Category Archives: Queen rearing

Brexit and beekeeping

The ‘oven ready’ deal the government struck with the EU in the dying hours of 2020 was a bit less à la carte and a bit more table d’hôte.

The worst of the predictions of empty supermarket shelves and the conversion of Essex into a 3500 km2 lorry park have not materialised 1.

But there are other things that haven’t or won’t appear.

And one of those things is bees.

Bee imports

There is a long history of bee imports into the UK, dating back at least a century. In recent years the number of imports has markedly increased, at least partially reflecting the increasing popularity of beekeeping. 

Going up! Imports of queens, nucs and packages to the UK, 2007-2020 (National Bee Unit data)

Queens are imported in cages, usually with a few attendant workers to keep them company. Nucs are small sized colonies, containing a queen, bees and brood on frames. 

Packages are the ‘new kid on the block’ (in the UK) with up to 2500 per year being imported after 2013. Packages are queenless boxes of bees, containing no frames or brood.

Empty boxes after installing packages of bees

They are usually supplied in a mesh-sided box together with a queen. The bees are placed into a hive with frames of foundation and the queen is added in an introduction cage. They are fed with a gallon to two of syrup to encourage them to draw comb.

Installing a package of bees

It’s a very convenient way to purchase bees and avoids at least some of the risk of importing diseases 2. It’s also less expensive. This presumably reflects both the absence of frame/foundation and the need for a box to contain the frames.

But, post-Brexit, importation of packages or nucs from EU countries is no longer allowed. You are also not allowed to import full colonies (small numbers of these were imported each year, but insufficient to justify adding them to the graph above).

Queen imports are still allowed.

Why are were so many bees imported?

The simple answer is ‘demand’.

Bees can be reared inexpensively in warmer climates, such as southern Italy or Greece. The earlier start to the season in these regions means that queens, nucs or packages can be ready in March to meet the early season demand by UK beekeepers.

If you want a nuc with a laying queen in March or April in the UK you have two choices; a) buy imported bees, or b) prepare or purchase an overwintered nuc.

I don’t have data for the month by month breakdown of queen imports. I suspect many of these are also to meet the early season demand, either by adding them to an imported package (see above) or for adding to workers/brood reared and overwintered in a UK hive that’s split early in the season to create nucleus colonies.

Some importers would sell the latter on as ‘locally reared bees’. They are … sort of. Except for the queen who of course determines the properties of all the bees in the subsequent brood 🙁

An example of being “economical with the truth” perhaps?

Imported queens were also available throughout the season to replace those lost for any number of reasons (swarming, poor mating, failed supersedure, DLQ’s, or – my speciality – ham-fisted beekeeping) or to make increase.

And to put these imports into numerical context … there are about 45,000 ‘hobby’ beekeepers in the UK and perhaps 200+ bee farmers. Of the ~250,000 hives in the UK, about 40,000 are managed by bee farmers.

What are the likely consequences of the import ban?

I think there are likely to be at least four consequences from the ban on the importation of nucs and packages to the UK from the EU:

  1. Early season nucs (whatever the source) will be more expensive than in previous years. At the very least there will be a shortfall of ~2000 nucs or packages. Assuming demand remains the same – and there seems no reason that it won’t, and a realistic chance that it will actually increase – then this will push up the price of overwintered nucs, and the price of nucs assembled from an imported queen and some ‘local’ bees. I’ve seen lots of nucs offered in the £250-300 range already this year.
  2. An increase in imports from New Zealand. KBS (and perhaps others) have imported New Zealand queens for several years. If economically viable this trade could increase 3.
  3. Some importers may try and bypass the ban by importing to Northern Ireland, ‘staging’ the bees there and then importing them onwards to the UK. The legality of this appears dubious, though the fact it was being considered reflects that this part of the ‘oven ready’ Brexit deal was not even table d’hôte and more like good old-fashioned fudge.
  4. Potentially, a post-Covid increase in bee smuggling. This has probably always gone on in a limited way. Presumably, with contacts in France or Italy, it would be easy enough to smuggle across a couple of nucs in the boot of the car. However, with increased border checks and potential delays, I (thankfully) don’t see a way that this could be economically viable on a large scale.

Is that all?

There may be other consequences, but those are the ones that first came to mind.

Of the four, I expect #1 is a nailed-on certainty, #2 is a possibility, #3 is an outside possibility but is already banned under the terms of the Northern Ireland Protocol which specifically prohibits using Northern Ireland as a backdoor from Europe, and #4 happens and will continue, but is small-scale.

Of course, some, all or none of this ban may be revised as the EU and UK continue to wrangle over the details of the post-Withdrawal Agreement. Even as I write this the UK has extended the grace period for Irish sea border checks (or ‘broken international law’ according to the EU). 

This website is supposed to be a politics-free zone 4 … so let’s get back to safer territory.

Why is early season demand so high?

It seems likely that there are three reasons for this early season demand:

  1. Commercial beekeepers needing to increase colony numbers to provide pollination services or for honey production. Despite commercials comprising only ~0.4% of UK beekeepers, they manage ~16% of UK hives. On average a commercial operation runs 200 hives in comparison to less than 5 for hobby beekeepers. For some, their business model may have relied upon the (relatively) inexpensive supply of early-season bees.
  2. Replacing winter losses by either commercial or amateur beekeepers. The three hives you had in the autumn have been slashed to one, through poor Varroa management, lousy queen mating or a flood of biblical proportions. With just one remaining hive you need lots of things to go right to repopulate your apiary. Or you could just buy them in.
  3. New beekeepers, desperate to start beekeeping after attending training courses through the long, dark, cold, wet winter. And who can blame them? 

For the rest of the post I’m going to focus on amateur or hobby beekeeping. I don’t know enough about how commercial operations work. Whilst I have considerable sympathy if this change in the law prevents bee farmers fulfilling pollination or honey production contracts, I also question how sensible it is to depend upon imports as the UK extricates itself from the European Union.

Whatever arrangement we finally reached it was always going to be somewhere in between the Armageddon predicted by ‘Project Fear’ and the ‘Unicorns and sunlit uplands’ promised by the Brexiteers.

Where are those sunlit uplands?

And that had been obvious for years.

I have less sympathy for those who sell on imported bees to meet demand from existing or new beekeepers. This is because I think beekeeping (at least at the hobbyist level) can, and should, be sustainable.

Sustainable beekeeping

I would define sustainable beekeeping as the self-sufficiency that is achieved by:

  • Managing your stocks in a way to minimise winter losses
  • Rearing queens during the season to requeen your own colonies when needed (because colonies with young queens produce brood later into the autumn, so maximising winter bee production) and to …
  • Overwinter nucleus colonies to make up for any winter losses, or for sale in the following spring

All of these things make sound economic sense. 

More importantly, I think achieving this level of self-sufficiency involves learning a few basic skills as a beekeeper that not only improve your beekeeping but are also interesting and enjoyable.

I’ve previously discussed the Goldilocks Principle and beekeeping, the optimum number of colonies to keep considering your interest and enthusiasm for bees and the time you have available for your beekeeping.

It’s somewhere between 2 and a very large number. 

For me, it’s a dozen or so, though for years I’ve run up to double that number for our research, and for spares, and because I’ve reached the point where it’s easy to generate more colonies (and because I’m a lousy judge of the limited time I have available 🙁 ).

Two is better than one, because one colony can dwindle, can misbehave or can go awry, and without a colony to compare it with you might be none the wiser that nothing is wrong. Two colonies also means you can always use larvae from one to rescue the other if it goes queenless.

And with just two colonies you can easily practise sustainable beekeeping. You are no longer dependent on an importer having a £30 mass-produced queen spare.

What’s wrong with imported bees?

The usual reason given by beekeepers opposed to imports is the risk of also importing pathogens.

Varroa is cited as an example of what has happened. 

Tropilaelaps or small hive beetle are given as reasons for what might happen.

And then there are usually some vague statements about ‘viruses’. 

There’s good scientific evidence that the current global distribution of DWV is a result of beekeepers moving colonies about.

More recently, we have collaborated on a study that has demonstrated an association between honey bee queen imports and outbreaks of chronic bee paralysis virus (CBPV). An important point to emphasise here is that the direction of CBPV transmission is not yet clear from our studies. The imported queens might be bringing CBPV in with them. Alternatively, the ‘clean’ imported queens (and their progeny) may be very susceptible to CBPV circulating in ‘dirty’ UK bees. Time will tell.

However, whilst the international trade in plants and animals has regularly, albeit inadvertently, introduced devastating diseases e.g. Hymenoscyphus fraxineus (ash dieback), I think there are two even more compelling reasons why importation of bees is detrimental.

  1. Local bees are better adapted to the environment in which they were reared and consequently have increased overwintering success rates.
  2. I believe that inexpensive imported bees are detrimental to the quality of UK beekeeping.

I’ve discussed both these topics previously. However, I intend to return to them again this year. This is partly because in this brave new post-Brexit world we now inhabit the landscape has changed.

At least some imports are no longer allowed. The price of nucs will increase. Some/many of these available early in the season will be thrown together from overwintered UK colonies and an imported queen.

These are not local bees and they will not provide the benefits that local bees should bring.

Bad beekeeping and bee imports

If imported queens cost £500 each 5 there would be hundreds of reasons to learn how to rear your own queens. 

But most beekeepers don’t …

Although many beekeepers practise ‘passive’ queen rearing e.g. during swarm control, it offers little flexibility or opportunity to rear queens outside the normal swarming season, or to improve your stocks.

In contrast, ‘active’ queen rearing i.e. selection of the best colonies to rear several queens from, is probably practised by less than 20% of beekeepers.

This does not need to involve grafting, instrumental insemination or rows of brightly coloured mini-nucs. It does not need any large financial outlay, or huge numbers of colonies to start with.

But it does need attention to detail, an understanding of – or a willingness to learn – the development cycle of queens, and an ability to judge the qualities of your bees.

Essentially what it involves is slightly better beekeeping.

But, the availability of Italian, Greek or Maltese queens for £20 each acts as a disincentive.

Why learn all that difficult ‘stuff’ if you can simply enter your credit card details and wait for the postie?

Overwintering 5 frame poly nuc

Overwintering 5 frame poly nuc

And similar arguments apply to overwintering nucleus colonies. This requires careful judgement of colony strength through late summer, and the weight of the nuc over the winter.

It’s not rocket science or brain surgery or Fermat’s Last Theorem … but it does require a little application and attention.

But, why bother if you can simply wield your “flexible friend” 6 in March and replace any lost colonies with imported packages for £125 each?

Rant over

Actually, it wasn’t really a rant. 

My own beekeeping has been sustainable for a decade. I’ve bought in queens or nucs of dark native or near-native bees from specialist UK breeders a few times. I have used these to improve my stocks and sold or gifted spare/excess nucs to beginners.

I’ve caught a lot of swarms in bait hives and used the best to improve my bees, and the remainder to strengthen other colonies.

The photographs of packages (above) are of colonies we have used for relatively short-term scientific research. 

I’m going to be doing a lot of queen rearing this season. Assuming that goes well, I then expect to overwinter more nucs than usual next winter. 

I then hope that the bee import ban remains in place for long enough until I can sell all these nucs for an obscene profit which I will use to purchase a queen rearing operation in Malta. 😉

And I’m going to write about it here.


 

A New Year, a new start

The short winter days and long dark nights provide ample opportunity to think about the season just gone, and the season ahead.

You can fret about what went wrong and invent a cunning plan to avoid repetition in the future.

Or, if things went right, you can marvel at your prescience and draft the first couple of chapters of your book “Zen and the Art of Beekeeping”.

But you should also prepare for the normal events you expect in the season ahead.

In many ways this year 1 will be the same as last year. Spring build-up, swarming and the spring honey crop, a dearth of nectar in June, summer honey, miticides and feeding … then winter.

Same as it ever was as David Byrne said.

That, or a pretty close approximation, will be true whether you live in Penzance (50.1°N) or Thurso (58.5°N).

Geographical elasticity

Of course, the timing of these events will differ depending upon the climate and the weather.

For convenience let’s assume the beekeeping season is the period when the average daytime temperature is above 10°C 2. That being the case, the beekeeping season in Penzance is about 6 months long.

