Category Archives: Problems

Superinfection exclusion

Alpha

Beta

Gamma

Delta 

The majority of readers will identify these as the current circulating variant strains of Covid 1. The World Health Organisation decided upon this naming system as being easier and more practical to discussed (sic) by non-scientific audiences 2.

All viruses vary, and viruses with genomes made from ribonucleic acid (RNA) vary more than those which have deoxyribonucleic acid (DNA) genomes. This is because the enzymes that replicate RNA virus genomes do not have an error correction facility. 

The virus that causes Covid-19 is an RNA virus and so is Deformed wing virus (DWV), probably the most significant virus to bees and beekeepers (other than those who have Covid that is).

Virus variation

Why does virus variation matter? 

Or, asking the same question in a more roundabout way, why don’t RNA viruses evolve error correcting enzymes (after all, the DNA viruses have these … can it be that difficult?).

If the enzyme makes errors then that’s surely a bad thing?

Actually … for the particular ‘lifestyle’ that these viruses practice, errors and variation are a good thing.

They benefit the virus.

But you know all this already, even if you don’t think you do.

Covid cases caused by the delta variant

The SARS-Cov2 delta variant now accounts for at least 99% of cases of Covid-19 in the UK. It accounted for just 0.1% of cases in late February.

The delta variant is much more transmissible. It carries errors, or mutations as they’re more correctly known, that – for whatever reason – means it can be passed from person to person much more efficiently 3.

These mutations benefit the virus and allow it to spread further and faster 4.

If the mutations (‘errors’) that the virus acquires are beneficial – by increasing transmission, by expanding the cell, tissue or host range, by helping evade the immune response in a partially vaccinated population (!) for example – then the virus will successfully replicate and produce more viruses carrying the same mutation.

And a bunch of additional ones as well … acquired during the last round of replication.

Strains and types

At some point a virus acquires sufficient mutations from an earlier incarnation that it’s identified as a distinct strain.

For example, SARS-Cov2 is ~82% identical at the genome level to SARS-Cov1 which caused the SARS pandemic in 2003 5. SARS-Cov2 did not evolve from SARS-Cov1, but they share a common ancestor. They are different strains or types of coronavirus.

There are no hard and fast rules that define when a virus is considered a different strain or type. Often it’s historical, reflecting the geographic origin, or the source from which the virus was isolated. Different strains may exhibit different phenotypes – host range, transmission, disease etc. – but don’t have to.

For example (before we get back to honey bee viruses) there are three ‘types’ of poliovirus that are about 80% identical at the genetic level. They all cause exactly the same disease (poliomyelitis) and they replicate in an identical manner – same cell, tissue and host range for example. However, to the human immune system they ‘look’ different. The immune response to poliovirus type 1 will not protect you from infection with poliovirus type 3. That’s why the poliovirus vaccine contains a mixture of all three types, to protect you from all polioviruses.

You can even get infected with two types of poliovirus simultaneously, and the virus can replicate in the same cells in your gut … or, if you’re unfortunate, your brain.

I’ll return to dual infections shortly as it’s an important topic … and related to the study I’m going to discuss.

Deformed wing virus

There are two types of DWV, designated type A and type B 6.

Originally these had names that reflected their original isolation.

DWV type A was termed Deformed wing virus and was isolated from honey bees displaying the characteristic symptoms of developmental deformities shown in the image below.

DWV type B was termed Varroa destructor virus type 1 and was isolated from the ectoparasitic mite Varroa destructor.

It “does what is says on the tin” … DWV symptoms in a recently emerged worker

These viruses are very similar to each other. They are something like 85% identical at the level of the RNA genome. More importantly than this genetic identity (or perhaps similarity would be a better term to use here) is the fact that they appear to cause very similar diseases in honey bees.

Although early studies suggested there were some differences in their virulence, more recent work from Prof. Rob Paxton in Germany, from my lab, and from Dr. Eugene Ryabov and colleagues in the USA suggests these two types of DWV are actually very similar, at least in pathogenesis.

There do appear to be some differences, with the suggestion that type A does not replicate in Varroa whereas (full disclosure, the following study is from my lab) type B does. Undoubtedly, the other genetic differences between the types will confer some subtle variation in phenotype (effectively what they ‘do’), but – as far as beekeeping is concerned – they should probably be considered the same.

A protective, non-lethal type A DWV? 

All of which made a 2015 colony-level study 7 of DWV infection rather intriguing.

This reported the survival of colonies that were infected with a “non-lethal” type B strain which were protected from infection with the “lethal” type A strain 8. The authors summarised the significance of this study like this:

We propose that this novel stable host-pathogen relationship prevents the accumulation of lethal variants, suggesting that this interaction could be exploited for the development of an effective treatment that minimises colony losses in the future.

At the time there was a flurry of excitement and discussion about this 9

Superinfection exclusion

They proposed that the mechanism that prevented the infection with the type A strain was superinfection exclusion.

Virologists love mechanisms … 🙂 

Which brings me back to virus variation. 

Imagine a population of variant viruses trying to infect a new host … like a bee.

Survival of the fittest – selection for better replicating viruses from a mixed population

In mixed infections, a virus that has an advantage over the others in the population ends up ‘winning’ the competition for the resources of the host. They therefore make more progeny viruses.

One of the advantages could be that the virus simply replicates faster

Another – more subtle, but the same outcome – is that the virus prevents other viruses from infecting the same cell (and, by extension, host).

By excluding competing viruses it effectively ‘wins’ the competition.

Superinfection exclusion – one virus (type) can prevent infection by related but different viruses

And some viruses do exactly this using a variety of cellular mechanisms.

For example, many viruses turn off the expression of the cellular receptor (think of this as the door) they use to enter the cell. If there’s no receptor (no door) then another virus cannot enter.

With no other viruses to compete with in the same cell there can only be one type of virus produced 10.

There are other mechanisms as well, but we’ll stick with the receptor one as it’s easy to comprehend.

Superinfection of a cell containing a virus that has the ability to turn off the cellular receptor it uses will effectively exclude the second virus from replicating … hence superinfection exclusion.

What don’t we know about DWV?

A lot 🙁

We don’t know how it gets into cells. We don’t know a huge amount about its replication and we know precious little about the way it interacts with the cellular machinery (the ‘stuff’ in the cell that the virus hijacks to make more viruses) of the host. 

However, since 2015 we do know that all the types of DWV that have been carefully studied appear to be more or less equally virulent. None appear to be ‘non-lethal’ as claimed for the type A virus in the superinfection exclusion paper.

This prompted us to look in a bit more detail at the consequences of dual or sequential infections with DWV in the laboratory. 

Is there a precedence at work?

In mixed infections, does one virus always ‘win’ and predominate in the new virus population?

In sequential infections, does it matter the order in which the viruses are acquired?

And mixed infections are pretty much the norm for DWV infection. All bees, whether previously exposed to Varroa or in Varroa-free regions, appear to have low levels of DWV already present. If parasitised by the mite, these bees must experience a mixed infection.

In addition, studies we published several years ago 11 showed that recombinant viruses – essentially hybrids between type A and type B DWV – often predominated in heavily Varroa-infested colonies 12.

If mixed infections cannot occur, how do such hybrids form?

Some of the answers to these questions are in our recently paper published in the ISME Journal. Gusachenko, O. et al., (2021) First come, first served: superinfection exclusion in Deformed wing virus is dependent upon sequence identity and not the order of virus acquisition. ISME J (2021). https://doi.org/10.1038/s41396-021-01043-4

Mixed DWV infections

I don’t propose to give a pupa-by-pupa account of the studies we conducted. You can read the paper – it’s open access and (because Olesya ‘Alex’ Gusachenko, the lead author, did most of the writing) relatively easy to comprehend 😉

But here are a few highlights.

Over the last few years we have produced reagents that allow us to produce almost ‘pure’ stocks of type A, type B or hybrid type A/B 13 strains of DWV.

At least 99.99% of these stocks are of one DWV type. Note that there will still be variation within this population as the replication errors probably generate one mutation per virus in the population. We therefore refer to these virus stocks as near clonal.

Injection of honey bee pupae with any of these viruses resulted in very similar levels and kinetics of replication – all the viruses replicate as far and as fast as each other.

In mixed injections, when two viruses were administered simultaneously, both replicated to equivalent levels. 

We therefore found no evidence for the dominance of one strain over another.

Sequential DWV infections

But it got more interesting when we did sequential injections. We did these by injecting with one virus, waiting 24 hours and then injecting with a second virus.

Using type A and type B DWV both viruses had replicated to similar high levels (billions of virus per pupa) within 48 hours, irrespective of the order of addition. 

If superinfection exclusion was operating we would have perhaps expected type A to have prevented or reduced the replication of type B. However, that didn’t appear to be the case.

Competition between sequential infecting DWV isolates. VVV is type B, VDD is type A and VVD is a hybrid between them.

But, when we looked at sequential infections between type A and a type A/B hybrid we did see that replication of the second virus was delayed.

Delayed, but not stopped altogether.

It would take a complete post to describe the figure above 🙁 . We’ve quantified the virus present 5-7 days after sequential injections with type A (VDD), type B (VVV) or a hybrid virus (VVD) 14.

The columns labelled VDD→VVV or VVV→VDD show the viruses and order of addition. The dots represent the amount of virus present at 5 or 7 days post injection. When the viruses were more similar to each other – for example, the VVV→VVD or VVD→VVV pairs on the right – there was a greater impact on the replication of the virus added second.

The same but different

We extended these studies to look at sequential infections with two viruses that only differed by 4 nucleotides (the building blocks) of the 10,140 nucleotides in the RNA genome of DWV i.e. 99.6% identical.

Cunningly, these four differences allowed us to unambiguously identify which virus was replicating.

In this part of the study the virus added second did not replicate to detectable levels. 

So … our data clearly demonstrates that viruses that were more similar to each other were more likely to inhibit replication during sequential infections.

In addition, no individual virus type was dominant over any others. 

This didn’t look much like classic superinfection exclusion to us.

Red or green viruses

Not content with generating graphs and tables we went on to take photographs of virus infected pupae. 

You can’t beat a nice colour image when trying to impress the peer reviewers 😉 .

Remember that DWV is too small to see with even the most powerful light microscope. You could fit several billion on the head of a pin.

We therefore engineered the virus genome to ‘show’ us where it was replicating.

Green bees

We did this by introducing an additional gene that fluoresced green or red when under UV light. I’ve discussed green viruses before … the red version uses similar technology, but using a different fluorescent reporter gene.

DWV replication (showed by green fluorescent signal) in the head, wing and abdomen of honey bee pupae

Using the red or green viruses we showed very similar results to those described above. When we superinfected with a genetically similar virus, its replication was inhibited. When it was genetically more divergent it could replicate (and we could visualise it as red or green foci of infection in a variety of tissues of the developing pupa).

Red and green viruses

We also infected bee with the red and green viruses simultaneously. Most of the fluorescent foci of infection were red or green, but a small number were both red and green. 

Green (EGFP) and red (mCherry) expressing DWV coinfecting a honey bee pupa. Arrow indicates dually infected cells.

The most likely explanation for having both colours overlapping in the photograph was because the virus were replicating in the same cells in the honey bee pupa.

Since this was exactly the sort of situation that was needed to generate recombinants (hybrids or chimeras) between the two different DWV viruses we specifically looked for them 15.

And there were lots and lots of recombinants …

Recombination between DWV viruses. The size and position of ‘bubbles’ indicate the location and number of junctions.

The bubble plot above shows the location and frequency of junctions. One virus is plotted on the horizontal and one on the vertical axis. It’s a sort of two-dimensional map of the virus. Think of a junction as where one virus ends and the other starts. They are located throughout the DWV genome – hundreds of them.

This suggests that pupae infected with both type A and type B DWV will act as ‘factories’ for the production of thousands more different hybrid variants between the two viruses.

Most of these hybrids will grow poorly.

Many will be uncompetitive.

But some – like the delta variant of SARS-Cov2 – might be more transmissible.

And some could be more pathogenic.

Or – the nightmare scenario – both 😯 .

What’s this got to do with practical beekeeping?

Every time a beekeeper moves bees about s/he is also moving viruses about.

This happens when you move bees to out apiary, when selling a nuc or when importing a queen.

Double brood ...

Moving viruses (and bees) to a new apiary …

This will contribute to the constant mixing of DWV variants that occurs when bees drift between hives, when drones mate with queens, when phoretic Varroa jumps onto a bee that is robbing a collapsing colony.

There’s a difference of course.

All those bee-driven mixing events are local and small scale … a few bees and a few miles.

But if you import a nuc from Greece via Northern Ireland both the distance and number of bees (and hence number of viruses) is much greater.

Of course, most of this mixing will just generate more mixtures of viruses.

It will also generate more recombinants.

But there’s always the possibility it might throw up a highly virulent, highly transmissible variant.

Which would not be a ‘good thing’.

