A queen that has just started laying sometimes lays more than one egg per cell. However, she settles down fast and will usually lay in a reasonably tight pattern in the centre of one of the middle frames in the brood nest.

Have patience

Add all those timings up and you have a minimum of two weeks between the capping of the queen cell and the day when she starts laying.

To be sure, you need to know when the queen cell was capped which is difficult if you’re dealing with a colony that swarmed. Was the cell capped on the day the colony swarmed (not unusual), or was it capped during the lousy weather a few days earlier that then delayed the emergence of the swarm?

It is unwise to disturb a virgin queen.

All sorts of things can go wrong. You might inadvertently crush her during an inspection 4 or scare her into taking flight and getting lost in the long grass.

Equally calamitous would be inspecting the colony on the nice, calm, warm mid-afternoon when she decides to go off on her mating flight. She’ll be off consorting with the local drones for about 10 – 30 minutes, and may go on more than one flight on subsequent days. If she returns to find the roof and supers off, the brood frames out and smoke being puffed everywhere she may never find the hive entrance.

Inspecting a colony

None of the above ends well.

Minima and maxima

The two weeks detailed above is the absolute minimum. I don’t check these things routinely but think the only time I’ve really seen it taking that short a period (from cell sealing to a mated laying queen) is when queen rearing using mini-mating nucs.

Mini-nucs …

Queens tend to get mated in these very fast if the weather is suitable. I don’t know why 5.

But, if the weather is unsuitable, irrespective of the hive type, mating will be delayed.

By ‘unsuitable’ I mean lousy. If it’s raining persistently or blowing a hoolie the queen will not venture forth.

If it’s cool (16 – 18°C) and cloudy she might, particularly if she’s of the darker Apis mellifera mellifera strain. 

But then again, she might not 🙁 

All of which means that the two weeks quoted really is a minimum.

What if it rains for a month? The virgin queen has a ‘shelf life’. If she does not get mated within ~26-33 days of emergence she is unlikely to get successfully mated at all.

Here we go again ...

No queen mating today …

To summarise, it will take a minimum of two weeks from queen cell capping to having a laying queen in the hive. If 40 days elapse before the queen is mated (again from cell capping) it is likely that she will be a dud.

Three weeks

Assuming the weather has been OK for queen mating I usually leave a minimum of three weeks between closing the hive up with a capped queen cell and looking for the mated queen. 

There’s little to be gained by rummaging around the hive before then … and a whole lot to be potentially lost.

If you do open the hive up too early – assuming none of the nightmare scenarios above occur – what can you expect to see?

Lift the dummy board out, prise out the last frame and then split the hive somewhere in the middle of the remaining frames i.e. don’t work through frame by frame, this isn’t a routine inspection, it’s a Royal Checkup.

If you look around the middle of the face of the central frames you can often see polished cells. These have been cleaned and prepared by the workers for the queen to lay in. They’re particularly obvious if the comb is a bit old and dark – then they really do look polished and shiny.

If there are polished cells present, but no eggs, I’m then reasonably confident that there’s a queen in the hive but that she’s not started laying yet (but is probably mated).

There’s no point in looking for her. Close the hive up and leave it another week.

Brood frame with a good laying pattern

If she is laying, leave her be. Wait until she’s laid up a few frames and you can tell she has a good laying pattern of worker brood i.e. look at the appearance of the sealed brood, then find her and mark her 6.

Breathe a sigh of relief … your colony is again queenright.

Five weeks

If five weeks 7 have elapsed between leaving a freshly capped cell in the hive and the non-appearance of eggs I start to fear the worst.

The colony will now have no brood – it all emerged about two and half weeks ago – and the lack of brood pheromone means there’s a possibility that the colony will develop laying workers

Laying workers ...

Laying workers …

There may be a queen present, but she’s rapidly becoming an ageing spinster

In this situation it is probably wise to decide what Plan B is … how will you ‘rescue’ the colony?

If you leave the colony for another week or fortnight you might find a laying queen, but you probably won’t. During this period the colony will dwindle further in size and strength 8

Plan B

You effectively have four choices:

  1. Unite the colony with a known queenright colony.
  2. Requeen the colony with a mated, laying queen 9.
  3. Add a mature capped queen cell to the colony. Start nervously pacing the apiary again waiting for her to emerge, mature, mate and start laying.
  4. Allow the colony to rear their own queen by providing a frame of eggs (see below).

It is important to find and dispatch the ‘failed’ queen if you are going to do 1, 2 or 3. The queen may have failed to get mated but she might still be able to kill a challenger queen in the hive. 

Uniting the colony is often the best and safest option. It’s quick. It uses the bees remaining in the colony immediately and it strengthens another hive. It’s my preferred option … but I have quite a few colonies to work with. If you have just one (and you shouldn’t have) it’s clearly a non-starter. 

An Abelo/Swienty hybrid hive ...

An Abelo/Swienty hybrid hive …

Adding an expensive purchased mated laying queen (or a cheap one) can be risky. Terminally queenless and broodless colonies are often tricky to requeen. The most successful way I’ve found to do this 10 is to use a large cage pinned over a frame of emerging brood. And even then it doesn’t always work 🙁 

If you already have laying workers it is not worth trying to requeen the colony – they’ll almost certainly kill her. I usually try once to ‘rescue’ a laying worker hive (details here), but then shake them out.

Adding a capped queen cells can work if the colony is queenless but you will have another long wait ahead of you … and all the time the colony is dwindling in size.

She emerges into a population of geriatric workers. Far from ideal.

But what if you can’t find the queen?

Is the colony really queenless?

Perhaps she mated quite late because of poor weather and is about to get started?

Perhaps she failed to mate and is just lurking in there waiting to slaughter the £40 Buckfast queen you’re about to add 🙁 

Frame of eggs

Most of these questions can be answered by adding a ‘frame of eggs’.

A queenless colony will start to rear a new queen if presented with eggs and larvae.

A queenright colony will not.

If you are unsure whether a colony is queenright add a frame containing a good number of eggs. I usually like to use a full brood frame also containing some larvae and sealed brood. The brood pheromone will help hold back laying worker development. The new young bees that emerge will bolster the hive population and will be there to help the new queen when she returns from getting mated.

If you have the luxury of choosing a frame of eggs on relatively new fresh comb the bees will find it easier to draw queen cells. However, don’t worry if you don’t … if they’re queenless they’ll be thankful for anything.

Check the colony a few days after adding the frame of eggs. If they’ve started queen cells 11 then I just let them get on with it and check again in about a month or so for a laying queen. They won’t swarm or generate casts as – by this time – bee numbers are significantly depleted. 

However, if they don’t start queen cells it means there’s a queen somewhere in the hive. Check the other frames in the hive for eggs. It’s not at all unusual to find the original queen has now started laying. Again, leave her to get on with it.

