Category Archives: Problems

Autumn cleaning

Over the last fortnight, despite some occasional warm and sunny days, the autumn has made its presence known. 

Flaming autumn aspen

The aspen down the road are a stunning colour at this time of the year. Although I’ve planted a couple of dozen, they’re still not more than thigh-high and it will be quite a few years until they can match the display shown above.

Almost overnight hundreds of redwing have arrived from Scandinavia and many of the rowan have already been stripped bare 1.

In Fife, the leaden skies are filled with skeins of geese forming raggedy V’s as they fly in from the North Sea. It’s an evocative sight … it reminds me of my first weeks as an undergraduate student at Dundee University half a lifetime ago

And it also emphasises that the beekeeping season is over.

Of course, there will be jobs to do in the winter, but the bees are pretty much on their own for at least the next five months.

Apivar

The final essential task of the season for me is to remove the Apivar strips that went into the hives in August. Initially the strips were placed on either side of the – still large – brood nest. A few weeks ago I removed the strips, scraped them free of propolis and wax and re-inserted them around the, now shrunken, brood nest.

Mid-autumn and time for the Apivar strips to be removed

You can just about see them in the photo above, flanking the four central frames.

It is important to remove the strips. Although Apivar has a relatively short half-life, some residual activity will remain. If you leave them in the hive any surviving Varroa – and there will be survivors 2 – will continue to reproduce in the presence of trace levels of amitraz, the active ingredient in Apivar. 

With reduced – and possibly borderline for killing – levels of amitraz present, these are ideal conditions in which resistance may develop. Although this has been reported it does not appear to be widespread. 

Therefore, to ensure that Apivar remains an effective miticide it is important to remove any remaining strips before the winter.

Your next adventure in Glenrothes awaits!

Tragic isn’t it?

That’s the subject line on the emails I receive from Travelodge where I stay when I’m doing my beekeeping in Fife. 

Have you ever been to Glenrothes?

‘Adventure’ isn’t the word most people associate with Glenrothes. 

Good morning Glenrothes

GetMeOuttaHere is. 

This is a town where every third car being driven late at night has a raucous exhaust, lowered shocks, tinted windows and a spoiler. The drive-in queue for McDonald’s sounds like the pit lane at the Indianapolis 500 and there are more donuts in the car park than in the fast food outlets 3.

But none of that usually bothers me as, by the time I get to the hotel, I’ll have been driving for 5 hours and will have spent about the same amount of time inspecting colonies or lifting cleared supers. I may also have squeezed in a couple of hours of meetings at work.

The environment might be noisy, but the beds are comfortable. 

But visits in late autumn are a bit different.

No colonies to inspect, no grafting to do, no nucs to check for mated queens and no supers to remove.

All I need to do is gently lift a few crownboards and pull out the Apivar strips now that treatment is complete.

So, what do I do for the rest of the day?

Long range weather forecasting

Is that an oxymoron?

I book my trips to Fife to fit in with what the bees need. To make the hotel affordable I book many weeks in advance.

I therefore put up with whatever the weather throws at me. Usually it works out OK.

Furthermore, as regular reader know, several hives are in a bee shed, so the weather is largely irrelevant.

But ~60% of them are outside.

And Monday was really wet. 

Having driven for four hours through increasingly heavy rain – stopping en route to make a honey delivery – I fortified myself with a cappuccino and excellent almond croissant from Taste, the best independent coffee shop in St Andrews 4.

Essential fortifications

I then sat in the shed enjoying my late breakfast listening to the rain hammering on the roof.

I needed something to occupy me until either:

  • the rain stopped
  • it got so late in the day that I’d just have to open the hives and remove the strips anyway

And the obvious thing to do was a bit of spring autumn cleaning. 

During the season the bee shed is used on a daily or weekly basis depending upon the experiments underway. In addition, we have a storage shed on the same site and a number of additional hives in the same apiary. I also do most of my queen rearing in this apiary (the bee shed provides a near-perfect environment for grafting), distributing the nucs to other apiaries for mating.

And all that beekeeping tends to leave a bit of a mess. At least, it does where I’m involved.

Super job

For the last couple of years I’ve not bothered returning the extracted supers to the hives for the bees to recover the last of the honey.

