Category Archives: Queen rearing

Getting answers

Synopsis : Getting good answers involves asking good questions. But remember that the bees have no concept of what is ‘best’, or of the calendar.

Introduction

If you’re just starting beekeeping it’s likely you will have a never-ending list of questions about the somewhat arcane and often perplexing hobby you are embarking on.

And if you’ve been beekeeping for years (or even decades) you might have the same number of questions, albeit somewhat more specialised or esoteric ¹. You’ll also probably be involved in answering some of the questions from less experienced beekeepers.

Hive tools … which is the best?

Getting good quality and appropriate answers broadly depends upon three things:

who (or what) the question is directed at,
the wording of the question, and
whether the answer is simply factual or involves a subjective assessment

Let’s take a simple example …

Q. What is the scientific name of a bee?
A. The scientific name for the order of bees is Hymenoptera, but this encompasses many different species of bees, not just a single type of bee. To specify a particular species, a scientific name would be necessary, for example, the scientific name for a honey bee is Apis Mellifera.

… so, let’s be more specific …

Q. What is the scientific name of a honey bee?
A. The scientific name for a honey bee is Apis Mellifera.

By asking a more specific question you have received a better answer.

Unfortunately, it’s still not completely correct.

There is a convention on the capitalisation of scientific names; the name of the genus (in this case Apis) is always capitalised, but the name of the particular species (mellifera; meaning honey-bearing) should start with a lowercase letter.

The correct answer should therefore be Apis mellifera.

Or what?

You may have noticed the who (or what) in the list above. The ‘or what’ was deliberate because, increasingly these types of questions will be answered, not by humans or beekeepers ², but by a machine.

In fact, the two questions and answers shown above were answered by a machine … in this case one called ChatGPT. In this post, ChatGPT responses are shown in red text. The only reason you may not have heard of ChatGPT ³, released just 2 months ago, was because you’ve been living as a hermit in the Tora Bora cave complex for the last few years.

If ChatGPT is new to you then you’re going to have ample opportunity to experience it – or its successors – in the next few months and years. ChatGPT is described as a chatbot which has been trained through supervised and reinforcement learning. It can do a range of things that are a lot more impressive than not quite getting the scientific name of honey bees correct … it can compose music, write computer code and pass written exams for medicine. It can produce essays and it is causing considerable concern in education, where students are already using it for homework ⁴.

Perhaps I’ll save myself some late night writing by asking ChatGPT to do my homework …

Write a 500 word article for beekeepers about hive tools and Taranov swarm control, remembering to include a handful of poor quality puns and some truly shocking, punctuation.

I asked it to do this … it generates the answer in less than a minute. I’ve added the answer as a separate page as I don’t want to get even more distracted from the topic of the post today.

Factual inaccuracies

ChatGPT does OK on the poor puns (beeeing, buzz-worthy etc.) but less well on the shoddy punctuation, unless you count the Oxford comma.

It does much less well on Taranov swarm control. Indeed, I had started by asking it to “Describe how to conduct swarm control using the Taranov method” and received the following answer:

The Taranov method is a technique used for swarm control in beekeeping that involves manipulating the hive to prevent swarming. The process begins by identifying a colony that is likely to swarm and removing any queen cells, which are the cells that contain developing queen bees. Next, the beekeeper will divide the colony by removing a section of comb with eggs and young larvae, and placing it in a new hive box. This creates a population imbalance in the original hive, which triggers the bees to start raising new queen bees. Finally, the beekeeper will monitor the hive to ensure that no new queen cells are forming, and will continue to remove them as they appear. This method aims to reduce the likelihood of swarming, by keeping the colony balanced and preventing the development of new queen bees.

Which is almost completely wrong. I’m not going to describe Taranov swarm control, but it’s not that. If you want to know (and it is an intriguing method, though not one I have used) I suggest you ask an experienced beekeeper, not a machine. It requires a simple, correct, factual answer.

Subjective or qualified answers

The other thing ChatGPT is not good at is answering questions that require some experience and/or may be subjective. For example …

Q. What is the best type of beehive for a beginner?
A. For a beginner, the Langstroth beehive is considered the best option due to its standardized dimensions, ease of management, and wide availability of equipment and accessories. Additionally, it is commonly used by many beekeepers and has a strong community of support and resources available.

… OK, what about something even more important?

Q. What is the best type of hive tool for beekeeping?
A. A J-shaped hive tool is widely considered as one of the best for beekeeping. It is versatile, durable and can be used for a variety of tasks such as prying apart beehive boxes, scraping propolis and wax, and removing frames.

The first answer actually isn’t bad, particularly if you are in the US. However, the justification is a little odd and could equally well be applied to a National, Dadant ⁵ or Layens hive.

The ‘best hive’-type question is one commonly asked by beginners and one where a better question improves the answer. If you instead ask ChatGPT for the best beehive for a beginner in the UK it recommends a National hive (justified by compatibility).

However, the answer ChatGPT gives to the ‘best hive tool’ question is clearly wrong.

Simply the best … no contest!

Anyone with even passing experience of opening a hive knows that the claw-type hive tool is by far and away the most practical, comfortable and good looking 😉 .

The ’widely considered’ in ChatGPT’s answer is the giveaway. It’s (obviously) never used a hive tool and so cannot speak from experience.

Flame wars

The examples above are trivial but they do show both the abilities and shortcomings of ChatGPT. But it will get better – more accurate, more factually correct, better at providing qualified subjective answers (and hiding the fact that it has no direct experience of any of the things it is comparing).

You’d better get used to it as it will revolutionise our interactions – direct or indirect – with computers; websites, discussion forums, computer programming, teaching, student assessment and – increasingly – creative work as well.

It’s going to put a lot of people out of work 🙁 .

I’m already out of work, so I don’t feel too threatened, but perhaps it’s also going to eventually replace the beekeeping blogger.

As an aside, I thought a ChatGPT-powered ‘user’ on a beekeeping discussion forum like Beesource in the US, or the BeekeepingForum here in the UK would be – at least briefly – entertaining. Some of the discussion threads on these can get really out of hand, even with strict moderation (a thankless task).

ChatGPT can already interact conversationally, its command of the subject and of English (and, of course, a range of other languages) is already better than many readers/contributors, and it would be a whole lot more persistent in an online argument (though it is currently not particularly creative when it comes to insults).

It’s going to render many of these discussion forums worthless and is already banned from some of the computing forums. The scientific journal Nature has established ground rules for its use – it cannot ‘share’ authorship (!), its use in data analysis must be documented etc.

I already find many discussion forums unrewarding … they might get a lot worse, at least in the short term. Since many beginners use them a lot, I thought it was worth mentioning.

Better answers

Although I’ve been wandering some way off topic there are some important points embedded in the first half of this post.

The question you ask influences the answer you get.

In beekeeping, your local environment and your latitude are particularly influential in the lives of your bees.

I gave a talk this evening on queen rearing. One of the questions was ‘How early in the season can I start?’

That’s a perfectly good and valid question, but answering it requires knowing something about the local climate and colony development.

It also varies from year to year … for example, a cold spring delays things.

Some might simply answer ‘mid-May’ or ‘late-April’ … indeed, questions like that may be asked by someone wanting a calendar-based answer.

Unfortunately, beekeeping isn’t that simple. Beekeepers on the UK south coast can often start queen rearing two months before I can here in north west Scotland.

My answer involved something about drone availability. A drone takes 24 days to develop and a few days after that to become sexually mature. Studies have shown that the peak of drone brood production occurs about one month before swarming (Page and Erickson, 1988), though production starts earlier.

And the correct answer is …

So a better answer is to keep an eye on your colonies, observe drone brood production increasing and – 3-4 weeks later (or perhaps a little before ⁶ ) – start your queen rearing with every expectation the bees will have got the timing about right.

One of many …

In a cold spring they’ll start producing drones later, at a more southerly latitude they’ll produce drones much earlier than they will in northern Scotland.

Yes, the answer is more difficult to understand than ‘the 19^th of April’, but it’s much more likely to be correct because it is based on an understanding of the biology of the bees.

It’s also likely to be correct most years. You can test this by keeping notes. You’ll then have something to refer back to next year and the one after that, and you’ll be able to answer, with compelling authority, anyone who asks you the same question 😉 .

In about 30 years you’ll be able to review your notes – of drone production and queen rearing successes – and see whether the timing needs revising 🙂 .

In doing that you’ll have completed the transition from seeking a calendar-based response, to understanding the drivers that determine colony development and reproduction, and end up with an answer that is generically applicable, qualified and based upon personal experience.

Expect biased answers

My preference for a particular type of hive tool is based upon personal experience (and unrelated to the fact that I bought 20 of them very cheaply a decade ago). Do not underestimate the importance of personal experience in answering beekeeping questions … or its ability to generate biased, unqualified or even completely incorrect answers.

Ask three experienced beekeepers a question and you’ll get five answers … one will be completely wrong, another will involve ‘brood and a half’ (also wrong … obviously), a third will answer a different question altogether and the final two will express diametrically opposing views about whether the J-shaped or claw-shaped hive tool is ‘best’.