But if there are no eggs on other frames and no queen cells (on the frame you added) you need to find the non-functioning queen … and we’ll deal with that sometime in the future 😉

Good luck


The usual dictionary definition of queenright just references a colony of bees that contains a queen. The OED has references going back to 1911 (When a colony is found that is not queen-right, it is remorselessly broken up, and distributed among other colonies, or united with a weak colony having a good queen, C.C. Miller in Fifty Years among Bees) including some from Wedmore and E.O Wilson.  

However, none specifically state whether the queen is laying. Or what she’s laying. A queenless colony is easy to define. But what about a colony containing a virgin queen? Or a drone laying queen? 

I’d argue that in these situations the colony contains a queen, but things aren’t really ‘right’ (as in correct). In my view, queenright means a mated, laying queen. 

Please, no pedantic questions or comments about a colony containing a well mated queen that, because there’s a nectar dearth, has stopped laying … 😉

Polyandry and colony fitness

Honey bees are polyandrous. The queen mates with multiple drones during her mating flight(s). Consequently, her daughters are of mixed paternity.

In naturally mated queens there is a relationship between the number of patrilines (genetically distinct offspring fathered by different drones) and the ‘fitness’ of a colony.

Colony fitness

A ‘fit’ colony is one that demonstrates one or more desirable traits (those that benefit the colony … and potentially the beekeeper) such as better population growth, weight gain, resistance to pathogens or survival.

If you analyse the molecular genotype of the worker offspring you can determine which patriline they belong to. If you genotype enough workers you start to see the same patrilines appearing again and again. The more patrilines, the more drones the queen mated with 1.

Shallow depth of field

One of many …

Naturally mated queens mate with ~13 drones. Depending upon the study a range from as low as 1 to as high as 40 (and exceptionally into the high 50’s) has been demonstrated, though different studies all tend to produce an average in the low- to mid-teens.

There is a well-established link between polyandry and colony fitness 2. Essentially, the more genetically diverse a colony i.e. the larger the number of patrilines, the fitter that colony is.

The benefits of polyandry

Why should colonies with increased genetic diversity be fitter?

There are a number of hypotheses that attempt to explain why intracolonial genetic diversity is beneficial. These include the increased behavioural repertoire of the worker bees, a reduced production of diploid drones (which would otherwise be produced due to the single-locus sex determination system) and an increased resistance to a wide range of parasites and pathogens 3.

Parasites and pathogens are an extremely effective evolutionary selective pressure. Several studies from David Tarpy and Thomas Seeley have shown that increased polyandry results in better resistance to chalkbrood and American foulbrood.

But what about Varroa? It’s a new pathogen (evolutionarily speaking) to honey bees and there is evidence that the resistance mechanisms observed are genetically determined 4.

Does polyandry contribute to Varroa resistance? 

Would increased polyandry result in improved resistance to mites?

Limits of polyandry and natural resistance

Why is the average number of drone matings in the low teens?

If polyandry is beneficial – and there’s no doubt it is – then surely more patrilines (hyperpolyandry) would be even more beneficial?

How could this be tested?

Naturally mated queens only very rarely exhibit 30+ drone matings. Not only are these colonies hard to find, but they are so rare that doing any sort of statistical analysis of the improved (or otherwise) fitness is probably a non-starter.

Perhaps there’s an alternative way to approach the question? Rather than look at individual colonies within a mixed population, why not study the overall level of polyandry within a population that demonstrates resistance?

For example, do queens that head colonies of untreated feral bees that exhibit a demonstrated enhanced resistance to Varroa, the most important pathogen of honey bees, exhibit higher levels of polyandry?

Two relatively recent scientific papers have tackled these questions. Both have produced clear answers.

Drones : if more is better, is lots more better still?


Keith Delaplane and colleagues used instrumental insemination (II) of virgin queens to produce queens ‘mated’ with 15, 30 or 60 drones. Sperm was collected from 1, 2 or 4 drones from 15 donor colonies, mixed thoroughly and used for queen insemination.

Full-sized colonies were requeened with the II queens and left for 6 weeks 5 after which sampling started. Over two seasons a total of 37 colonies (with 11, 13 and 13 colonies respectively headed by queens ‘mated’ with 15, 30 and 60 drones) were tested at approximately monthly intervals.

Testing involved visual analysis of colony strength 6 and comb construction. Mite levels were measured using standard alcohol wash of ~300 bees at mid- or late-summer timepoints.

Brood frame with a good laying pattern

The results of this study are commendably brief … just 8 lines of text and two tables. I’ll summarise them in just a couple of sentences.

Colonies headed by queens ‘mated’ with 30 or 60 drones produced significantly more brood than the colony headed by the queen ‘mated’ with only 15 drones. Conversely, significantly more colonies headed by queens mated with only 15 drones had a higher level of mite infestation 7.

Natural Varroa resistance and polyandry

One of the best studied populations of feral bees co-existing with Varroa are those in the Arnot Forest in New York State. These are the bees Thomas Seeley and colleagues study.

These colonies live in natural holes in trees at low density through the forest. The colonies are small and they swarm frequently. Their spatial distribution, size and swarminess (is that a word?) are all evolutionary traits that enable resistance, or at least tolerance, to Varroa and the pathogenic viruses the mite transmits.

I’ve discussed Seeley’s studies of the importance of colony size and swarming previously. I don’t think I’ve discussed his work on spatial separation of colonies, but I have described related studies by Delaplane and colleagues.

Essentially, by being well-separated, mite transmission between colonies (e.g. during robbing) is minimised. Similarly, by existing as small colonies that swarm frequently Iwith concomitant brood breaks) the mite population is maintained at a manageable level.

Marked queen surrounded by a retinue of workers.

Her majesty …

Do the Arnot Forest Varroa-resistant 8 bees exhibit especially high levels of polyandry, suggesting that this contributes their survival?


Seeley and colleagues determined the number of patrilines in 10 Arnot Forest colonies using the same type of genotyping analysis described earlier. They compared these results to a similar analysis of 20 managed honey bee colonies located nearby.

On average, Arnot Forest queens had mated with ~18 drones (17.8 ± 9.8) each. In contrast, queens in managed colonies in two nearby apiaries had mated with ~16 and ~21 drones. These figures are not statistically different from each other or from the natural mating frequencies reported for honey bees in other studies.

Hyperpolyandry and colony fitness

The first of the studies confirms and extends earlier work demonstrating the polyandry (and in this instance hyperpolyandry i.e. at an even greater level than seen normally) increases colony fitness – at least in terms of colony strength and Varroa resistance.

Delaplane and colleagues hypothesise that the increased mite resistance in hyperpolyandrous (30 or 60 drones) colonies may be explained by either:

  • the importance of extremely rare alleles (gene variants), which would only be present in colonies in which the queen had mated with a very large number of drones.
  • the presence of beneficial non-additive interactions between genetically-determined traits e.g. grooming and hygienic behaviour and reduced mite reproduction.