In contrast, in Thurso it’s only about 4 months long.

More or less the same things happen except they’re squeezed into one third less time.

Once you have lived in an area for a few years you become attuned to this cycle of the seasons. Sure, the weather in individual years – a cold spring, an Indian summer – creates variation, but you begin to expect when particular things are likely to happen.

There’s an important lesson here. Beekeeping is an overtly local activity. It’s influenced by the climate, by the weather in an individual year, and by the regional environment. You need to appreciate these three things to understand what’s likely to happen when.

OSR ... can you believe it?!

Late April 2016, Fife … OSR and snow

Events are delayed by a cold spring, but if there’s oil seed rape in your locality the bees might be able to exploit the bounteous nectar and pollen in mid-April.

Mid-April 2014, Warwickshire

Foraging might extend into October in an Indian summer and those who live near moorland probably have heather yielding until mid/late September.

Move on

You cannot make decisions based on the calendar.

In this internet-connected age I think this is one of the most difficult things for beginners to appreciate. How many times do you see questions about the timing of key events in the beekeeping season – adding supers, splitting colonies, broodlessness – with no reference to where the person asking, or answering, the question lives?

It often takes a move to appreciate this geographical elasticity of the seasons at different latitudes.

When I moved from the Midlands to Scotland 3 in 2015 I became acutely aware of these differences in the beekeeping season.

When queen rearing in the Midlands my records show that I would sometimes start grafting in the second week in April. In some years I was still queen rearing in late August, with queens being successfully mated in September.

Locally bred queen ...

Locally bred queen …

In the last 5 years in Scotland the earliest I’ve seen a swarm was the 30th of April and the latest I’ve had one arrive in a bait hive was mid-July. Here, queen rearing is largely restricted to mid-May to late-June 4.

All of this is particularly relevant as most of my beekeeping is moving from the east coast to the west coast of Scotland this year.

I’m winding down my beekeeping in Fife and starting afresh on the west coast.

The latitude is broadly the same, but the local environment is very different.

And so are the bees … which means there are some major changes being planned.

What are local bees?

I’m convinced about the benefits of local bees. The science – which I’ve discussed in several previous posts – shows that locally-reared bees are physiologically adapted to their environment and both overwinter and survive better.

But what is local?

Does it mean within a defined geographical area?

If so, what is the limit?

Five miles?

Fifty miles?

What is local? Click to enlarge and read full legend.

I think that’s an overly simplistic approach.

The Angus glens are reasonably ‘local’ to me. Close enough to go for an afternoon walk, or a summer picnic. They’re less than 40 miles north as the bee flies 5.

However, they’re a fundamentally different environment from my Fife apiaries. The latter are in intensively farmed, low lying, arable land. In Fife there’s ample oil seed rape in Spring, field beans in summer and (though not as much as I’d like) lime trees, clover and lots of hedgerows.

The Angus glens

But the Angus glens are open moorland. There’s precious little forage early in the season, but ample heather in August and September. It’s also appreciably colder in the hills due to the altitude 6.

I don’t think you could keep bees on the Angus hills all year round. I’m not suggesting you could. What I’m trying to emphasise is that the environment can be dramatically different only a relatively short distance away.

My bees

I don’t name my queens 7 but I’m still very fond of my bees. I enjoy working with them and try and help them – by managing diseases, by providing space or additional food – when needed.

I’ve also spent at least a decade trying to improve them.

Every year I replace queens heading colonies with undesirable traits like running on the comb or aggression or chalkbrood. I use my best stocks to rear queen from and, over the years, they’ve gradually improved.

They’re not perfect, but they are more than adequate.

When I moved from the Midlands to Scotland I brought my bees with me.

Forgot the scythe

Delivering bees from the Midlands to Fife

I ‘imported’ about a dozen colonies, driving them up overnight in an overloaded Transit van. The van was so full of hive stands, empty (and full) beehives and nucs that I had a full hive strapped down in the passenger seat. Fortunately the trip went without a hitch (or an emergency stop 🙂 ).

Passenger hive

Passenger hive

They certainly were not ‘local’ but I’d invested time in them and didn’t want to have to start again from scratch. In addition, some hives were for work and it was important we could start research with minimum delay.

But I cannot take any of my bees to the west coast 🙁

Treatment Varroa free

Parts of the remote north and west coast of Scotland remain free of Varroa. This includes some of the islands, isolated valleys in mountainous areas and some of the most westerly parts of the mainland.

It also includes the area (Ardnamurchan) where I live.

Just imagine the benefits of not having to struggle with Varroa and viruses every season 🙂

Although I don’t feel as though I struggle with managing Varroa, I am aware that it’s a very significant consideration during the season. I know when and how to treat to maintain very, very low mite levels, but doing so takes time and effort.

It would certainly be preferable to not have to manage Varroa; not by simply ignoring the problem, but by not having any of the little b’stards there in the first place 😉

Which explains why my bees cannot come with me 8. Once Varroa is in an area I do not think it can be eradicated without also eradicating the bees.

A green thought in a green shade … Varroa-free bees on the west coast of Scotland

I’ve already got Varroa-free bees on the west coast, sourced from Colonsay.

Is Colonsay ‘local’?

Probably. I’d certainly argue that it’s more ‘local’ to Ardnamurchan than the Angus glens are to Fife, despite the distance (~40 miles) being almost identical. Both are at sea level, with a similar mild, windy and sometimes wet, climate.

Sometimes, in the case of Ardnamurchan, very wet 🙁

My cunning plans

Although the season ahead might be “same as it ever was”, the beekeeping certainly won’t be.

My priorities are to wind down my Fife beekeeping activities (with the exception of a few research colonies we will need until mid/late 2022) and to expand my beekeeping on the west coast.

Conveniently, because it’s something I enjoy and also because it’s not featured very much on these pages recently, these plans involve lots of queen rearing.

Queen rearing using the Ben Harden system

In Fife I’m intending to split my colonies to produce nucs for sale. I’ll probably do this by sacrificing the summer honey crop. It’s easier to rear queens in late May/June and the nucs that are produced can be sold in 2021, or overwintered for sale the following season.

If I leave the queen rearing until later in the summer I would be risking either poor weather for queen mating, or have insufficient time to ensure the nucs were strong enough to overwinter.

It’s easier (and preferable) to hold a nuc back by removing brood and bees than it is to mollycoddle a weak nuc through the winter.

And on the west coast I’ll also be queen rearing with the intention of expanding my colonies from two to about eight. In this case the goal will be to start as early as possible with the aim of overwintering full colonies, not nucs. However, I’ve no experience of the timing of spring build up or swarming on the west coast, so I’ve got a lot to learn.

Something old, something new

I favour queen rearing in queenright colonies. This isn’t the place to spend ages discussing why. It suits the scale of my beekeeping, the colonies are easy to manage and it is not too resource intensive.

I’ve written quite a bit about the Ben Harden system. I have used this for several years with considerable success and expect to do so again.

I’ve also used a Cloake board very successfully. This differs from the Ben Harden system in temporarily rendering the hive queenless using a bee-proof slide and upper entrance.

Cloake board ...

Cloake board …

Using a Cloake board the queen cells are started under the emergency response, but finished in a queenright hive. It’s a simple and elegant approach. In addition, the queen rearing colony can be split into half a dozen nucs for queen mating, meaning the entire thing can be managed starting with a single double brood colony.

One notable feature of the Cloake board is that the queen cells are raised in a full-sized upper brood box. During the preparation of the hive this upper box becomes packed with bees 9. This means there are lots of bees present for queen rearing.

Concentrating the bees ...

Concentrating the bees …

It’s definitely a case of “the more the merrier” … and, considering the size of my colonies, I’m pretty certain I can achieve even greater concentrations of bees using a Morris board.

A Morris board is very similar to a Cloake board except the upper face has two independent halves. It’s used with a divided brood box (or two 5 frame nucleus boxes) and can generate sequential rounds of queen cells. I understand the principle, but it’ll be a new method I’ve not used before.

Since the bees are concentrated into half the volume it should be possible to get very high densities of bees using a Morris board.

And since I like building things for beekeeping 10, that’s what I’m currently making …

Which explains why I’ve got bits of aluminium arriving in the post, chopped up queen excluders on my workbench and Elastoplast on three fingers of my left hand 🙁

Happy New Year!


Notes

I’m rationalising my beekeeping equipment prior to moving. I have far too much! Items surplus to requirements – currently mainly flat-pack National broods and supers – will be listed on my ‘For Sale‘ page.

 

Winter bee production

There are big changes going on in your colonies at the moment.

The summer foragers that have been working tirelessly over the last few weeks are slowly but surely being replaced. As they die off – whether from old age or by being eaten by the last of the migrating swallows – they are being replaced by the winter bees.

Between August and late November almost the entire population of bees will have changed. The strong colonies you have now (or should have) will contain a totally different workforce by the end of the year.

Forever young

The winter bees are the ones responsible for getting the colony from mid/late autumn through to the following spring. They are sometimes termed diutinus bees from the Latin for “long lived”.

These are the bees that thermoregulate the winter cluster, protecting the queen, and rearing the small amounts of brood during the cold, dark winter to keep the colony ticking over.

Midwinter cluster

A midwinter colony

Physiologically they share some striking similarities with so-called hive or nurse bees 1 early in the summer.

Both hive bees and winter bees have low levels of juvenile hormone (JH) and active hypopharyngeal glands. Both types of bee also have high levels of vitellogenin, high oxidative stress resistance and corpulent little bodies.

But early summer nurse bees mature over a 2-3 week period. Their JH levels increase and vitellogenin levels decrease. This induces additional physiological changes which results in the nurse bee changing into a forager. They sally forth, collecting nectar, pollen and water …

And about three weeks later they’re worn out and die.

Live fast, die young

And this is where winter bees differ. They don’t age.

Or, more accurately, they age   v  e  r  y    s  l  o  w  l  y.

In the hive, winter bees can live for 6 months if needed. Under laboratory conditions they have been recorded as living for up to 9 months.

They effectively stay, as Bob Dylan mumbled, forever young.

Why are winter bees important?

Although not quite eternal youth … staying forever young is useful as their longevity ensures that the colony does not dwindle and perish in the middle of winter.

With little or no nectar or pollen available in the environment the colony reduces brood rearing, and often stops altogether for a period.

But what about the kilograms of stores and cells filled with pollen in the hive? Why can’t they use that?

Whilst both are present, there’s nothing like enough to maintain the usual rate of brood rearing. If they tried the colony would very quickly starve.

Evolution has a very effective way of selecting against such rash behaviour 🙁

If you doubt this, think how quickly hives get dangerously light during the June gap. With no nectar coming in and thousands of hungry (larval) mouths to feed the colony can easily starve to death during a fortnight of poor weather in June.

The winter bees ‘hold the fort’, protecting the queen and rearing small amounts of brood until the days lengthen and the early season pollen and nectars become available again.

And, just as the winter bees look after the viability of the colony, the beekeeper in turn needs to look after the winter bees … we rely on them to get the colony through to spring.

Lots of bees

Can you identify the winter bees?

But before we discuss that, how do you identify and count the winter bees? How can you tell they are present? After all, as the picture above shows 2, all bees look rather similar …

Counting the long lived winter bees

The physiological changes in winter bees, such as the JH and vitellogenin levels, are only identifiable once you’ve done some rather devastating things to the bee. These have the unfortunate side effect of preventing it completing any further bee-type activities 🙁

Even before you subject them to that, their fat little bodies aren’t really sufficiently different to identify them visually.

But what is different is their longevity.

By definition, the diutinus or winter bees are long lived.

Therefore, if you record the date when the bee emerged you can effectively count back and determine how old it is. If it is more than ~6 weeks old then it’s a winter bee.

Or the queen 😉

And, it should be obvious, if you extrapolate back to the time the first long lived bees appear in the hive you will have determined when the colony starts rearing winter bees.

The obvious way to determine the age of a bee is to mark it upon emergence and keep a record of which marks were used when. Some scientists use numbered dots stuck to the thorax, some use combinations of Humbrol-type paint colours.

I’m not aware that anyone has yet used the barcoding system I discussed recently, though it could be used. The winter bee studies I’m aware of pre-date this type of technology.