Instead I’ve just stacked them ‘wet’ in the shed, protected from wasps, mice and robbing bees, by covering the top of the stack with a well-fitting roof.

Or a snug-fitting crownboard and a badly fitting roof.

Stacked ‘wet’ supers

Experience has taught me that the floor of the shed isn’t level and/or has gaps between the planking. Rather than seal all these gaps I simply stand the stack of boxes on the sort of closed cell foam sheeting used for packing furniture, or – when I run out – on double thicknesses of cardboard 5. This stops the wasps, ants and bees from getting access. 

So I started by tidying the stacks of supers. Inevitably this necessitated moving them first, sweeping the floor clean, laying out the foam/cardboard and then restacking them. There’s not enough space in the shed to move ~60 supers so they went out in the rain.

So I got wet 🙁

Floors, roofs, boards, unidentifiable objects and wax moth

Once they were back I could turn my attention to the other side of the storage shed which houses spare roofs, nuc feeders, floors, boards (split, crown, surf, Morris, Snelgrove etc. 6 ), a breeding colony of queen excluders 7 and a motley collection of other items that:

  • might come in useful
  • don’t logically belong anywhere else
  • appear valuable and/or difficult to make … but I don’t know what they are
  • are essential and were needed several times in the season … but I’d lost them 🙁

Sorting this lot out took another hour or two, and involved a further soaking as I needed to clear the space before I could refill the space.

Early on in the process … 

Is beekeeping the largest volume hobby?

… and when at least partial order had been restored …

Floors from Abelo, Pete Little and some homemade abominations

I also found several brood boxes full of drawn comb or sealed stores.

Excellent 🙂

And I found a nuc box lurking in the far corner containing comb riddled with wax moth 🙁

Wax moth larvae and damage

Aargh!

DiPel DF

Wax moth are something I’ve largely avoided or ignored for most of the last decade. The cold winters in Scotland seem to keep their numbers down.

Not this time … 

All of the infested frames were bagged up for burning at the earliest opportunity. The remaining brood frames were treated with DiPel DF, a suspension of Bacillus thuringiensis kustaki spores and toxins. If ingested by the larvae of wax moths, the δ-endotoxin component dissolves in the alkaline environment of the gut, is activated following cleavage by gut proteases and then ‘punches’ a hole through the gut wall.

Ouch.

And the spores germinate, allowing the bacteria to grow inside the larva.

As I wrote in a post several years ago about this treatment:

This isn’t good for the moth larva. Not good at all. Actually, it’s probably a rather grisly end for the moth but, having seen the damage they can do to stored comb, my sympathy is rather limited.

DiPel DF is non-toxic for bees.

DiPel DF

I’ve not had problems with wax moth infesting supers stored ‘wet’ … they’re after the old cocoons and other rubbish that accumulates in brood frames.

Vita used to sell a product called B401 – also a suspension of Bacillus thuringiensis spores and proteins – which was withdrawn from sale in 2019. Despite assurances that a replacement – imaginatively labelled B402 – would be available ‘soon’ it appears to only currently be sold in the US.

Out with the old … and the not fit for purpose

I was on a roll … 

All this organisation meant I discovered things that I’d lost … like a small stack of contact feeders hiding in the corner that had not been used this season as I hadn’t done any shook swarms.

There they are! Contact feeders lurking shyly in the furthest corner (unlike those brazen frame feeders at the front)

I also found some mini-nucs I’d built for queen mating almost 10 years ago. They were made of ply and housed a tri-fold full-size brood frame (you can now buy these, but couldn’t when I built them). 

Tri-fold brood frame

However, the ply was starting to delaminate and it was pretty clear that they wouldn’t survive a Scottish summer season so they were unceremoniously binned.

And I finally bit the bullet and got rid of all my XP Plus queen excluders. These were bought from Thorne’s a few years ago and had been used only when I ran out of everything else.

In principle they are a good idea. A white plastic queen excluder with bee space on the underside provided by a raised rim and a series of small X-shaped spacers that stand on the top bars.

XP Plus queen excluder (the plus must mean ‘plus warp’)

However, in practice, they’re rubbish. They were the ‘ugly’ in my 2017 description of queen excluders that included the phrase ‘the good, the bad and the ugly’.  

They warp really badly. The photo above – if anything – obscures the warp because the QE is not being held flat. When you place them under a super the centre bows up and contacts the underside of the super frames.