The answer you get is based upon the experience of whoever you ask … and how willing they are to answer.

Be warned, it’s not unusual for the most (usefully) experienced and the most vociferous beekeepers to be different individuals. In fact, it’s not unusual for the most vociferous to be much less experienced than they sound.

For a subject as practical as beekeeping, practical experience is far, far more valuable than ‘knowledge’ gleaned from the internet (after all, you might have been reading something written by ChatGPT).

I know the difference between the Miller and Hopkins methods for queen rearing. I’ve not used either (yet) so I don’t know which is better – either outright, or in particular circumstances.

This can all be a bit overwhelming as a beginner … use your judgement, listen, check some of the answers in a good book ⁷ or a reputable online source, ask a follow-up question.

Nobody knows all the answers and it sometimes feels as though the more knowledge you acquire, the more questions appear.

Answer your own questions; observation and understanding

There is no ‘best’ hive, or ⁸ hive tool. The bees don’t care and – through experience – you’ll find what suits your beekeeping.

It’s likely that the ‘best’ anything in beekeeping – bee, hive tool, hive, smoker, forage, honey, hive stand, extractor, queen excluder etc. – is a meaningless concept.

It’s an irrelevant question as far as bees are concerned. There may be good ones and bad ones, but it’s surprising how tolerant and accommodating the bees – and a beekeeper – can be.

I’d strongly recommend that anyone starting beekeeping ignores articles with the word ‘best’ in them – except perhaps this one.

Ask meaningful questions and look for insightful answers.

The ‘when to start queen rearing?’ is a good – albeit incomplete – question and I suggested how I would (or did) answer it above.

In that example it is really by observing and understanding the bees that you answer the question. I think those are two of the most important skills to acquire as a beekeeper; doing so will always help you get better answers, not least because they help answer them yourself.

Asking an experienced beekeeper gets you part way there but it doesn’t come close to working something out yourself.

Do an experiment

I used the word ‘arcane’ in one of the opening sentences. It means mysterious, obscure or little understood. Despite sounding a little like the word archaic – meaning old fashioned or belonging to an earlier period – it has a totally different etymology. Arcane is derived from the Latin arcānus meaning ‘closed or shut up’, whereas archaic is from the Greek ἀρχαϊκός for ‘ancient’.

Nevertheless, bits of beekeeping are both arcane and archaic.

Sometimes they’re ‘old fashioned’ because experience has shown that a particular method works reliably well, so is promoted and becomes widely used. However, sometimes it’s because ”it’s always been done like that” and everyone unquestioningly follows the approach without asking whether there are other – perhaps better – ways of achieving something.

Let’s take a trivial example … starter strips in foundationless frames.

Beautiful …

A foundationless frame is a frame containing no foundation (helpfully, the clue is in the name) . Every foundationless frame you use saves you about £1.40 based upon the current price of foundation. What’s more, because commercial foundation contains miticide residues, every frame you use reduces traces of miticides in your hive.

And, if you read online about making foundationless frames you’ll find lots of descriptions, many of which include instructions to provide a wax, or waxed, starter strip attached under the top bar for the bees to start drawing comb from.

When I started using foundationless frames I – unquestioningly – followed these instructions, cutting 2 cm strips of commercial foundation and nailing them in place in my frames.

Foundationless frame

And they often fell out … which prompted me to ask a question I should have done in the first place, and to do an experiment.

Ask the bees

Where do bees naturally start drawing comb?

Wherever the swarm ends up. If it’s in a previously unoccupied tree hole, or loft space, there are no convenient strips of wax foundation to be used as guides. The cavity may be completely empty.

Clearly they don’t need a starter strip to work from.

However, the beekeeper does not want the bees to build comb totally haphazardly. We want them to stick within the confines of the frame or it becomes impossible to manipulate. It’s therefore useful to provide them with ‘guides’ on where to start … they might not follow them, but they almost always do.

So, by understanding what the bees do naturally, qualified by our own selfish interests in managing the colony, we can ask the bees what suits them … a strip of wax foundation, a waxed wooden guide or a lollipop stick.

And, having worked out which the bees prefer, we can make a decision based upon what also suits us.

Take your pick …

I therefore did a simple experiment. I built a dozen or so foundationless frames, each containing three randomly positioned starter strips under the top bar – one third wax foundation, one third waxed wood and one third plain wooden lollipop stick (or ‘tongue depressors’).

Over the course of a season I used these in a variety of colonies and observed which of the starter strips was preferred … indicated by which the bees chose first, or which they avoided.

And the results are in …

It made no difference at all. There was no correlation between the type of starter strip and use by the bees.

They just don’t care.

So I only now use plain wooden starter strips. They are quicker and easier to prepare, impervious to the steam wax extractor and much more robust. I use them in full hives and in my mini-nucs for queen mating, simply gluing the wood in place.

Kieler mini-nuc topbar frames – no need for foundation or waxing

A little understanding of what the bees do, coupled with a little experimentation and some observation, allowed me work out the answer to the question ’What is the best material for starter strips’ ⁹.

I should note that many other have reached exactly the same conclusion independently. I’m not claiming to have discovered this first, but I did formally test what the bees preferred.

Some ChatGPT creativity to end with

To finish I thought I’d ask ChatGPT to do something creative.

Rather than eliciting incorrect answers about Taranov swarm control I asked it for a poem, in the form of a haiku ¹⁰, about honey bee swarms;

Golden bees swarm forth
Honeyed scent on the breeze
Nature’s sweet bounty.

Not bad 🙂

References

Page, R.E., and E. H. Erickson, Jr. (1988) Reproduction by Worker Honey Bees (Apis mellifera L.). Behavioral Ecology and Sociobiology 23: 117–126 https://www.jstor.org/stable/4600197. Accessed February 3, 2023. Note: this might not be accessible online, so you could try this instead – Page, R.E. Jr (1982) The seasonal occurrence of honey bee swarms in north-central California. American Bee Journal 121:266-272.

Learn from my mistakes

12 Replies

Synopsis : “It’s good to learn from your mistakes. It’s better to learn from other people’s mistakes.” Here are some related ones of mine and the lessons I (again) learned from them.

Introduction

This is the 50th post of the year. Usually ¹ this is the week in which I write some erudite words ² about the season in retrospect.

What went well, what could have gone better and what was an unmitigated disaster. The highs and the lows, together with a smattering of the in-between bits that actually constitute the majority of the beekeeping year.

Writing these review-type posts is quite enjoyable ³. I have to go back through my hive notes which inevitably brings back a flood of memories of warmer days, heavy supers, lost hive tools and unmated queens.

Laden foragers returning …

These notes are succinct – rarely more than ~10-20 words – littered with acronyms but surprisingly informative. At least they are when you take into account their dates and the timing of the development cycle of workers, drones and queens.

And, as a bonus, they are also legible.

Hive records

I’ve mentioned before that I take my hive inspections records on a digital recorder in the apiary and then transfer them to a spreadsheet with the company of a coffee ⁴ later.

Testing, testing, one, two three

Whilst this approach misses the immediacy of a scribbled note on a sheet of paper under the hive roof, it does allow me to easily refresh my memory before the next apiary visit.

It also means they don’t blow away, get damp, eaten by slugs or require me to have a functioning pen in my beesuit … the recorder lives in the bee bag, so is always available.

By looking at the hive records in advance I’m reminded that – for example – I need more supers for hives #23 and #27, that the former had a good open queen cell on frame 7 (OC/7 in the notes) which should now be sealed ⁵ and that nuc #13 has a recently mated queen who will need finding (!), clipping and marking.

And by looking at the same records in mid-December, with the temperature plummeting outside, I can see how the season progressed week-by-week in my three main apiaries.

The nuc was sold a fortnight later, the queen in #23 eventually emerged but was lost before she got mated (or perhaps on her way back) and supers filled so rapidly that I regretted not adding more at once.

It was a great year for honey, the best since returning to Scotland in 2015.

But that can wait until the review of the season … which will now be next week ⁶.

Impressions vs. reality

I give a lot of talks on beekeeping. Whilst a talk might be about bait hives, swarm control or queen rearing, the questions afterwards can be more wide-ranging. It is quite fun to get questions on apparently random beekeeping topics (I’ve done ’ask the expert’ ⁷ panels at beekeeping conventions which are enjoyable; for one question and four panel members you get six different answers and a good-natured argument … no wonder we run out of time).

Some of the more ‘left field’ questions can be a challenge and require a degree of lateral thinking. I’m more than happy to acknowledge that I’m sometimes stumped for an answer … I understand the question, but I cannot explain why whatever happened ⁸, er, happened.

However, the very fact that I’m asked for an opinion at all suggests that I give the impression I know what I’m talking about ⁹ when it comes to almost any subject about honey bees.

I don’t.

Hive inspections

Even if you just restrict the topic to the hive, its contents and what’s happening in it on a week-to-week basis, the apparent impression some have of my understanding can be wildly different from reality.