Neither of which are mutually exclusive and both fit at least some of the extant data on natural mite resistance. Discriminating between these two hypotheses and teasing apart the variables will not be straightforward.

Absence of hyperpolyandry in naturally mite-resistant colonies

At first glance, the absence of the hyperpolyandry in the mite-resistant Arnot Forest bees studied by Thomas Seeley and colleagues appears to contradict the studies using the instrumentally inseminated queens.

The Arnot Forest bees exhibit the same level of polyandry as nearby managed colonies, and for that matter, as colonies studied elsewhere. They are mite-resistant but the queen has not mated with an increased number of drones.

In other studies 9, naturally mated colonies exhibiting different levels of polyandry (within the normal range) showed no correlation between Varroa levels and queen mating frequency.

Perhaps it’s surprising that the Arnot Forest queens hadn’t mated with fewer drones considering the extreme separation of the colonies (when compared with managed colonies). The colony density within the Forest is approximately one per square kilometre.

However, at least during the peak swarming and mating period in the season, drone availability is rarely limiting.

This is because drones are not evenly spread in the environment. Instead, they accumulate in drone congregation areas (DCA) to which the queen flies for mating.

What limits polyandry?

Polyandry is beneficial and, apparently, hyperpolyandry is more beneficial. However, queens mate with 10 – 20 drones, rather than 50 or more. Why is this?

Queen mating is a risky business. The queen has to fly to the DCA, mate with multiple drones and then return to the hive. She may make one or several mating flights.

I’ve discussed how far drones and queens fly to reach the DCA previously. Most drones fly less than 3 miles and 90% of matings occur within about 5 miles of the virgin queen’s hive. The queen probably flies further to the DCA.

All the time she is travelling to and from the DCA, and all the time she is present within it mating, she’s potentially at risk from hungry house martins, swallows, bee eaters (!) or from thunderstorms.

Or simply from getting lost.

Additionally, a number of honey bee pathogens are transmitted between drones and queens during mating. Hyperpolyandrous queens 10 are therefore at risk from these sexually transmitted diseases 11.

It’s therefore likely that the level of polyandry observed in honey bees has evolved as a consequence of the beneficial pressures polyandry brings balanced by the risks associated with mating multiple times.

Practical beekeeping

Although the two studies described here don’t have an immediate relevance to day-to-day practical beekeeping, it’s worth remembering that poor queen mating is regularly blamed for queen failures e.g. queens that develop into drone layers during the winter.

I’m going to write about drones later this year so for the moment will just make these points:

  • drone production is maximised to generate sexually mature drones for the swarming season
  • after eclosion, drones need to mature before being able to mate
  • drones live about 30 days and their sperm volume, though not necessarily viability, decreases as they age

Together this means that late in the season – perhaps late July or early August (though this will vary depending upon location) – the number of drones will decrease.

More significantly, the drones will be ageing.

In turn this means that late-mated queens may not mate with as many drones, or that the matings may not result in insemination.

Most beekeepers will be aware of queens that apparently ‘run out of sperm’ and become drone layers.

However, there may be less obvious problems with late-mated queens. I’m not aware of any studies on seasonality of queen mating and polyandry. However, I would not be at all surprised if they exhibited a reduced level of polyandry.

And, as described above, these colonies are likely to exhibit reduced fitness.

Something else to consider when deciding whether to unite a colony late in the season or hope the last of your virgin queens mates successfully …


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 🙁

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% ± 9.9% more sealed cells) than the loss observed in the reverse exchange (-8.0% ± 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 🙂


And they’re off …

I posted last week on the relative lateness of the start of the beekeeping season here in Scotland 1. Having been away for a few days I was both surprised and disconcerted to find this waiting for me when I arrived at the apiary to conduct the first inspections of the year.

When is a swarm not a swarm?

When is a swarm not a swarm?

Surprised because I’d missed all the seasonal clues that indicated swarming might be imminent.

Disconcerted because, in the interests of full disclosure, I’d have to admit to it 😉

The colony behind the near-invisible one inch entrance hole through the bee shed wall is a double brood colony in an Abelo poly hive. It was headed by a 2018 queen (or had been 🙁 ) and had a nice temperament and good manners.

The queen was marked blue and one wing was clipped to prevent her flying off.

But it wouldn’t have stopped her trying to fly off. Instead she would have ignominiously spiralled to the ground 2.

Usually what then happens is she attempts to climb back up and the swarm gathers around her. In a standard hive this is often this is underneath the hive stand.

My guess was that she’d made it up to the landing board and stopped or got stuck there.

I had a gentle prod about in the beard of well-tempered bees but could see no sign of her.

With about 20 more hives to inspect I quickly decided to walk them into a fresh hive … I’d let them do this while I got on with other colonies in the apiary.

Don’t think, do

Walk this way

Walk this way

I put together a new floor and a brood box of mostly foundationless frames. I put one or two frames of drawn comb in and gently dislodged a couple of clumps of bees into the box.

Within a very short time more bees were marching down the wall of the shed and clustering between the frames of drawn comb in the brood box.

What started as a trickle became – if not a torrent – then certainly a determined stream of bees taking up residence in the new box.

To encourage them I balanced a split board across the tops of the frames to provide a welcoming dark ‘cavity’ for them to occupy. Very soon you could see bees fanning strongly at the opening between the split board and the shed wall.

Fanning workers

Fanning workers

I interpreted this as meaning the queen had entered the box and the workers were encouraging others to join her.

After an hour or so I moved the hive a few inches away from the shed wall, placed a crownboard and roof on and carried on inspecting other hives in the apiary. By this time about 75% of the bees had left the ‘swarm’ and entered the brood box.

Not so fast

And that’s when everything ground to a halt.

There were no bees fanning at the hive entrance. No more bees entered the box through the entrance. Instead they started leaving in dribs and drabs.

I’ve hived swarms like this before, or done the classic ‘walk them up a sheet’ having dumped them from a skep outside a hive. Other than this being a real spectacle, one of the striking features is that what starts as a mass of bees ends being an absence of bees … they all enter the hive.

'Walking' a swarm into a hive

‘Walking’ a swarm into a hive

Clearly something was wrong and I was beginning to suspect that there wasn’t a queen in the ‘swarm’ at all.

So I did what I should have done in the first place. I had a look in the original hive.

Hello there!

Blue skinny queen

Blue skinny queen

I smoked the double brood box gently from the bottom, intending to encourage the queen (if she was there) into the upper box.

The box was busy but not packed with bees 3, there were good amounts of sealed brood (and a really nice tight laying pattern on many frames).

There were quite a few ‘play cups’ and a few had eggs in them. This is one of the early signs of swarming.

I found the queen on the 19th of 22 frames.

Perhaps I was too gentle with the smoke 🙄

She was the queen I was expecting. Marked blue, though the paint was beginning to rub off a bit, and with the left wing clipped.

She looked like she had lost a bit of weight.

Big fat queens in full laying mode (which they should be getting to by late April) aren’t very aerodynamic so workers slim the queen down before swarming to improve her flying ability.