Rubbish. 

Out they went.

The little things

There’s something rather poignant about the death throes of the beekeeping season. It can end with a bang as autumn storms roll in, or it can end in a protracted stutter as intermittent good days allow the bees to forage late into October. 

Of course, it’s au revoir 8 and not a final goodbye

It forms such a large part of my life for six months of the year that little things found during the clear-out bring back a flood of memories …

Nicot cup and (partly squidged) queen cell amongst the debris on the shed floor

A Nicot cup and vacated queen cell reminded me what a good queen rearing season we’d had on the east coast. Although the first round of grafting was a near-total failure, successive rounds were excellent, and queen mating was very successful. One of the best seasons in memory 9.

Coffee stirrer … or AFB test kit

Not all the memories were good ones though. I received one of the dreaded ‘AFB alert’ warnings for the apiary and spent a very long couple of days checking every cell on every brood frame in every colony, and testing any that looked suspicious.

I don’t take sugar, and the coffee stirrer shown above is provided in the AFB LFD kit to lift the dodgy-looking larva into a tube for analysis. Everything looked clear, but it gave me a few very stressful days.

And … after all that tidying, and repeated trips to the industrial-scale bins, it finally stopped raining.

Finally … some practical beekeeping

I fired up the smoker and quickly, but gently, removed all the Apivar strips. The crownboards on all the hives were very firmly stuck down with propolis and the bees, although calm, weren’t exactly overjoyed to see me.

Autumn still life – smoker, hive tool, Varroa trays and Apivar strips

I still had another apiary to visit. With rain threatening there wasn’t time to monitor the level of brood present so I slipped cleaned Varroa trays under the hives. This will allow me to inspect both residual mite drop and look for the presence of the characteristic biscuit-coloured cappings when brood is uncapped.

And then, after about half an hour of practical beekeeping, I set off back to the west coast as the rain started again.

The Moidart hills – An Stac, Rois-Bheinn and Sgùrr na Ba Glaise

Two days later the Moidart hills had their first dusting of snow.

It’s official, autumn is here and the beekeeping season is over.


 

Dancing in the City

Beekeeping is an increasingly popular pastime. Since ~84% of the UK population live in urban areas (up from ~70% in 1950’s) it is not unsurprising that the number of hives kept in cities is increasing.

Of course, not everyone who lives in a town or city keeps their bees in the back garden. When I lived in the Midlands I lived on a small estate that was indisputably ‘urban’, although there was farmland within sight 1. My bees were on the nearby farmland and I just kept a few mating nucs and a bait hive in the back garden 2

Hive in a field margin

I kept my bees in the farmland because 3 I reasoned that there were both larger amounts and a greater range of forage available for them there.

But I was probably wrong.

It wouldn’t be the first time, and it certainly won’t be the last.

Too many bees?

Before discussing urban bees and forage in more detail I’ll digress a minute to mention the suggestions that the inexorable rise and rise of urban beekeeping is threatening our native pollinators.

Actually … more than suggestions.

There are a number of scientific reports and reviews that indicate that urban beekeeping harms – by outcompeting – native pollinators like solitary bees. A recent report by the Royal Botanic Gardens at Kew states:

‘Campaigns encouraging people to save bees have resulted in an unsustainable proliferation in urban beekeeping. This approach only saves one species of bee, the honeybee, with no regard for how honeybees interact with other, native species.’

‘In some places, such as London, so many people have established urban hives that the honeybee populations are threatening other bee species.’

Our bees (Apis mellifera) are generalists. They are not particularly well adapted to any flower or nectar/pollen source.

They are equal opportunists.

Individually, there are many solitary bee species or non-hymenopteran pollinators, that are ‘better’ adapted. They pollinate more efficiently, or collect more nectar, faster.

But honey bees arrive in the environment mob handed.

Thousands of them.

Actually, tens or hundreds of thousands of them if there are several hives in an apiary. They are a formidable force and can easily outcompete other pollinators that are either solitary or only live in small colonies.

Save the bees, save humanity

When you see the phrase “Save the bees” what it usually means is save the honey bees.

Save the bees ...

Save the bees …

What it should be encouraging is “Save anything but the honey bees, because they don’t need saving … actually, there are possibly too many of them altogether”.