I know that because of my hive notes.

And I’m acutely aware of this because I’ve just re-read them … 😉 .

Perhaps I’m not being entirely fair. If you read the entries for any individual week in isolation it’s clear I sometimes don’t have a scooby ¹⁰ what’s happening in the hive.

It’s obvious because as well as acronyms there’s a dusting of ??? throughout the notes ¹¹.

Not every hive and certainly not every week (thankfully), but more often than I’d like.

However, the regular repeat visits combined with the utter predictability of honey bee development, and comparison with adjacent hives, usually allows me to work out what’s happening with some certainty.

I think the take home message ¹² here is that hive notes are very useful. Even if you don’t quite understand what’s happening in the hive, record what you see.

At the next visit, or the one after that, it should start to make sense.

And, when you re-read them in midwinter with the benefit of ~~knowing what happened~~ time, tea and a roaring stove, you’ll at least learn what to do in the same situation next year.

Or what not to do 🙁 .

Train wreck

But, there remains this disconnect between that impression of insightful understanding and the day-to-day reality of some of my apiary visits.

These are are not always as polished and assured as I’d like …

Photo by Roberto Prusso from 1913

… frankly, they can sometimes be a bit of a train wreck.

I trip over or drop stuff ¹³ .

I reassemble the hive leaving the queen in a JzBz cage in my pocket.

Hives are inspected in the wrong order, meaning I might have to re-open one again to the understandable irritation of the bees.

I leave the hive tool inside a hive and can’t find another, or put it in the wrong pocket of my beesuit so it falls through the hole, down the inside of the leg and into my boot.

An elusive unmarked queen is found on my third run through the brood box … at exactly the same time as I find a suicidally psychotic worker inside my veil. I have insufficient hands to hold the frame, the queen and the worker (though, truth be told, I’m probably not intending to just ’hold’ the worker if she’s that agitated). The queen is abandoned until the following week.

When I fail to find her altogether.

Or, I do and she dies as I pick her up.

Composed and unruffled

Thankfully, not every apiary visit is like that.

Many are reassuringly ‘composed and unruffled’; I open the box, the bees are wonderfully calm. The queen is on the second frame I check, but the first had eggs anyway so I didn’t need to see her. However, since I found her and she’s mated and laying well, I mark her and clip her wing. The colony barely notice her absence and I run her gently back in between two brood frames to get on with her business.

Returning a marked and clipped queen – not distressed, just sauntering down between the frames

This is the beekeeper I’d like to always be … not that other ham-fisted muppet in an ill-fitting beesuit.

Here are a couple of queen-related examples from the season just gone. At the time I was flummoxed. In retrospect I’ve learned a few important lessons.

Or re-learned those that I had been taught already … and subsequently forgotten 😉 .

A tale of several queens

Almost all of my swarm control these days uses the nucleus method. In comparison to the widely-taught Pagden artificial swarm it only requires an additional nuc box. In my experience, it is almost totally foolproof as long as:

you take care to only leave a single queen cell in the hive; remember, this requires a visit one week after removing the queen. They cannot swarm if you leave just one cell.
you don’t make the nuc too strong; if you do there’s a chance it might swarm anyway.

My colony #7 contained an ageing but really lovely queen. I’d used her last year for queen rearing and hoped to do so again. At the start of May I’d obviously tempted fate by noting … ’Strong. Lovely BP. PC only … graft next week’ ¹⁴.

Lovely BP (brood pattern)

And, of course, the next week (the 8th of May) there were a couple of big, fat, 3-4 day old (i.e. open) queen cells and it was clear they were making plans to vamoose. I made up a nuc (number #47) with the old queen and moved her to a distant apiary.

One cell was left in box #7.

This box had overwintered with a nadired super and the original queen had snuck down in April and laid it up. Irritatingly, she’d chosen the frames of drone comb … not the end of the world because the queen was so good, but I didn’t want the new queen to do a repeat performance so moved the super above the queen excluder (QE). I discussed this in Early season inspections back in April – go there for all the gory details.

Upper entrances

Drones emerging above a QE is a recipe for carnage as they try and squeeze through ¹⁵, so I added a thin eke that included an upper entrance. The drones could fly from this and, in due course, they did.

Hive with an upper entrance

In the meantime, the new queen should have emerged and got mated … but she was nowhere to be found. There were polished cells in the brood box and the colony was behaving as though it was queenright.

On the 12th of June I decided to give it ’one more week’ and, eventually, I then found her in the supers on the 19th, together with several frames of open brood.

D’Oh!

Since there was no sealed brood she had presumably been slow getting mated and only started within the last week or so. I clipped and marked her and returned her to the brood box.

But she didn’t stay long.

My notes on the 21st of July include the comment ’no eggs’ (there had been on the 12th) and at the end of the month I found a new laying queen in the box.

She’s still there … or was when I last checked the colony (’lovely, well-tempered bees’ I was pleased to record in my notes) in mid-September. I presume the previous one hadn’t been ‘up to scratch’ and so was superseded sometime in mid-July .

Lessons learned

When I removed the supers (unsurprisingly disappointingly light 🙁 ) and the plastic QE I found the latter had a couple of cracks in it. Although these didn’t result in any gaping holes, it’s possible the QE deformed sufficiently to let the queen through.

Lesson No. 1 – try and avoid using those ~~inexpensive~~ cheap plastic QE’s. A framed, wired excluder costs less than three jars of honey and is a good investment. I bought more at the start of this season, but still didn’t have enough.

You know it makes sense

However, I suspect the queen didn’t get into the supers through the QE.

I expect she instead returned from her last mating flight and entered the hive via the upper entrance. It’s not at all unusual for returning queens to end up in the wrong hive altogether, so simply mistaking an upper for the lower entrance is understandable.

Lesson No. 2 – ideally avoid upper entrances altogether on hives containing virgin queens, or at least orientate them to the opposite side of the hive.

What happened to the nuc?

The original queen was moved to another apiary in nuc #47 created during swarm control. A fortnight later (22/5/22) they were moved to a full hive ¹⁶ and three weeks later (12/6/22) I found a single charged queen cell in the middle of a central frame.

Although I suspected supersedure (ageing queen, single QC etc.) I’d seen the queen and she was still laying well. I therefore transferred the frame with the open cell to a neighbouring queenless and poorly tempered hive in the same apiary. She emerged, was mated and laying well by late July … and is heading the colony through this winter.

However, back to hive #47 … they attempted to swarm ~10 days later. Since the queen was clipped the swarm returned to the hive, but the queen was lost 🙁 .

I knocked back all but one of the charged open cells and guesstimated that she would emerge on the 6/7th of July. On my next inspection (on the 5th), the queen was quacking in the cell ¹⁷. My maths suggested she should be mated around the 20th but I didn’t open the box again until the last day of July when I found her laying well.

And it was about then that things started to go ‘pear-shaped’ ¹⁸.

Hands like feet

I fumbled the first couple of attempts at picking up the queen to mark her.

In my defence … it had been a long afternoon. This was the last of about 15 colonies I’d inspected and the supers were very heavy. It was hot, the sun was bright and I was really tired.

I’d removed the supers and set them aside, lifted the QE and found the unmarked laying queen after just a couple of minutes.

The first attempt at picking her up was pretty reasonable, albeit unsuccessful, but the second was badly botched. She skittered across the frame, clambering over and under the workers without giving them a chance to move aside.

Lesson No. 3 – when it comes to picking up queens, if at first (or at the most, second attempt) you don’t succeed, do not ’try, try and try again’. Close the box, leave her for another day when both she and you will be calmer.

Of course, I persevered 🙁 .

On the third attempt she took off, seemingly in slow motion, but still too fast for me. She circled around, briefly landed on my shoulder, lifted off again and then – like Baron von Richthofen – was obscured by the dazzling sun.

I never saw her again.

But it gets worse …

I’ve had queens fly off before. In my experience they often (perhaps over 50% of the time) manage to return to the hive. They fly poorly and often crash land in the grass. It’s therefore important not to stampede about like a herd of wildebeest, but instead to calmly reassemble the hive and quietly retreat.

Ideally without moving your feet.

I managed the former but not the latter, cursing my failure to learn from ‘Lesson No. 3’ … it is not the first time that I’ve attended that lesson 🙁 .

I rarely inspect hives in August. This hive was on the other side of Scotland and I didn’t open it again until I removed the supers during the summer honey harvest at the end of the month. With only one and a bit filled supers there were relatively few bees above the QE … but there was a queen.

At least, there was evidence that there had been a queen present as there was a ~5 cm patch of brood on opposing faces of two of the super frames.

Small patch of brood – all of about the same age – in a super

I didn’t see the queen, and only discovered the brood as I shook the bees off the frames to remove them for extraction.

It was a patch of similarly aged brood; there were no younger larvae or eggs, and only a few cells were capped. It must have been laid 7-9 days previously.

Remember, I’d not opened the hive in the last 23 days.