Actually, some of these studies date back almost 50 years, though the resulting papers were published much more recently.

This is painstaking and mind-numbingly repetitive work and science owes a debt of gratitude to Floyd Harris who conducted many of the studies.

Colony age structure – autumn to winter

Here is some data showing the age structure of a colony transitioning from late summer into autumn and winter. There’s a lot in this graph so bear with me …

Colony age structure from August to December - see text for details

Colony age structure from August to December – see text for details

The graph shows the numbers and ages of bees in the colony.

The ages of the bees is indicated on the vertical axis – with eggs and brood (the youngest) at the bottom, coloured black and brown respectively. The adult bees can be aged between 1 and ~100 days old 3. The number of bees is indicated by the width of the coloured bar at each of the nine 12 day intervals shown.

All of the adult bees present in the hive at the end of August are coloured blue, irrespective of their age. There are a lot of these bees at the end of August and almost all of them have disappeared (died) by mid-November 4.

The remaining colours indicate all the bee that emerge within a particular 12 day interval. For example, all the bees that emerge between the 31st of August and the 12th of September are coloured yellow.  Going by the width (i.e. the numbers of bees of that age) of the yellow bars it’s clear that half to two-thirds of these bees die by mid October, with the rest just getting older gracefully.

But look at the cohort that emerge between the end of September and early October, coloured like this 5. The number of these bees barely changes between emergence and early December. By this time they are 72 days old i.e. an age that most summer bees never achieve.

Brood breaks and climate

In the colony shown above the queen continued laying reduced numbers of eggs – the black bars – until mid-October and then didn’t start again until the end of November. During this period the average age of the bees in the colony increased from ~36 days to ~72 days and the strength of the colony barely changed.

The figure above comes from a BeeCulture article by Floyd Harris. The original data isn’t directly referenced, but I suspect it comes from studies Harris conducted in the late 70’s in Manitoba, some of which was subsequently published in the Journal of Apicultural Research. In addition, Harris co-authored a paper presenting similar data in a different format in Insectes Sociaux which describes the Manitoban climate as having moderate/hot summers and long, cold winters.

My hives in Scotland, or your hives in Devon, or Denmark or wherever, will experience a different climate 6.

However, if you live in a temperate region the overall pattern will be similar. The summer bees will be replaced during the early autumn by a completely new population of winter bees. These maintain the colony through to the following spring.

The dates will be different and the speed of the transition from one population to the other may differ. The timing of the onset of a brood break is likely to also differ.

However, the population changes will be broadly similar.

And, it should be noted, the dates may differ slightly in Manitoba (and everywhere else) from year to year, depending upon temperature and forage availability.

Colony size and overwintering survival

Regular readers might be thinking back to a couple of posts on colony size and overwintering survival from last year.

One measured colony weight, showing that heavier colonies overwintered better 7. A second discussed the better performance of local bees in a Europe-wide study of overwintering survival. In this, I quoted a key sentence from the discussion:

“colonies of local origin had significantly higher numbers of bees than colonies placed outside their area of origin”

I can’t remember when during the season those studies recorded colony size, but I’m well aware that large colonies in the winter survive better.

The colonies that perish first in the winter are the pathetic grapefruit-sized 8 colonies with ageing queens or high pathogen loads.

In contrast, the medicine ball-sized ‘boomers’ go on and on, emerging from the winter strongly and building up rapidly to exploit the early season nectar.

But what the graph above shows is that the bees in a strong colony in late summer are a completely different population from the bees in the colony in midwinter.

The strength of the midwinter colony is determined entirely by when winter bee rearing starts and the laying rate of the queen, although of course both may be indirectly influenced by summer colony strength.

The influence of the queen

Other than this potential indirect influence, it’s possibly irrelevant how large the summer colony is in terms of winter colony size (and hence survival).

After all, even if the summer bees were three times as numerous, their fate is sealed. They are all going to perish six weeks or so after emergence.

Are there ways that beekeepers can influence the size of the overwinter colony to increase its chances of survival?

I wouldn’t pose the question if the answer wasn’t a resounding yes.

It has been known for a long time 9 that older queens stop laying earlier in the autumn than younger queens. As explained above, the longer the queen lays into the autumn the more winter bees are going to be produced.

Mattila et al., 10 looked at the consequences of late season (post summer honey harvest) requeening of colonies. In these they removed the old queen and replaced her with either a new mated or virgin queen, or allowed the colony to requeen naturally.

Using the ’12 day cohort’ populations explained above, the authors looked at when the majority of the winter bees were produced in the colony, and estimated the overall size of the winter colony.

The influence of new queens on winter bee production.

The influence of new queens on winter bee production. Note shift to the right in B, C and D, with new queens.

With the original old queen, 53% of winter bees were produced in the first two cohorts of winter bees. With the requeened colonies 54-64% of the winter bees were produced on average 36 days later, in the third and fourth cohorts of winter bees.

This indicates that young queens produce winter bees later into the autumn.

This is a good thing™.

In addition, though the results were not statistically significant, there was a trend for colonies headed by new queens to have a larger population of bees overwinter.

Perhaps one reason the requeened colonies weren’t significantly larger was that the new queens delay the onset of winter bee rearing. I’ll return to this at the end.

The influence of deformed wing virus (DWV)

Regular readers will know that this topic has been covered extensively, and possibly exhaustively, elsewhere on this site … so I’ll cut to the chase.

DWV is the most important virus of honey bees. When transmitted by Varroa destructor there is unequivocal evidence that it is associated with overwintering colony losses. The reason DWV causes overwintering losses is that it reduces the longevity of the winter bees.

The virus might also reduce the longevity of summer bees but,

  1. there’s so many of them to start with
  2. there’s loads more emerging every day, and
  3. they only survive a few weeks anyway,

that this is probably irrelevant in terms of colony survival.

Dainat et al., (2012) produced compelling evidence showing that DWV reduces the longevity of winter bees 11. The lifespan was reduced by ~20%.

A consequence of this is that the winter bees die off a little faster and the colony shrinks a little more. At some point it crosses a threshold below which it cannot thermoregulate the cluster properly, further limiting the ability of the colony to rear replacement bees (assuming the queen is able to lay at a low rate).

This colony is doomed.

Even if they stagger through to the longer days of spring they contain too few bees to build up fast. They’re not dead … but they’re hardly flourishing.

Winter bees and practical beekeeping

I think there are three ways in which our understanding of the timing of winter bee production should influence practical beekeeping:

Firstly … The obvious take-home message is that winter bees must be protected from the ravages of DWV. The only way to do this is to minimise the mite population in the colony before the winter bee rearing starts.

The logical way to do this is to treat using an approved miticide as soon as practical after the summer honey is removed 12.

I discuss the importance of the timing of this treatment in When to treat?, which remains one of the most-read posts on this site.

Secondly … Avoid use of miticides (or other colony manipulations) that reduce the laying rate of the queen in early autumn.

When I used to live at lower latitudes I would sometimes use Apiguard. This thymol-containing miticide is very effective if used when the temperature is high enough. However, in my experience a significant proportion of queens stop laying when it is being used. Not all, but certainly more than 50%.

I don’t know why some stop and others don’t. Is it genetic? Temperature-dependent?

Whatever the reason, they stop at exactly the time of the season you want them to be laying strongly.

Thirdly … consider requeening colonies with young queens after the summer honey is removed. This delays the onset of winter bee production and results in the new queen laying later into the year. The later start to winter bee production gives more time for miticides to work.

A win-win situation.


 

Bigging up nucs

The phrase bigging up [somebody or something] means saying they are very good, usually in public 1. It is slang and used informally and usage has increased significantly in the last couple of decades.

The term bigging in bigging up meaning promotion, is relatively new. However, the same word can be traced back to Middle English and (a bit more more recently) obsolete Scottish, when it meant build.

Two days work bigging a brick wall in the Braidfoots house 2.

Anyone who has used nucs, for queen mating or swarm control for example, is likely to big them up … as in sing their praises. Small enough to need only limited resources to start them, large enough to function as a self-contained and resilient colony etc.

However, in this post I’m going to discuss bigging up nucs in the older meaning of the phrase … building them up from a nuc to a full colony.

Which could also, of course, be considered as promoting them 😉

Problems with history and latitude

One of the perils of writing about beekeeping in the UK is the variation in the season between the south and the north of the country.

Just as you can’t be prescriptive in any one location about when certain events in a particular beekeeping year occur – e.g. swarming, winter bee production, broodlessness – it’s also pretty obvious that the season is longer 3 at lower latitudes.

It’s therefore not possible to say ‘in late May’ or ‘by mid-June’ nucs will start to be overcrowded 4. Not only does this depend upon the local climate, but it is also significantly influenced by how the nucs were prepared.

If the nuc was established for swarm control, started with the old queen and 1-2 frames of brood, it is likely to have built up rapidly and will quickly overrun the box if not dealt with promptly.

Alternatively, if the nuc was used for queen mating, started with a sealed cell (or virgin queen) and a frame of emerging brood, it will build up less fast as the queen has to get out and mate and then start laying.

Overcrowding

Whatever the history (or the latitude), at some point the colony will grow to be too large for the box. Then, but ideally earlier (so you are prepared), you need to decide what you are going to do with them.

With experience you can judge overcrowding by gently popping the lid up and peering through the thin plastic or polycarbonate crownboard. 

I use Thorne’s Everynucs which have an integral feeder at one end of the box. When they start building brace comb in the feeder they need to be given more space.

Here's one I prepared earlier

Here’s one I prepared earlier

The colony above is overwintered and very clearly overcrowded. The photo was taken in the third week of April (in Scotland). By mid-season, a colony that crowded would have probably swarmed.

Comb in feeder

The photo immediately above was taken in late June this year. The nuc was set up in mid-May for swarm control with the queen and just one frame of emerging brood.

However, in the intervening six weeks I had already removed two or three frames of sealed brood (but not adhering bees) to boost other colonies, replacing the frames with a mix of drawn comb and foundation, all of which had been drawn and filled again.

Nucs can build up very fast … be warned.

Decision time

Nucs are really versatile. Your choice includes (but isn’t restricted to):

  1. Overwintering the nuc
  2. Expanding the nuc into a full hive
  3. Uniting the nuc with a queenless colony
  4. Removing the queen and uniting the nuc with a queenright colony
  5. Leaving it too late and letting them swarm 🙁

I’m not going to discuss the last option, but it is an inevitability if the colony is healthy and there’s a reasonable amount of forage in the area. 

One more week’ for a nuc is usually not worth risking.

Overwintering nucs deserves a post of its own (and has been covered some time ago 5). It’s worth noting that nucs started in May for swarm control or for queen mating require a lot of maintenance if they are not to outgrow their accommodation by the end of the season. You need to regularly remove bees and brood or the colony will swarm.

It is much better to start nucs later in the season for overwintering.

Before doing anything with the nuc it is worth confirming that the queen appears well mated and is laying well 6. She should be producing frame after frame packed with brood. In new(ish) comb you can easily tell her quality based upon the presence of even sheets of brood, with relatively few missed cells.

Good laying pattern from queen in 5 frame nucleus

The frame above is from a nuc this spring. The majority of the missed cells, at least at the top of the frame, are due to the wires in the foundation.

Returning a marked and clipped queen to a nuc

And, while you’re at it, use this opportunity of the last inspection of the nuc to mark and clip the queen (if she isn’t already). It’s always easier to find a queen in a nuc – fewer bees, less frames to hide on the other side of etc.

From nuc to a full brood box

This is about as easy as it gets and should take no more than 5 minutes if you have everything to hand.