This queen looked to me like she’d been on the F-plan diet (but remember I’d not seen her since last August). In addition, the number of eggs in the colony was relatively low. This would also be expected if the colony had been preparing to swarm as queens reduce their laying rate in the few days before swarming.

What else could be seen?

Stores and pollen levels were good.

The notable absence from the hive was of well developed, sealed or unsealed queen cells.

A colony will normally swarm once developing queen cells are capped. A colony with a clipped queen often delays swarming for a few more days. It’s therefore usual to find sealed queen cells in a swarmed colony. There may also be unsealed cells as well.

~3 day old queen cell ...

~3 day old queen cell …

There wasn’t anything close to a sealed queen cell in the colony 4. The best developed were, at the very most, a couple of days old.

So what happened?

Other than the absence of well developed queen cells the colony looked as though it had swarmed.

If it walks like a duck etc.

Since the queen was clipped she had eventually clambered back to the hive and re-entered, leaving many of the workers who had left with her clustered around the hive entrance.

That’s currently my best guess 5.

If that was the case, notwithstanding the current lack of well-developed queen cells, they’d be trying again as soon as the weather was good enough. I therefore decided to preempt them by doing a classic artificial swarm.

I moved the queen on a frame with a small patch of brood into the box I’d used to try and ‘walk’ the swarm into. I then moved the – now queenless – double brood box a couple of metres off to one side in the shed. Finally I placed the queenright box in the place the original colony had occupied.

And what will happen?

Full details are in the description of Pagden’s artificial swarm. The flying bees from the double brood box will return to the box with the queen. The hive bees in the double brood box will start to rear one or more new queens.

And at that point I’ll intervene.

The double brood box has lots of brood and stores spread across 21 frames. The bees are well tempered, stable on the comb and have no significant signs of chalkbrood or other diseases (and Varroa and virus levels are exceptionally low – I’d measured both 6).

They are a good stock to make increase from.

I’ll check them in a  few days and see how queen cells are developing. Once there are good sealed cells I’ll split the colony into several 3-5 frame nucleus colonies. The final number will depend upon the number of good queen cells and the number of bees left in the colony.

It should be possible to generate half a dozen good nucleus colonies from a suitable double brood colony without too much of a problem.

First inspection summary

I got through all my colonies (eventually). With a reasonable number to compare it’s easy to define the good, the bad and the indifferent ones.

It’s much easier to do this once the season is properly underway, which is a good reason not to inspect too soon in the year. Some colonies are very early-starters, others lag bit. If you inspect too early you might consider the slow ones are dud or failed queens.

I was pleased to see that most were good or at least indifferent, with only a couple clearly exhibiting undesirable personality traits – aggression, laziness, running, following – or, in one case, disease (rather too much chalkbrood). These will be destined for prompt requeening and drone brood will be removed to reduce their contribution to the gene pool.

My overwintered 5 frame nucs looked excellent, with a couple needing re-hiving immediately.

Here's one I prepared earlier

Here’s one I prepared earlier

The first inspection is really little more than a check that things are all OK. It doesn’t matter whether I see the queen. If there are eggs present I’m happy.

Eggs? Overt disease? Stores? Brood? Space? … next please!

Overwintering losses

I lost 10% of my colonies this winter – two from 20. This includes both full colonies and overwintered 5 frame nucs.

One colony drowned. The lid and crownboard blew away in a severe storm and they were subjected to a three-day deluge over a long weekend when I was away.

Mea culpa. I should have had more bricks on the roof.

Spot the drone laying queen

Spot the drone laying queen

In the second colony the queen failed and turned into a drone laying queen (DLQ). This had been my worst-tempered colony last year and was scheduled for requeening. However, the queen I found wasn’t the clipped and marked one I’d left there in August. Clearly there had been a late-season supercedure and the replacement queen was poorly mated.

Although she was a bee I didn’t keep it is great to be beekeeping again 🙂


And they’re off! is the phrase used by horse racing commentators at the start of a race. It is also the title of a song composed by William Finn from the musical A New Brain. The song is about the damage gambling does to families. There’s a good cover version by Philip Quast on YouTube.

Spot the queens

A little over a month ago I opened a recently-united hive to be faced with this …

Spot the queen

Spot the queen …

That’s a ‘bit of a stooshie’ as they might say in Glasgow 1. Somewhere in the middle of that brawling mass of workers is a queen.

She’s unmarked and not clipped.

This was a surprise as the queen I had expected to find in the box should have been marked blue 2.

A potted history

The colony had overwintered with a 2016 white marked and clipped queen. I’d conducted a vertical split on the colony in mid-May and by early July I had two queens in the box, one above and one below the split board.

Neither of them was marked white.

A few days after setting up the split the queenright half looked very much like it was preparing to either supercede the queen or swarm. The white marked queen was still there but there were also charged queen cells present.

Either supercedure or swarming should have eventually resulted in the queen being replaced. However, the quality of her successor could not be relied upon … she might have been great, but she might have been poor. The white clipped queen was pretty good and I didn’t want to lose her 3.

I therefore made up a nuc with the ‘old’ white marked queen for safety and left the box with one charged queen cell.

The upper and lower boxes of the split both eventually – by early July – contained new queens, both of which I’d marked blue and clipped.

On the 7th of July I made up a nuc for overwintering with one of the blue clipped queens. The remainder of the – now queenless – colony I united with the queenright colony below it from the original vertical split. This formed one good strong colony.

We had an excellent nectar flow in July and I got two full supers from the colony by the end of the month.

Pining for the fjords

Nine days after making up the nuc and uniting the colonies I conducted a follow-up inspection. The newspaper was chewed away and most of the bees were behaving as normal. So far, so good.

However, on one frame the bees were agitated and formed a gobstopper-sized clump. I gently teased apart the melee with my forefinger to see if there was a queen buried in the middle … there was.

Unmarked and unclipped. Puzzling.

A little further across the same frame was another queen. To paraphrase Monty Python, this queen was not ‘stunned’ or ‘pining for the fjords’, rather she was very much an ex-queen. And probably relatively recently.

Regicide ...

Regicide …

So, as expected, I’d found the 2018 blue-marked and clipped queen in the united colony.

Unexpectedly, she was a corpse 🙁

If in doubt … wait

What was going on in the colony? Frankly, I didn’t have a Scooby’s 4.

Was the (apparently) new, unmarked queen mated or a virgin? Presumably the latter. However, other colonies in the apiary were requeening and it’s not unknown for a queen to go to the wrong hive when returning from a mating flight.

Would she survive the aggravation she was receiving from the workers in the colony?

Where had the new queen come from? If not from outside she must have come from a queen cell in the split hive. However, both sides of the split had new 2018 mated queens, and the timing wasn’t really right.

Under these circumstances the best thing is often to do nothing. I closed the hive up. My notes simply state “Dead BMCLQ! Virgin?? Left them to it.” 5.