And if it does, a ‘non-lethal type A strain’ (should such a thing actually exist) is not going to help prevent infection by the mechanism of superinfection exclusion I’m afraid 🙁 .

Without doubt the best way to prevent infection is to minimise the mite numbers in your colonies. This is a subject I’ll be tackling again in a couple of weeks.

But, before I go, do we understand how the more closely related strains of DWV prevent superinfection? 

Yes … probably, and it’s all to do with the immune response of the bee

I’ll discuss this in the future as it’s a mechanism that could be exploited to produce bees immune to the ravages of DWV.


 

Fainting goats … and queens

Myotonia congenita is a genetic disorder that affects the muscles used for movement. Myotonia refers to the delayed relaxation of these skeletal muscles, resulting in a variety of obvious symptoms including temporary paralysis, stiffness or transient weakness.

In humans these symptoms are often manifest as difficulty in swallowing, gagging and frequent falls. Children are affected more than adults. One of the most dramatic manifestations are the falls (‘fainting’) that can occur as a result of a hasty movement. 

Although physiologically distinct, ‘fainting’ is a reasonably accurate description of the sudden loss of movement and the transient nature of the disorder. Like fainting, loss of movement is usually quickly resolved. However, unlike fainting, myotonia congenita involves muscular rigidity or stiffness, so more closely resembles catalepsy.

Genes

There are two types of myotonia congenita, termed Thomsen disease and Becker disease, both of which are usually associated with mutations in the gene CLCN1 1. This encodes a chloride channel (a ‘hole’ through the cell membrane that allows the transfer of chloride ions) critical for muscle fibre activity. 

Cartoon of a transmembrane chloride channel.

With loss-of-function mutations in CLNC1 the muscle fibre continues to to be activated. When stimulated, for example if the fibre is triggered to suddenly contract for jumping or running (or  to stop a fall), the muscle fibre is hyper-excitable and continues to contract, and shows delayed relaxation

Around 1 in 100,000 people exhibit myotonia congenita, though it is about ten times more common in northern Scandinavia. Treatment involves use of a number of anticonvulsant drugs.

The same loss-of-function CLCN1 mutation in humans is seen in symptomatically similar horses, dogs … and goats.

Goats

In the late 19th century four goats were imported to Marshall County, Tennessee. Their strange behaviour when startled was first described in 1904 and defined as a congenital myotonia by Brown and Harvey in 1939. 

The eponymous Tennessee fainting goat

These pre-war studies formed the basis of of our understanding of both the physiology and genetics of myotonia congenita, though the specific mutation in the CLCN1 gene was only confirmed several years after it had been identified in humans.

Since then myotonic goats have become an internet staple, with any number of slightly distressing (for me at least, if not for the goats) YouTube videos showing their characteristic fainting when surprised or frightened 2.

Don’t bother watching them.

If you want to see a fainting goat in action watch little ‘Ricky’ jump up onto a swinging seat on the National Geographic website.

It’s a perfect example.

He jumps up, gets a mild fright as the swing moves, goes stiff legged and simply rolls over and falls to the ground. A few moments later he’s back on his feet again, looking slightly shaken perhaps, but none the worse for wear.

Queens

All of that preamble was to introduce the topic of fainting queens. 

A fainting queen

This was a subject I’d heard about, but had no experience of until last week.

Periodically it gets discussed on Beesource or the Beekeepingforum – usually the topic is raised by a relatively small-time amateur beekeeper (like me) and it gets a little airtime before someone like Michael Palmer, Michael Bush, Hivemaker or Into the Lion’s Den 3 shuts down the conversation with a polite “Yes, I see it a few times a year. They recover”, or words to that effect.

Since these commercial guys handle hundreds or perhaps thousands of queens a year I think we can safely assume it’s a relatively rare phenomenon. 

Since I don’t handle hundreds or thousands of queens a year – and you probably don’t either – I thought the incident was worth recounting, so you know what to expect should it ever happen.

And to do that I have to first explain the fun I had with the first of the two queens in the hive I was inspecting.

A two queen colony

It was late afternoon and I was inspecting the last of our research colonies in the bee shed.

The hive had two brood boxes and a couple of supers. Nothing particularly surprising in that setup at this time of the season; the colony was quite strong, the spring honey had been extracted and a couple of supers had been returned to the hive for cleaning.

However, it wasn’t quite that straightforward. 

The lower brood box had been requeened ~3 weeks earlier with a mature queen cell from one of my queen rearing attempts. I’d seen that the virgin had emerged and restricted her to the lower box at my last visit. 

I’d added a queen excluder (QE) over the lower box with the intention of removing all the old frames above the QE once the brood had emerged.

However, at that last visit I’d ended up with a good looking 4 ‘spare’ virgin queen. Although I had no need for her at the time, and no time to make up a nuc 5, I decided to put her in a fondant-plugged introduction cage in this upper box.

This ‘upper’ queen couldn’t fly and mate in the week I was away, but I reasoned that I could merge the colony with the bottom box if the ‘lower’ queen failed to mate 6.

So, after adding the virgin queen to the top box I added a second QE and the two supers.

She can fly …

Having removed the supers and the upper QE I carefully inspected the upper box looking for the virgin queen who had been released from the cage

No sign of her 🙁

I went through the box again.

Time to try some of the ‘queen finding tricks’.

I moved three frames out of the way having examined them very carefully. The remaining 8 frames were then spaced out as four, well separated, pairs. I let the colony settle for a few minutes and then looked at the inner face of each pair of frames.

No sign of her 🙁

I looked again … nada, rien, niets, nunda, dim byd and sod it 7.

The obvious conclusion was that the colony had killed the queen after releasing her from cage. 

How uncharitable.

I reassembled the upper brood box and lifted it off the lower QE, in preparation to leave it outside the shed door while I went through the lower box. 

As I carried the brood box to the door I briefly looked up and saw a 8 virgin queen climbing up the inner pane of one of the shed windows, flapping frantically and fast approaching the opening that would allow her escape.

For obvious reasons I have no photographs of the next few minutes.

Bee shed window ...

Bee shed window …

For those unfamiliar with the bee shed windows, these have overlapping outer and inner panes, so are always open. They provide a very effective ‘no moving parts’ solution to clearing the shed of bees very quickly.

Which was the very last thing I wanted at that moment 😉

… rather well

I had a brood box and hive tool in my hands, the shed door was wide open, there was all sorts of stuff littering the floor and the virgin queen was inches away from making a clean getaway.

It’s worth noting that when virgin queens are disturbed and fly they almost always return to the hive. However, the hives in the shed have a single entrance and all the hives were already occupied with queens. I couldn’t let her fly and hope for the best … it probably wouldn’t end well.

By balancing half the brood box on an unoccupied corner of an adjacent hive roof I made a largely ineffective swipe for the queen, but disturbed her enough she flew away from the window in spirals around my head.

I    s  t  r  e  t  c  h  e  d    to reach the shed door and pulled it close, so reducing the possible exits from eight to seven. A small victory.

I put the brood box safely on the floor, leaning at an angle against the hive stand 9, and abandoned the hive tool.

The next 5 minutes were spent ineptly trying to catch the queen. When she wasn’t flying around the shed (where the lighting isn’t the best) she usually made for the same window.

The one behind the hive with four supers stacked on top 🙁

After a few more laps of the shed, dancing around the precariously balanced brood box and reaching around the hive tower for the window, I finally caught her.

And caged her 10.

I’m looking for publisher for my latest book, ‘Slapstick beekeeping’. If any readers know of a publisher please ask them to contact me.

After all that I should have had a little rest. I’d had enough excitement for the afternoon 11.

But there was still the queen in the bottom box to find and mark.

Feeling faint

The queen in the bottom box was mated and laying well. 

I made a near-textbook example of finding her 12.

After moving aside a few frames I should have announced (to the non-existent audience), She’s on the other side of the next frame … ” (the big reveal) ” … ah ha! There you are my beauty!”.

Holding the frame in one hand I checked my pockets for my marking cage 13.

All present and correct.

I then calmly picked her up by her wings. She was walking towards me, bending slightly as she crossed over another bee, so her wings were pushed up and away from her abdomen.

A perfect ‘handle’.

I didn’t touch her abdomen, thorax or head.

A swooning queen

And, as soon as I lifted her from the frame, she fell into a swoon and ‘dropped dead’.

This is an ex-parrot

Her wings were extended to the sides, her abdomen was curled round in a foetal position and she appeared completely motionless.

It is pining for the fjords

I dropped her into the marking cage and took the photo further up the page.

It was 6:49 pm.

For several minutes there was no obvious movement at all. Her legs and antennae were immobile. She showed no sign of breathing.

I gently shook her out onto a small piece of Correx on a nuc roof to watch and photograph her. I picked her up by the wing and held her in my palm … perhaps she needed some warmth to ‘come round’.

Was that a twitch?

Or was that me shaking slightly because I’d inadvertently killed her? 

Several more minutes of complete catatonia 14 passed … and then a gentle abdominal pulsing started.

This was now 10-11 minutes after I’d first picked her up.

Which got a bit stronger and was accompanied by a feeble waggle of the antennae.

And was followed a minute or so later by a bit of uncoordinated leg flexing.

And after 15 minutes she took her first steps.

It looked like she’d been on an ‘all nighter’ and was still rather the worse for wear.

I slipped her into a JzBz queen cage, sealed it with a plastic cap, and left it hanging between a couple of brood frames.

From picking her up to placing the caged queen into the brood box had taken 24 minutes.

Caged queen after fainting (and recovering … more or less)

I reasoned that if …

  • she fully recovered they’d feed her through the cage and I could release her the following morning
  • I’d released her immediately and she’d acted abnormally the colony might have killed her off
  • she did not recover I would at least be able to find the corpse easily ( 🙁  ) and so could confidently requeen the colony (with the virgin I’d tucked away safely in my pocket)

The following morning the cage was covered in bees and she looked just fine, so I released her. 

Somewhere under that lot is the recovered queen – still caged

She walked straight down between the frames as though nothing untoward had happened.

I didn’t have the heart to mark and clip her … I didn’t want to risk her ‘fainting’ again and, if she had, didn’t have the time to hang around while she recovered 15.

So was this ‘fainting’ myotonia congenita?

I suspect not.

Another name for the Tennessee fainting goat is the ‘stiff-legged’ goat. This reflects the characteristic rigidity in the limbs when the muscles fail to relax. The queen’s legs were curled under her, rather than being splayed out rigidly.

However, this interpretation may simply reflect my near complete ignorance of the musculature of honey bees 😉

However, I do know that the basics of muscle contraction and relaxation are essentially the same in invertebrate and vertebrate skeletal muscle. There are differences in the innervation of muscle fibres, but the fundamental role of chloride channels in allowing muscle relaxation is similar.

Therefore, for this fainting queen to be affected by myotonia congenita she should have a mutation in the CLCN1 gene encoding the chloride channel.

Although the honey bee genome has been sequenced a direct homolog for CLCN1 appears not to have been identified, though there are plenty of other chloride channels present 16

The majority of the 60 or so mapped mutations associated with myotonia congenita (in humans) are recessive. Two copies of the mutated gene (in diploids, like humans or female honey bees) are needed for the phenotype to occur.

Of course, drones are haploid so it should be easier to detect the phenotype.

I’ve never heard of drones ‘fainting’ when beekeepers practise their queen marking skills on them. Have you?

Repeated fainting

I’ll try to mark and clip this queen again.

It will be interesting to see if she behaves in the same way 17.

A quick scour of the literature (or what passes for the ‘literature’ on weird beekeeping phenomena i.e. the discussion fora) failed to turn up examples of the same queen repeatedly fainting.

Or any mention of daughter queens showing the same behaviour.

All of which circumstantially argues against this being myotonia congenita.

However, there are many other causes of sudden fainting (from the NHS website):

  • standing up too quickly – (low blood pressure)
  • not eating or drinking enough
  • being too hot
  • being very upset, angry, or in severe pain
  • heart problems
  • taking drugs or drinking too much alcohol

… though I can exclude the last one as my bees are teetotal 😉

So, there you have it, a brief account of a cataleptic queen … and her recovery.


Notes

A fortnight after the events described above I clipped and marked the queen. I did everything the same – picked her up by the wings in the shed (so again not exposed to bright sunlight – which may be relevant, see the comment by Ann Chilcott).

She (the queen) didn’t faint. She behaved just like the remaining 4 queens I marked on the same afternoon.

So no repeat of the ‘amateur dramatics’ 🙂

Little dramas

This post was originally titled Drama queens.

Apposite … it’s mostly about queens.

However, the term drama queen refers to someone who overreacts to a minor setback 1 … which is almost the complete opposite of what I’m intending to discuss.

Instead, this post is about the – sometimes unseen – little dramas in the apiary. Things that go wrong, or could go wrong but eventually go OK because you gently intervene … or often because you don’t intervene at all 😉

It’s also about observing rather than doing. It’s sometimes surprising what you see, and – with a little application – you can learn something about your bees 2.