  1. Move the nuc a metre or so away from its original location.
  2. Place a new floor and a brood box on the original site.
  3. The brood box should contain a couple of frames of drawn comb if you have them, or frames with fresh foundation. Place one next to each side wall (see note below for comment on warm and cold way).
  4. If the floor has open mesh I slide in the Varroa tray. I do not want the bees to be distracted by smells from other ‘potential’ routes into the hive.
  5. Open the nuc using a very small amount of smoke 7.
  6. Remove the dummy board from the nuc and gently separate the frames if they’re propolised together.
  7. Transfer each frame to the new brood box maintaining their position and orientation relative to the neighbouring frames. Arrange the frames from the nuc close to the new hive entrance (see below).
  8. Ideally , make sure the queen is seen … just to give you confidence 🙂
  9. Move the second new frame of drawn comb or foundation to ‘sandwich’ the frames from the nuc.
  10. Fill the rest of the box with frames containing drawn comb or new foundation.
  11. Replace the dummy board removed in #6 above.
  12. Add syrup if needed – see below.
  13. Replace the crownboard and roof.
  14. Reduce the entrance to help the colony defend their new, much larger, residence.

Feeding

If there is a good nectar flow you may not need to feed the colony. If you’ve used new foundation rather than drawn comb then they probably will need feeding. It’s important they draw new comb so the queen can continue laying uninterrupted. This ensures they build up rapidly.

Use thin syrup (1:1 by weight of sugar and water) in a contact feeder. 

I usually give nucs a gallon or so of syrup to help them draw comb. They use this surprisingly fast. Check them every 48 hours. 

Welcome to your new home … nuc ‘promoted’ to hive with contact feeder in place

My crownboards lack holes, so I place the contact feeder directly above the top bars, separated by a couple of spare frame bottom bars. I add a super to ‘house’ the contact feed and then close the hive up.

Defending the hive

All of my full-sized hives are arranged warm way. This means the frames are parallel with the entrance of the hive. The alternative, cold way, has the frames perpendicular to the entrance.

To help the small colony defend the new large box they are in, the nucleus frames should be located close to the hive entrance.

The hive entrance is on the left with the frames arranged ‘warm way’.

Initially, these are the frames that are covered in bees, so providing a deterrent to any potential robbers.

It may also help to reduce the size of the hive entrance so the bees only need to defend an inch wide hole, rather than the full width of the box.

If your hives are organised cold way’ the same requirements apply – arrange the bees near to the entrance and reduce the entrance width. For example, place the frames in the centre of the hive, flanked on each side by three new frames, and leave a narrow central entrance open.

Finally, do not slop syrup around all over the place when feeding them. It’s a near-certain way to encourage robbing (particularly if there’s a shortage of nectar).

Uniting the nuc with a queenright or queenless colony

I can deal these two together because the only difference is where the queen is in the stacked boxes at the end of the procedure.

Collect together the things you will need:

  • A new brood box
  • Two sheets of newspaper
  • Six frames of drawn comb or foundation

Queens

If the hive and the nuc are both queenright you must remove the unwanted queen 8.

Typically this is when you have used the nucleus method of swarm control. The colony has reared a good new queen and the old queen in the nuc is now surplus to requirements.

Alternatively, the colony might have generated a sub-standard or poorly mated queen and you want a single united colony headed again by the original queen.

If the old(er), unwanted queen is still laying OK consider offering her to someone else in your association. Remove the queen, does not necessarily mean sacrifice her. 

Caged queen with attendants

Place the queen in a introduction cage with some attendant workers and some candy. Put her somewhere safe (the breast pocket works for me) and give her to someone who needs her more than you do … perhaps in exchange for a nice bottle of merlot 9 😉

Don’t risk leaving two queens in the same box and hoping the ‘better’ one (i.e. the one you want) will survive the ruckus that will happen. 

Sod’s Law dictates that the queen you want will not make it … particularly if it’s late in the season, she’s particularly good or she’s otherwise precious.

Uniting

I generally move the nuc to the hive it is being united with. Waft some smoke at the hive entrance, remove the roof and gently lift the corner of the crownboard. Add a second gentle puff of smoke into the gap and let the bees move down.

Remove the crownboard and gently lay two intact sheets of newspaper flat over the tops of the frames. It helps to remove brace comb from the top bars as it can puncture the newspaper and lead to premature mixing and a bit of a melee.

In the good old days a single page from a broadsheet 10 newspaper was sufficient. These days I think you have to read the Financial Times to achieve this

Assuming you’re not Gordon Gekko, a hedge fund manager or derivatives trader you will probably need two slightly overlapping sheets. Don’t bother about moving all the bees off the top bars – they’ll move down soon enough once you put the newspaper on.

If it’s windy use your initiative, recruit a helper or evolve at least one additional limb to hold the newspaper in place.

Add a second empty brood box on top.

Make a small hole (about the size of the o in hole) in the sheet using your hive tool, somewhere near the middle, above a gap between two frames. You can just see the hole above the curve of the hive tool here …

Newspaper, second brood box and a very small hole

Add two or three frames of drawn comb or foundation. Transfer all the frames from the nuc to the new brood box, as before, maintaining their order and orientation. Fill the rest of the box with frames, shake in the last bees from the nuc box and close the hive up.

Just checking!

As before, if you are uniting a queenright nuc with a queenless hive, it’s always good to be certain the queen was on one of the frames transferred to the new box.

Have patience

Hives usually have sufficient stores at this time of the season. If both boxes are light you might have to feed them syrup (to help them draw comb) or fondant (just to tide them over until the nectar flow starts).

Leave them to it. There’s nothing to be gained by ‘having a peek’. The bees will chew their way through the newspaper in 24-48 hours.

Successful uniting ...

Successful uniting …

Look out for a pile of shredded newspaper falling through the open mesh floor and, after a week, continue inspections as normal.

Miscellaneous final thoughts

If the recipient hive is broodless it will end up with lots of space and empty frames. Under those circumstances I usually unite them down to a single box. Rather than adding additional frames to the top box I use a fat dummy to fill the space.

Uniting a nuc with a full colony

Uniting a nuc with a full colony …

A block of polystyrene tightly wrapped in a bin bag works just as well 11.

A week after uniting them rearrange the brood-containing frames with pollen and stores into a single box and remove the empty frames and unwanted second brood box.

Lost bees

How will the bees reorientate to the new location?

Don’t worry. The bees from the nuc will discover that everything is changed when they have to muscle their way through the lower brood box to reach the hive entrance. They will quickly reorientate to the new hive.

Some of the workers from the nuc will have been out foraging when you rudely removed their home. They will, in time, move to a nearby hive and blag their way in 12

Where has the house gone?

You can speed this process up by removing the hive stand the nuc was on. With nowhere to land they quickly find an adjacent hive. If I unite colonies in poor weather (or just before rain starts 13) I’ll try and minimise the number of stranded bees by doing this.

For the same reason I prefer not to unite late in the afternoon to give the bees time to relocate. 

Supers

When I was younger and much better organised I’d clear the supers in advance on the recipient hive. I’d visit the apiary 24 hours before I intended to unite them and add a clearer board. When preparing the recipient colony I’d put the (now emptied) supers aside, unite the colonies and then add the supers back on top (all on the same visit). 

These days I’m definitely older and usually less well organised 🙁

Newspaper and queen excluder

If I’ve forgotten to clear the supers I’ll also unite the bees in the supers over the nuc. I separate them with newspaper as before and add a queen excluder to stop the queen moving up into the supers.

All that then remains to do is tidy up the apiary and go home for a cup of tea.

Time to tidy up and go home


 

If it quacks like a duck …

Quack

… it might be a trapped virgin queen.

I discussed the audio monitoring of colonies and swarm prediction last week. Whilst interesting, I remain unconvinced that it is going to be a useful way to predict swarming. 

And, more importantly, that replacing the manual aspects of hive inspections is desirable. I’m sure it will appeal to hands off beekeepers, though I’m not sure that’s what beekeeping is about.

However there was a second component to what was a long and convoluted publication 1 which I found much more interesting.

Listening in

If you remember, the researchers fitted hives with sensitive accelerometers and recorded the sounds within the hive for two years. Of about 25 colonies monitored, half swarmed during this period, generating 11 prime swarms and 19 casts.

In addition to the background sounds of the hive, with changes in frequency and volume depending upon activity, some colonies produced a series of very un-bee-like toots and quacks.

Have a listen …

The audio starts with tooting, the quacking starts around 8-9s, and there’s overlapping tooting and quacking from near the 21s mark.

Queen communication

I’ve previously introduced the concept of pheromone-based communication within the hive. For example, the mated queen produces the queen mandibular and queen footprint pheromones, the concentrations of which influence the preparation and development of new queen cells.

Tooting and quacking is another form of queen communication, this time by virgin queens in the colony.

It’s not unusual to hear some of these sounds during normal hive inspections, but only during the swarming season and only when the colony is in the process of requeening.

If you rear queens, and in my experience particularly if you use mini-mating nucs, you will regularly hear “queen piping” – another term for the tooting sound – a day or so after placing a mature charged queen cell into the small colony.

But we’re getting ahead of ourselves. 

How does the queen make these sounds?

Queen piping or tooting

Queen tooting has been observed. The queen presses her thorax tight down against the comb and vibrates her strong thoracic wing muscles. Her wings remain closed. The comb acts as a sounding board, amplifying the sound in the hive (and presumably transmitting the vibrations through the comb as well).

This doesn’t happen just anywhere … the virgin queen is usually near the cell she has recently emerged from. 

And this swarm cell is usually on the periphery of a frame.

This is because the laying queen only rarely ventures to the edges of frames, so the concentration of her footprint pheromone is lower in this area, eventually resulting in queen cells being produced there

In their study, accelerometers embedded in the periphery of comb were able to detect much stronger tooting and quacking signals, supporting the conclusions of Grooters (1987) 2 who had first published studies on the location of piping queens.

Queen tooting and quacking

Queen piping is usually recorded at around 400 Hz and consists of one or more 1 second long pulses, followed by a number of much shorter pulses. In previous studies the frequency of tooting had been shown to be age-related. It starts at ~350 Hz and rises in frequency to around 500 Hz as the virgin queen matures over several days.

Compare the image above with the audio file linked further up the post. The tooting is followed by an extended period of quacking, and then both sounds occur at the same time.

Going quackers

The duck-like quacking is presumably also made by queens vibrating their flight muscles while pressed up against the comb.

I say ‘presumably’ as I don’t think it has been observed, as opposed to heard.

The reason for this is straightforward, the queens that are quacking are still within the closed queen cell.

Quacking is a lower frequency sound (is this because of the confines of the queen cell, the way the sound is produced, or the ‘maturity’ of the queen’s musculature?) but has also been shown to increase in frequency – from ~200 Hz to ~350 Hz – the longer the queen remains within the cell.

Afterswarms = casts

Before discussing the timing of tooting and quacking we need to quickly revisit the process of swarming. I’ve covered some of this before when discussing the practicalities of swarm control, so will be brief.

  1. Having “decided” to swarm the colony produces swarm cells. Usually several.
  2. Weather permitting, the prime swarm headed by the original laying queen leaves the hive, on or around the day that the first of the maturing queen cells is capped.
  3. Seven days after the cell was capped the first of the newly developed virgin queens emerges. 
  4. If the colony is strong, this virgin also swarms (a cast swarm). Some texts, including the publication being discussed, call these afterswarms.
  5. Over the following hours or days, successively smaller cast swarms may leave the hive, each headed by a newly emerged virgin queen.

Not all colonies produce multiple cast swarms, but initially strong colonies often do.

From a beekeeping point of view this is bad news™. It can leave the remnants of the original colony too weak to survive and potentially litters the neighbourhood with grapefruit, orange and satsuma-sized cast swarms. 

Irritating 🙁

Whether it’s good for the bees depends upon the likelihood of casts surviving. The very fact that evolution has generated this behaviour suggests it can be beneficial. I might return to this point at the end of the post.

Tooting timing

The Grooters paper referred to earlier is probably the definitive study of queen tooting or piping. The recent Ramsey publication appears to largely confirm the earlier results 3, but has some additional insights on colony disturbance during inspections 4.

Here is the acoustic trace of an undisturbed colony producing a prime swarm and two casts.

Timing of tooting and quaking in a swarming colony

I’ve added some visible labels to the image above indicating the occurrence of tooting and quacking in an undisturbed naturally swarming colony.

  • The prime swarm exited the hive on the afternoon of the 13th. No tooting had been recorded before that date.
  • On the 17th tooting starts and increases in frequency over the next two days.
  • Quacking starts 6 hours after the tooting starts.
  • The first cast swarm (afterswarm) exits the hive on the 19th and is followed by a three hour break in tooting.
  • Tooting and quacking then continue until the second cast swarm on the afternoon of the 21st.