There was no upper entrance and the QE was wired.

And the final lesson today

I ended up uniting the hive over a QE with a known queenright colony. I simply laid some newspaper over the top bars of the latter, added a QE to hold it in place and added the ’is-there-or-isn’t-there-a-queen-in-this-box?’ upper brood box onto which I shook the bees from the remaining super frames.

Newspaper and queen excluder

I do not understand the brood in the super.

The brood pattern indicated it was laid by a queen (rather than laying workers – in which case it would be scattered all around the frame). However, the age of the brood suggested she had only laid for a day or so. The brood looked like worker, but I didn’t photograph the frame until it was back in the extracting room and it was a bit bashed about by then.

My best guess is that the queen that flew away landed back on top of the open stack of supers next to the hive. However, that was over three weeks earlier. If it was the original laying queen that I’d failed to mark (or even pick up) then why didn’t she continue laying at the same rate?

Why the fortnight’s hiatus?

And why did she stop laying having only just started?

I’m afraid we’ll never know.

Which doesn’t mean that there’s not a lesson from this sorry debacle:

Lesson No. 4 – if a mated and laying queen does fly off, check the hive again one week later. If she’s back she’ll be laying and all will be well. If she’s missing in action there will be queen cells, but no remaining larvae young enough to rear queens from. Knock back all the cells you don’t want and let them get on with things ¹⁹.

One final queen

If you rewind about 1500 words you might remember that I found two good open queen cells in box #7. The one I didn’t leave in that hive I used to requeen an adjacent hive (#6).

Waste not, want not 😉 .

A week previous I’d already started swarm control on colony #6 by removing the queen to a nuc. I’d intended to let them requeen themselves ²⁰ but the availability of a good queen cell from much better stock was too good an opportunity to miss.

I knocked back all the queen cells in hive #6, confident that there were no larvae young enough start as queens.

I added the frame from #7 with the queen cell and she emerged on about the 17th of May. She was mated and laying well by the 12th of June … probably earlier but I hadn’t checked at the intervening visit as I’d been taking three full supers of spring honey from the hive. I got a further three full supers of summer honey 9 weeks later.

The bees are well tempered and productive. They went into the winter very strong … headed by the same queen.

Sometimes, actually most of the time, everything ’just works’ 🙂 .

And a final word

The tale of misadventures recounted above was probably the only truly shambolic part of my beekeeping year in 2022 ²¹. It was just a coincidence that it all originated from one hive. Some of it was self-inflicted, but other bits were just dumb luck.

Shambolic, but not catastrophic, and it all ended OK.

And I learned some valuable lessons.

Again 🙂 .

Notes

The opening quote, “It’s good to learn from your mistakes. It’s better to learn from other people’s mistakes”, is attributed to Warren Buffet.

A few readers signed up to receive notifications either about new posts or comments added to the post last week will have received repeat emails sometime on Tuesday evening. These were seemingly due to a hiccup in the server software and outside my control. With thanks to those who brought this to my attention, and apologies to the recipients. The current commenting and notification software is becoming ‘unfit for purpose’ due to subscriber numbers. I have plans to change, both along with some other developments, sometime next year.

The most fun you can have in a beesuit

9 Replies

Synopsis : Queen rearing is enjoyable and educational. Don’t let the experts put you off. You don’t need to graft day-old larvae to rear queens.

Introduction

A long time ago ¹ I bought, read and re-read Ted Hooper’s excellent book Guide to Bees and Honey. Every time I read it I’d find something I’d missed the last time and, even now, there are nuanced comments I think I am only now beginning to understand.

I’m exaggerating slightly when I say ’read and re-read’ as there was one chapter I pretty-much skipped over each time.

That was the chapter on queen rearing.

What put me off?

It was probably his description of opening queen cells with the tip of a penknife to check how far development had progressed, re-sealing the cell and returning the frame to the hive.

She’s gone …

I knew enough about bees to know that the future success of the hive depended upon it successfully requeening after swarming.

But I didn’t know enough to stop them swarming 😉 .

I’d also already had to ‘borrow’ a frame of cells from a friend to rescue a terminally queenless colony of mine. ’Enthusiastically clumsy’ defined my beekeeping skillset, and was probably the comment the ² examiner made in his notes during my BBKA Basic assessment.

The prospect of meddling with developing queens, with something so precious, seemed like total madness.

Surely it’s better to let them get on with it?

For the first couple of years of beekeeping, I thought of queens as an exquisitely fragile – and by implication valuable – resource. The prospect of rearing them, handling them, putting them in little boxes or – surely not? – prising a cell open to see if they’d developed sufficiently, was an anathema to me.

Consequently, I repeatedly skipped the chapter on queen rearing.

Too difficult … not for me … nope, not interested.

The BBKA Annual Convention

Before they moved the event to Harper Adams, the BBKA used to hold its spring convention at the Royal Agricultural showground just outside Warwick. My (then) local association provided stewards for the event and I was asked – or volunteered – to help the late Terry Clare run the queen rearing course one year.

I’d never done any queen rearing … and still hadn’t completely read that chapter in Hooper’s book.

I’d like to take this opportunity to apologise to those who paid to attend the course … at least those who received any ‘help’ from me, though everything else about the course was very good.

Checking grafted larvae

After an introductory lecture from Terry, we spent a warm afternoon in a poorly lit room practising grafting larvae. A thin cloud of disorientated bees circled our heads before being ushered out through the windows. Most of the larvae on the frames were visible from across the room ³ but at least they didn’t turn to mush with our neophyte fumblings as we transferred them from comb to plastic queen cups.

Terry moved from table to table, checking progress. He explained things well. Very well. The preparation and procedures seemed a whole lot more accessible than they had in Hooper’s book.

I’m a reasonably quick learner and that afternoon convinced me I should, and could, at least try it on my own.

The session ended with a wrap-up lecture in which Terry encouraged us all to ‘have a go’, and not be put off by an initial lack of success.

He assured us it would be worthwhile and enjoyable.

We dispersed into the late afternoon sun, talking of bees and queens and our plans for the season ahead.

Balmy April weather

There was an early spring that year, colonies had overwintered well and were strong. The Convention was held in early April if I remember and the good weather continued for at least another 2-3 weeks.

Well before the end of the month I had my first successfully grafted larvae being reared as queens.

Success!

It wasn’t an overwhelming success.

I probably grafted a dozen, got half accepted, lost more during development ⁴ and ended with just two virgins. I don’t have notes from those days, but I’m pretty sure only one got successfully mated.

So, success in a very limited way, but still success 🙂 .

It still makes me smile.

Terry’s presentation had clarified the mechanics of the process. It no longer seemed like witchcraft. It was all very logical. He’d made it clear that the little specialised equipment needed was either ’as cheap as chips’ ⁵ or could easily be built at home by someone as cack-handed as I ~~was~~ am ⁶.

The practical session had given me confidence I could see and manipulate ~~huge fat larvae that were far too old to be reared as queens~~ larvae. Even with my ’hands like feet’ moving a delicate larva from comb to plastic queen cup seemed possible, if not entirely natural.

JzBz plastic queen cups

I scrounged some JzBz cups from someone/somewhere, built a cell bar frame and some fat dummies ⁷ the week after the Convention and used one of my colonies as a cell raiser and the other as the source of larvae.

And, at a first approximation, everything sort of worked.

I could rear queens from larvae I had selected 🙂 .

Try, try and try again

I repeated it again the following month. I was more successful. The nucs I produced were either overwintered or built up strongly enough to be moved into full hives.

I think one went to my mentee. My association encouraged relative newcomers to mentor, probably one of the best ways to improve your beekeeping (other than queen rearing).

Within a year I had 6-8 colonies or nucs and twice than number the year after that.

Almost all were headed by queens I had reared … ‘almost’ as my swarm control skills were still developing 😉 .

Now, over a decade later, my swarm control skills have improved considerably … as has my queen rearing.

I remain resolutely cack-handed but I’m now a lot more confident in my hamfistedness.

I still mainly use the same technique Terry Clare taught on that course in Stoneleigh, though I’ve now also used a number of other approaches and successfully reared queens using most of them. Even the cell bar frame I built is still in use, though I’ve built some fancier fat dummies.

Fat dummy … with integral feeder and insulation

Queen rearing has taught me more about keeping bees than any other aspect of the hobby … more about judging the state of the colony, the quality of the bees, the suitability of the environment, the weather, the forage etc.

Queen rearing has improved the quality of my bees, year upon year, so that they suit my environment and colony management.

But – more importantly and perhaps a little selfishly – queen rearing has given me more enjoyment than any other aspect of beekeeping.

I’d prefer to rear queens than get a bumper honey crop … but because I rear queens that suit me and the environment, I do pretty well for honey as well.

10%

I give 20-30 talks a season to beekeeping associations. When I’m talking about queen rearing I usually ask the organisers about the number in their association that actively rear queens.