Of course, in the end some things do not end well … but there’s no point in being a drama queen about it 😉

Swarmtastic

There’s a certain predictability to the beekeeping year. It’s dictated by the climate and latitude, by the forage available, by the need for bees to reproduce (swarm) and by our efforts as beekeepers to corral them and keep them producing honey 3.

All of which means that June has been pretty manic. 

After a record-breakingly cold spring things finally warmed up. Here in Scotland this was 2-3 weeks into May.

Since then it’s been a near-constant round of queen rearing, swarm control, making up nucs and adding supers. Most of the OSR supers are now off, meaning that I’ll be hunched over the extractor for hours when I’m not with the bees 🙁

All the OSR near my bees is well and truly over – this lot is sadly just out of range

The rapid warming in late spring triggered a lot of swarming activity. I found my first charged queen cell on the 18th of May and, in at least one or two colonies, at every subsequent inspection since then.

Visits to the apiaries have been hard work. Inspecting a double brood colony with four full supers involves a lot of lifting 4.

And the lifting is necessary because I need to check whether there are any queen cells in the brood chamber.

I know some beekeepers simply prise the two brood boxes apart and expect to see queen cells at the junction.

That certainly works … sometimes.

However, I’ve found several colonies with queen cells in the middle of frames, or otherwise in positions I would not see them if I just looked at the interface between the boxes. 

Queen cell … and what else?

And I would still have to remove the supers to prise the brood boxes apart.

Although I’ve invested in some better quality hive tools, I’d need a crowbar to separate the boxes if there was 80 kg of supers on top 5.

So, if I have to take the supers off, I might as well look through the box carefully.

More haste, less speed

But before I fire up the smoker and start rushing around prising off crownboards I always try and simply observe what’s happening in the apiary.

Are all the colonies equally busy? If it’s the time of day when the new foragers are going on orientation flights are any colonies much less active? Have they had a brood break?

Which direction are the bees flying off or returning from? Has the main forage changed?

Are there any drones on orientation flights yet?

What’s happening at the hive entrances?

Is there pollen going in?

Any sign of fighting?

Or robbing?

It’s surprising what a few minutes observation can tell you about the local forage, the state of the colonies and their relative strength.

If you’ve not already read it (and even if you have) it’s worth finding a copy of At the Hive Entrance by Prof. H. Storch 6. The book’s strap-line is “How to know what happens inside the hive by observation on the outside”. Recommended.

And, now and again, you notice something unusual …

Queen under the open mesh floor

Like – in my peripheral vision – a single bee flying out from underneath an open mesh floor.

My queens are generally clipped. If the colony swarms the queen often finds her way back to the hive stand after crashing – very unregally 7 – to the ground. She crawls up the leg of the stand and ends up underneath the open mesh floor (OMF).

The bees then join her. It’s not unusual to find a large cluster of bees under the hive floor, with lots of activity, and lots of bees flying to and fro from underneath the OMF 8.

But last Friday, by chance I noticed a single bee and this prompted me to investigate.

A quick peek confirmed that there wasn’t a swarm under the OMF.

But there was a queen.

I spy with my little eye … you can just see the marked and clipped queen under this Abelo floor.

Almost completely alone.

I presume the colony had swarmed, the queen had got as far as she could and the swarm had eventually abandoned her and returned to the hive. 

When I inspected the colony I found a single sealed queen cell and confirmed that the queen I found was the one that was missing.

This colony was one of my ‘middle third’ ones 9i.e. destined for requeening with better stock if I had any spares.

There’s a near-to-eclosion queen cell under there …

I did.

I had half a dozen ‘spare’ queen cells almost ready to emerge from grafting at the start of June. I removed the queen cell in the hive and carefully checked I’d not missed any others. I then added the grafted cell, seating it in a thumb-sized depression over some brood. She will have emerged the following day and might even be mated when I check early next week.

Had I not seen the bee emerge from under the floor I’d have never otherwise checked. There are always a few bees under an OMF, but it’s rare to find a queen all alone there.

Queen in the grass

In another apiary the previous week I’d found a satsuma-sized cluster of bees in long grass about 10 metres from the hives. The application of a little gentle smoke and some prodding around with my index finger resulted in a clipped and marked queen calmly walking up onto my hand.

Microswarm? … or more likely the remains of a much larger one …

Again, I wouldn’t have seen this had I not been taking my time checking the hive entrances and the activity in the apiary. I was being even more leisurely than normal as there was rain threatening and I was trying to decide whether to start the inspections or not

Because of the known state of other colonies in the apiary – most were nucs with virgin or recently-mated queens – it was obvious which colony the queen had come from. 

The ‘threatening rain’ looked like it would soon become a certainty. I ran the queen in through the front entrance of the hive and the remaining bees eventually returned to the hive, fanning madly at the entrance.

Bees fanning at the entrance

When I next checked the hive the queen had gone 🙁

There was no sign the colony had swarmed, but there was a recently opened queen cell in there. I assumed there’s a newly emerged virgin queen running about in there with ‘blood on her hands’ having done away with the original queen.

We’ll find out next week.

Again, a few minutes just watching things in the apiary meant I found the queen. Had I not done so I’d have only seen the end result – a queenless colony – not the events that led to it.

Preventative and reactive swarm control

I should emphasise that the majority of my colonies are a little more under control than the two described above, both of which clearly attempted to swarm.

In both cases the clipped queen saved the day, even though she may not have lived to fight another day.

My swarm control (and success thereof) this season has been in stark contrast to last year’s ‘lockdown beekeeping’.

Then the priority was minimising travel and guaranteeing I wasn’t haemorrhaging swarms that might cause problems for the the public or other beekeepers.

I therefore used the nucleus method of swarm control on all my colonies. I implemented it well in advance of the peak swarming period. By doing so, I undoubtedly weakened my colonies. I produced less honey and did no queen rearing.

But I didn’t lose a single swarm 🙂

This year the priority has been to maintain strong colonies. Some are being used for honey production 10 and others are being split to make up nucs.

Inevitably a few have got a little ‘overcooked’ … but the clipped queen has usually ensured the bees remain in the hive.

I don’t think I’ve lost a swarm, but I have lost a few queens.

Queen in the cage

One of my colonies went queenless in mid May. This was well before I’d got any spare queens – mated or otherwise. I’d hoped that they would rear another, but it was too cold for the new queen to mate and the colony started to look a little pathetic.

I considered uniting them but, for a variety of reasons, never got round to it.

When I finally had a spare mated queen (in early June) I popped her into a JzBz introduction cage. I’d already plugged the tube with candy and placed a plastic cap over the end. 

The bees could feed the queen through the cage, but could not release her.

This is my usual method for queen introduction. I check the cage a day or so after hanging it between the frames. If the bees are showing aggression to the queen I leave it and check again 24 hours later.

Once they’re no longer showing any aggression I remove the plastic cap. The bees chew through the candy and release the queen.

Job done 🙂

I then leave the colony at least a week before inspecting, by which time I expect to see eggs and larvae.

JzBz queen introduction & shipping cage with removable plastic cap

On returning a week after removing the plastic cap I was dismayed to find the queen still in the cage. Most of the candy had gone, but there was a plug at one end that was rock hard. Clearly the bees had been unable to release her.

The colony had now been broodless almost a month. Brood pheromone is really important in suppressing laying worker activity in the hive. Queen pheromone is no substitute for brood pheromone 11 and I was very concerned about the additional lost week due to my stupidity 12.

But there was no point in being a drama queen … I opened the cage and gently released the queen onto a seam of bees. Five days later there are eggs and larvae (and the queen) in the hive, though I also think there are a few laying workers as there’s a smattering of drone pupae in worker cells (a classic sign).

Fingers crossed 🙂

Queen failure

The final ‘little drama’ was played out in full view over almost two months. Its eventual unsatisfactory conclusion was largely due to my procrastination … though I suspect a swallow or house martin may have hastened events at the end.

In late April, during one of the rare warm days it was possible to actually open a colony, I noticed some strange egg laying behaviour in one hive. 

The colony was queenright. The queen was marked and clipped and laying. However, although she was laying single eggs in worker comb, she was laying multiple eggs in about 10% of cells, almost all of which were in drone comb.

A fortnight or so later she was still doing the same thing. Even if it wasn’t obvious to me, it was clearly obvious to the bees that the queen was failing as they started a couple of queen cells. Here’s an enlargement of an earlier photo in this post – blue arrows mark single eggs, red arrows indicate multiples.

SIgns of a failing queen

I removed the queen and added a near-mature queen cell from my first round of grafting. She had emerged when I next checked, but was not yet laying (and I didn’t bother looking for her).

But, unlike the queen stuck in the cage, this didn’t have a happy ending.

By early June there was no sign of the queen and I fear she failed to return from a mating flight. There’s a big pond bear the apiary and it’s a magnet for swallows and house martins 13.

I added a frame of open brood (including both young larvae and eggs) in the hive, but they ignored it 14.

Frames showing the characteristic dispersed bullet brood of laying workers

When I next checked it was clear there were laying workers and I cut my losses and shook the colony out. 

In retrospect what should I have done? 

I should have united the colony in mid-May.

It was obvious then – at least to the bees – that the queen was failing. I’d never seen a queen laying singles in worker comb 15 but multiples in drone cells. 

Uniting would have immediately provided both brood pheromone and a laying queen. This would have suppressed the development of laying workers.

My notes go something like:

  • 18/5 – Still laying singles in worker and multiples in drone. Weird. QC looks like supersedure. Give them a week.
  • 26/5 – Q out. Didn’t check further. Decision time next week.
  • 3/6 – Strange. Increasing drone brood. Behaving queenright. Decision time next week.
  • 12/6 – Laying workers. Shook them out. Will I ever learn? EEJIT 16

The second rule of beekeeping

Anytime I write Decision time next week (or variants thereof, like Give them another weekin two successive weeks then it’s almost always going to end in tears 🙁

If it happens three times in succession it’s a nailed on certainty.

The first rule is – of course – Knocking off queen cells is not swarm control 😉


 

More from the fun guy

Great fleas have little fleas upon their backs to bite ’em,
And little fleas have lesser fleas, and so ad infinitum.

Augustus de Morgan’s quote from A Budget of Paradoxes (1872) 1 really means that everything is preyed upon by something, which in turn has something preying on it.

The Flea, engraving from Robert Hooke’s Micrographia (1665)

As a virologist I’m well aware of this.

There are viruses that parasitise every living thing.

Whales have viruses and so do unicellular diatoms. All the ~30,000 named bacteria have viruses. It’s likely that the remaining 95% of bacteria that are unnamed also have viruses.

There are even viruses that parasitise viruses. The huge Mimivirus that infects amoebae 2 is itself parasitised by a small virophage (a fancy name for a virus that infects viruses) termed Sputnik.

Whether these interactions are detrimental depends upon your perspective.

The host may suffer deleterious effects while the parasite flourishes.

It’s good for the latter, but not the former.

Whether these interactions are detrimental for humans 3 also depends upon your perspective.

The deliberate introduction of rabbit haemorrhagic disease virus to Australia benefitted sheep farmers who were plagued with rabbits … but it was bad news for rabbit farmers 4.

Biocontrol

Beneficial parasitism, particularly when humans use a pathogen to control an unwanted pest, is often termed biocontrol, a convenient abbreviation for biological pest control.

There are numerous examples; one of the first and best known is control of greenhouse whitefly infestations with the parasitoid wasp Encarsia formosa.

Tomato leaf with whitefly nymphs (white) parasitized by E. formosa (black).

One of the benefits of biocontrol is its self-limiting nature. The wasp will stop replicating once it runs out of whitefly to parasitise.

A second benefit is the specificity of the interaction between the host and whatever is administered to control it; by careful selection of the biocontrol agent you can target what you want to eradicate without lots of collateral damage.

Finally, unlike toxic chemicals such as DDT, the parasitoid wasp – and, more generally, other biocontrol agents – do not accumulate in the environment and cause problems for the future.

And, with all those benefits, it’s unsurprising to discover that scientists have investigated biocontrol strategies to reduce Varroa mite infestation of honey bee colonies.

It’s too early for an aside, but I’ll make one anyway … I’ve discussed the potential antiviral activity of certain fungi a couple of years ago. That wasn’t really biocontrol. It was a fungal extract that appeared to show some activity against the virus. Although that story has gone a bit quiet, one of the authors – Paul Stamets – is also a co-author of the Varroa control paper discussed below.

Biocontrol of Varroa using entomopathogenic fungi

Entomopathogenic means insect killing 5. There are several studies on the use of insect killing fungi to control Varroa 6, with the most promising results obtained with a variety of species belonging to the genus Metarhizium

Metarhizium produces asexual spores termed mitospores. The miticidal activity is due to the adhesion of these mitospores to Varroa, germination of the spore and penetration by fungal hyphae 7 through the exoskeleton of the mite and proliferation within the internal tissues.