So, in summary, tooting starts after the prime swarm leaves and stops temporarily when the first cast leaves the hive. Quacking starts after the tooting starts and then continues until the last swarm leaves the hive.

Why all the tooting and quacking?

The timing of queen tooting is consistent with it being made by a virgin queen that has emerged from the cell. The cessation of tooting upon swarming (the first afterswarm) suggests that the virgin left with the swarm. The restarting of tooting a few hours later suggests a new virgin queen has been released from another cell and is announcing her presence to the colony.

In previous studies, Grooters had shown that replaying the tooting sound to mature virgin queens actively chewing their way out of a queen cell delayed their emergence by several hours. This delay allowed the attendant workers to reseal the cell and obstruct her emergence for several days.

These timings and the behaviour(s) they are associated with suggest they are a colony-level communication strategy to reduce competition between queens. 

The newly emerged virgin queen toots (pipes) to inform the workers that there is ‘free’ queen in the colony. The workers respond by holding back emergence of other mature queens. 

If all (or several) of the virgin queens emerged and ran around the hive simultaneously they would effectively be ‘competing’ for the hive resources needed for successful swarming i.e. the workers. 

By controlling and coordinating a succession of queen emergence, a strong colony has the opportunity to generate one, two or more cast swarms whilst sufficient workers remain in the hive. It presumably helps ensure the casts are of a sufficient size to give them the best chance of survival.

At what point does this succession stop or break down? One possibility is that this happens when there are insufficient workers to prevent additional virgin queens from emerging.

Unanswered questions

Why do mature virgin queens within the cell quack? It is clearly a response to tooting, rather than being standard behaviour of a soon-to-emerge queen. 

Hear! Hear the pipes are calling, Loudly and proudly calling (from Scotland the Brave)

Is the quacking to attract workers to help reseal the cell?

I suspect not. At least, I suspect there is a more pressing need to attract the workers. After all, wouldn’t it be easier for the queen to simply stop chewing her way out for a few hours? 

Isn’t there a risk that a quacking cell-bound queen might attract the virgin queen running around ‘up top’ who might attempt to slaughter her captive half-sister? 

Possibly, so perhaps the workers that are attracted to the quacking cell also protect the cell, preventing the loose virgin queen from damaging the yet-to-emerge queens.

This would make sense … if the virgin leaves with a cast, the workers that will remain must be sure that there will be a queen available to head the colony

And finally, back to the tooting. I also wonder if this has additional roles in colony communication. For example, what other responses does it induce in the workers? 

Does the increasing frequency of tooting inform the workers that the virgin is maturing and that they should ready themselves for swarming? Perhaps tooting above a certain frequency induces workers to gorge themselves with honey to ensure the swarm has sufficient stores?

In support of this last suggestion, studies conducted almost half a century ago by Simpson and Greenwood 5 concluded that a 650 Hz artificial piping sound induced swarming in colonies containing a single mobile (i.e. free) virgin queen.

Casts

The apparently self-destructive swarming where a colony generates a series of smaller and smaller casts seems to be a daft choice from an evolutionary point of view.

Several studies, in particular from Thomas Seeley, have shown that swarming is a risky business for a colony … and that the majority of the risk is borne by the swarm, not the parental colony. 

87% of swarmed colonies will rear a new queen and successfully overwinter, but only 25% of swarms survive. And the latter figure must only get smaller as the size of the swarms decrease. 

One possibility is that under entirely natural conditions a colony will not undergo this type of self-destructive swarming. Perhaps it is a consequence of the strength of colonies beekeepers favour for good nectar collection or pollination?

Alternatively, perhaps it reflects the way we manage our colonies. Ramsey and colleagues also record tooting and quacking from colonies disturbed during hive inspections. In at least one of these their interpretation was that there were multiple queens ‘free’ in the hive simultaneously, presumably because workers had failed to restrict the emergence of at least one virgin queen.

So, perhaps hive inspections that (inadvertently) result in the release of multiple virgin queens are the colonies that subsequently slice’n’dice themselves to oblivion by producing lots of casts.

I can only remember one colony of mine doing this … and it started days after the previous inspection, but that doesn’t mean the disturbance I created during the inspection wasn’t the cause.

I’d be interested to know of your experience or thoughts.


Colophon

The title of this post is derived from the Duck Test:

If it looks like a duck, swims like a duck, and quacks like a duck, then it probably is a duck.

This probably dates back to the end of the 19th Century. It’s a form of abductive 6 reasoning or logical inference. It starts with an observation or set of observations and then seeks to find the simplest and most likely conclusion from those observations. In comparison to deductive reasoning, logical inference does not lead to a logically certain conclusion. 

Inevitably, Monty Python stretched the logical inference a little too far in the Witch Logic scene from Monty Python and the Holy Grail:

What do you do with witches? Burn them! And what do you burn apart from witches? Wood! So, why do witches burn? ‘cos they’re made of wood? So; how do we tell if she is made of wood? Build a bridge out of ‘er! Ah, but can you not also make bridges out of stone? Oh yeah. Does wood sink in water? No, it floats! It floats! Throw her into the pond! What also floats in water? Bread! Apples! Very small rocks? Cider! Gra-Gravy! Cherries! Mud! Churches? Churches! Lead, Lead. A Duck! Exactly. So, logically… If she weighs the same as a duck, she’s made of wood… and therefore… a witch!

The million drones fiasco

Accidents happen.

Sometimes they are due to stupidity, sometimes to forgetfulness, or sometimes they are just the result of plain dumb luck.

They’re also often caused or at least exacerbated by ‘local’ factors – like a rainstorm or a cancelled train preventing timely inspections. 

Or a countrywide lockdown necessitated by a global viral pandemic.

With the exception of the cancelled train my excuse for what follows is “all of the above” 😉

Social distancing

Beekeeping, like other activities involving livestock management, has been a permitted activity during lockdown. Beekeepers have been allowed to travel to their apiaries and to move bees for pollination etc

I was away when lockdown was imposed and opted 1 to stay where I was. For the first half of the season I’ve had to forego weekly colony inspections. I’ve not had the pleasure of watching the colonies build up, of queen rearing or of sweating profusely when shifting nectar-filled supers 🙁

Instead all my beekeeping – the first inspection of the season, the swarm prevention and the swarm control – have been squeezed into two visits, each of a few frantically busy days, in late April and mid-May.

And, inevitably, mistakes have been made.

Well, one mistake … that I’m currently aware of.

First inspections and swarm prevention

We’re late starters in Fife.

It’s not unusual to delay the full first inspection until the very end of April in this part of Scotland. A couple of years ago we had knee-high oil seed rape (OSR) ankle deep in snow at the end of April.

There seems little point in disturbing the colony if it’s too cold to have a leisurely look through the brood box. The bees get tetchy, the brood gets chilled and you don’t have time to look for the important things – like disease, or that elusive queen you failed to mark last autumn.

However, this season started well and I should have started colony inspections in the second week of April.

But by that time the world had changed dramatically …

I finally snatched a couple of days around the 25th of April to do the first inspections and swarm prevention all rolled into one … and coupled this with reducing my colony numbers by 50% to make management over the coming months easier 2.

I’ll discuss how I did all this in a couple of full-on days some other time. The end result was about a dozen united colonies, each topped with three supers, containing a good marked laying queen. Many of the colonies were very strong, with up to 15 frames of brood after uniting 3.

The colonies were strong and healthy. All were headed by a laying queen. I saw all but a couple of the queens 4 and clipped and marked all those I found that weren’t already 5.

Safely back in the hive

Three supers were overkill for the usual spring nectar flow. However, there was already a reasonable flow on and I wanted to give the colonies a good amount of space in the hope of delaying swarm preparations. 

Swarm control

Colonies usually start making clear their intent to swarm in the second half of May here. It varies a bit depending upon how advanced or otherwise the season is – one of those unknown knowns.

I kept in email contact with beekeeping friends about their own colony build up. By the time I received the first email saying charged queen cells were present (~16th of May) I was travelling back to do my own swarm control.

I decided to use the nucleus method whether queen cells were present on not.

Effectively I was going to implement preemptive swarm control on some colonies. By taking the queen out into a nuc the colonies would be forced to requeen, I’d then leave a single charged/capped queen cell and let them get on with it.

All looking good …

And for eleven of the colonies that’s precisely what happened. 

I removed the queen on a frame of emerging brood and shook some of the bees from a second frame into the nuc box. These were to be relatively small nucs but made sure each had a full frame of capped stores (saved from colonies at the first inspection). I also added a frame of drawn comb and two foundationless frames.

I sealed the nucs and moved them to another apiary.

Three of many … and hive number 29

Most of the brood boxes had play cups with eggs and about 50% had charged queen cells. There were no capped cells. I marked frames containing promising looking charged cells and closed the boxes up.

… and still looking good six days later

Six 6 days later I went carefully through every frame in the de-queened colonies.

One good queen cell, an old play cup and some rather old comb

All the boxes had good looking queen cells and I made sure I left just one in each colony. 

The nucs also all looked great when I checked them on the same day. 

New comb with queen already laying it up

The queens were laying well and the bees were drawing new comb. They would be fine for another few weeks. 

Come in Number 29, your time is up

One of the colonies proved more problematic.

Hive #29 … this had been left as a strong single brood colony on the 25th of April.

Three weeks later it was – unsurprisingly – still a strong single brood colony. The bees were busy and the supers were already filling nicely 7.

What was missing from the brood box in mid-May were eggs, larvae or capped brood 🙁

Had I inadvertently killed the queen 8 at the last inspection? The 21-22 day interval would have meant that all worker brood would have matured and subsequently emerged 9.

However, the temperament of the colony suggested it wasn’t queenless. The bees were calm, they were foraging well and bringing in good amounts of OSR pollen.

With a sense of dread I had a look in the supers …

Let there be drones

About 75% of my many super frames are drawn on drone cell foundation. For the same amount of wax – by weight – you store more honey. I also think there may be advantages when spinning it out in terms of honey recovery 10.

In addition, if you use drone cell comb immediately over the brood box, you dissuade a strong colony from storing an arch of pollen over the brood nest in the super … 

Drone comb in super

… though they do often leave cells empty, ready for the queen to lay.

But she can’t do that because she’s trapped under the queen excluder. 

Right?

Wrong 🙁

The middle few frames of the lower couple of supers were wall to wall capped drone brood and drone larvae. The queen was busy laying up some of the remaining space that wasn’t already filled with nectar.

I found the marked and clipped queen on the very first super frame I removed.

Sod it.

Snatching victory from the jaws of defeat

Perhaps.

Here was the dilemma. Hive #29 was strong and healthy but effectively queenless. Time was against me. I didn’t have the luxury of simply plonking her beneath the QE and checking the colony didn’t make swarm preparations in another three or four weeks 11

I’d already united all my other colonies and made up the nucs. I didn’t want to disassemble any of these to accommodate this colony.

With bad weather approaching in a few days I decided to make up a nuc with the queen and, in due course, donate a queen cell from another colony.

Which is what I did. 

An adjacent colony helpfully raised several very good looking cells which I knew were charged. One of these, on a frame holding a sideplate-sized patch of brood, was added to the colony just before the rain arrived.

Open the box, open the box

But on the same day I added the queen cell I also checked the supers thoroughly.

I wanted to make sure that every frame was drone foundation and that I’d not missed a queen cell drawn from any worker comb in the supers. That might have resulted in a virgin queen running about in my supers and, knowing my luck, squeezing through the QE and slaughtering the queen from the cell I’d just introduced. 

There were lots of “queen cells” in the supers. However all were little more than play cups drawn along the top edge of the drone comb, against the top bar. 

Lots of drone brood … but no real queen cells

None contained eggs. It was as though the bees, sensing the colony was now truly queenless, had known what to do but had no primary material to work with.

Over the next fortnight or so this hive was going to generate hundreds thousands lots of drones. Not in itself a bad thing – this was a good colony and the positve influence on local bee genetics might be beneficial.

However, all the drones would emerge in the supers and be prevented from exiting the hive due to the queen excluder.