By actively I mean that do more than simply allow colonies to requeen themselves during swarm control. Don’t get me wrong, I’m not denigrating this essential aspect of beekeeping. We all (have to) do it.

To me ‘active’ queen rearing doesn’t necessarily mean grafting larvae, incubators, mini-nucs and all that palaver. But it does mean:

preparing a colony to be in a suitable state to rear new queens ⁸.
rearing queens from larvae selected (though not necessarily individually selected) from a colony with desirable characteristics e.g. good temper, productivity, frugality.
rearing more than one queen at a time, with the excess used for making increase, for sale, for ’just in case’ situations etc.

There’s perhaps a slightly grey area where you split a hive (with desirable characteristics) that’s making swarm preparations into multiple nucs, each of which gets an immature queen cell.

But, let’s not get bogged down in definitions … that’s not the point of this post (which, although it might not be obvious yet, is to encourage you to ’have a go’).

And, when I ask ⁹, I’m regularly told that only a small number, perhaps ~10%, of association members actively rear queens.

Why so few?

Enjoyable, educational, useful … choose any three

Of course, there’s no requirement that a beekeeper gets involved in queen rearing. You can keep bees for years without rearing queens, other than during swarm control and by making up splits. I know a few beekeepers who have been keeping bees like this for decades … by many criteria they are skilled and successful beekeepers.

But sometimes, which might mean ‘often’, being able to rear queens and having some of those ‘spare’ queens available is extremely useful.

Spare queens, heading nucs in the apiary, can be overwintered to make up losses. These can be sold or donated in Spring to meet the enormous ¹⁰ demand for bees early in the season. The availability of a queen can ‘fix’ an aggressive colony, can rescue an otherwise doomed colony, or can effectively ‘gain’ a month of brood rearing and nectar collection should the old queen fail.

And that extra month of brood might make the difference between successful overwintering or not.

In my view, once you can rear your own queens you are pretty-much self-sufficient … there are very few situations that cannot be rescued.

And all of those benefits are before you even consider the two other things I mentioned above:

that successful queen rearing will inevitably improve your more general skills as a beekeeper, and
you will get a lot of satisfaction and enjoyment from doing it … literally ’the most fun you can have in a beesuit’ ¹¹.

Why so few?

Beekeeping, like many other hobbies, can appear an esoteric pastime. Weird terminology, hierarchical organisation ¹², specialised equipment, unusual costumes and a tendency to still use arcane practises.

And queen rearing – probably like candle making or the production of excellent mead ¹³ – is a specialised niche within what is already a rather niche activity.

It has its own terminology, equipment and methods.

To the uninitiated – even to another beekeeper, like me reading Ted Hooper’s book – it can appear fiendishly difficult.

And, unfortunately, some practitioners make it sound esoteric, specialised and difficult.

It’s a sort of one-upmanship.

They promote methods that may not suit the beginner, that require lots of resources, or that involve techniques that sound exceptionally skilful, even when they’re not. Not deliberately perhaps, but that’s what happens.

All of which means that:

people are dissuaded from trying it in the first place
those that do try (with trepidation because, you know, ”it’s difficult”) and that achieve only limited success, have their initial impression reinforced and are unlikely to try again

It’s very easy to talk yourself out of trying something you think will be difficult and/or you are unlikely to succeed at.

Actually, it’s not only easy, it’s also entirely understandable.

Why go to all that trouble if it’s unlikely to work?

After all, you can usually buy queens ‘next day delivery’ for £50 … surely that would be easier?

Perhaps … if they’re available when you want them. Really early in the season? Think again. During the peak swarming season when everyone else wants to requeen their colonies they accidentally destroyed all the queen cells in. Nope.

But, as Terry Clare so ably instructed … it is not that difficult to rear your own.

There’s more than one way to do it

I’ve written an entire post on this topic and it applies as much to queen rearing as it does to other aspects of our hobby.

If not more.

There are many different ways of successfully achieving the three key components of the process:

preparing the colony to receive larvae
presenting the larvae
getting the resulting virgin queens mated

Today’s post isn’t an introduction to queen rearing … it’s meant instead as an encouragment to try queen rearing.

If you’ve got a year or two of beekeeping experience and one, or preferably two, colonies you have the essentials you need to start. It’s what I started with … and look how that ended 😉 .

Over the next three months I’ll write two or three more posts on the basics, in good time for you to ’have a go’ in 2023.

Preliminary setup for Ben Harden queen rearing

If you’re impatient to read more, I’ve already written about two methods I have used extensively – the Ben Harden system and queen rearing with a Cloake board.

However, throughout these descriptions I’ve emphasised the use of individual grafted larvae.

Grafting is the transfer of larvae from the comb where the egg hatched to a wax or plastic queen queen cup. For best results the larvae should no more than ~18 hours old.

A suitable larva may well be no bigger than the egg it hatched from.

Already I can feel beginners switching off … “Too difficult … not for me … nope, not interested.”

Although grafting is an easily learned and reasonably straightforward technique it can appear very daunting to the beginner.

Perhaps I’m therefore also guilty of making queen rearing sound ‘esoteric, specialised and difficult’.

Am I guilty as well?

Indubitably, m’lud.

But … in my defence please consider the two recent posts on ’Picking winners’.

The purpose of those posts was to highlight – for people (like me) that already routinely use grafting as part of their queen rearing – that the bees may choose different larvae to rear as queens than the beekeeper might choose.

The beekeeper is essentially non-selective, whereas the bees are very selective.

I think this is interesting and it’s got me wondering about the qualities the bees select and whether they’d be beneficial for my beekeeping.

But there’s another equally important ‘take home message’ from these two posts. This is relevant to beekeepers who do not already rear queens (but who would like to) but that are put off by the thought of grafting.

And that is that you can easily produce excellent quality queen cells without grafting or ‘handling’ larvae at all.

If you refer back to that three point list above, point 2 ( ‘presenting the larvae’) can be as straightforward as simply adding a frame of eggs and larvae to a suitably prepared hive.

That’s it.

What could be easier?

No magnifying glasses, no headtorch, no treble ‘0’ sable paintbrush, no JzBz plastic cups, no cell bar frame, no ’do I or don’t I prime the cups with royal jelly?’, no desperate searching around the frame for larvae of the right size, no worries about larvae getting chilled, or drying out …

Pick a frame, any frame

As long as it has eggs and young larvae … and comes from a donor colony that has the characteristics you like in your bees.

Eggs and young larvae

There’s little point in rearing queens from poor quality bees.

For starters I’d suggest you select a frame from a colony of calm, well behaved bees.

If none of your colonies are dependably calm and well behaved you definitely need to learn to rear queens, but you should ask a friend or mentor ¹⁴ for a frame of eggs and larvae from a good colony.

Bees are very good at picking larvae suitable for rearing into queens. Let them do the ‘heavy lifting’. Once the queen cells are ready you cut them out of the frame and use them in the same way as you would use cells from grafted larvae.

So, having hopefully convinced you that you don’t need to graft larvae to produce queen cells, that seems like a logical place to end this post.

In future posts I’ll discuss points 1 and 3 in that numbered list above.

You already know almost everything you now need to know about point 2 😉 .

Picking winners, part 2

16 Replies

Synopsis : Some larvae are nutritionally deprived and may produce sub-optimal queens. Grafting may miss the ‘best’ larvae the colony would select for rearing as emergency queens.

Introduction

A fortnight ago I discussed the preference colonies show for heavy eggs – or more accurately for larvae reared from heavy eggs – when producing queens under the emergency response.

Why might this be interesting?

The longevity of the queen and the absolute dependence the colony has on her quality means that the choice of larvae they rear new queens from is of fundamental importance.

These are the larvae that develop to produce queens with the traits that benefit the colony – in fecundity, disease resistance and a range of other characteristics.

It cannot be random.

If they make the right choice the colony will flourish, swarm and their genes will be perpetuated.

That is a significant evolutionary selective pressure. Its application over the 90 million years or so since the evolution of eusociality has resulted in the honey bees we have today.

Now, these traits favoured by the bees might not all benefit our beekeeping, but some of them should. Longevity, fecundity and disease resistance are likely to be evolutionarily favourable traits, and will also be useful for beekeepers.

Defensiveness and swarminess … er, not so much 😉 .

But the bees have little time to select the best larvae. They have 6 days from the day the egg is laid until a larva is too old ¹ to reliably develop into a queen.

In practice they select very young larvae (or even 3 day old eggs) so ensuring the resulting queen is fed for the maximum time with royal jelly, thereby producing a larger queen with more ovarioles.

So what do the bees choose?

Given the choice, which larvae are selected by the bees to rear new queens?

Artificial experiments and nepotism

The ‘heavy eggs’ experiment I discussed a fortnight ago was primarily designed to study kin selection and nepotism in honey bees. The study was conducted over a decade ago ², but wasn’t published until 2021, though the results were known before then ³.

If you remember, nepotism in honey bees is a nice idea; particular patrilines of workers (fathered by the same drone) should favour larvae of the same patriline. However, there have been no convincing studies that actually support this, and there are compelling theoretical arguments why nepotism could actually be detrimental to the colony.