A gruesome end no doubt.

And thoroughly deserved 🙂

Although Metarhizium is entomopathogenic it has a much greater impact on Varroa than it does on honey bees. This is the specificity issue discussed earlier.

It is for this reason that scientists have continued to explore ways in which Metarhizium could be used for biocontrol of Varroa.

But there’s a problem …

Although dozens of strains of Metarhizium have been screened, the viability – and therefore activity – of the mitospores is significantly reduced by the relatively high temperatures within the colony.

The spores would be administered, they’d show some activity and some Varroa would be slaughtered. However, over time treatment efficacy would reduce as spores – either administered at the start of the study, or resulting from subsequent replication and sporulation of Metarhizium on Varroa – were inactivated.

As beekeepers you’ll be familiar with the limitation this would impose on effective control of mites.

Varroa spend well over half of their life cycle capped in a cell while it feeds on developing pupae. Anything added to kill mites must be present for extended periods to ensure emerging mites are also exposed and killed.

This is why Apiguard involves two sequential treatments of a fortnight each, or why Apivar strips must be left in a hive for more than 6 weeks.

In an attempt to overcome these limitations, scientists are using directed evolution and repetitive selection to derive strains of Metarhizium that are better able to survive within the hive, and so better able to control Varroa than the strains they were derived from.

Good news and bad news

Like many scientific papers on honey bees 8 those with even a whiff of ‘saving the bees’ get a lot of positive press coverage.

This often implies that the Varroa ‘problem’ is now almost solved, that whatever tiny, incremental advance is described in the paper represents a new paradigm in bee health.

This is both understandable and disappointing in equal measure.

It’s understandable because people (not just beekeepers) like bees. News publishers want ‘good news’ stories to intersperse with the usual never-ending menu of woe they serve up.

It’s disappointing because it’s a variant of “crying wolf”. We want the good news story to describe how the impact of Varroa can now be easily mitigated.

It gets our hopes up.

Unfortunately, reality suggests most of these ‘magic bullets’ are a decade away from any sort of commercial product.

They will probably get mired in licensing problems.

And they may not be any better than what we currently use.

You finally end up as cynical as I am. This might even force you to read the original manuscript, rather than the Gung ho press release or the same thing regurgitated on a news website.

And, if you do that, you’ll better understand some of the clever approaches that scientists are applying to the development of effective biocontrol for Varroa.

We’re not there yet, but progress is being made.

V e r y   s l o w l y.

The paper I’m going to discuss below is Han, J.O., et al. (2021) Directed evolution of Metarhizium fungus improves its biocontrol efficacy against Varroa mites in honey bee colonies. Sci Rep 11, 10582.

It’s freely available should you want to read the bits I get wrong 😉

Solving the temperature-sensitivity problem of mitospores

The strain of Metarhizium chosen for these studies was M. brunneum F52. This had previously been demonstrated to have some efficacy against Varroa. Almost as important, it can be genetically manipulated and there was some preliminary evidence that its pathogenicity for Varroa – and hence control potential – could be improved.

Genetic manipulation covers a multitude of sins. It could mean anything from selection of pre-existing variants from a population to engineered introduction of a toxin gene for destruction of the parasitised host.

In this study the authors used directed evolution of a population of Metarhizium to select for strains with more heat tolerant spores.

Directed evolution of Metarhizium to select mitospores with increased thermotolerance

This is not genetic engineering. They grew spores under stressful conditions and increasing temperatures. Hydrogen peroxide (H2O2) , a mild mutagen, was added in some cases. Nutritional stress also increases population variation. Spores selected using nutritional stress are better able to withstand UV and heat stress.

The optimal growth temperature for the strain of Metarhizium they started with was 27°C. By repeated selection cycles at increasing temperatures they derived spores that grew at 35°C, the temperature within a colony.

Ladders and snakes

A well known phenomena of repeated selection in vitro (i.e. in a test tube in the laboratory, though you actually grow Metarhizium on agar plates) is that a pathogen becomes less pathogenic.

It was therefore unsurprising that – when they eventually tested the thermotolerant spores – only about 3% of the Varroa that died did so due to Metarhizium infection.

Field selection after directed evolution of Metarhizium in the laboratory

They therefore modified the repetitive selection, but this time did it on Varroa-infested colonies in the apiary. Mites that died from Metarhizium mycoses 9 were used as a source to cultivate more Metarhizium.

They were therefore selecting for both thermotolerant (because the experiments were being conducted in hives at 35°C) and pathogenic fungi, because they only cultivated mitospores from Varroa that had died from mycoses.

And it worked …

Amplification of Varroa mycoses by Metarhizium. Black arrows indicate the treatment dates.

After four rounds of selection over 60% of the mites that died did so because they were infected with Metarhizium.

All very encouraging … but note I was very careful with my choice of words in that last sentence. I’ll return to this point shortly.

Before that, here’s the ‘proof’ that the strain selected by directed evolution (which they termed JH1078) possessed more thermostable spores.

Thermostable spores

They measured this by recording the percentage that germinated. At 35°C ~70% of JH11078 spores germinated compared to only ~45% of the M. brunneum F52 strain they started with.

But it’s not all good news

My carefully chosen “60% of the mites that died” neatly obscures the fact that you could get a significant increase in mites dying of Metarhizium, but still have almost all the mites in the hive surviving unscathed.

The authors continued repeated Metarhizium monthly treatments for a full season after the selection experiments described above. The apiary contained 48 colonies, 24 received Metarhizium JH1078 and the remainder received no treatment.

Did Metarhizium treatment stop the well documented increase in Varroa levels observed in colonies not treated with miticides?

Varroa levels in Metarhizium treated and untreated (control) colonies.

Er … no.

They describe this data (above) as showing a ‘delay’ in the exponential increase in Varroa … but acknowledge that it ‘did not totally prevent it’.

Hmmm … looking at the error bars in the last few timepoints I’d be hard pressed to make the case that there was any significant difference in Varroa increase caused by treatment.

And while we’re here look at the mite infestation rate … 10-25 mites per 100 bees.

These are catastrophically high numbers and, unsurprisingly, 42 (~88%) of the 48 colonies – whether treated or untreated – died by the end of 2018, succumbing to “Varroa, pathogen pressure and intense yellow jacket predation”

There was some evidence that colonies receiving Metarhizium treatment survived a bit longer than the untreated controls, but the end results were the same.

Almost every colony perished.

Metarhizium vs. oxalic acid

Typically a paper on a potential improved biocontrol method for Varroa would do a side-by-side comparison with a widely used, currently licensed treatment.

There’s only one comparative experiment between Metarhizium and dribbled oxalic acid treatment. It’s buried at the end of the Supplementary Data 10. In it they show ‘no significant difference’ between the two treatments.

Frankly this was a pretty meaningless experiment … it was conducted in June 2020 when colonies would have been bulging with brood. Consequently 90% of the mites would have been hidden under the cappings. They assayed mite levels only 18 days after a single application of Metarhizium or oxalic acid.

Although it showed ‘no significant difference’ – like the “60% of the mites that died” quote – it obscures the fact that most mites were almost certainly completely untouched by either treatment.

What does this study show?

This study involved a large amount of work.

The directed evolution in the laboratory is a very nice example of how the combination of phenotypic selection and natural variation can rapidly yield new strains with desirable characteristics.

Combination of this with in vivo selection for enhanced pathogenesis successfully produced a novel strain of Metarhizium with some of the features desirable for biocontrol of Varroa.

However, in the apiary-based studies the majority of the colonies, whether treated or not, died.

This shows that, although scientists might have made a promising start, they are still a very long way from having an effective biocontrol solution for Varroa.

Unmanaged Varroa replicates to unmanageable levels

One of the Supplementary Data figures illustrated the Varroa drop per month from colonies in the research apiary.

Cumulative mite drop per colony for the month of July 2018

This is relatively late in the study, July 2018. These hives were established from commercial packages of bees in April 2017. They were either treated with the experimental Metarhizium spores or were untreated controls for the 15 months between April 2017 and July 2018 11.

Look at those Varroa numbers!

This is the mite drop just after the peak of the season. Brood levels would be close to maximum in their short, warm summer 12. The majority of the mite population would have been safely tucked away feasting on developing brood.

This is not the mite drop after miticide treatment … it’s just the drop due to bee grooming, natural mite mortality and the general ‘friction’ in the hive.

The average is 2866 mites dropped per month per hive 😥

Maybe nothing could have saved hives as heavily infested as these? 13

Don’t wait for Metarhizium … be vigilant now

These numbers – of mites and dead colonies – are a stark warning of the replication potential of Varroa and the damage is causes our bees.

Left untreated, Varroa will replicate to very high levels.

Colony mortality – either directly due to the mite and viruses, or indirectly due to the weakened colonies succumbing to robbing – is a near-inevitable consequence.

I’ve discussed the importance of Varroa management repeatedly over the years. It’s a topic I’ll be returning to again – probably in August when it starts to become a necessity.

In the meantime, keep an eye on the mite levels in your own colonies as they get stronger during the season.

While you’re doing that think of the scientists who are looking for practical, effective and environmentally-friendly strategies to control Varroa. Understand that these studies are time-consuming, progress is glacial incremental … and they might not work anyway.

Of course, if we finally manage to develop a suitable Metarhizium-based mite control strategy then bees and beekeepers will not be the only beneficiaries.

Metarhizium has its own parasites. Some of the best characterised of these are small RNA viruses.

If beekeepers are sprinkling billions of Metarhizium spores over their colonies every year then these viruses will be having a great time 😉


 

It’s a drone’s life

What has a mother but no father, but has both a grandmother and grandfather?

If you’ve not seen this question before you’ve not attended a ‘mead and mince pies’ Christmas quiz at a beekeeping association. 

Drone

Drone … what big eyes you have …

The answer of course is a drone. The male honey bee. Drones are produced from unfertilised eggs laid by the queen, so formally they have no father. Drones are usually haploid (one set of chromosomes), whereas queens and workers are diploid 1

Anyway, enough quiz questions. With the relaxation in Covid restrictions we may all be able to attend in person this Christmas 2, so I don’t want to spoil it by giving all the answers away in advance.

The long cold spring has been pretty tough for new beekeepers, it’s been a struggle for smaller colonies and it’s been really hard for drones.

Spring struggles

New beekeepers have had to develop the patience of Job to either acquire bees in the first place or start their inspections. Inevitably new beekeepers are bursting with enthusiasm 3 and the cold northerlies, unseasonal snow (!) and low temperatures have prevented inspections and delayed colony development (and hence the availability and sale of nucs).

Small colonies 4 are struggling to rear brood and to collect sufficient nectar and pollen.

This is an interesting topic in its own right and deserves a post of its own 5. In a nutshell, below a certain threshold of bees, colonies are unable to keep the brood warm enough and have sufficient foragers to collect nectar and pollen.

As a consequence, smaller colonies are low on stores and at risk of starvation. 

It’s a Catch-22 situation … to rear sufficient brood to collect an excess of nectar (or pollen) the colony needs more adult workers. 

I don’t know what the cutoff is in terms of adult bees, but most of my colonies with <7 frames of brood have needed feeding this spring.

One feature of these smaller colonies is that, unless they have entire frames of drone comb 6, there is little if any drone brood in the hive.

There might be drones present in the colony, but I don’t know whether they were reared there or drifted there from another hive.

And, for those of us attempting to rear queens, drones are an essential indicator that queen mating will be timely and successful.

On a brighter note …

But it’s not all gloom and doom.

Strong colonies are doing very well.

Several of mine have a box packed full of brood and I’m relying on a combination of …

  • lots of space by giving them more supers than they need
  • low ambient temperatures
  • crossed fingers

… as my swarm prevention strategy 😉

Beginners take note … one of these is likely to help (space), one is frankly pretty risky (chilly) and the last is not a proven method despite being widely used by many beekeepers 😉

I’m pretty confident that colonies will not swarm at 13-14°C.

I am inspecting colonies every 7 days and have only seen two with charged queen cells. One was making early swarm preparations; I used the nucleus method of swarm control and then split the colony into nucs a fortnight ago 7.

The other colony contained my first attempt at grafting this year, which seems to have gone reasonably well 8.

Lots of brood, nectar and drones

A typical brood frame from one of these strong colonies contains a good slab of sealed or open brood, some pollen around the sides and an interrupted arc of fresh nectar above the brood. 

In the photo above you can see pollen on the right hand side of the frame and glistening fresh nectar in the top left and right hand corners.

Typically these strong colonies also have partially filled supers, though it’s pretty clear that the oil seed rape is likely to go over before the weather warms enough (or the colonies get strong enough) to fully exploit it.

Spring honey is going to be in short supply and my fantastic new honey creamer is going to sit idle 🙁

Drones

What you probably can’t really see in the picture above is that these strong colonies also contain good numbers of drones.

Strong colonies … ample drones

I can count about a dozen in the closeup above. 