When this happens the drones die in their droves stuck half way through the excluder.

This is a distressing sight and, for a drone, a demoralising experience (I would imagine 12).

Under normal circumstances I would simply return every 3-4 days, pop the lid off the hive and release them. This wasn’t possible living four hours away … 

… so I played the ‘get out of jail free’ card by adding a thin eke and upper entrance.

Upper entrance

When I next check the colony I expect the drone brood to have all emerged and, largely, left the supers. I hope there’s a mated laying queen in the bottom box and there should be some capped worker brood.

What there’s unlikely to be is three full supers of honey 🙁

With no worker brood being reared for at least 5 weeks the foraging workforce will be significantly depleted. I hope they manage to defend what they’ve already collected … time will tell.

What went wrong?

After finding the supers full of drone brood I wrote “dodgy” on both sides of the queen excluder frame as I replaced it with a plastic spare.

I assumed the queen had found a bent wire and   s  q  u  e  e  z  e  d  her way through to have a field day – actually three weeks – in the supers.

However, I think the explanation is more prosaic than that 13.

My notes indicated I’d not seen the queen in this hive during the April inspection. In this instance evidence of absence was not absence of evidence … there were lots of eggs and brood in all staged. The colony was queenright and the queen was in the right place.

At least before I opened the hive 😉

And this is where stupidity, forgetfulness and plain dumb luck played their part. I … 

  • stupidly botched the inspection, taking the strength and health of the colony as the most important signs that all was well, but …
  • forgot that the next inspection – when I would be making up nucs – would also need worker eggs in the brood box to rear new queens from.
  • There’s more … I also presumably forgot to thoroughly inspect the queen excluder before laying it to the side, allowing …
  • dumb luck to intervene when the queen scooted around to the other side of the excluder and so end up trapped in the supers when I reassembled the hive.

Mea culpa.

That’s my best guess anyway.

Did I do the right thing?

Hive #29 was the last to be inspected after a hard day of beekeeping in late April.

Coincidentally it was also the last to be checked in mid-May 14

This limited my options somewhat and I made a judgement call as to the best course of action. Doing what I describe above risks the queen failing to emerge or mate. It also potentially risks the box being robbed as the workforce diminish, particularly with the upper entrance I’ve added.

Both of these could lead to the loss of the hive, but the loss/problem would be all mine. At the time, standing there swearing sweating in my beesuit, gasping for a beer, it seemed like the safest bet. It also seemed like the responsible course of action in the middle of a global pandemic.

I chose not to just dump the queen back into the brood box, add the upper entrance and leave them to it. Had the colony subsequently swarmed 15 the problem might then have been someone else’s

Did I do the right thing?

We’ll know soon enough … 😉


 

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


Colophon

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 … 😉

Queen cells … quantity and quality

How many queen cells should I leave in my hive?

This question pops up year after year at this time of the season.

Up and down the country we’re all busy implementing swarm control because our swarm prevention, er, didn’t 🙁

The majority of swarm control methods leave part of the colony to rear a new queen. Once she has emerged, matured, mated and proved her worth by laying up a frame or two you can then decide what to do with the old queen. 

Irrespective of the swarm control method you use – e.g. Pagden, nucleus method or a vertical split – the colony often produces quite a few queen cells. 

Similarly, if both your swarm prevention and swarm control failed and a prime swarm disappeared over the fence, there are likely to be several (or possibly lots of) queen cells left in the colony.

Queen cells – the good, the bad and the ugly

How many of these queen cells should you leave in the hive? 

Which one(s) should you leave?

Assumptions

I’m in the middle of my own swarm control at the moment and so intend to keep this relatively short and simple 1.

I am going to assume you start with one hive and you want to finish with one hive at the end of the process (i.e. you do not want to make increase). I’ll briefly mention rescuing queenless colonies and stock improvement as it’s relevant.

I’m also going to keep this as generic as possible. It’s not going to depend upon the method of swarm control employed or – with some caveats to be discussed later – whether the colony has naturally swarmed.

Here’s the starting position.

Your hive is making preparations to swarm. You apply a swarm control method that removes the old queen from the original brood box 2. This box therefore contains brood in all stages (BIAS) – eggs, larvae and sealed brood. This brood probably occupies most of the frames in the brood box. 

Also in the box are a very large number of adult bees, both workers and drones 3

And there will probably be one, several or lots of unsealed queen cells 4 present as well 🙂

Why do anything? or What’s the worst thing that could happen?

When a colony swarms naturally about 75% of the adult bees leave with the old queen. This figure is similar whether the colony is large or small. 

If you start with a large double brood colony it might contain 60,000 bees. Let’s assume a large swarm leaves as the first queen cells are capped (which is when the swarm usually scarpers).

There are still 15,000 bees and perhaps 15-18 frames of brood, several frames of which are close to emerging. The queen laying rate 3 weeks prior to the swarm was probably 1,000 to 2,000 eggs per day, meaning that number of adult workers are now emerging per day. 

Honey bee development

Honey bee development

About eight days after the queen cells were capped and the swarm left the new virgin queens emerge (see the bottom row in the picture above). By this time the worker population in the hive might well be over 20,000 again (some adult worker will have died of old age in the intervening period).

20,000 bees is more than enough to swarm again if several queens emerge 5.

These secondary swarms are called casts. They are headed by a virgin queen. They can be quite large if the original colony was very strong. 

However, with a lot of virgin queens emerging around the same time a strong colony can produce several casts, one after another. These are usually successively smaller and smaller 6. Not only are these casts too small to form an effective colony, but the originating colony can be weakened sufficiently to make its survival doubtful.

What’s the alternative?

Imagine the same double-brood colony. The old queen heads for the hills with 75% of the workforce. A week later the colony strength has been boosted by the emergence of a further 7 – 10 thousand workers … but this time there is only one capped queen cell developing.

The queen emerges.

If this queen also disappeared in a cast swarm the original colony would inevitably perish.

Why?

Because a week after the original swarm leaves there are no eggs or larvae in the colony young enough to be reared as new queens. 

She’s gone …

Swarming is reproduction of the honey bee ‘superorganism’. The survival of natural swarms is low (~25%) whereas the survival of swarmed colonies is reasonably high (>75%).

From an evolutionary perspective it makes no sense for the only queen to also leave, heading a cast swarm. The colony would have ‘traded’ a ~1:5 chance of producing two viable colonies for a 1:16 chance 7

It’s a no brainer as they say 8.

So, you can probably see where this is going now …

Swarm control

The three relatively generic and representative swarm control methods –  Pagden, a nucleus method or a vertical split – all involve manipulation of the hives one week after the initial intervention.

In the ‘classic’ Pagden method the original hive is moved from one side of the artificial swarm to the other. This has the effect of ‘bleeding off’ some of the workforce, so weakening the hive. The resulting reduced worker population often tear down all but a small number of queen cells. The reduced bee numbers also make the production of casts less likely as the colony is weaker.

Pagdens' artificial swarm ...

Pagdens’ artificial swarm …

In a vertical split the hive is reversed on the stand after 7 days, achieving exactly the same outcome on a much smaller footprint with less equipment 🙂 9

In both these methods the flying bees that have reoriented to the initial new position of the queenless hive return to find the hive moved. They then enter the nearest hive, which is the queenright component (i.e. the artificial swarm). 

I’ll get to the nucleus method in a moment.

Sometimes you will see it recommended that you also check the queenless colony at this one week timepoint to ensure that there are not large numbers of queen cells still present 10. It’s not usually necessary but – assuming you are careful – it does not cause any harm. As I explain below, it can help give you confidence.

If you don’t perform the one week hive manoeuvre you really should check for queen cells and reduce the number present.

In the nucleus method I describe the beekeeper must manage queen cell numbers in the queenless hive. Not doing so almost certainly risks losing multiple casts when the queens emerge together.

How many queen cells should you leave?

The queenless component of your swarm control only needs one queen cell

Any less than that and the colony will be non-viable without further intervention from the beekeeper.

Any more and there’s a risk that the colony will generate one or more casts. 

A very strong queenless colony with large numbers of queen cells is a recipe for disaster … or, if not a disaster, then a lot of frustration as you scurry around trying to catch the casts and/or rescue the colony from swarming itself to destruction.

Workers in very strong colonies can ‘hold back’ queens, effectively trapping them in the cell, so that emergence is more-or-less simultaneous. Should you chance to open a colony in this situation all hell breaks loose, with virgin queens dashing about all over the place.

Been there, got the T-shirt 🙂

Although entertaining – at least is retrospect – it’s better to avoid this sort of situation by restricting queen cell numbers.

All your eggs in one basket

And this is where the beginner starts to experience some trepidation.

They have to reduce queen cell numbers … to one.

That queen will head the colony for the next year or three. She’ll mother tens of thousands of workers who will make countless foraging trips and collect tens or (hopefully) hundreds of pounds of honey.

Choosing that one queen cell feels like a lot of responsibility.

The consequences of choosing a dud feel very serious indeed.

Surely leaving two or three would be a ‘safer’ bet? 

Backups, if you will … just in case the first one turns out to be a dud.

How do you know which one to pick?

Trust the bees

And this is where you need to trust the bees. They’ve been doing this pretty well for several million years.

You don’t need to choose a single egg from the thousands possibly present in the colony. The one egg that will be cared for, fed copious amounts of royal jelly and eventually emerge to head the colony.

The bees have already made those decisions 11.

They’ve started several queen cells, the majority of which are likely to be suitable. You just need to choose one of those queen cells to leave in the hive. 

It’s not a one in thousands chance of choosing a ‘winner’, it’s more like one in ten … in which any of the ten would probably be OK.

With a few caveats …

What are the features of a good queen cell?

You open the hive and find a number of sealed and unsealed queen cells.

Which to choose?

What are the features you are looking for?

What are the features you can see?

Sealed queen cell ...

Sealed queen cell …

Size, shape and appearance are the obvious ones. Position on the comb might also influence your choice.

What are the features you cannot see?

Is is a charged cell i.e. does it contain a developing pupa? Has that pupa been well fed as a larva?

Size, shape, appearance and position

Mature queen cells are large, about 3 cm long. The position on the comb – whether on the face or edge can influence the apparent size. They are generally conical, more or less evenly tapering to a neatly rounded tip. Queen cells that have been well-tended by the bees are often heavily sculpted on the outside. This is generally taken to be a “good thing”, but note that this doesn’t happen until after the queen cell is capped (see the photo above). Uncapped cells are usually smooth (see the next photo).

I think the position on the frame is irrelevant in terms of queen cell quality, but it does influence which I choose. The cell should be drawn from worker comb (!) 12 and – particularly if I’m likely to be either cutting the cell out or moving the entire frame – I like it to be in a position unlikely to get damaged as I manipulate the frames in the hive.

The edge of drawn comb, with space below and to the side, makes things easy. The central face of the comb, especially if it’s on fresh comb and not near a wire in the foundation, is also a good bet. 

The position is more important if you’re going to do something with the cell or frame other than let it emerge in situ.

Charged cells

How do you know there’s a well-fed pupa in the cell?

Ted Hooper (in his Guide to Bees and Honey) describes gently prising the cap off a sealed queen cell to check it is occupied, then re-sealing it to let development run its course. He finishes discussing how to re-seal the cell with the words “you have to do a good job or the bees will tear it down.”

I bet 😉

There are easier ways.

Firstly you can be pretty sure that any well-shaped sealed cell with a good, well sculpted appearance is likely to be occupied. Alternatively, you can identify these cells in advance and only allow those you know contain a developing larva sitting on a thick bed of royal jelly to mature.

A practical example

A few days ago I used the nucleus method for swarm control in all my colonies in one apiary. Due to work constraints and lockdown some colonies were only just starting to make preparations to swarm. None of the colonies had well developed, charged queen cells. Some had ‘play cups’ with eggs present.

Three days after making up the nucs I checked the queenless parent colonies. All had a few developing queen cells.

Here is the same photograph as above, with some cells numbered on the frame.

Queen cells – capped, open and just plain dodgy

Which do you choose?

Here is the view from below of the same frame.