But that’s a lot less catchy … and it won’t let the supermarket, or food manufacturer or whatever, illustrate their campaign with cute photos of pollen-laden honey bees returning to quaint white-painted WBC hives in some sort of idyllic rural scene 4.

I suppose a photo of a honey bee would be better than drawing of a wasp though … which is what the NRDC used for a (mis)information campaign on CCD (colony collapse disorder) a few years ago.

D’oh!

Anyway … saving the bees is usually “greenwash” or bee-washing as it was termed by MacIvor and Packer in a 2015 paper on bee hotels 5

I’ll return to this topic, and urban beekeeping, later in the winter 6.

The Town mouse bees and the Country mouse bees

Where was I?

Oh yes, my – as will shortly be clear – incorrect assumption that country bees have access to more diverse and richer sources of forage than their poor relatives living in the town.

There are many sorts of countryside and many sorts of urban environments.

A hive in an intensively farmed arable landscape could be located in hundreds of acres of wheat fields, where all of the hedgerows were grubbed up years ago and replaced – if they weren’t just removed altogether – with barbed wire 7

How different is that environment to the ‘concrete jungle’ of a modern city? 

Tokyo

Surely that must be a terrible environment for bees?

In contrast, the suburban sprawl that surrounds most cities is possibly a good place for bees to live. Lots of neat little gardens, each 8 with a profusion of flowering plants, each chosen to provide vibrant colour for a much longer period than native plants.

And I’m sure we can all think of forage-rich rural environments. Here the bees gorge on early crocus, then gorse, willow, oil seed rape, clover, lime, blackberry, fireweed and himalayan balsam, before finishing the season will full crops and corbiculae from the ivy.

Now, in a recent publication 9 scientists have compared the forage available to town and country bees, and the results are quite surprising.

Let the bees tell you

If you live in a country with enlightened right to roam laws (like Scotland) you could wander around the countryside recording all the forage available to bees.

But the laws in England are much less enlightened. However, your right to roam in either country does not extend to the land around a private house or building. 

So, although you might be able to determine the forage available in the Scottish countryside, you can’t be certain you would have good enough access to do the same in England. And in a city you could only map what forage was available by peering over fences from public roads. 

So Elli Leadbetter and colleagues let the bees do the work.

They established 20 observation hives. Ten were in the the centre of London and 10 in agricultural land to the North East and South West of London. The hives were 5 km from each other to avoid overlapping areas of forage. They used observation hives so that they could watch and record the waggle dances of foragers in the hives.

I’ve discussed the waggle dance before. It is used to communicate three important pieces of information about a forage source:

  • direction
  • distance
  • quality

The first two bits of information are encoded in the angle of the waggle run to the perpendicular and the duration of the waggle run. The quality is conveyed by the number of circuits a dancing working performs. It’s a case of ‘the more, the better’ … energetically higher quality resources 10 result in more circuits.

Having recorded thousands of these waggle dances, they used the direction and distance information to ‘map’ where the bees were foraging.

GIS data, land use and foraging bees

For many locations there exists a wealth of land use data (GIS data; Geographic Information Systems). Much of this is at high resolution. For each of the 20 observation hives they produced a map of land use within 2.5 km of the hive at a resolution of 25 m. 

Land use was defined in broad categories such as 11 buildings, woodland, arable, pasture, fruit, OSR (all in agricultural areas) and dense or sparse residential, parks, amenity grassland or railways (all in urban landscapes). 

They then used some clever mathematics to decode the waggle dances 12 to work out where the bees had been, converting the distance and direction components to geographic locations.

Urban (top) and rural foraging heat maps, overlaid on GIS land use maps (5 km diameter)

These locations were overlaid on the land use maps to produce ‘heat maps’ showing where the bees were foraging. The image above shows these heat maps. In the spring the urban bees (top left) were foraging intensively to the east and west of the hive and the rural bees (bottom left) were mainly foraging in two large areas to the south east and south west of the hive.

Foraging distance and nectar quality

Even a cursory look at the image above shows that the urban bees tended to forage closer to the hive than the rural bees. But remember, that’s just three snapshots during the season.

Waggle run duration – rural bees fly further

However, more detailed analysis confirmed that this was the case. Throughout the season, the bees in the agricultural landscape foraged further from the hive. I’m showing the log-transformed median waggle run duration (above) as this allows slightly easier comparison across the season. The further the wiggly ‘best fit’ line is above the horizontal axis, the longer the duration of the waggle dance run, and so the further the bees are flying to find the forage.