I like seeing drones in a strong, healthy colony early(ish) in the season 9.

Firstly, the presence of drones indicates that the colony (and presumably others in the neighbourhood which are experiencing a similar environment and climate) will soon be making swarm preparations. This means I need to redouble my efforts to check for queen cells to avoid losing swarms 🙁  … think of it as a long-range early warning system.

But it also means I can start thinking about queen rearing 🙂

Secondly, although these drones are unlikely to mate with my queens, you can be sure they’re going to have a damned good go at mating with queens from other local apiaries.

In addition to being strong and healthy, this colony is well-tempered, steady on the comb and pleasant to work with. The production of a few hundred thousand frisky drones prepared to lay down their lives 10 to improve the local gene pool is my small act of generosity to local beekeepers 11.

How many drones?

Honey bee colonies that nest in trees or other natural cavities produce lots of drone comb. Studies of feral colonies on natural comb show that about 17% of the comb is dedicated to rearing drones (but also used for storing nectar at other times of the season).

Foundationless triptych ...

Foundationless triptych …

Similarly, beekeepers who predominantly use foundationless frames regularly see significantly greater amounts of drone comb (and drone brood and drones) in their colonies. With the three-panel bamboo-supported frames I use it’s not unusual for one third of some frames to be entirely drone comb.

In contrast, beekeepers who only use standard worker foundation will be used to seeing drone comb occupying much less of the brood nest. Under these circumstances it’s usually restricted to the edges or corners of frames.

However, given the opportunity e.g. a damaged patch of worker comb or if you add a super frame into the brood box, the workers will often rework the comb (or build new brace comb) containing just drone cells.

The bees only build drone comb when they need it.

A newly hived swarm will build sheet after sheet of new comb, but it will all be for rearing worker brood. If you give them foundationless frames they only build worker comb and if you provide worker foundation they don’t rework it to squeeze in a few drone cells.

The colony will also not build new drone comb late in the season. Drone comb is drawn early in the season because the drones are needed before queens are produced.

The timing of drone production

Studies in the late 1970’s 12 demonstrated that drone brood production peaks about one month before the the main period of swarming. Similar studies in other areas have produced similar results.

Why produce all those drones when there are no queens about?

The timing is due to the differences in the development time (from egg to eclosion) of drones and queens, together with the differences in the time it takes before they are sexually mature.

Drones take 50% longer to develop than queens – 24 days vs. 16 days. After emergence the queen take a few days (usually quoted as 5-6) to reach sexual maturity before she embarks on her mating flight(s).

In contrast, drones take from 6-16 days to reach sexual maturity.

Swarming tends to occur when charged queen cells in the hive are capped. These cells will produce new virgin queens about a week later and – weather permitting – these should go on mating flights after a further six days. 

Therefore a colony that swarms in very early June will need sexually mature drones available 12-14 days later (say, mid-June) to mate with the newly emerged queen that will subsequently return to head the swarmed colony. These drones will have to have hatched from eggs laid in the first fortnight of May to ensure that they are sexually mature at the right time.

Decisions, decisions

How does the colony know to produce drones at the right time? Is it the workers or the queen who makes this decision?

I’ve recently answered a question on this topic for the Q&A pages in the BBKA Newsletter. In doing some follow-up reading I’ve discovered that (inevitably) it’s slightly more complicated than I thought … which was already pretty complicated 🙁

The workers build the comb and therefore determine the amount of drone vs. worker comb the brood nest contains.

I don’t think it’s known how the workers measure the amount of brood comb in the nest, but they clearly can. We do know that bees can count 13 and that they have some basic mathematical skills like addition and subtraction.

Perhaps these maths skills 14 include some sort of averaging, allowing them to sample empty cells, measure them and so work out the proportion that are drone or worker.

Whatever form this ‘counting’ takes, it requires direct contact of the bees with the comb. You cannot put a few frames of drone comb in the hive behind a mesh screen and stop the bees from building more drone comb. It’s not a volatile signal that permeates the hive.

However they achieve this, they are also influenced by the amount of capped drone brood already present in the colony. If there’s lots already then the building of additional drone comb is inhibited 15.

Colonies therefore regulate drone production through a negative feedback process.

So … does the queen simply lay every cell she comes across, trusting the worker population has provided the correct proportions of drone and worker comb?

Not quite

Studies by Katie Wharton and colleagues 16 showed that the queen could also regulate drone production.

Wharton confined queens on 100% drone or worker comb in a frame-sized queen ‘cage’ for a few days.

Frame sized queen ‘cage’ …

She then replaced the comb in the cage with 50:50 mix of drone and worker comb and recorded the number of eggs laid in drone or worker cells over a 24 hour period (and then allowed the eggs to develop).

Queens that had only been able to lay worker brood for the first four days of confinement laid significantly more drone brood when given the opportunity.

The scientists showed reasonably convincingly that this was a ‘decision’ made by the queen, rather than influenced by the workers e.g. by preparing biased number of drone or worker cells for eggs to be laid in, by preferentially ‘blocking’ certain cell types with honey or by selectively cannibalising drone or worker eggs.

Interestingly, queens initially confined on worker comb laid significantly (~25%) more eggs on the 50:50 comb than those confined on drone comb. I’m not sure why this is 17.

Wharton and colleagues conclude “these results suggest that the regulation of drone brood production at the colony level may emerge at least in part by a negative feedback process of drone egg production by the queen”.  

So it seems likely that drone production in a colony reflects active decisions made by both workers and the queen.

Why has this spring been really hard for drones?

To be ready for swarming, colonies therefore need to start drone production quite early in the season – at least 4-5 weeks before any swarms are likely.

Late May ’21 forecast. Swarmy weather? I don’t think so …

But with consistently poor weather, these drones are unlikely to be needed. Colonies will not have built up enough to be strong enough to swarm.

Producing drones is a high energy process – they are big bees and require a lot of carbohydrate and protein during development.

Under natural conditions 18 a colony puts as many resources into drone production over the season as it does into swarms.

Thomas Seeley has a nice explanation of this in The Lives of Bees – if you take the dry weight of primary swarms and casts produced by a colony it’s about the same as the dry weight of drones produced throughout the season. 

Rather than waste energy in drone production the workers remove unwanted drone eggs and larvae. The queen lays them, but the workers prevent them being reared.

How do the workers decide the drones aren’t going to be needed?

Do workers have excellent long-range weather forecasting abilities?

Probably not 19

If the weather is poor the colony will be unable to build up properly because forage will be limited. As a consequence, the colony (and others in the area) would be unlikely to swarm and so drones would not be needed for queen mating.

Free and Williams (1975) demonstrated that forage availability was the factor that determined whether drones were reared and maintained in the colony. 

Under conditions where forage was limited, drone eggs and larvae were rejected (cannibalised) and adult drones were ejected from the hive.

Unwanted drone ejected from a colony in early May

Beekeepers are familiar with drones being ejected from colonies in the autumn (again, a time when forage becomes limiting), but it also happens in Spring.

And at other times when nectar is in short supply …

Those of you currently enjoying a good nectar flow from the OSR should also look at colonies during the ‘June gap’. With a precipitous drop in nectar available in the environment once the OSR stops yielding, colonies can be forced to eject drones.

It’s tough being a drone … which may explain why one of my PhD students has the name @doomeddrone on Twitter 😉


 

Acting on Impulse

Men just can’t help acting on Impulse … 

This was the advertising strapline that accompanied the 1982 introduction of a new ‘body mist’ perfume by Fabergé. It was accompanied by a rather cheesy 1 set of TV commercials with surprised looking (presumably fragrant) women being accosted by strange men proffering bouquets of flowers 2.

Men just can’t help acting on Impulse …

And, it turns out that women – or, more specifically, female worker honey bees – also act on impulse

In this case, these are the ‘impulses’ that result in the production of queen cells in the colony.

Understanding these impulses, and how they can be exploited for queen rearing or colony expansion (or, conversely, colony control), is a very important component of beekeeping.

The definition of the word impulse is an ‘incitement or stimulus to action’.

The action, as far as our bees are concerned, is the development of queen cells in the colony.

If we understand what factors stimulate the production of queen cells we can either mitigate those factors – so reducing the impulse and delaying queen cell production (and if you’re thinking ‘swarm prevention‘ here you’re on the right lines) – or exploit them to induce the production of queen cells for requeening or making increase.

But first, what are the impulses?

There are three impulses that result in the production of queen cells – supersedure, swarm and emergency.

Under natural conditions i.e. without pesky meddling by beekeepers, colonies usually produce queen cells under the supersedure or swarm impulse.

The three impulses are:

  1. supersedure – in which the colony rears a new queen to eventually replace the current queen in situ
  2. swarm – during colony reproduction (swarming) a number of queen cells are produced. In due course the current queen leaves heading a prime swarm. Eventually a newly emerged virgin queen remains to get mated and head the original colony. In between these events a number of swarms may also leave headed by virgin queens (so-called afterswarms or casts).
  3. emergency – if the queen is lost or damaged and the colony rendered queenless, the colony rears new queens under the emergency impulse.

Many beekeepers, and several books, state that you can determine the type of impulse that induced queen cell production by the number, appearance and location of the queen cells.

And, if you can do this, you’ll know what to do with the colony simply by judging the queen cells.

If only it were that simple

Wouldn’t it be easy?

One or two queen cells in the middle of frame in the centre of the brood nest? Definitely supersedure. Leave the colony alone and the old queen will be gently replaced over the next few weeks. Brood production will continue uninterrupted and the colony will stay together and remain productive.

A dozen or more sealed queen cells along the bottom edge of a frame? The colony is definitely  in swarm mode and – since the cells are already capped – has actually already swarmed. Time to thin out the cells and leave just one to ensure no casts are also lost.

But it isn’t that simple 🙁

Bees haven’t read the textbooks so don’t necessarily behave as expected.

I’ve found single open queen cells in the middle of a central frame, assumed it was supersedure, left the colony alone and lost a swarm from the hive a few days later 🙁

D’oh!

Or I’ve found loads of capped queen cells on the edges of multiple frames in a hive, assumed that I’d missed a swarm … only to subsequently find the original marked queen calmly laying eggs as I split the brood box up to make several nucleus colonies  🙂

Not all queen cells are ‘born’ equal

It’s worth considering what queen cells are … and what they are not. And how queen cells are started.

There are essentially two ways in which queen cells are started.

They are either built from the outset as vertically oriented cells into which the queen lays an egg, or they start their life as horizontally oriented 3 worker cells which, should the need arise, are re-engineered to face vertically.

Play cup or queen cell?

Play cup or are they planning their escape …?

Queen cells started under the supersedure or swarming impulse are initially created as ‘play cups‘. A play cup looks like a small wax version of an acorn cup – the woody cup-like structure that holds the acorn nut. In the picture above the play cup is located on the lower edge of a brood frame, but they are also often found ‘centre stage‘ in the middle of the frame.

Play cups

A colony will often produce many play cups and their presence is nothing to be concerned about. In fact, I think it’s often a rather encouraging sign that the colony is sufficiently strong and healthy that it might be thinking of raising a new queen. 

Before we leave play cups and consider how emergency queen cells start life it’s worth emphasising the differences between play cups and queen cells.

Play cups are not the same as queen cells

Until a play cup is occupied by an egg it is not a queen cell.

At least it’s not as far as I’m concerned 😉

And, even if it contains an egg there’s no guarantee it will be supported by the workers to develop into a new queen 4.

However, once the cell contains a larva and it is being fed by the nurse bees – evidenced by the larva sitting in an increasingly thick bed of royal jelly – then it is indisputably a queen cell.

Charged queen cell ...

Charged queen cell …

And to emphasise the fundamental importance in terms of colony management I usually refer to this type of queen cell as a ‘charged queen cell’.

Once charged queen cells appear in the colony, all other things being equal, they will be maintained by the workers, capped and – on the 16th day after the egg was laid – will emerge as a new queen.

And it is once charged queen cells are found in the colony that swarm control should be considered 5.

But let’s complete our description of the queen cells by considering those that are produced in response to the emergency impulse.

Emergency queen cells

Queen cells produced under the emergency impulse differ from those made under the swarm or supersedure impulse. These are the cells that are produced when the colony is – for whatever reason – suddenly made queenless. 

Without hamfisted beekeeping it’s difficult to imagine or contrive a scenario under which this would occur naturally 6, but let’s not worry about that for the moment 7

The point is that, should a colony become queenless, the workers in the colony can select one or more young larvae already present in worker cells and rear them as new queens.

So, although the eggs are (obviously!) laid by the queen 8, they have been laid in a normal worker cell. To ensure that they get lavished with attention by the nurse bees, feeding them a diet enriched in Royal Jelly, the cell must be re-engineered to project vertically downwards.

Location, location

Queen cells can occur anywhere in the hive to which the queen has access.

Queen cell on excluder

Queen cell on underside of the excluder …

But they are most usually found on the periphery of the frame, either along the lower edge …

Queen cells ...