Parts of the study I’m going to discuss this week were also designed to test for nepotism. I’m going to ignore these ⁴ and instead focus on some more interesting results that I think have practical relevance for beekeeping.

In addition, the study this week uses methods that are more typical of those used by beekeepers and that avoid the artificiality of rearing larvae in vitro before reintroducing them to a queenless colony.

In this regard I’d argue that they more closely resemble what’s happening in a colony rearing emergency queens. Furthermore, they should be easier for beekeepers to understand, and to repeat … not for experimental purposes, but when rearing queens.

Sagili et al., (2018)

The majority of the studies I’m going to discuss are from Sagili et al., (2018). The title ’Honey bees consider larval nutritional status rather than genetic relatedness when selecting larvae for emergency queen rearing’ neatly summaries their conclusions, but some of the detail is worth discussing in a bit more detail.

It’s always interesting to know what goes on in the hive.

During inspections we see frames of brood – capped and open cells. Other than the larvae getting bigger as they get older they all look much of a muchness … but they’re not.

All larvae are equal, but some are more equal than others 😉 .

Hungry mouths

The queen lays an egg in an empty cell. Other than the egg, the cell remains empty for 3 days when the egg hatches to release the larva. Without prompt and regular feeding the larva will starve or suffer setbacks in development.

Unrealised potential … a frame with eggs and young larvae

For this reason the nurse bees make frequent visits to the occupied cells to determine their content and needs.

Is it an egg or a larva?

Is it hungry?

And these visits continue during the 5 days of larval development.

How many visits do they make, how often is a larva fed, and are all larvae treated equally?

Sagili and colleagues used observation hives and video cameras to record nurse bees visiting cells containing larvae of precise ages ⁵. They recorded visits over 4 hours to 2 day old larvae, and one hour observations of 5 day old larvae.

In four separate hives, 4-8% of the young (2 day old) larvae did not receive a visit from a nurse bee during the 4 hour period they were filmed.

Of the 5 day old larvae, again ~10% didn’t receive a visit during the observation period and, of those fed, the longest interval between feeds was ~36 minutes. However, over one hour, the older larvae that were being fed were visited very regularly; the median interval between feeds was a little under 4 minutes and they were fed for a total of ~7 minutes over one hour.

Clearly some larvae, for whatever reason, get little or no attention for extended periods, whereas those that are visited, are fed very frequently.

Nutritionally deprived and non-deprived larvae

Larvae that are infrequently visited are likely to be nutritionally deprived … or, using the technical jargon beloved of beekeepers and scientists alike, hungry ⁶.

Do nurse bees respond differently to nutritionally deprived and non-deprived larvae?

Which are visited first and fed first?

The scientists caged the queen on a frame and allowed her to lay eggs for 24 hours. They then removed the queen (caging her elsewhere in the hive) and waited for the eggs to hatch. 24 hours later the larvae were caged under either 13 mm mesh or 3 mm mesh. Workers can access the larvae through the 13 mm mesh, but cannot get through 3 mm mesh. Cages were left in place for four hours to create two populations of larvae on the same frame; nutritionally deprived and non-deprived.

Small and large mesh cages over day-old larvae

They then again used video recording of randomly selected larvae to record and quantify the attention and feed visits they received.

The purpose of this part of the study was to determine whether the nurse bees could discriminate between larvae that were nutritionally deprived and those that were not.

And they could …

Inspections and feeding visits to nutritionally deprived and non-deprived larvae

Deprived larvae were visited (inspected) sooner, the black bars in the graph above, and fed earlier. You can just about determine this from the graph; the stats are more convincing (but less comprehensible 😉 ).

In addition, deprived larvae received more frequent inspections, more frequent feeds and were fed for longer.

Acceptance of larvae for queen rearing

A colony rendered suddenly queenless will attempt to rear a replacement under what is called the emergency response. Suitable young larvae are selected, fed a diet rich in royal jelly and the cell is reshaped to be orientated vertically.

This vertical orientation is a major inducement for the workers to continue to feed the developing larva with royal jelly (She et al., 2011). This is exploited in queen rearing techniques that involve the grafting of young larvae into wax or plastic cups which are then placed, open end down, in a queenless colony.

Sagili et al., investigated whether nutritionally deprived and non-deprived larvae were favoured when queens were reared under the emergency response.

Interestingly, they did so using larvae in natural comb and following grafting into plastic queen cups.

Which were favoured for queen rearing? Grafted or natural, nutritionally deprived or non-deprived?

In both instances the larvae were presented to a queenless and broodless colony, using 6 recipient colonies in each case.

Larval acceptance for queen rearing using two different methods – grafted and in natural comb

In the case of grafting, 12 of each type of larvae were presented on a cell bar frame. When transferring comb (prepared as described before using caged larvae) the entire frame was introduced.

The recipient colony did not discriminate between the nutritionally deprived and non-deprived larvae when they were grafted, but they showed a marked preference for the non-deprived larvae in natural comb.

In addition, they reared significantly more queens from grafted larvae than they did from larvae in comb.

All larvae are equal, but some are more equal than others …

Since I’m enthusiastic about queen rearing, this last set of experiments was by far the most interesting part of the study.

There are two results that are particularly striking.

Firstly, more queens were reared from grafted larvae than were reared following the transfer of a frame of larvae. The difference was significant, with almost twice as many queens being produced following grafting. It’s also worth noting that the bees only had 24 grafted larvae to choose from, compared to a much larger number of larvae on the transferred natural comb.

More is better … right?

Secondly, the workers showed no preference between nutritionally deprived and non-deprived grafted larvae, but showed a strong preference for the well-fed larvae in natural comb.

So, what do these results mean?

Let’s have a quick recap:

developing larvae receive different amounts of ‘attention’ from nurse bees
about 10% of developing larvae received no (or only a limited number of) visits during an extended observation period. The presumption is that these larvae are likely to be nutritionally deprived (though this was not demonstrated)
nurse bees can readily distinguish between nutritionally deprived and non-deprived larvae; the former receive earlier inspections, are fed sooner and more frequently
when rearing emergency queens, workers preferentially select larvae in natural comb that are not nutritionally deprived. In contrast, they make no distinction between nutritionally deprived and non-deprived grafted larvae

We know from numerous studies that high quality queens must be well fed during larval development. The best queens are produced from very young larvae (or even 3 day old eggs) that are then fed for an extended period with copious amounts of royal jelly in a strong hive full of nurse bees.

Queens produced under these conditions are larger and have more ovarioles, so should lay more eggs for longer.

It makes sense that nurse bees can distinguish between ‘hungry’ and replete larvae … the former need feeding or they won’t develop properly. The former may already have been held back developmentally … not an ideal start for a new queen.

Since nurse bees can determine the nutritional status of very young larvae, logic would dictate that they would select those that are not nutritionally deprived to rear new queens from.

After all, the future of the colony, and any resulting swarms will depend on it.

So, why don’t they make a similar distinction when presented with grafted larvae?

Selection vs. maintenance of larvae for queen rearing

The key difference between the grafted larvae and those in natural comb was the orientation of the ‘cells’ in which the larvae were presented to the queenless and broodless colony.

Grafted larvae were presented in a vertically orientated plastic queen cup, whereas the cells in natural comb are horizontal ⁷.

Cell bar frame with vertically orientated plastic Nicot queen cups

The interpretation is that larvae in vertically orientated cells are not selected by the nurse bees, but are instead just maintained as developing queens.

In contrast, larvae in horizontal(ish) natural comb are selected as the starting material for new queens, and the resulting reshaping of the comb to form the queen cell leads to their maintenance as developing queens.

Significance for beekeeping and queen rearing

The increased number of queens reared from grafted larvae probably reflects this ‘maintenance’ response being triggered in the nurse bees. ’Any’ larva presented in a suitably orientated cell must have been preselected as suitable … so the bees feed them up with royal jelly.

The nurse bees don’t know that these grafted larvae were selected by the beekeeper, not on the basis of them being nutritionally replete, but more likely because they were visible, about the right size, and in an accessible part of the comb.

But … think back to the first experiment. This suggested that ~10% of all larvae are nutritionally deprived because they have received, at best, infrequent visits over the last few hours. If the beekeeper hadn’t picked them during grafting, it’s unlikely the bees would have selected them as being suitable for producing queens.

Are 10% of grafted queens sub-optimal? Remember, the differences may be subtle.

The other point I found interesting is that the bees reared fewer queens in natural comb than from grafted larvae.

Why?

Charged queen cell

One possibility is that the reshaping of the comb is a physical limitation and restricts queen production. Perhaps this is why so many are at the edge of combs? ⁸

Another is that the bees only rear ’enough’ queens for their needs, but that they can only determine what ‘enough’ is using larvae in natural comb.

Whilst these are certainly possible I think there’s an intriguing alternative … only a small proportion, even of well fed larvae, are considered suitable by the colony for queen rearing and so selected by the nurse bees.