Queen cells – practical example

  1. A sealed cell, perhaps a bit small 13
  2. Is a nice looking unsealed cell with a thick bed of royal jelly supporting a larva inside.
  3. Also unsealed and with a good space underneath for the cell to be drawn out as it develops.
  4. Is very similar to #2. Smooth exterior as it’s only 3 days old and unsealed.
  5. A thickened play cup from a previous season. There is no egg, larva or royal jelly inside it.

Remember that this is only 3 days after implementing swarm control.

I destroyed the sealed cell #1. Since it was already sealed it was probably made from an older larva. Cells are sealed on the eighth day after the egg is laid. Since this was only 72 hours after removing the queen the larva was probably two days old before being reared as a queen – i.e. 8 minus 3 days since queen removal minus the three days it would have already spent as an egg. Alternatively, it might have been present when I removed the queen, though I did check reasonably thoroughly.

I couldn’t be sure of the contents of this cell and I suspected that it may not have been fed on copious amounts of royal jelly during the very early days after hatching from the egg.

Cell #3 was also squidged. If you look closely from below you can clearly see the larva but no thick bed of royal jelly. I doubted it had been fed well enough in the early days. Here’s an enlargement …

Cells #1 to #4 enlarged.

Why risk it? There are better cells on the frame.

I ignored #5. It’s not a queen cell and never will be.

Uncapped cells #2 and #4 were retained. They are the right size, have a good appearance and are well placed on the frame.

I marked the top of the frame with a queen marking pen to remind me where to check, and more importantly where to be careful, when I inspect the colony a week after making up the nuc.

X marks the spot

Note that the photo above is a different hive to the numbered photo of queen cells (which I forgot to photograph).

Hold on … not so fast

Go back and look again at the numbered photo of queen cells.

There is another cell, uncapped and filled with royal jelly, to the left and a little higher than the sealed queen cell #1.

This cell is actually pretty obvious. There are relatively few bees on the frame and it is not particularly well ‘hidden’. 

Miss a couple more like that in a very strong hive and there’s a chance the colony will throw off several casts when the queen emerge. The unlabelled cell, and cells #2 and #4 are all very similar in age and appearance and would likely emerge within hours of each other.

Seven days after implementing swarm control

The hives are checked again 14.

I know which frames have good, charged developing queen cells. They are the ones that are marked. I therefore :

  • treat these frames very carefully. Do not shake the bees off the frame!
  • make sure the cells are now capped and starting to be sculpted by the bees.
  • gently inspect the remainder of the frame for other queen cells.
  • destroy any new cells that I find

I choose one of the queen cells and destroy any others on the frame. If there is more than one marked frame and I don’t need the cell for another colony (see below) then I destroy the cells on the other marked frame as well.

I then thoroughly inspect every frame in the brood box, shaking all the bees off the frames and checking for any queen cells I may have missed previously. There will be some.

All I find are destroyed.

I close the hive up and leave it undisturbed for the queen to emerge, mate and start laying. I’ll discuss this – apparently interminable – period in the future sometime.

I’m confident the cell contains a well fed pupae. It was the the bees that really selected the queen … all I did was whittle down their selection to the final choice.

Using ‘spare’ queen cells

In the photo above there are two marked frames. This is a good colony. Frugal, productive, well behaved etc. 15

There is another colony in the apiary which is poorly tempered. They are also requeening and are at the same stage.

Assuming the cells on both marked frames are good I’ll transfer one to the badly behaved colony when I conduct the seven day inspection. You can transfer the entire frame or you can gently cut the queen cell out and use it directly 16

All of the developing queen cells in the badly behaved colony will first be destroyed. Since there are no eggs or young larvae in that colony (and no queen as she was removed a week ago) they cannot rear another from their own genetic material.

The new queen will be better quality.

Similarly, you can use a ‘spare’ queen from a good hive to rescue a terminally queenless colony, or to replace an underperforming or substandard queen.

A really dodgy queen cell

I wanted to squeeze in a picture of what not to choose. 

Bride of Frankenstein queen cell

There are so many things wrong with this.

Where to start?

It’s drawn from drone comb and is not neatly tapering and conical. It’s poorly sculpted considering its age and size, which is far too big.

Whatever emerges from this cell, if anything, will not be any use to me or the bees 🙁

Seven day only inspection

The process described above involves an additional inspection 3-4 days after implementing swarm control. I think this is a modest amount of additional work for:

  • the peace of mind it gives when selecting the final cell to leave
  • the time saved when going through the colony at the seven day inspection

However, often it’s not possible. In that case I refer you back to the description of what a good sealed queen cell looks like.

Choose one of those.

Just one 😉


Notes

With gale force winds predicted for the next 2-3 days I ended up checking the ‘example’ colony (above) on day 6 after implementing swarm control measures. Here is the same frame:

Just one!

I removed two less convincing queen cells on either side of the one selected (#2 in the labelled photograph further up the page). There were a small number of queen cells elsewhere in the colony. All were removed. I’m leaving just one cell sealed, I know it contains a well fed larva. She’ll emerge in about a week and should be mated – weather permitting – a week or so after that.

And now the wait begins … 😉

 

Darwinian beekeeping

A fortnight ago I reviewed the first ten chapters of Thomas Seeley’s recent book The Lives of Bees. This is an excellent account of how honey bees survive in ‘the wild’ i.e. without help or intervention from beekeepers.

Seeley demonstrates an all-too-rare rare combination of good experimental science with exemplary communication skills.

It’s a book non-beekeepers could appreciate and in which beekeepers will find a wealth of entertaining and informative observations about their bees.

The final chapter, ‘Darwinian beekeeping’, includes an outline of practical beekeeping advice based around what Seeley (and others) understand about how colonies survive in the wild.

Differences

The chapter starts with a very brief review of about twenty differences between wild-living and managed colonies. These differences have already been introduced in the preceding chapters and so are just reiterated here to set the scene for what follows.

The differences defined by Seeley as distinguishing ‘wild’ and ‘beekeepers’ colonies cover everything from placement in the wider landscape (forage, insecticides), the immediate environment of the nest (volume, insulation), the management of the colony (none, invasive) and the parasites and pathogens to which the bees are exposed.

Some of the differences identified are somewhat contrived. For example, ‘wild’ colonies are defined fixed in a single location, whereas managed colonies may be moved to exploit alternative forage.

In reality I suspect the majority of beekeepers do not move their colonies. Whether this is right or not, Seeley presents moving colonies as a negative. He qualifies this with studies which showed reduced nectar gathering by colonies that are moved, presumably due to the bees having to learn about their new location.

However, the main reason beekeepers move colonies is to exploit abundant sources of nectar. Likewise, a static ‘wild’ colony may have to find alternative forage when a particularly good local source dries up.

If moving colonies to exploit a rich nectar source did not usually lead to increased nectar gathering it would be a pretty futile exercise.

Real differences

Of course, some of the differences are very real.

Beekeepers site colonies close together to facilitate their management. In contrast, wild colonies are naturally hundreds of metres apart 1. I’ve previously discussed the influence of colony separation and pathogen transmission 2; it’s clear that widely spaced colonies are less susceptible to drifting and robbing from adjacent hives, both processes being associated with mite and virus acquisition 3.

Abelo poly hives

50 metres? … I thought you said 50 centimetres. Can we use the next field as well?

The other very obvious difference is that wild colonies are not treated with miticides but managed colonies (generally) are. As a consequence – Seeley contends – beekeepers have interfered with the ‘arms race’ between the host and its parasites and pathogens. Effectively beekeepers have ‘weaken[ed] the natural selection for disease resistance’.

Whilst I don’t necessarily disagree with this general statement, I am not convinced that simply letting natural selection run its (usually rather brutal) course is a rational strategy.

But I’m getting ahead of myself … what is Darwinian beekeeping?

Darwinian beekeeping

Evolution is probably the most powerful force in nature. It has created all of the fantastic wealth of life forms on earth – from the tiniest viroid to to the largest living thing, Armillaria ostoyae 4. The general principles of Darwinian evolution are exquisitely simple – individuals of a species are not identical; traits are passed from generation to generation; more offspring are born than can survive; and only the survivors of the competition for resources will reproduce.

I emphasised ‘survivors of the competition’ as it’s particularly relevant to what is to follow. In terms of hosts and pathogens, you could extend this competition to include whether the host survives the pathogen (and so reproduces) or whether the pathogen replicates and spreads, but in doing so kills the host.

Remember that evolution is unpredictable and essentially directionless … we don’t know what it is likely to produce next.

Seeley doesn’t provide a precise definition of Darwinian beekeeping (which he also terms natural, apicentric or beefriendly beekeeping). However, it’s basically the management of colonies in a manner that more closely resembles how colonies live in the wild.

This is presumably unnnatural beekeeping

In doing so, he claims that colonies will have ‘less stressful and therefore more healthful’ lives.

I’ll come back to this point at the end. It’s an important one. But first, what does Darwinian mean in terms of practical beekeeping?

Practical Darwinian beekeeping

Having highlighted the differences between wild and managed colonies you won’t be surprised to learn that Darwinian beekeeping means some 5 or all of the following: 6

  • Keep locally adapted bees – eminently sensible and for which there is increasing evidence of the benefits.
  • Space colonies widely (30-50+ metres) – which presumably causes urban beekeepers significant problems.
  • Site colonies in an area with good natural forage that is not chemically treated – see above.
  • Use small hives with just one brood box and one super – although not explained, this will encourage swarming.
  • Consider locating hives high off the ground – in fairness Seeley doesn’t push this one strongly, but I could imagine beekeepers being considered for a Darwin Award if sufficient care wasn’t taken.
  • Allow lots of drone brood – this occurs naturally when using foundationless frames.
  • Use splits and the emergency queen response for queen rearing i.e. allow the colony to choose larvae for the preparation of new queens – I’ve discussed splits several times and have recently posted on the interesting observation that colonies choose very rare patrilines for queens.
  • Refrain from treating with miticides – this is the biggy. Do not treat colonies. Instead kill any colonies with very high mite levels to prevent them infesting other nearby colonies as they collapse and are robbed out.

Good and not so good advice

A lot of what Seeley recommends is very sound advice. Again, I’m not going to paraphrase his hard work – you should buy the book and make your own mind up.

Sourcing local bees, using splits to make increase, housing bees in well insulated hives etc. all works very well.

High altitude bait hive …

Some of the advice is probably impractical, like the siting of hives 50 metres apart. A full round of inspections in my research apiary already takes a long time without having to walk a kilometre to the furthest hive.

The prospect of inspecting hives situated at altitude is also not appealing. Negotiating stairs with heavy supers is bad enough. In my travels I’ve met beekeepers keeping hives on shed roofs, accessed by a wobbly step ladder. An accident waiting to happen?

And finally, I think the advice to use small hives and to cull mite-infested colonies is poor. I understand the logic behind both suggestions but, for different reasons, think they are likely to be to the significant detriment of bees, bee health and beekeeping.

Let’s deal with them individually.

Small hives – one brood and one super

When colonies run out of space for the queen to lay they are likely to swarm. The Darwinian beekeeping proposed by Seeley appears to exclude any form of swarm prevention strategy. Hive manipulation is minimal and queens are not clipped.

They’ll run out of space and swarm.

Even my darkest, least prolific colonies need more space than the ~60 litres offered by a brood and super.

Seeley doesn’t actually say ‘allow them to swarm’, but it’s an inevitability of the management and space available. Of course, the reason he encourages it is (partly – there are other reasons) to shed the 35% of mites and to give an enforced brood break to the original colony as it requeens.

These are untreated colonies. At least when starting the selection strategy implicit in Darwinian beekeeping these are likely to have a very significant level of mite infestation.

These mites, when the colony swarms, disappear over the fence with the swarm. If the swarm survives long enough to establish a new nest it will potentially act as a source of mites far and wide (through drifting and robbing, and possibly – though it’s unlikely as it will probably die – when it subsequently swarms).

A small swarm

A small swarm … possibly riddled with mites

Thanks a lot!

Lost swarms – and the assumption is that many are ‘lost’ – choose all sorts of awkward locations to establish a new nest site. Sure, some may end up in hollow trees, but many cause a nuisance to non-beekeepers and additional work for the beekeepers asked to recover them.