Queen cells …

… or a vertical side edge of the frame …

Sealed queen cells

… but they can also be found slap, bang in the middle of a brood frame.

Single queen cell in the centre of a frame

And remember that bees have a remarkable ability to hide queen cells in inaccessible nooks and crannies on the frame … and that finding any queen cells is much more difficult when the frame is covered with a wriggling mass of worker bees.

Location and impulses

Does the location tell us anything about the impulse under which the bees generated the queen cell?

Probably not, or at least not reliably enough that additional checks aren’t also needed 🙁

Many descriptions will state that a small number (typically 1-3) of queen cells occupying the centre of a frame are probably supersedure cells. 

Whilst this is undoubtedly sometimes or even often true, it is not invariably the case.

The workers choose which larvae to rear as queens under the emergency impulse. If the only larvae of a suitable age are situated mid-frame then those are the ones they will choose.

In addition, since generating emergency cells requires re-engineering worker cells, newer comb is likely more easily manipulated by the workers.

Some beekeepers ‘notch’ comb under suitably aged larvae to induce queen cell production at particular sites on the frame 9. The photograph shows a frame of eggs with a notch created with the hive tool. It’s better to place the notch underneath suitably aged larvae, not eggs. Clearly, the age of the larvae is more critical than the ease with which the comb can be reworked. Those who use this method [PDF] properly/extensively claim up to a 70% ‘success’ rate in inducing queen cell placement on the frame. This can be very useful if the plan is to cut the – well separated – queen cells out and use them in mating nucs or for requeening other colonies.

Eggs in new comb ...

Eggs in new comb …

Comb at the bottom or side edges of the frame often has space adjacent and underneath it. Therefore the bees might favour these over sites mid-frame (assuming ample suitable aged larvae) simply because the comb is easier to re-work in these locations.

And don’t forget … under the emergency impulse the colony preferentially chooses the rarest patrilines to rear as new queens 10.

Not all larvae are equal, at least when rearing queens under an emergency impulse.

Active queen rearing and the three impulses

By ‘active’ queen rearing I mean one of the hundreds of methods in which the beekeeper is actively involved in selecting the larvae from which a batch of new queens are reared.

This doesn’t necessarily mean grafting , towering cell builders and serried rows of Apidea mini nucs.

It could be as simple as taking a queen out of a good colony to create a small nuc and then letting the original colony generate a number of queen cells.

Almost all queen rearing methods use either the emergency or supersedure impulses to induce new queen cell production 11.

For example, let’s consider the situation described above.

Active queen rearing and the emergency impulse

A strong colony with desirable traits (calm, productive, prolific … choose any three 😉 ) is made queenless by removing the queen on a frame of emerging brood into a 5 frame nucleus hive. With a frame of stores and a little TLC 12 the queen will continue to lay and the nuc colony will expand.

Everynuc

Everynuc …

But the, now queenless, hive will – under the emergency impulse – generate a number of new queen cells. These will probably be distributed on several frames if the queen was laying well before she was removed.

The colony will select larvae less than ~36 hours old (i.e. less than 5 days since the egg was laid) for feeding up as new queens.

If the beekeeper returns to the hive 8-9 days later it can be split into several 5 frame nucs, each containing a suitable queen cell and sufficient emerging and adherent bees to maintain the newly created nucleus colony 13.

Active queen rearing and the supersedure impulse

In contrast, queenright queen rearing methods such as the Ben Harden system exploit the supersedure impulse.

Queen rearing using the Ben Harden system

In this method suitably aged larvae are offered to the colony above the queen excluder. With reduced levels of queen pheromones present – due to the physical distance and the fact that queen cannot leave a trail of her footprint pheromone across the combs above the QE – the larvae are consequently raised under the supersedure impulse.

Capped queen cells

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

I’m always (pleasantly) surprised this works so well. Queen cells can be produced just a few inches away from a brood box containing a laying queen, with the workers able to move freely through the queen excluder. 

Combining impulses …

Finally, methods that use Cloake or Morris boards 14 use a combination of the emergency and supersedure impulses.

Cloake board ...

Cloake board …

In these methods the colony is rendered transiently queenless to start new queen cells. About 24 hours later the queenright status is restored so that cells are ‘finished’ under the supersedure response.

The odd one out, as it’s not really practical to use it for active queen rearing, is the swarming impulse. Presumably this is because the conditions used to induce swarming are inevitably rather difficult to control. Active queen rearing is all about control. You generally want to determine the source of the larvae used and the timing with which the queen cells become available.

Environmental conditions can also influence colonies on the brink of swarming … literally a case of rain stopping play.

Acting on impulse

If there are play cups in the colony then you don’t need to take any action 15, but if there are charged queen cells present then your bees are trying to tell you something.

Precisely what they’re trying to tell you depends upon the number and position of the queen cells, the state or appearance of those cells, and the state of the colony – whether queenright or not.

What you cannot do 16 is decide what action to take based solely on the number, appearance or position of the queen cells you find in the colony. 

Is the colony queenright?

Are there eggs present in the comb?

Does the colony appear depleted of bees?

If there are lots of sealed queen cells, no eggs, no sign of the queen and a depleted number of foragers then the colony has probably swarmed. 

Frankly, this is pretty obvious, though it’s surprising the number of beekeepers who cannot determine whether their colony has swarmed or not.

But other situations are less clear … 

If there are a small number of charged queen cells, eggs, a queen and a good number of bees in the hive then it might be supersedure.

Or the colony might swarm on the day the first cell is sealed 🙁

How do you distinguish between these two situations? 

Is it mid-May or mid-September? Swarming is more likely earlier in the season, whilst supersedure generally occurs later in the season.

But not always 😉

Is the queen ‘slimmed down’ and laying at a reduced rate?

Much trickier to determine … but if she is then they are likely to swarm.

Decisions, decisions 😉 … and going by the number of visits to my previous post entitled Queen cells … don’t panic! there are lots of beekeepers trying to make these decisions right now 🙂


 

Winter losses

I lost 10% of my colonies this winter.

It’s always disappointing losing colonies, but it’s sometimes unavoidable.

I suspect the two I lost were unavoidable … though, as you’ll see, they weren’t completely lost.

April showers frosts

Late April may seem like mid-season for many beekeepers based in southern England. While they were adding their second super, the bees here in Scotland were only just starting to take their first few tentative flights of the year.

This April has been significantly cooler in Fife 1 than any year ‘since records began’.

However, the records I’m referring to are from the excellent Auchtermuchty weather report 2 which only date back to about 2013 … I like it because it’s local, not because it’s historically comprehensive 😉

The average April temperate has only been 5.5°C with 15 nights with frost in the first three weeks of the month. In contrast, the same month in 2019 and 2020 averaged over 9°C with only 3-4 nights with frosts 3

In both 2019 and 2020 swarming started at the end of April. Several colonies had queen cells when I first inspected them and I hived my first swarm (not lost from one of my colonies 😉 ) on the last day of the month.

First inspections and winter losses

Unsurprisingly, with appreciably lower temperatures, things are less well advanced this season. None of the colonies I inspected on the 19th were making swarm preparations. Instead, most were 2-4 frames of brood down on the strength I’d expect them to have before they started thinking about swarming.

Nevertheless, most were busy on a lovely spring day … lots of pollen (mainly gorse and some late willow by the looks of things) being delivered by heavily-laden foragers, and fresh nectar in some of the brood frames.

Fresh nectar glistening in a brood frame

The first inspection of the season is an opportunity to not only check on the strength and behaviour of the colony, but also to do some ‘housekeeping’. This includes:

  • swapping out old, dark brood frames (now emptied of stores) and replacing them with new foundationless frames
  • removing excess stores to make space for brood rearing
  • removing the first sealed drone brood in the colony to help hold back Varroa replication

And, as the winter is now clearly over, it’s the time at which the overall number of winter losses can be finally assessed.

Winter losses

Winter losses generally occur for one of four reasons:

  • disease – in particular caused by deformed wing virus (DWV) vectored by high levels of Varroa in the hive. DWV reduces the longevity of the diutinus winter bees, meaning the colony shrinks in size and falls below a threshold for viability. There are too few bees to thermoregulate the colony and too few bees to help the queen rear new larvae. The colony either freezes to death, dwindles to the size of an orange, or starves to death because the cluster cannot reach the stores 4.
  • queen failure – for a variety of reasons queens can fail. They stop laying altogether or they only lay drone brood. Whatever the reason, a queen that doesn’t lay means the colony is doomed.
  • natural disasters – this is a bit of a catch-all category. It includes things like flooded apiaries, falling trees and stampeding livestock. Although these things might be avoidable – don’t site apiaries in flood risk areas, under trees or on grazing land – these lessons are often learnt the hard way 5.
  • unnatural disasters – these are avoidable and generally result from inexperienced, or bad 6 , beekeeping. I’d include providing insufficient stores for winter in this category, or leaving the queen excluder in place resulting in the isolation of the queen, or allowing the entrance to be blocked. These are the things that the beekeeper alone has control over. 

The BBKA run an annual survey of winter losses in the UK. This is usually published in midsummer, so the graph below is from 2020.

BBKA winter survival survey

Over the 13 years of the survey the average losses were 18.2% 7. Long or particularly hard winters result in higher levels of losses.

Lies, damn lies and statistics

I’ve no idea how accurate these winter loss surveys are.

About 10% of the BBKA membership reported their losses, and the BBKA membership is probably a bit over 50% of UK beekeepers. 

I would expect, with precious little evidence to back it up, that the BBKA generally represents the more ‘engaged’ beekeepers in the UK 8. It also probably represents a significant proportion of new beekeepers who were encouraged to join while training.

So, like Amazon reviews, I treat the results of the survey with quite a bit of caution. I suspect beekeepers who have low losses complete it enthusiastically to ‘brag’ about their success (despite its anonymity), while those with large losses either keep quiet or are happy to share their grief. 

Unlike Amazon reviews, I’d be surprised if there are many fake submissions to the BBKA and I’m not aware there’s a living to be made from selling fake colony survival reviews in bulk online.

For comparison, the Bee Informed Partnership in the USA runs a similar survey every year.

Bee Informed Partnership loss and management survey

This survey covers about 10% of the colonies in the USA. Again it is voluntary and likely subject to the same inherent biases that may affect the BBKA survey.

The USA winter colony losses average ~28% over the same 13 year survey period.

Are US beekeepers less good at keeping their colonies alive than beekeepers in the UK?

Perhaps the US climate is less suited to honey bees?

Or, possibly, US beekeepers are simply more honest than their UK counterparts?

I doubt it 9.

Running on empty

My two colony losses were due to queen failures.

Old winter bees and no brood

In the first colony there was no evidence the queen had laid any brood since the previous autumn. There were about 6 seams of bees in the hive, but the outer 2-3 frames were solid with untouched winter stores.

Unused winter stores

This is usually a dead giveaway … literally. The colony hasn’t used the stores because they’ve not had any hungry mouths to feed. With no new brood the colony is doomed.

This queen appears to have simply run out of sperm and stopped laying. She was present (a 2019 marked queen and the same one I’d seen in August last year) and ambling around the frame, but she wasn’t even going through the pretence of inspecting cells before laying.

I removed the queen and united what remained of the colony over a nearby strong colony.

Strong colony ready for uniting

Assuming the queen stopped laying at the end of year all the bees in the hive – and there were a good number – were old, winter bees. These won’t survive long, but will provide a temporary boost to the colony I united them with. 

Every little bit helps 🙂

Even more valuable than the bees were the frames they were on.

Most of the comb in the colony with the failed queen was relatively new. By uniting them I can quickly swap out the old comb (from the stronger hive #34) when I next inspect the hive. At the same time I’ll rescue the frames of sealed stores for use when making up nucs during queen rearing.

Drone laying queen

The second failure was a drone laying queen (DLQ).

These are usually unmistakeable … the brood is clustered, with drone pupae occupying worker brood cells. If the queen has been drone laying for some time there may be lots of undersized ‘runt’ drones present in the hive as well.

Drone laying queen ...

Drone laying queen …

Again, this colony was doomed. With no new queens available and a lot of pretty old bees in the hive they could not be restored to a functioning colony.

However, many of the bees could be saved …

The colony wasn’t overrun with drones. Going by the amount of stores consumed it had probably been rearing worker brood since the winter solstice.

The queen was unmarked and unclipped. I strongly suspect she was a late-season supersedure queen who was very poorly mated.

The 3-4 weeks of drone brood rearing 10 had wrecked quite a few of the frames, but the bees were worth saving.

Under these circumstances I decided to shake the colony out.

When I do this I like to move the original hive and the stand it’s on. If you don’t move the stand the displaced bees tend to cluster near the original hive entrance, festooned from the hive stand. 

In poor weather, or late in the afternoon, this can lead to lots of bees unnecessarily perishing.

However, the stand was shared with two other colonies, so couldn’t be removed. It was also late morning and the weather was excellent.