No happy ending

Much as I’d like to be able to report that now, a month later, the colony is headed by a new mated queen laying frame after frame of worker brood … I can’t.

At the last inspection 6 the colony only contained several hand-sized patches of brood. However, it was all drone brood in worker cells.

The combination of drone brood in worker cells, with their characteristically domed cappings on sealed brood, coupled with the clustered arrangement of the brood clearly indicated that the colony contained a drone-laying queen (DLQ).

Drone laying queen ...

Drone laying queen …

Within the patches of drone brood were one or two attempts to make queen cells. These were abnormally shaped – either short, fat and unsculpted or overly long – and are often seen in colonies with DLQs or laying workers.

It’s too late in the year (here in Fife) to get a new queen mated – at least with any certainty. The bees in the colony were old and their numbers were much reduced. I therefore cut my losses and shook the colony out in front of a row of strong hives.

In retrospect

It’s difficult to see where things went wrong with this colony, or what I could have done to rescue the situation.

Perhaps the timing of my inspection – presumably very soon after the blue queen was killed – distressed the colony, causing them to ball (and possibly also kill) the new queen. Sufficient time then elapsed for the colony to rear a new queen (~16 days) from the eggs or larvae originally laid by the blue queen. However, this queen – who I never found – was either unmated or unsuccessfully mated and was a drone layer.

So, if in doubt … wait.

Particularly if it’s not clear what else to do.

But it still might end badly 🙁


Downstairs? Upstairs?

Colony inspections usually concentrate on the brood box. This is where all the action is. This is where the queen is and where there needs to be sufficient space for the colony to expand.

Or, if times are lean, sufficient stores and pollen to survive.

In contrast, the honey supers get no more than a cursory glance. There’s little of interest going on up there until it’s time to harvest the honey for extraction.

If the supers are light there’s nothing more to do other than hope for a good nectar flow in the future. In contrast, if they’re really heavy they might be ready to remove for extraction. If the frames are all capped the honey is ready.

Usually the supers are not heavy enough (a full super weighs something like 25kg) and they often don’t even get a glance, instead being bodily lifted off and left in a pile while the brood box is inspected.

Checking supers

Nectar has a high water content which the bees evaporate off during the production of honey. If they didn’t get rid of the water the stores would ferment. Since honey is hygroscopic they then add a wax ‘cap’ to the honey-filled cell to protect their stores for the winter.

Nectar is generally stored in the supers, starting in the middle of the middle frames and moving towards the periphery. This is the warmest part of the hive and presumably the easiest to evaporate water from. Therefore, the central frames in the super are most likely to contain capped honey stores.

Ready to extract

Ready to extract …

All I do when checking a heavy super is to first briefly look at the central frame to see if the stores are capped. If they are not then there’s no point in looking anywhere else in the super.

If the central frame is capped then it’s worth looking to see if the outside frames are as well. If so a clearer board can be placed below the super and you can take the honey for extraction.

Actually, there’s a bit more complexity as sometimes the honey is ready to extract, but isn’t capped. I’ll deal with that another time. The point I’m (slowly) trying to make is that supers are rarely checked in any detail … until they’re full.

It’s therefore interesting what turns up when you do remove them for extraction.

Pollen and stores-free area

With a strong colony, the bottom super i.e. the one immediately above the queen excluder, often has no honey stored in a semi-circular area immediately above the brood nest. Sometimes the edge of this clear area, adjacent to the honey, contains a band of stored pollen.

This clear area indicates that the colony need more space. The workers are keeping it clear for the queen to lay, but the queen excluder prevents her from accessing it. Sometimes you can get the bees to backfill this area by switching the super with one higher in the stack.

“Billy no mates” brood

It’s not unusual to find a very few scattered capped pupae in a stack of supers. These are almost invariably drone pupae, irrespective of whether the drawn super comb is on worker or drone foundation. In ~24 supers I extracted last weekend I saw three or four.

Billy no-mates ...

Billy no-mates …

I’ve always assumed that these were due to laying worker activity. There are always a few laying workers in a colony, but their numbers are suppressed by a pheromone produced by unsealed brood. Laying workers can be a significant problem in queenless and broodless colonies.

Since workers are unmated, the eggs that laying workers produce are unfertilised and so develop as drones 1.

There may be other explanations for these singleton pupae e.g. workers moving eggs up from the brood box. However, this doesn’t explain why they are almost always drones 2.

Clustered brood

Sometimes you’ll find a super packed with brood in all stages … wall to wall eggs, open and sealed brood. This happens when the queen has somehow sneaked above the queen excluder.

When this has happened to me I usually put it down to a lack of attentiveness in checking the underside of the queen excluder when opening the box. If the queen was on the underside and the QE is leant against the hive stand she can easily wander round to the other side, thereby giving her access to the supers.

Spot the queen

While checking supers for extraction last month I found one box – the lowest super of a stack of three – contained two or three frames with small amounts of clustered brood 3.

Another example of inattentiveness? Possibly, but there were some oddities about this colony.

Eggs and sealed brood ...

Eggs and sealed brood …

Firstly, there was no open brood … just eggs and sealed brood. I uncapped a few cells and the pupae were all just at the purple eyed stage. This is day 15 for workers and day 16 for drones. Since eggs hatch after 3 days this means that there had been a gap of at least 12 days when the queen wasn’t laying.

Half-sisters of the same age ...

Half-sisters of the same age …

Secondly, there was both worker and drone sealed brood present, but it was on separate frames. There was no drone brood in worker cells, which have characteristically domed caps 4.

Finally, I checked the brood box. There was plenty of brood in all stages – eggs, larvae and sealed pupae – in a busy hive. However, I didn’t see the queen (who was nominally marked and clipped) but by this time I was in a bit of a rush.

A partial solution

Some of these apparent oddities have a straightforward explanation.

The separation of drone and worker brood is because I use a range of different frames in my supers – worker foundation, drone foundation and foundationless. They start as matched boxes, but over the years have got completely mixed up.

All the drone brood was in a super frame originally drawn from drone foundation.

That was easy 😉

However, why was there brood at all in the super if the brood box contained the laying queen?

Or should that read a laying queen?

Perhaps there was another queen in the super?

Aside from speculating about how she got there, or – if she was the original queen in the box – where the one ‘downstairs’ came from, there’s also the puzzle about why she’d taken a 12 day holiday from egg laying.

And where the hell was she now?

She’d been in the top box sometime in the last 3 days (because there were eggs present). However, although I’m reasonably good at finding queens, I searched in vain in this super (and the two above) and couldn’t find her.

Time to be pragmatic

Carefully looking through ~30 super frames takes time and I was running out of both time and patience. These three supers were ready for extraction and I still had half a dozen colonies to check.

I could continue looking and eventually find her … if she was there at all.

If she wasn’t, I’d obviously never find her.

What did I do?

I shook all the bees off the super frames – directly over the brood box5 – and took them away for extraction.