In my view allowing uncontrolled swarming of untreated colonies is irresponsible. It is to the detriment of the health of bees locally and to beekeepers and beekeeping.

Kill heavily mite infested colonies

How many beekeepers reading this have deliberately killed an entire colony? Probably not many. It’s a distressing thing to have to do for anyone who cares about bees.

The logic behind the suggestion goes like this. The colony is heavily mite infested because it has not developed resistance (or tolerance). If it is allowed to collapse it will be robbed out by neighbouring colonies, spreading the mites far and wide. Therefore, tough love is needed. Time for the petrol, soapy water, insecticide or whatever your choice of colony culling treatment.

In fairness to Seeley he also suggests that you could requeen with known mite-resistant/tolerant stock.

But most beekeepers tempted by Darwinian ‘treatment free’ natural beekeeping will not have a queen bank stuffed with known mite-resistant mated queens ‘ready to go’.

But they also won’t have the ‘courage’ to kill the colony.

They’ll procrastinate, they’ll prevaricate.

Eventually they’ll either decide that shaking the colony out is OK and a ‘kinder thing to do’ … or the colony will get robbed out before they act and carpet bomb every strong colony for a mile around.

Killing the colony, shaking it out or letting it get robbed out have the same overall impact on the mite-infested colony, but only slaying them prevents the mites from being spread far and wide.

And, believe me, killing a colony is a distressing thing to do if you care about bees.

In my view beefriendly beekeeping should not involve slaughtering the colony.

Less stress and better health

This is the goal of Darwinian beekeeping. It is a direct quote from final chapter of the book (pp286).

The suggestion is that unnatural beekeeping – swarm prevention and control, mite management, harvesting honey (or beekeeping as some people call it 😉 ) – stresses the bees.

And that this stress is detrimental for the health of the bees.

I’m not sure there’s any evidence that this is the case.

How do we measure stress in bees? Actually, there are suggested ways to measure stress in bees, but I’m not sure anyone has systematically developed these experimentally and compared the stress levels of wild-living and managed colonies.

I’ll explore this topic a bit more in the future.

I do know how to measure bee health … at least in terms of the parasites and pathogens they carry. I also know that there have been comparative studies of managed and feral colonies.

Unsurprisingly for an unapologetic unnatural beekeeper like me ( 😉 ), the feral colonies had higher levels of parasites and pathogens (Catherine Thompson’s PhD thesis [PDF] and Thompson et al., 2014 Parasite Pressures on Feral Honey Bees). By any measurable definition these feral colonies were less healthy.

Less stress and better health sounds good, but I’m not actually sure it’s particularly meaningful.

I’ll wrap up with two closing thoughts.

One of the characteristics of a healthy and unstressed population is that it is numerous, productive and reproduces well. These are all characteristics of strong and well-managed colonies.

Finally, persistently elevated levels of pathogens are detrimental to the individual and the population. It’s one of the reasons we vaccinate … which will be a big part of the post next week.


 

Who’s the daddy?

I’ve recently discussed the importance and influence of polyandry for honey bee colonies. Briefly, polyandry – the mating of the queen with multiple (~12-18) drones – is critical for colony fitness e.g. ability to resist disease, forage efficiently or overwinter successfully.

Hyperpolyandry, for example resulting from instrumental insemination of the queen with sperm from 30+ drones, further increases colony fitness and disease resistance.

How do you measure polyandry?

Essentially, you genetically analyse the worker bees in the colony to determine the range of patrilines present. Patrilines are genetically distinct offspring fathered by different drones. Essentially they are subfamilies within the colony.

With a finite number of patrilines – which there must be, because the queen does not mate with an infinite number of drones – there will be a point at which the more workers you screen the fewer new patrilines will be detected.

Search and ye shall find – detecting rare patrilines

The more you screen, the more you are likely to have detected all the patrilines present.

However, the queen uses sperm randomly when fertilising worker eggs. This compounds the difficulty in determining the full range of different patrilines present in a population. In particular, it makes detecting very rare patrilines difficult.

For example, if 20% of workers belong to one patriline you don’t need to sample many bees to detect it. In contrast, if another patriline is represented by 0.0001% of randomly selected workers you would probably have to screen thousands to be sure of detecting it.

Consequently, rare patrilines in the honey bee worker population are very difficult to detect. Inevitably this means that the number of drones the queen mates (~12-18) with is probably an underestimate of the actual number 1.

Half-sisters and super-sisters

Worker bees are often described as ‘half sisters’ to each other. They share the same mother (the queen), but different fathers.

Actually, as you should now realise, that’s an oversimplification because – with only ~12-18 different fathers contributing to the genetics of the colony – some workers are going to be more related to each other because they share the same father and mother.

Half-sisters share the same mother but have different fathers and share about 25% of their genes.

Super-sisters share the same mother and father and so share about 75% of their genes (25% from the queen and 50% from the drone).

Super-sisters are more likely to help each other in the colony 2.

Emergency queens and nepotism

What’s the most important decision a colony makes?

If the queen is killed (or removed) the workers rear new queens under the so-called ’emergency response’. They feed selected young larvae copious amounts of Royal Jelly to rear a replacement queen.

Arguably, the most important decision the workers make is the selection of the day-old larvae to rear as new queens.

If they get it wrong the colony is doomed. If they get it right the colony will flourish 3.

But as described above, workers are more or less related to each other genetically.

To ensure the continued propagation of at least some of their genes it might be expected that the nurse bees making this selection 4 would choose larvae more closely related to themselves.

Do worker bees exhibit nepotism when rearing emergency queens?

If workers were nepotistic you’d expect the most common patrilines in the nurse/worker bee population would also predominate in the queens reared.

However, for at least 20 years evidence has been accumulating that indicates bees are not nepotistic. On the contrary, emergency queens appear to be reared from some of the rare patrilines in the colony.

A recent paper from James Withrow and David Tarpy has provided some of the best evidence for the existence of these so-called royal patrilines in honey bee colonies 5.

Royal patrilines

Evidence for these goes back to at least 1997 6, with about half a dozen publications in the intervening period. Essentially all used broadly the same approach; they genetically screened worker bees and the emergency queens they reared to determine which patrilines were present in the two groups. 

With certain caveats (size of study, number of microsatellites screened, colony numbers etc.) all concluded that colonies rear emergency queens from some of the rarest patrilines in the colony.

The recent study by Withrow and Tarpy is well explained and probably the most comprehensive, so I’ll use that to flesh out the details.

Experimental details

Six double-brood colonies were each split into a three separate colonies; a queenright single-brood colony and two five-frame nucs. The latter contained eggs and young larvae and so reared emergency queens.

Seven days later the developing emergency queens were all harvested for future analysis. One or two frames from the nucs were then exchanged with frames containing eggs and day-old larvae from the matched queenright colony.

The nucs then started rearing new queens … again.

And again … and again.

This process was repeated until the nucs failed.

In total over 500 queens were reared (to 7 days old) from these six original colonies. These queens were analysed genetically by microsatellite analysis, as were over 500 workers from colonies.

Within the 6 experimental colonies the authors identified a total of 327 patrilines (or subfamilies as Withrow and Tarpy describe them), ranging from 34-77 per colony. 108 patrilines (4-40 per colony) were exclusively detected in worker bees and 130 patrilines (5-55/colony) were exclusively detected in queens.

Cryptic “royal” subfamilies

Over 40% of queens raised per colony were produced from the patrilines exclusively detected in the queen population.

Subfamily distribution per colony.

As shown in the figure above, many queens (black bars) were reared from subfamilies (patrilines) not represented in the worker bee population (grey bars, sorted left to right by abundance).

Since there were different numbers of patrilines per colony (34-77), the bias towards the rarer patrilines is more apparent if you instead split them into tertiles (thirds) based upon worker abundance.

Are the queens predominantly reared from the most common tertile, the intermediate tertile or the rarest tertile?

Frequency distribution of subfamilies.

It’s very clear from this graph that workers select queens from the rarest patrilines within the colony.

It is therefore very clear that worker bees do not exhibit nepotism when choosing which larvae to rear emergency queen from.

Implications for our understanding of honey bee reproduction

Two points are immediately apparent:

  • there is a cryptic population of queen-biased patrilines that have largely been overlooked in genetic studies of honey bee polyandry
  • honey bee queens mate with more drones than conventional studies of worker bee patrilines indicate

Colony 5 had at least 77 distinct subfamilies (there might have been more detected had they screened more than the 94 workers and 135 queens from this colony). By extrapolation it is possible to determine that the effective queen mating frequency (me; the number of drones the queen had mated with) was ~32 if all the samples (worker and queen) were taken into account. If only the worker or queen samples were used for this calculation the effective queen mating frequency would be ~12 or ~65 respectively.

The average effective queen mating frequency over the six colonies was ~33 (total), significantly higher than the oft-quoted (including at the top of this page) me of ~12-18.

So perhaps honey bees really are hyperpolyandrous … or even extremely hyperpolyandrous as the authors suggest.

It’s worth noting in passing that routine mating frequencies over 30 are almost never quoted for honey bees 7, but that the ‘normal’ me ~12-18 is rather low when compared with other species within the genus Apis. The giant honey bee, Apis dorsata, exhibits mating frequencies of greater than 60.

Who’s the daddy?

So, when it comes to emergency queens , although we might not know precisely who the daddy is, we can be pretty certain the particular patriline selected by the workers is most likely to be one of the rare ones in the colony.

Mechanistically, what accounts for this?

Are these larvae selected solely because they are rare?

That seems unlikely, not least because it would require some sort of surveying or screening by nurse bees. Not impossible perhaps, though I’m not sure how this would be achieved.

Perhaps it is not even worker selection?

An alternative way to view it is larval competition. A better competing larvae would be fed Royal Jelly and would be much more likely to pass on her genes to the next generation.

We don’t know the answers to these questions … yet.

Or whether they’re the wrong questions entirely.

Swarming and supercedure

The colony rears a new queen under three conditions; enforced queenlessness (as described above) which induces emergency queen rearing, prior to swarming and during supersedure.

These are fundamentally different processes in terms of the larvae used for queen rearing.

During swarming and supersedure 8 the queen lays the egg in a ‘play cup’ which is subsequently engineered into a queen cell in which the new queen develops.

Play cups

However, it is known that the patrilines of queens reared during the swarming response are similar to those of workers in the same colony 9, implying that there is no overt selection by the workers (or the parental queen).

Queen rearing

Does this insight into how bees rear new queens have any implications for how beekeepers rear new queens?

There are about as many queen rearing methods as there are adult workers in a double-brood colony in late June. Many  exploit the emergency queen rearing response by a colony rendered temporarily or permanently queenless.

Beekeepers often comment on the differential ‘take’ of grafted larvae presented to queenless cell raising colonies.

Sometimes you get very good acceptance of the grafted larvae, other times less so.

Of course, we only show the ones that worked well!

3 day old QCs ...

3 day old QCs …

Differential ‘take’ is often put down to the state of the cell raising colony or the nectar flow (or the cackhandedness of the grafter, or the phase of the moon, or about 100 other things).

I have never heard of beekeepers comparing the ‘take’ of larvae originating from the cell raising colony with those from another colony. The latter are always going to be ‘rare’ if you consider the patrilines present in the cell raising colony. However, grafts taken from the same colony as used for cell raising 10 are likely to reflect the predominant patrilines.

Are these accepted less well by the nurse bees?

I suspect not … but it is testable should anyone want to try.

My expectation would be that the presentation of larvae in a vertically oriented cell bar frame would likely override any genetic selectivity by the colony. They’re desperate to raise a new queen and – thank goodness – here’s a few that might do.

Alternatively, differential acceptance is more likely to reflect use of larvae of an unsuitable age, or that have been damaged during grafting.

As I listen to the wind howling outside it seems like a very long time until I can test any of these ideas … 🙁


Colophon

Ray Winstone (as Carlin) 1979

Who’s the daddy? is British slang for who, or what, is the best. It originated in a line by Ray Winstone’s character Carlin from the 1979 film Scum. This was not a romantic comedy and I’m certainly not recommending viewing it. Nevertheless, the phrase became widely used over the subsequent couple of decades and seemed appropriate here because the colony is dependent on selecting high-quality larvae for colony survival.