Yes, the bees favour larvae that are not nutritionally deprived, but perhaps there are additional characteristics that are also desirable (and that vary between larvae).

Quantity and quality

Grafting larvae is a well established method of producing large numbers of queens. If the donor colony is good quality there’s every reason to expect that the resulting queens will be good.

Most, if not all, commercially reared queens come from grafted larvae (where quantity is paramount and quality might be a secondary concern) … probably hundreds of thousands of queens a year are produced like this. It’s the method I’ve used for many years.

Queen cells from grafted larvae …

But this paper raises two or three interesting ideas:

about 10% of larvae selected at random for grafting are likely to be nutritionally deprived and so would not have been chosen by the nurse bees. The presumption is that these will produce sub-standard queens.
nurse bees might be a lot more selective in the larvae they choose for emergency queens, only favouring a subset of even those not nutritionally deprived.
queen rearing methods that present larvae in natural comb might produce fewer queens but those queens may have the desirable characteristics selected by the bees (potentially resulting in better quality but a smaller quantity).

If this last point is correct, it’s worth noting that queen rearing methods – like the Hopkins method – that use larvae in frames placed horizontally over the colony ⁹, may trigger the queen maintenance response rather than allowing the selection of larvae by the nurse bees.

It would be very interesting to determine whether the bees would discriminate between nutritionally deprived and non-deprived larvae presented on a horizontal frame.

Two final thoughts;

Grafting works well, but that doesn’t mean it’s the best way to produce top-quality queens.
The desirable characteristics nurse bees favour (for colony survival and reproduction) may not be beneficial for beekeeping.

But I’d be surprised if they weren’t 🙂

Note

I’ve not had a chance to discuss it, but Free et al., (1989) previously demonstrated that nutritionally deprived larvae received more attention from nurse bees. I’ll deal with how the workers detect the nutritional status of larvae in the future.

References

Free, J.B., Ferguson, A.W., and Simpkins, J.R. (1989) The Effect of Different Periods of Brood Isolation on Subsequent Brood-Cell Visits by Worker Honeybees (Apis Mellifera L.). Journal of Apicultural Research 28: 22–25 https://doi.org/10.1080/00218839.1989.11100815. Accessed November 22, 2022.
Sagili, R.R., Metz, B.N., Lucas, H.M., Chakrabarti, P., and Breece, C.R. (2018) Honey bees consider larval nutritional status rather than genetic relatedness when selecting larvae for emergency queen rearing. Sci Rep 8: 7679 https://www.nature.com/articles/s41598-018-25976-7. Accessed November 21, 2022.
Shi, Y.Y., Huang, Z.Y., Zeng, Z.J., Wang, Z.L., Wu, X.B., and Yan, W.Y. (2011) Diet and Cell Size Both Affect Queen-Worker Differentiation through DNA Methylation in Honey Bees (Apis mellifera, Apidae). PLOS ONE 6: e18808 https://journals.plos.org/plosone/article?id=10.1371/journal.pone.0018808. Accessed November 22, 2022.

Picking winners, part 1

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Synopsis : Queenless colonies prefer to rear new queens from heavy eggs. How was this determined and what are the implications for our queen rearing?

Introduction

Arguably the most important decision a colony will ever make is the selection of the eggs or larvae from which a new queen is raised. Other decisions are obviously important, such as the nest site a swarm occupies, but if the choice of ’starting material’ for the new queen is poor then the resulting colony is unlikely to thrive.

Actually, I suspect this isn’t arguable at all; whether it’s a replacement queen to take over after the colony swarms, or a supersedure queen to replace the ageing matriarch as she runs out of sperm or energy, a poorly chosen larva will – sooner or later – result in the demise of the colony.

Let’s hope they’ve chosen a good ‘un (they will have!)

Conversely, a good larva, fed well by nurse bees, that mates with enough drones and evades marauding swallows on the return to the hive – and the clumsily wielded hive tool of the beekeeper – will end up heading a strong colony. This strong colony will collect a surfeit of pollen and nectar, so ensuring good overwintering survival. It will be better able to defend itself against wasps or other robbing bees, and will be less susceptible to disease ¹.

Reproduction

A strong, healthy colony will build up well in the spring and produce one or more swarms ². If these survive – undoubtedly also helped by having good genetics – the colony will have reproduced and can be considered successful.

Honey bee reproduction in action

This type of ’success’ is what evolution selects for, so you can be absolutely certain that the choice of eggs/larvae from which new queens are reared is not random.

Cooperation vs. nepotism

Rearing new queens involves cooperation. In fact, as eusocial insects, almost everything that happens in the colony is cooperative. Multiple nurse bees feed the developing queens, hundreds of scout bees survey the environment for new nest sites and thousands of related workers provision the hive with pollen and nectar.

It’s often stated that these workers are ‘half sisters’ … they share the same mother (the queen) but different fathers (drones).

And there are quite a lot of fathers … .

The queen mates with at least a dozen drones during the mating flights she takes. Some calculations suggest it’s significantly more than a dozen drones. Whatever the number, workers fathered by the same drone will be more related to each other than they will be to workers fathered by a different drone.

On average workers within a single patriline (i.e. fathered by the same drone) are supersisters and share 75% of their genes. In contrast, workers in different patrilines (i.e. different drones) only share 25% of their genes.

And this is potentially a problem for cooperation.

It might be expected that nurse bees would select their supersister larvae when rearing new queens. Doing so would help ensure the propagation of their genes in subsequent generations, rather than those of their half sisters.

This would be an example of nepotism; ’showing special favour or unfair preference to a relative’ ³.

Lots of studies have attempted – largely unsuccessfully – to demonstrate nepotism in social insects, but that doesn’t mean it’s not worth looking again.

Do worker honey bees exhibit nepotism when selecting larvae to rear new queens?

Nepotism vs. colony diversity

It’s easy to talk yourself out of an experiment.

You have a good idea, do a bit of reading, discuss it with your friends and collaborators and then – belatedly – consider the underlying theory.

At which point it all sort of falls apart and you find numerous reasons not to do the experiment in the first place.

It was a daft idea because of x, y and z.

Think of all the time and money you’ve saved … back to the drawing board.

And there are good theoretical reasons why nepotism is unlikely to be seen in social insects like honey bees.

The most compelling of these is that genetic diversity within the colony is beneficial.

And nepotism, by definition, reduces diversity.

A quick recap on the diversity story … colonies with limited genetic diversity e.g. those headed by poorly mated queens, are less ‘fit’ than colonies with extensive genetic diversity. Fitter colonies are bigger, stronger, healthier and more likely to reproduce. The seminal study on this was by Mattila and Seeley (2007) which I discussed briefly in Polyandry and colony fitness.

So, theoretically, nepotism is a ‘bad thing’ … don’t bother doing the experiment.

But hold on a second, we also know that different patrilines of workers ‘smell’ very different to each other because they produce distinct cuticular hydrocarbons (CHC).

If nepotism is such a ‘bad thing’ why retain the (evolutionarily ‘expensive’) genetic machinery to generate all these different CHC’s? Why not just make all workers from one queen distinct from those derived from a different queen?

Individual colonies need to have distinct CHC’s to prevent robbing, but why are different patrilines distinct in their CHC profile?

Maybe nepotism occurs after all?

Better do the experiment.

Nepotism and larval selection

The study I’m going to briefly discuss was recently published by AL-Kahtani and Bienefeld (2021). It’s interesting and reasonably definitive in my view. However, whilst it addresses the ”Do bees exhibit nepotism during larval selection?” question ⁴ I think there are features of the study that are somewhat artificial which might restrict the generality of the conclusions they reach.

More interestingly, and of relevance to practical beekeeping, they show that bees are highly selective in their choice of eggs/larvae.

Can beekeepers exploit this to produce better quality queens?

The experiment was very simple.

Simplified diagram of the experimental method (see text for details)

Unmated queens from diverse areas of Germany were instrumentally inseminated with sperm from 10 drones, each selected from different unrelated geographic areas.

Six colonies were established (only three shown above) which were subsequently split into a queenright egg-producing colony (EPC; presumably a nuc, though it’s not stated) and a queenless larvae-rearing colony (LRC).

Eggs laid within a 6 hour window were incubated for 48 hours in an incubator, weighed and then allowed to hatch. For the first 48 hours after hatching the larvae were artificially reared by feeding them a sugar/protein diet ⁵.

This artificial rearing was done to avoid any bias from non-genetic colony odours e.g. due to pollen/nectar.

After 48 hours, 30 larvae, 10 from the matched EPC and 10 from each of the unrelated EPC’s were grafted into plastic queen cups and presented to the LRC for rearing as queens.

The larvae selected were obvious as these were fed and wax was deposited to create the surrounding queen cell.

Did LRC’s preferentially select larvae from the matched EPC?

No.

This larval transfer was done several times to get statistically meaningful results, using six colonies, repeated either twice or three times in successive years. In total 450 grafted larvae were presented to the LRC’s.