I’m a great believer in Occam’s Razor. The simplest explanation is probably the correct one.

I reasoned that there was probably one queen in the box. Any other explanation was going to get convoluted.

If there was only one queen she was either in the brood box or the supers.

If she was in the brood box then all was well.

If she was in the supers she’d hopefully end up in the brood box.

There was little point in using a clearer board if the queen was in the supers. Firstly, with brood present many bees would probably remain. Secondly, if the queen was present in the supers, they’d definitely not clear.

Super frames with brood ...

Super frames with brood …

And … what happened?

I got well over 60 lb of honey from the colony 🙂

There was a blue marked and clipped queen in the bottom box when I checked the colony a few days later.

She was (still) laying well.

Unsatisfactory explanation

I suspect that the queen excluder was faulty or damaged. It was a wooden-framed wire one. If the wires were prised apart during cleaning or through carelessness the queen could get up into the super.

She could also therefore return to the brood box.

The 12 day gap in laying was probably explained by the queen returning to the brood box during this period.

The two short stints when she’d been ‘upstairs’ hadn’t noticeably left gaps in the brood pattern in the brood box – she might have only nipped up for a few hours or so. There were only a few hundred cells with eggs or pupae in the super.

And the most unsatisfactory thing of all … I thoughtlessly stacked the queen excluder with five others from the same apiary and so now need to carefully inspect all of them for damage 🙁


In perpetuity

Yet more frames ...

Yet more frames …

As I write this we’re approaching midsummer of one of the best years beekeeping I’ve had in a decade. In Fife we’ve had excellent weather, and consequently excellent nectar flows, for weeks. Queen mating has been very dependable. I’ve run out of supers twice and have been building frames like a man possessed.

I’m not complaining 😉 1

In a few short weeks it will be all over. The season won’t have ended, but this non-stop cycle of inspections, adding supers, building frames, splitting colonies, making up nucs, taking off laden supers, extracting and more inspections will be largely finished.

We’re in clover

Busy bees ...

Busy bees …

Literally, as it’s been yielding really well recently.

I’ve written previously about The Goldilocks principlenot too much, not too little – and bees. As an individuals’ competence improves over successive seasons, colony numbers can quickly change from too few to too many.

A single production 2 colony in a good year should probably also be able to generate a nuc for overwintering and possibly a new queen for re-queening without significantly compromising honey production.

That’s certainly been the case this year. I’ve got a few colonies that produced nucs in May, were requeened (through vertical splits) in late June or early July and that have produced several supers of honey, either from spring or summer flows.

Or in a few cases, from both. And it’s not quite over yet 🙂

But, there’s always a but …

I said in the opening paragraph it’s an exceptional year. The ability to produce a surfeit of both bees and honey requires some skill, some luck and some good timing.

In a bad year, just getting one of the three – a new nuc, a new queen or a honey surplus – from a colony should be regarded as a major success.

How do you cope with problems encountered in these bad years?


I’m a strong supporter of self-sufficiency in beekeeping. Although I’m not fundamentally opposed to purchasing queens or nucs, I do have concerns about importation of new virus strains and other ‘exotics’ that do or will threaten our beekeeping. However, buying in high quality bees for stock improvement is understandable, expensive at times and the foundation of at least some commercial (and amateur, but commercially viable) beekeeping.

I See You Baby

I See You Baby

What I’m far less keen on is purchasing bees – a significant proportion of which are imported – to compensate for lazy, slapdash or negligent beekeeping.

And there’s too much of that about … anyone who has been keeping bees successfully will have heard these types of comments:

  • Surely I can get away with less frequent inspections? I always have six weeks sailing in May and June … but I do want to make my own honey and mead
  • They all died from starvation sometime last year but I’ll buy some more in March from that online supplier of cheap bees (Bob’s Craptastic Nucs … Bees for the Truly Impatient)
  • Varroa treatment? Nope, not in the last couple of years mate. I’ve never seen one of them Verona, er, Verruca thingies so I don’t think my bees are infected with them anyway
  • I knocked off all the queen cells to stop them swarming in June and July. They just might be queenless. I know it’s early October but do you have a mated queen spare?

I’ve heard variants of all the above in the last few months.

In perpetuity

This stop-start beekeeping is not really beekeeping. I’ve discussed this in Principles and Practice extensively. I’ve called them beehadders before but perhaps the term ‘serial ex-beekeeper’ might be more accurate.

The reality is that, with a little skill, a little luck and just reasonable timing you can have bees in perpetuity … the real topic of this post.

In perpetuity meaning you are self-sufficient for stock and for spares.

You’re able to exploit the good years and survive the bad. You only need to buy in bees for stock improvement or to increase genetic diversity (which may be the same thing).

Once you’ve got bees, you’ve always got bees.

It’s a good position to be in. It gives you security to survive accidents, self-inflicted snafu’s and even the odd fubar 3. You are no longer dependent upon the importer, the supplier or your mate in the local association to bail you out. It gives you confidence to try new things. It means you can cope with vagaries in the weather, forage availability or simple bad luck.

How is this nirvana-like state of beekeeping self-sufficiency achieved?

I think it can be distilled to just two things – one is easy, the other slightly more challenging.

Firstly, you need to maintain a minimum of two hives. Secondly, you need to develop an appreciation of how the colony develops and understand when interventions and manipulations are most likely to be successful.

One is not enough

I’ve discussed the importance of a second hive previously. With one hive, beekeeping errors (or just plain bad luck) that result in a queenless, broodless and eggless colony might well be a catastrophe.

With two hives, you can simply take a frame of eggs from the second colony and voila, they’ll raise a new queen and your imminent categorisation as an ex-beekeeper is postponed.

Two are better than one …

The benefits of two colonies far outweigh the expense of the additional equipment and time taken to manage them. In a good year you’ll get twice as much honey to impress your friends and neighbours at Christmas, or to sell in the village fete. In a bad year, the ability to unite a weak colony headed by a failing queen in late September, might mean the difference between being a beekeeper and being an ex-beekeeper the following Spring.

Maintaining two colonies in the same apiary significantly increases your chances of having bees in perpetuity.

The art of the probable 4

Beekeeping isn’t really very difficult. You provide the colony with somewhere to live. You give them sufficient extra space to dissuade them from swarming (swarm prevention), or intervene in a timely manner to stop them swarming (swarm control). If you harvest some or all of the honey you provide them with more than they need of an alternative source of sugar(s) at the right time. Finally, you monitor and control the pathogens that afflict them and apply appropriate treatments, at the right time, to minimise their impact.

As you can see, timing is important. Do things at the right time and they work … at the wrong time they don’t.

Timing is also important in terms of the frequency of inspections (which I’ve briefly discussed before, so won’t repeat here), and in the manipulations of the colony.

These colony manipulations include – but aren’t restricted to – providing them space to expand, spreading the brood nest, making nucs, rearing queens or at least getting queens mated, adding supers, uniting weak colonies and feeding them up for the winter.