I moved the hive away and shook the bees out. 

Sure enough … they returned to their original location.

They then marched along the hive stand to the entrance of the adjacent hive.

This way sisters!

And, by the time I left the apiary in mid-afternoon there were only a few diehard bees clustered near where the original hive entrance was.

Why didn’t I just unite them as I’d done with the other failed queen?

Drone brood is a Varroa magnet

Varroa replicate when feeding on developing pupae. The longer development time of drone pupae (when compared with worker pupae) means that you get ~50% more Varroa from drone brood 11

Unsurprisingly perhaps (or not, because that’s the way evolution works) Varroa have therefore evolved to preferentially infest drone brood. When given the choice between a drone or worker pupa to infest, Varroa choose the drone about 10 times more frequently than the worker.

And that ~10:1 ‘preference ratio’ increases when drone brood is limiting … as it is early in the season.

What this means is that the first burst of drone brood production in a colony is very attractive to Varroa.

Unless there are compelling reasons to keep this very early drone brood – for example, a colony with stellar genetics I’d like to contribute as much as possible to the local gene pool – I often try and remove it.

Drone-worker-drone

Drone-worker-drone …

If you use foundationless frames this is often as easy as simply cutting out a single panel of drone brood.

But, in the case of this drone laying queen, it meant that the logical action was to discard all of the drone brood to ensure I discarded the majority of the Varroa also present in the colony 12.

Which is why I shook the colony out, rather than uniting them 🙂

Boxes of bees

Several colonies in one apiary went into the winter on double brood colonies. Inevitably, with the loss of bees during the winter months, the colony contracts and the queen almost invariably ends up laying in the upper box.

The first inspection of the season is often a good time to remove the lower box. It can be removed altogether, or replaced (above the other box) for a Bailey comb change if the weather is suitable.

At this stage of the year the lower box is often reasonably empty of bees and totally empty of brood. 

Emptying a box of bees

If the comb in the lower box is old and dark (see the picture above) I place the upper box on the original floor and add an empty super on top. I then go through the lower box, shaking the bees into the empty super. Good frames are retained, the rest are destined for the wax extractor and firelighters.

Using an empty super helps ‘funnel’ the bees into the brood box.

Sometimes the queen has already laid up a frame or so in the lower box. Under these circumstances – particularly if the comb is relatively new – I’ll simply reverse the boxes, placing the lower box on top of the upper one. This results in the queen quite quickly moving up and laying up the space in the upper brood chamber.

It’s then time to add a queen excluder and the first super.

The beekeeping season has definitely started 🙂


Notes

I commented a fortnight ago about the apparent lateness of the 2021 spring. I’m adding this final note on the afternoon of the 23rd and have still yet to see or hear either cuckoo or chiffchaff on the west coast. Last year they were here in the middle of the month. This, combined with the temperature data (see above) show that everything is a week or two behind events last year.

Which means I can expect to start doing some sort of swarm prevention and control in the next fortnight.

Quick thinking & second thoughts

I gave my last talk of the winter season on Tuesday to a lovely group at Chalfont Beekeepers Society. The talk 1 was all about nest site selection and how we can exploit it when setting out bait hives to capture swarms.

It’s an enjoyable talk 2 as it includes a mix of science, DIY and practical beekeeping.

Nest sites, bait hives and evolution

The science would be familiar to anyone who has read Honeybee Democracy by Thomas Seeley. This describes his studies of the features considered important by the scout bees in their search for a new nest site 3.

Under offer ...

Under offer …

The most important of these are:

  • a 40 litre cavity (shape unimportant)
  • a small entrance of 10-15cm2
  • south facing
  • shaded but in full view
  • over 5m above ground level
  • smelling of bees

All of which can easily be replicated using a National brood box with a solid floor. Or two stacked supers.

And – before you ask – a spare nuc box is too small to be optimal.

That doesn’t mean it won’t work as a bait hive, just that it won’t work as well as one with a volume of 40 litres 4.

Evolution has shaped the nest site selection process of honey bees. They have evolved to preferentially occupy cavities of about 40 litres.

Presumably, colonies choosing to occupy a smaller space (or those that didn’t choose a larger space 5 ) were restricted in the amount of brood they could raise, the consequent strength of the colony and the weight of stores they could lay down for the winter.

Get these things wrong and it doesn’t end well 🙁

A swarm occupying a nuc box-sized cavity would either outgrow it before the end of the season, potentially triggering another round of swarming, or fail to store sufficient honey.

Or both.

Over thousands of colonies and thousands of years, swarms from colonies with genetics that chose smaller cavities would tend to do less well. In good years they might do OK, but in bad winters they would inevitably perish.

Bait hive compromises

If you set out a nuc box as a bait hive, you’re probably not intending to leave the swarm in that box.

But the bees don’t know that. Their choices have been crafted over millenia to give them the best chance of survival.

All other things being equal they are less likely to occupy a nuc box than a National brood box.

Another day, another bait hive, another swarm …

For this reason I don’t use nuc boxes as bait hives.

However, I don’t recapitulate all the features the scout bees look for in a ‘des res’.

I studiously ignore the fact that bees prefer to occupy nest sites that are more than 5 metres above ground level.

This is a pragmatic compromise I’m prepared to make for reasons of convenience, safety and enjoyment.

Bees have probably evolved to favour nest sites more than 5 metres above ground level to avoid attention from bears. The fact that there are no bears in Britain, and haven’t been since the Middle Ages 6, is irrelevant.

The preference for high altitude nest sites was ‘baked into’ the genetics of honey bees over the millenia before we hunted bears 7 to extinction.

However, I ignore it for the following reasons:

  • convenience – I usually move occupied bait hives within 48 hours of a swarm arriving. It’s easier to do this from a knee height hive stand than from a roof ladder.
  • safety – I often move the bait hive late in the evening. Rather than risk disturbing a virgin queen on her mating or orientation flights (assuming it’s a cast that has occupied the bait hive) I move them late in the day. In the ‘bad old days’ when I often didn’t return from the office until late, this was sometimes in the semi-dark. Easy and safe to do at knee height … appreciably less so at the top of a ladder.
  • enjoyment – I can see the scout bees going about their business at a hive near ground level without having to get the binoculars out. Their behaviour is fascinating. If you’ve not watched them I thoroughly recommend it.

Scout bee activity

The swarming of honey bees is a biphasic process. In the first phase the colony swarms and forms a temporary bivouac nearby to the original nest site.

The two stage process of swarming

The scout bees search an area ~25 km2 around the bivouacked swarm for suitable nest sites. They communicate the quality and location of new nest sites by performing a waggle dance on the surface of the bivouac.

Once sufficient scouts have been convinced of the suitability of one of the identified nest sites the second phase of swarming – the relocation of the swarm – takes place.

Swarm of bees

Swarm of bees

However, logic dictates that the scout bees are likely to have already identified several potential new nest sites, even before the colony swarms and clusters in a bivouac.

There are only a few hundred scout bees in the swarmed colony, perhaps 2-3% of the swarm.

Could just a few hundred scouts both survey the area and reach a quorum decision on the best location within a reasonable length of time?

What’s a reasonable length of time?

The bivouacked swarm contains a significant amount of honey stores (40% by weight) but does not forage. It’s also exposed to the elements. If finding sites and reaching a decision on the best nest site isn’t completed within a few days the swarm may perish.

Which is why I think that scout bees are active well before the colony actually swarms.

Early warning systems

If scout bees are active before a colony swarms they could be expected to find and scrutinize my bait hive(s).

If I see them doing this I’m forewarned that a colony within ~3 km (the radius over which scout bees operate) is potentially making swarm preparations.

Since I’ll always have a bait hive or two within 3 km of my own apiaries I’ll check these hives at the earliest opportunity, looking for recently started queen cells.

Whether they’re my colonies or not, it’s always worth knowing that swarming activity has started. Within a particular geographic area, with similar weather and forage, there’s usually a distinct swarming period.

If it’s not one of my colonies then it soon might be 😉

So, in addition to just having the enjoyment of watching the scout bees at work, a clearly visible – ground level – bait hive provides a useful early warning system that swarming activity has, or soon will, start.

Questions and answers

Although talking about swarms and bait hives is enjoyable, as I’ve written before, the part of the talk I enjoy the most is the question and answer session.

And Tuesday was no exception.

I explained previously that the Q&A sessions are enjoyable and helpful:

Enjoyable, because I’m directly answering a question that was presumably asked because someone wanted or needed to know the answer 8.

Helpful, because over time these will drive the evolution of the talk so that it better explains things for more of the audience.

Actually, there’s another reason in addition to these … it’s a challenge.

A caffeine-fueled Q&A Zoom session

It’s fun to be ‘put on the spot’ and have to come up with a reasonable answer.

Many questions are rather predictable.

That’s not a criticism. It simply reflects the normal range of topics that the audience either feels comfortable asking about, or are interested in. Sometimes even a seemingly ‘left field’ question, when re-phrased, is one for which there is a standard answer. The skill in this instance is deciphering the question and doing the re-phrasing.

But sometimes there are questions that make you think afresh about a topic, or they force you to think about something you’ve never considered before.

And there was one of those on Tuesday which involved biphasic swarming and scout bee activity.

Do all swarms bivouac?

That wasn’t the question, but it’s an abbreviated form of the question.

I think the original wording was something like:

Do all swarms cluster in a bivouac or do some go directly from the original hive/location to the new nest site?

And I didn’t know the answer.

I could have made a trite joke 9 about not observing this because my own colonies swarm so infrequently 🙄

I could have simply answered “I don’t know”.

Brutally honest, 100% accurate and unchallengeable 10.

But it’s an interesting question and it deserved better than that.

So, thinking about it, I gave the following answer.

I didn’t know, but thought it would be unlikely. For a swarm to relocate directly from the original nest site the scout bees would need to have already reached a quorum decision on the best location. To do this they would need to have found the new nest site (which wouldn’t be a problem) and then communicate it to other scout bees, so that they could – in turn – find the site. Since this communication involves the waggle dance it would, by definition, occur within the original hive. Lots of foragers will also be waggle dancing about good patches of pollen and nectar so I thought there would be confusion … perhaps they always need to form a bivouac on which the scout bees can dance? Which explains why I think it’s unlikely.

In a Zoom talk you can’t ponder too long before giving an answer or the audience will assume the internet has crashed and they’ll drift off to make tea 11.

An attentive beekeeping audience … I’d better think fast or look stupid

You therefore tend to mentally throw together a few relevant facts and assemble a reasonable answer quite quickly.

And then you spend the rest of the week thinking about it in more detail …

Second thoughts

I still don’t know the answer to the question Do all swarms bivouac?”, but I now realise my answer made some assumptions which might be wrong.

I’ll come to these in a minute, but first let me address the question again with the help of the people who actually did the work.

I’ve briefly looked back through the relevant literature by Seeley and Lindauer and cannot find any mention of swarms relocating without going via a bivouac. I may well have missed something, it wouldn’t be the first time 12.

However, their studies are a little self-selecting and may have overlooked swarms that behaved like this.

Both were primarily interested in the waggle dance and the decision making process, they therefore needed to be able to observe it … most easily this is on the surface of the bivouac.

Martin Lindauer mainly studied colonies that had naturally swarmed, naming them after the location of the bivouac, and then studied the waggle dancing on the surface of the clustered swarm. In contrast, Tom Seeley created swarms by caging the queen and adding thousands of very well fed bees.

Absence of evidence is not evidence of absence.

So, what were the assumptions I made?

There were two and they both relate to confusion between waggle dancing foragers and scout bees.

  1. Swarming usually occurs during a strong nectar flow. Therefore there are likely to be lots of waggle dancing foragers in the hive at the same time the scouts are trying to persuade each other – using their own fundamentally similar – waggle dances.
  2. Bees ‘watching’ are unable to distinguish between scouts bees and foragers.

So, what’s wrong with these assumptions?

A noisy, smelly dance floor

Foragers perform the waggle dance on the ‘dance floor’. This is an area of vertical comb near the hive entrance. It’s position is not fixed and can move – further into the hive if the weather is cold, or even out onto a landing board (outside the hive) in very hot weather 13.

So, although the dance floor occupied by foragers isn’t immovable, it is defined. There’s lots of other regions of the comb that scouts could use for their communication i.e. there could be spatial separation between the forager and scout bee waggle dances.

Secondly, foragers provide both directional and olfactory clues about the identity and location of good sources of pollen and nectar. In addition to two alkanes and two alkenes produced by dancing foragers 14 they also carry back scents “acquired from the environment at or en route to the floral food source” which are presumed to aid foragers recruited by the waggle dancer to pinpoint the food source.

Importantly, non-dancing returning foragers do not produce these alkanes and alkenes. Perhaps the dancing scouts don’t either?

A dancing scout would also lack specific scents from a food source.