Larval acceptance rates were ~48-60%, a figure often exceeded when grafting for queen rearing.

Capped queen cells produced using the Ben Harden queenright queen rearing system

I suspect this rather mediocre acceptance rate reflects the in vitro rearing of the larvae for the first 2 days, potentially compounded by the age of the larvae which – at 48 hours – are at least 30 hours older than optimal.

But the acceptance rate doesn’t really matter as it was similar whether the larvae were derived from the matched or unmatched EPC. This therefore ’contradicts the hypothesis that kinship plays a central role in the selection of larvae for queen breeding’ (to quote the authors verbatim).

Larval selection is not nepotistic.

But certain larvae were preferentially selected

Despite the fact that the bees didn’t appear to care whether the eggs were from a related colony or not, they did preferentially select larvae produced by certain queens.

And I’ve already given you a clue of the characteristic favoured by the workers … though the characteristic per se wasn’t directly selected by the workers bees.

The six different queens used in this study produced eggs that differed slightly in weight. On average, the heaviest and lightest eggs varied in weight by ~5%.

There was a significant and direct correlation between the average weight of eggs produced by a queen and the likelihood that the resulting larvae would be selected by the larval rearing workers.

Heavier eggs produced larvae that were favoured by the cell raising colony.

Relationship between average egg weight and whether they were accepted for queen rearing

Of course, the cell raising colony never saw the eggs … these were hatched in an incubator and fed for two days before grafting and introduction.

Nevertheless, there was something about heavy eggs that the larval rearing colony favoured.

A total of 248 virgin queens were produced from the 450 larvae grafted (55%). These virgins were weighed and subsequently naturally mated, resulting in 190 egg-laying queens (42% of grafts, or 77% of virgins). Of these, 147 came to a grisly end as they were dissected two months after they started laying to count the number of ovarioles (the sub compartments of the ovaries in which the developing oocytes are produced).

Queen weight and ovariole number have previously been considered as markers of queen quality. Perhaps disappointingly, there were no significant differences in terms of virgin queen weight, ovariole number or the delay in onset of egg laying between queens produced from heavy or light eggs.

Crude criteria of what’s best

I’m not unduly concerned that the crude criteria we use to judge the quality of these queens (weight and ovariole number) failed to demonstrate significant differences. These criteria may not be the same as the ones selected by the bees ⁶. The fact that we cannot measure differences in the resulting queens does not mean that there were not qualitative differences in queens reared from heavy eggs that would benefit the colony.

Or would have been if they hadn’t been dissected 🙁 .

Bigger AND better … or just bigger?

It just means we were probably not measuring the right things.

However, extending this experiment from the relatively straightforward ‘heavy eggs are favoured’ observation is not trivial. If the scientists cannot see a difference in the queens then they might have to look at colony performance over time, or in adverse years, or when swarming, or in hard winters etc. Each of these may directly or indirectly act as a selective pressure on the queen quality, and hence the choice the bees make in the initial eggs or larvae that are selected for queen rearing.

Caveats

The relationship between egg weight and larval acceptance shown above is based upon the average egg weight produced by each of the 6 queens used in the study.

These average weights varied by about 5% (154.9 to 162.7 micrograms). That’s not a lot, and is narrower than the range of egg weights produced by an individual queen. Unfortunately, this data isn’t presented, but it can be inferred from the standard deviation of the mean egg weight.

For example, the average weight of the heaviest eggs was 162.7 micrograms with a standard deviation of 22.2 micrograms. With some basic assumptions of the distribution of weights, that means that 68% of the eggs were between 140.5 and 184.9 micrograms, but the remaining 32% were heavier or lighter.

Clearly, queens produce eggs that vary considerably in weight … and this has also been shown in previous studies (e.g. AL-Kahtani et al., 2013).

I would have liked to see a graph of the weight of individual eggs and an indication of whether or not the resulting larva was accepted as starting material for a new queen.

Secondly, there are methodological problems – acknowledged by the authors – in the relationship between queen quality and egg weight. So few queens were reared from lightweight eggs that it was difficult to determine if these produced poor quality flyweight queens with low numbers of ovarioles.

You can only work with what’s available.

Emergency response and egg/larval selection

The other two caveats I have are to do with the experimental design. The study involved rearing queens under the emergency response; larvae were presented to queenless and broodless colonies. For survival, they had to rear queens from the material presented (but still exhibited a preference).

However, I’d suggest that the vast majority of queens reared by honey bees – over the millions of years that have shaped the evolutionary choices we are now testing – are produced under either the swarming or supersedure responses.

Is egg/larval choice under the emergency response the same?

We don’t know ⁷.

The non-random construction of queen cells.

Finally, it has been shown that colonies prefer to rear queens from 3 day old eggs rather than 48 hour old larvae. I understand why the authors reared the eggs in vitro, but it does rather ignore the known preferences of the colony (see The bees know best for more on this topic).

Yes … they had no option other than to choose between the offered 48 hour old larvae … but would they have made the same choice if they had been given the eggs in the first place?

Why are heavier eggs preferred … ?

This is where we get to speculation and I’m going to save the discussion for after a follow-up post ⁸ in the next fortnight or so.

The bottom line is we don’t really know, but we have some pretty good ideas (though some are extrapolated from other social insects).

However, there’s a related question; ”How are the heavier eggs/larvae selected?” … and I think it’s fair to say this remains unclear ⁹.

… and is this relevant to our queen rearing?

When I rear queens I select larvae from a colony that shows some or all of the traits that I favour in my bees.

I’m a simple beekeeper and I have very simple needs … I want my bees to be calm, well-tempered, steady on the comb and frugal in winter. The best colonies that exhibit these traits are used as a source for grafting larvae when queen rearing.

Nice bees and a nice queen

In contrast, colonies that exhibit lots of chalkbrood, have poor temper, run about the comb or – worst of all – ‘follow’ are never used for queen rearing. Nor are they allowed to replace their queen during swarm control. Instead, these are requeened (as early as practical) from better stock.

I’ve described before my ‘rule of thirds’. When comparing the sum total of the various traits I care about, the best third are used for queen rearing. These queens are used to requeen the worst third and – if there are spares – the intermediate quality third as well.

However, if I run out of queens I’m reasonably happy to let the middle third requeen themselves (for example, during swarm control).

You’d be surprised how quickly the average quality of your bees improves using a strategy like this.

Grafting larvae vs. letting the bees choose

But the ‘best third’ are defined solely by my criteria.

I ignore any preferences the bees might have by choosing the larvae when grafting.

Assuming the queens that head these top third ‘good’ colonies produce a range of egg sizes (which they will), the bees would preferentially select the larvae from the largest eggs.

I just pick the larvae of the right age that I can see ¹⁰ and transfer them to a Nicot plastic queen cup.

Eggs and young larvae

Not the same thing at all.

Perhaps it doesn’t matter? After all, thousands of apparently satisfactory queens are reared by grafting every season.

Perhaps the characteristics the bees select for – whatever they are – are irrelevant for beekeeping? We don’t know, but I’d bet that some of the criteria that benefit the bees – and are evolutionarily selected – might well benefit beekeeping.

Poor ‘take’

Sometimes I get 100% ‘take’ i.e. all the grafted larvae accepted and reared as queens ¹¹.

Sometimes it’s less, a few times it’s almost none 🙁 .

Cell bar frame with three day old queen cells, The Apiarist.

3 day old queen cells …

In the latter instance I usually assume that the cell raising colony is not sufficiently ‘receptive’ but perhaps I’ve chosen undersized larvae (for their age), or perhaps the donor queen only produces undersized larvae (again, for their age)?

In the best tradition of “If at first you don’t succeed, try, try and try again” I usually just have another go. Almost always, I have another go in exactly the same way.

Perhaps if I used larvae from a different (but still good) colony the take would be improved?

Or perhaps if I presented the larvae in a manner that allowed the bees to select those to be developed into queens I might get improved acceptance? ¹²

I could end up with more queens and – potentially – better queens.

It’s blowing a hoolie tonight

And, as another autumn storm winds itself up to come barreling in from the Atlantic, dreaming about balmy May afternoons in the apiary and improved ways to produce better queens is about as close as I can get to beekeeping 😉 .

References

AL-Kahtani, S.N. and Bienefeld, K. (2021) ‘Strength surpasses relatedness–queen larva selection in honeybees’, PLOS ONE, 16(8), p. e0255151. Available at: https://doi.org/10.1371/journal.pone.0255151.
Al-Kahtani, S.N., Wegener, J. and Bienefeld, K. (2013) ‘Variability of Prenatal Maternal Investment in the Honey Bee (Apis mellifera)’, Journal of Entomology, 10(1), pp. 35–42. Available at: https://doi.org/10.3923/je.2013.35.42.
Mattila, H.R. and Seeley, T.D. (2007) ‘Genetic Diversity in Honey Bee Colonies Enhances Productivity and Fitness’, Science, 317(5836), pp. 362–364. Available at: https://doi.org/10.1126/science.1143046.