Again, if you do the manipulations at the right time they will probably work. Hence the ‘art of the probable’.

The time is right

For many of these manipulations, the ‘right time’ essentially depends upon the development of the colony and weather. And, of course, colony development is itself very much influenced by the weather.

Consider queen mating. Of the various manipulations listed above, this is one upon which the future viability of the colony is absolutely dependent.

Queen mating usually occurs mid-afternoon during dry, preferably sunny weather, on days with relatively light winds and temperatures of at least 18°C. Therefore if there’s a mature virgin queen in your hive 5, the weather is suitable and there are drones flying, she’ll probably get mated.

Good laying pattern ...

Good laying pattern …

Days like this occur pretty dependably in late May and June. It’s no coincidence that this is the peak swarming season.

Conversely, if through carelessness or neglect your colony goes queenless in late September, the probability of getting a warm, dry, calm afternoon are much less. It’s therefore less probable (and potentially highly improbable) that the new queen will get mated.

That’s not to say it won’t happen … it might, but it is less probable 6.

Beekeeping nirvana

In re-reading this post I feel as though I’ve skirted around the core of the issue, without satisfactorily tackling it.

Having bees in perpetuity is readily achievable if you have a backup hive and you understand how colony development and the weather determines what you can and cannot do to the colony during the season 7.

Having two hives but inadvertently damaging both queens in March during heavy-handed inspections will not provide bees in perpetuity.

Conversely, irrespective of your best efforts, a single terminally broodless and queenless colony at the peak of the swarming season cannot magically create a new queen … meaning you’re about to become an ex-beekeeper.

Another one for the extractor ...

Another one for the extractor …

I’ve used queen mating as an example because it’s a binary event … she’s mated successfully or she’s not, and colony survival absolutely depends upon it.

However, the timing of many of the other manipulations can also influence the strength, health and robustness of the colony. Providing too much space in cold weather delays expansion as there are too few bees to keep the brood warm. Trying to feed syrup very late in the season may mean it’s too cold for them to access the feeder, leading to starvation. Finally, using the wrong miticide at the wrong time is a guaranteed way to ensure more mites survive to damage the colony in the future.

Learn to do the right thing at the right time … to both your colonies. The recipe to having bees in perpetuity.


In (for or to) perpetuity means “for all time, for ever; for an unlimited or indefinitely long period” and  has origins in Latin and French with English usage dating back to the early 15th Century.

‘Unlimited or indefinitely long’ could also refer to the length of this post or the delay to my flight last Sunday. You can thank EasyJet for providing me with more than ample time to write this magnum opus.

Or write and complain for the very same reason 😉

Old and new duds

The Beast from the East ...

The Beast from the East …

Despite the best efforts of the Beast from the East 1 Spring is definitely on the way.

The snowdrops and crocus have been out for some time, willow is looking good, large queen bumble bees are searching for nest sites and the temperature here in Fife has consistently reached double figures during the warmest part of the day for the last week.

Consistently … but only just and only briefly.

Pollen boost

Pollen boost …

Consequently it’s too cold for full inspections and the only colonies I’ve been ‘in’ are the two described below. However, I’ve not ignored the others. I’ve lifted the crownboard on most colonies to determine their approximate strength (or just peeked through those with perspex crownboards which is even less intrusive) and have continued to heft colonies to see if they have enough stores. Those that were feeling a bit light have had a fondant top up. I’ve also given several colonies a pollen boost to help them rear early season brood.

Other than that – and moving colonies to the new bee shed – I’ve left them well alone.

Early season checkups

On the warmest part of the warmest day of the week I visited the apiary to check the colony strength. With the exception of two, all were flying well with foragers returning laden with pale yellow pollen.

However, two were suspiciously quiet, with only a handful of bees going in and out 2.

A pretty small handful.

Almost none of the bees returning to these two colonies carried pollen.

One was a five frame poly nuc in the bee shed. This had been made up in mid/late summer while the parental colony was requeened. The old queen, a frame of emerging brood with the adhering bees and a frame of stores had gone into the nuc box. The little colony had built up reasonably well going by my infrequent peeks through the transparent crownboard, but not well enough to move them to a full hive for the winter.

The other suspiciously quiet colony was a full (or full-sized 🙁 ) hive headed by one of the older queens in my apiary. Most colonies are requeened annually or every other year, but this one was reared in my first year in Scotland (2015) 3.

I popped the lid off both colonies and examined them in greater detail. It wasn’t the recommended ‘shirtsleeve weather‘ by a long-shot, but I feared the worst and didn’t think a bit of cold would do these two any further damage.

Unfulfilled promise

The nuc contained about a cup full of bees and a small, unclipped pale queen.

Overwintered virgin queen?

Overwintered virgin queen?

This definitely wasn’t the queen I’d put in the box last August. For whatever reason, the colony had clearly replaced the queen late in the year. It hadn’t swarmed, so it looks like they’d tried to supercede the old queen. Going by the total absence of worker brood I presume the new queen hadn’t mated successfully, or at all, and that she was a virgin.

She wasn’t running about skittishly like new virgin queens do, but she wasn’t doing anything very useful either.

There were a few drones in the colony and one or two sealed drone cells. Whether these were from unfertilised eggs laid by the queen, or laying workers, is largely irrelevant 4. The colony was doomed …

Worn out

The full sized colony was only full sized in terms of the hive it occupied. Inside there was another rather pathetic cupful of bees together with a very tatty, marked and clipped queen 5. There was more paint on her head than her thorax and I remember marking her with a very ‘blobby’ Posca pen. This was the queen I’d expected to find in the box.

Old and tired ...

Old and tired …

There were no drones in this colony, but no eggs either. There was also no sign of a second queen or evidence of attempted supercedure. I suspect the ageing queen simply ran out of sperm, stopped laying and never got started again.

Sometimes old queens turn into drone layers and sometimes they just stop. I’m not sure why they exhibit this different behaviour. It might actually reflect when they’re detected. I think I usually find drone laying queens a bit later in the Spring. Perhaps a failed queen starts laying (unfertilised) eggs only once the ambient temperature has risen sufficiently to help the much-reduced numbers of workers keep the brood nest warm enough?

That’s guesswork. It’s still cold here, with frost most nights. The small number of bees in the colony would have been unable to maintain the mid-30’s temperatures required for brood rearing. It’s surprising they’d survived this long.

Health check

Neither colony had any obvious signs of disease. The floor of the full hive was thigh-deep – if you’re a bee – in corpses.

Winter losses ...

Winter losses …

However, a good poke around through the cadavers failed to find any with signs of the deformed wings that are indicative of high viral loads. I hadn’t really expected to … the Varroa loads in this colony in the late-summer and midwinter treatments had been very low.

Corpses ...

Corpses …

Lose them or use them?