Therefore, at least theoretically, there’s probably a good chance that scout bees could communicate within the hive. Using spatially distant dances and a unique combination of olfactory clues (or their absence) scouts may well be able to recruit other scouts to check likely new nest sites.

All of which would support my view that bait hives provide a useful early warning system for colonies that are in the very earliest stages of swarm preparations … rather than just an indicator that there’s a bivouacked swarm in the vicinity.

But?

All this of course then begs the question … if the scout bees can communicate within the hive, why does the swarm need to bivouac at all?

The bivouac must be a risky stage in the already precarious process of swarming. 80% of wild swarms perish. At the very least it’s subject to the vagaries of the weather. Surely it would be advantageous to stay within the warm, dry hive until a new nest site is identified?

Apple blossom ...

Apple blossom … and signs that a bivouacked swarm perished here

This suggests to me that the bivouac serves additional purposes within the swarming process. A couple of possibilities come to mind:

  • the gravity-independent, sun-orientated waggle dancing 15 on the surface of the bivouac may be a key part of the decision making process, not possible (for reasons that are unclear to me) within the confines of the hive.
  • the bivouac acts to temporally coordinate the swarm. A swarm takes quite a long time to settle at the bivouac. Many bees leave the hive during the excitement of swarming but not all settle in the bivouac. Perhaps it acts as a sorting mechanism to bring together all the bees that are going to relocate, separate from those remaining in the swarmed colony?

Clearly this requires a bit more thought and research.

If your association invites me to discuss swarms and bait hives next winter I might even have an answer.

But, as with so many things to do with bees, knowing that answer will only spawn additional questions 😉


 

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.


Notes

BBKA statement made a day or two after this post appeared. The BBKA and other national associations are concerned about the potential import of Small Hive Beetle (SHB) into the UK via Northern Ireland. Whilst I still think this breaches the Northern Ireland Protocol, it doesn’t mean it won’t be attempted (and there’s at least one importer offering bees via this route). It’s not clear that the NI authorities have the manpower to inspect thousands of packages.

It’s worth noting that SHB was introduced to southern Italy in 2014 and remains established there. The most recent epidemiological report shows that it was detected as late as October 2020 in sentinel apiaries and is also established in natural colonies.

With a single exception – see below – every country into which SHB has been imported has failed to eradicate it. As I wrote in November 2014:

“Once here it is unlikely that we will be able to eradicate SHB. The USA failed, Hawaii failed, Australia failed, Canada failed and it looks almost certain that Italy has failed.”

And Italy has failed.

The one exception was a single import to a single apiary in the Portugal. Notably, the illegal import was of queens, not nucs or packages. Eradication involved the destruction of the colonies, the ploughing up of the apiary and the entire area being drenched in insecticide.

The Beekeepers Quarterly

This post also appeared in the summer 2021 edition of The Beekeepers Quarterly published by Northern Bee Books.

Going the distance

I’m going to continue with a topic related to the waggle dance this week.

This is partly so I can write about the science of how bees measure distance to a food source.

But it’s also to encourage those who didn’t read the waggle dance post to visit it. Weirdly it was only read by about 50% of the usual Friday/weekend readership and I suspect (from a couple of emails I received) that the weekly post to subscribers ended up in spam folders 1.

If you remember, the duration of the waggle phase of the dance – the straight-line abdomen-wiggling sashay across the ‘dance floor’ – indicates the distance from the nest to the desirable food source 2. The vigour of the wiggle indicates the quality of the source.

How do bees measure distance?

Karl von Frisch, the first to decode the waggle dance, favoured the so-called ‘energy hypothesis’. In this, the distance to a food source was determined by the amount of energy used on the outbound flight.

Does that seem logical?

Foragers forage randomly, but usually return directly

If correct, foragers would only be able to determine the energy used after their second trip to a food source. This presumes their first trip was longer as they searched the environment for something worth dancing about 3.

This would be an easy thing to test, though I’m not sure it was ever investigated 4.

As it happens, far better brains determined that the energy hypothesis was probably incorrect. Many of these studies explored how gravity influences the distances reported by dancing foragers.

Going up!

Bees use more energy when flying up. For example, when flying from ground level to the top of a tall building, when compared to level flight. Similarly, they use more energy flying if they have small weights attached to them 5.

A series of experiments, nicely reviewed by Harald Esch and John Burns 6, failed to provide good support for the energy hypothesis. There were lots of these studies, involving steep mountains, tall buildings or balloons, between the 1950’s and mid-80’s.

Interesting science, and no doubt it was a lot of fun doing the experiments.

For example, bees flying to a sugar feeder situated on top of a tall building dance to ‘report’ the same distance as bees from the same hive flying to a feeder at ground level adjacent to the same building.

Similarly, foragers loaded with weights do not overestimate the distance to a food source, as would be expected if the energy expended to reach it was being measured 7.

Interesting and entertaining science certainly, but none of it providing compelling support for the energy hypothesis

It’s notable that there is a rather telling sentence from the Esch & Burns review that states “While reading the original papers, one gains the impression that evidence supporting the energy hypothesis was favored over arguments against it”.

Ouch!

Splash landing

Although Von Frisch was a supporter of the energy hypothesis 8 he also published a study that provided evidence for our current understanding of how bees measure distance.

Bees generally don’t like flying long distances over water. Von Frisch provided two equidistant nectar sources, one of which was situated on the other side of a lake.

Bees flying over calm water underestimate distances

On very calm days the bees that flew across the lake under-reported the distance to the feeder. This underestimate was by 20-25% when compared to bees flying to an equidistant feeder overland.

Von Frisch commented “the bee’s estimation of distance is not determined through optical examination of the surface beneath her”.

He assumed that the mirror-like water surface provided no optical input as it contained no visual ‘clues’. After all, one calm patch of water looks much like any other. Von Frisch used this as an argument for the energy hypothesis.

He also noted that the bees generally flew very low over the water surface, often so low that they drowned 🙁

Perhaps these bees were flying dangerously low to try and find optical clues.

Such as their height above the surface?

Or perhaps the distance travelled?

Going with the flow

Having debunked the energy hypothesis, Esch & Burns proposed instead the optic flow hypothesis. This states that “foragers use the retinal image flow of ground motion to gauge feeder distance”.

Imagine optic flow as tripping a little odometer in the bee brain that records distance as her eyes observe the environment flashing past during flight. The clever thing about that is that the environment is variable. It’s not like counting off regularly spaced telegraph poles from a train window.

When flying, environmental objects that are nearby will move across her vision much faster than distant objects. Bees don’t have stereo vision, but instead use this speed of image motion to infer range.

Optic flow – the arrow size indicates the speed with which the object apparently moves, and hence its range

Esch & Burns returned again to tall buildings to provide supporting evidence for their optic flow hypothesis. They trained bees to fly between two tall buildings with 228 metres separating the hive and the feeder 9.

Returning foragers reported that the food source was only 125 metres away.

However, the bees didn’t make a direct flight. Instead they flew at altitude for 30-50 metres, descended to fly much lower, then ascended again to approach the feeder again at altitude.

Esch & Burns experiment to support the optic flow hypothesis

The interpretation here was that the high altitude flight provided insufficient optic flow to measure distance. The bees descend to get the visual input needed to judge distance, but it’s only for part of the flight … hence leading to under-reporting the distance separating the hive and feeder.

Tunnel vision

Jurgen Tautz 10 and colleagues trained bees to forage in a short, narrow tunnel 11. This elegant experiment provided compelling support for the optic flow hypothesis.

The tunnel was ~6 m long and with a cross sectional area of ~200 cm2 – big enough for a bee to fly along, but sufficiently narrow so that the bee would be closer to the ‘walls’ than in normal free flight. The walls and floor of the tunnel had a random visual texture. Only the end of the tunnel facing the hive was open.

The tunnel experiment.

These studies were conducted when the terms round and waggle were used to distinguish the dance induced by food sources <50 m and >50 m respectively from the hive 12. Rather than emphasise the shape of the dance I’ll just describe it as a >50 m or <50 m waggle dance.

‘Tunneling’ bees misreport distances

In the first tunnel experiment (1) the feeder was 35 m from the hive. 85% of dances indicated the feeder was <50 m away. However, when the feeder was moved to the opposite end of the tunnel (2) – still only 41 m from the hive – 90% of the dances indicated the feeder was >50 m away.

To test how the random pattern influenced the perceived distance the scientists used a third tunnel (3) lined with lengthwise stripes. In this instance – despite the feeder position being unchanged from experiment 2 – 90% of the dances indicated the feeder was <50 m away.

The stripes were predicted to ‘work’ in the same way as the smooth lake surface, providing no visual clues.

In the fourth experiment (4) the feeder was 6 m along a randomly patterned tunnel, which was placed just 6 m from the hive. Over 87% of dances indicated that the feeder was >50 m away.

Interpreting the waggle run

In open flight 13 there is usually an excellent correlation between the duration of the waggle run and the distance to a feeder (see the graph below 14 ). By extrapolation, the bees in experiments 2 and 4 ‘thought’ they had flown 230 m and 184 m respectively. In reality they had flown only 41 m and 12 m in these experiments.

Determining distances from waggle dance observation

How could the bees get it so wrong?

Increased optic flow

Tunnel-traversing bees fly just a few centimeters away from the visible ‘environment’.

As a consequence, at the same flight speed, they experience greater optic flow.

If, instead of driving around in your lumbering old van, you pack your hive tool in a Caterham 7 for the trip to the apiary you’d be well aware of what I mean.

Caterham 7 … check out that optic flow … then make another trip to collect the smoker

30 mph in a Toyota Hilux feels very much slower than 30 mph in a Caterham 7. This is largely because visual reference points, like the broken white lines between lanes in the road, appear in and disappear from your field of view much faster … because you’re much closer to them.

Because the tunnel dimensions were known it was possible to calculate the calibration of the bee’s odometer. Classically this would be defined in terms of metres of distance flown generating a particular waggle run length or duration.

These tunnel studies demonstrate that distance flown is not what calibrates the odometer. Instead it’s quantified indirectly in terms of the image motion experienced by the eye. Since environments vary the way to express this is the amount of angular image motion that generates a given duration of waggle.

And, using some mathematical trickery we don’t need to bother with 15, it turns out that this angular motion is only dependent upon distance flown, not the speed of flight.

This is important. Headwinds or tailwinds could change the speed of flight, but not the distance flown 16.

It’s all relative

It’s worth emphasising that the dance followers in experiments 2 (above) should still find the feeder.

The waggle dance would ‘instruct’ them to fly 230 m at the bearing indicated and they’d experience the same visual clues en route.

This means that they should still enter the narrow tunnel and experience increased optic flow because of the encroaching walls. But they’d be experiencing the same optic flow the initial dancing bee had experienced, so would not attempt to fly further down the tunnel.

This means that the optic flow experienced is context dependent. It is related to the environment the bees are foraging in.

This makes sense as the dancing bees and dance followers all occupy the same environment.

How do we know this? 17

Changing the environment

If we change the environment the dance followers search at the wrong distance.

I qualified the statement above when I said that the dance followers should still enter the tunnel and find the feeder.

Actually, most recruits will miss the tunnel entrance – remember it’s smaller that a sheet of A5 paper. At 35 m distance a bee would have to get the bearing correct to about 0.16° to enter the tunnel 18.

So the bees that do not enter the tunnel experience a different environment.

Where do they search for the feeder?

They search at the distance indicated by the waggle duration … so bees that missed the tunnel entrance in experiment 2 (above) would have searched for the feeder 230 m from the hive. Similarly, the dance followers in experiment 4 would have searched 184 m away 19

Context dependent dance calibration

And, finally, the calibration of the odometer depends upon the environment.

Odometer calibration depends upon the environment

If the environment experienced by the dancing bee en route to the feeder in experiments 2 and 4 is different, then it generates a different relationship between waggle run duration and distance.

For example, if one feeder was across a closely mown lawn and the other was across dense shrubby woodland, they would each generate a unique optic flow, so changing the image motion experienced, and hence the waggle run generated.

In the diagram above, you shouldn’t use dance calibration for bees trained to direction A to determine the distance bees going in direction B would forage.

Phew!

Optic flow, waggle dancing and implications for practical beekeeping

None 😉

At least, none that I can think of.

A Caterham 7 isn’t an ideal car for a beekeeper but would be a lot of fun to help you understand optic flow 😉

Most of us keep our bees in mixed environments. Your apiary isn’t situated with a cliff edge on one side and an unbroken prairie on the other. Since the environment is mixed, the waggle dance calibration is not going to be wildly different, whichever way the bees fly off in. You can therefore use an approximate figure of 1 second per kilometre to estimate the the distance at which your bees are foraging, irrespective of the direction they go.


Notes

Most of the referenced studies are at least two decades old. Honey bees have remained a fertile research tool for neurobiologists. Our understanding of honey bee vision continues to improve. However, I cannot discuss any of these more recent studies with reference to optic flow. Anyway, just because they’re old doesn’t make the experiments any less elegant or interesting 🙂