Clearly both queens had failed. Both were despatched. To keep them in the vain hope that they’d miraculously start laying again would have been a waste of time and, more importantly, other bees. The virgin would now be too old to get mated and there won’t be drones available here for at least 6 weeks.

This left the dilemma of what to do with the remaining bees. Both colonies were apparently healthy, but too small to survive. In the autumn the obvious thing to do is to unite small healthy colonies with large healthy colonies. This strengthens the latter further and helps them get through the winter.

However, this is the Spring. There were probably no more than 300 bees in either of the failed hives. All of these bees would have been at least 3 months old, and quite probably significantly older. They were unlikely to live much longer.

Furthermore, uniting these small colonies with larger colonies in the apiary would have caused disruption to the latter and increased the volume of the hive to be kept warm. Neither of these are desirable.

I therefore shook both small colonies out allowing the healthy flying bees to redistribute themselves around the half dozen strong hives in the apiary. Before shaking them out I either moved the original hive altogether or – in the case of the nuc from the shed – sealed the entrance, forcing them to look elsewhere for a colony to accept them.


The term dud is used these days to mean a “thing that fails to function in the way that it is designed to”, with this usage dating back to the 1914-18 war where it referred to shells that failed to explode. However, the word is much older. Its original meaning was a cloak or mantle, often of coarse cloth, with references to the word dudde dating back to the 14th Century. Over the next few hundred years the meaning, in the plural duds, evolved to mean clothes and – more rarely but more specifically – ragged, shabby clothes or scraps of cloth. This seemed appropriate considering the tatty state of the old marked queen …



The Autumn of the Matriarch

I’ve previously commented that weak colonies that build up very slowly in Spring are more trouble than they’re worth. The resources they need – syrup, frames of emerging brood, more TLC – are rarely reflected in the subsequent honey yield.

Quite the contrary, they’re often a lost cause and it could be argued that, from a purely efficiency point of view, it would be better if the colony succumbed during the winter than staggered on into the Spring.

Better still, assuming they’re disease free, use the bees in the autumn by sacrificing the queen and uniting the colony with a strong colony. You’ll boost the latter and strong colonies both overwinter better and build up better the following year.

Do as I say, don’t do as I do.

All the above makes perfect sense, but a combination of sentimentality and ill-placed optimism means that it’s not unusual – in late Spring – to find myself being reminded that “weak colonies that build up very slowly in Spring are more trouble than they’re worth”.

And it’s happened again.

One of my colonies was undersized in late autumn and had built up very slowly this Spring. The queen was a little older than most in the apiary but she’d done well in the past and I thought she might have another season in her. Varroa drops in late autumn and mid-winter had been very low and the bees were beautifully tempered, calm, steady on the comb and a pleasure to work with.

But in the first inspection of the year (10th of May) there just weren’t enough of them. The queen was laying, pollen was coming in, there were no signs of disease and the colony behaviour remained exemplary.

Lagging behind

Comparison between colonies is very informative. That’s why it’s easier to maintain two colonies than one. Other colonies in the same apiary were building up well. By late May I was starting swarm prevention measures on these, using pre-emptive vertical splits.

The small colony was largely forgotten or ignored. I peeked through the perspex crownboard a couple of times and could see they were building up.


I got distracted harvesting the early season honey from other colonies, running out of frames and with more swarm prevention and control. I finally completed a full inspection of the colony on the 17th of June, shortly before the summer solstice and the first official day of summer (so still technically Spring).

Queen failure … not epic, but failure nevertheless

The colony had only a couple of frames of brood and covered a frame or two more than that. The temper and behaviour was still very good. The queen was present and laying. She was being attended by a retinue of workers and not being ignored or harassed.

Failing queen ...

Failing queen …

But she was clearly losing her faculties. Many of the cells contained two or more eggs.

Multiple eggs in cells are often seen with laying workers and sometimes seen when a newly mated queen first starts laying. With laying workers the eggs are often placed on the sidewalls of cells and, as they’re unmated, they develop into drones. The brood pattern is scattered randomly around the frame. With newly mated queens the eggs are usually correctly placed in the base of the cell.

Occam’s razor

The colony was clearly doomed. They showed no sign of trying to replace the queen, without which the future was bleak. I needed to rescue something from the situation. The choice depended on my interpretation of what had gone wrong. The options were:

  1. Queen failure, plain and simple
  2. Laying workers in a colony with a failed queen still present (an unusual situation)
  3. A new, recently mated, queen was also present with the old queen (supercedure)

A thorough inspection of the colony failed to find another queen or any evidence of a recently vacated queen cell. Frankly this didn’t take long, the colony was simply too small to ‘hide’ either of these. Option 3 could therefore be discounted. The presence of another queen would be really important if I was considering requeening the colony or uniting it with a queenright hive – both these are likely to go badly if there was a queen still present.

There was no drone brood at all in the colony and the laying pattern was clustered as would be expected from eggs laid by a queen. Option 2 could therefore almost certainly be discounted. Fortunately again as it’s difficult to requeen a colony containing laying workers. As another aside, I can’t remember seeing a colony with laying workers that also contained a (failed) queen.

That left the most likely explanation for the multiple eggs (and the undersized colony) was the simple failure of the queen. For whatever reason, she was laying at a much lower rate than usual and had started laying multiple eggs in cells. Of the three possibilities, this is the most straightforward. Occam’s razor (William of Ockham, ~1287-1347) is the problem-solving principle that states that the simplest explanation is probably the correct one.

Better late than never

The queen was removed from the colony and it was united over newspaper on top of a strong hive in the same apiary. Two days later the Varroa board underneath the colony was covered in shredded paper indicating that the colonies were united successfully.

Successful uniting ...

Successful uniting …

Which is what I should have done in mid-autumn last year.

Better late than never  😉

A few days later I rearranged the colony, placing the two frames of brood into the bottom brood box and putting a clearer board underneath the top brood box. The resulting single colony, now a bit stronger, will be well-placed for the summer nectar flow and the nine frames of drawn comb vacated by the colony will be reused making up nucs for overwintering.

† Interestingly, I’ve never seen several larvae developing in cells after the multiple eggs hatch. Either the excess eggs or larvae must be removed by workers. I presume this means that the workers can’t count eggs, but may be able to count larvae – not literally of course, but by the amount of pheromones produced presumably. If they could count eggs they’d remove the excess and leave only one, making the identification of laying workers (or a recently mated misfiring queen) much more difficult. Something to be thankful for perhaps? They can, of course, identify the origin of eggs – this process is the basis of worker policing which was touched on in discussion of Apis mellifera capensis, and is of relevance to those using grafting for queen rearing.


The title of this post is a corruption of The Autumn of the Patriarch, a book by the Nobel laureate Gabriel García Márquez, written in 1975. The book is about the God-like power and status of a dictator, the General, and the awe in which he is held by the people. Of course, this isn’t the situation in matriarchal honey bee colonies, the structure of which is determined as much – if not more – by the workers, the brood and the circulating pheromones.