Interestingly, the median foraging distances were relatively short when compared with the maximum foraging distances from the decoded waggle dances. This applied wherever the bees lived. For urban bees the median was 492 m (max 9375 m) and for agricultural bees it was 743 m (max 8158 m 13 ).

Perhaps the agricultural bees were flying further because there was better quality nectar available at more distant sites?

Nectar concentration (% w/w) sampled from returning workers

They controlled for this by recovering returning foragers and robbing them of their nectar load before analysing the sugar content. There was no significant difference 14 between the nectar from agricultural or urban landscapes. The sugar content of the nectar was recorded as reducing through the season.

Foraging preferences

So where did the town bees and the country bees prefer to forage?

City rooftop bees

City rooftop bees …

In urban areas the bees exhibited a strong preference for residential gardens (the ‘sparse residential land’) … these are presumably the flower-rich urban gardens that the homeowners also tend to prefer.

In contrast, bees in an agricultural landscape showed an entirely unsurprising reliance upon mass-flowering crops like oil seed rape (in spring only). They also showed weaker preferences for arable land and fruit crops throughout the season.

Mid-April in the apiary ...

Mid-April in a Warwickshire apiary …

I’ve skipped over a host of additional observations from the study, and almost all of the controls. Two things that are worth mentioning though.

Firstly, hive strength had no influence on waggle dance duration (and hence foraging distance). It therefore wasn’t the case that stronger hives had a larger workforce that could survey and exploit more distant forage. 

Secondly, cities are warmer due to the urban heat island effect. However, temperature also did not affect waggle dance duration when it was factored in. So, the city bees aren’t foraging at shorter distances because the dance is truncated at higher temperatures.

Conclusions

So, although cities are predominantly filled with buildings and roads, city bees travel less far to forage when compared to bees in agricultural landscapes.

This strongly suggests that the urban landscape consistently provided more available forage 15

Conversely, the bees in agricultural landscapes had to fly further, not because there were better quality nectar sources available at long-range, but because there wasn’t enough nectar nearby.

There were a number of additional interesting points in the paper, some of which were known already from other studies. For example, the high sugar content in spring nectar was already known (and confirmed here). Similarly, foraging distances in midsummer were longer than those in spring or autumn. This could be predicted due to the reduced rainfall in summer, and consequently the reduced overall level of nectar available.

I need to think more about how this study contributes – if at all – to the ‘too many bees in cities’ argument. If anything, forage-rich towns should be able to support a greater number of bees 16 without the honey bees impacting on other species. In contrast, honey bees in agricultural landscapes might dominate the available nectar sources because they can forage at long distances and then communicate to their nestmates the location.

Perhaps it just shows that that heavily farmed land is actually very poor in terms of nectar availability? It’s either boom or bust … once the OSR has finished, or the clover has been ploughed in, or the fruit trees stop flowering then there’s nothing left 🙁

It would be interesting to conduct a similar study in a forage-rich, non-agricultural, rural landscape.

Access all areas

Finally, I think a particularly neat thing about this study is the use of bees to ‘map’ the forage availability using what the authors term “a large-scale search effort that has no access limitations”. The scientists interpreted the waggle dance to get information that would otherwise have been difficult or impossible to determine.

However, using the bee’s own perception of the distances flown might actually lead to inaccuracies in the calculations. As discussed previously, bees measure distance by optic flow. Optic flow increases in complex landscapes … and cities are likely (at least to us, and certainly when compared with arable farmland, to bees) to be complex landscapes. Increased optic flow leads to a perception of increased distance, and hence to longer waggle runs.

This means that bees in complex landscapes might overestimate the distance they have actually flown to forage. Conversely, those in uniform landscapes might underestimate.

Which means that the results of this study may be a conservative estimate of the differences in foraging distance.

And therefore a conservative estimate of the differences in forage availability in urban and agricultural landscapes.


Notes

Dancing in the City was a pretty cheesy song that reached #3 in the charts for the rock-pop duo Marshall Hain in 1978. Having remembered the track when thinking up a title for this post, I made the mistake of listening to it on Spotify.

I now can’t get the damned thing out of my head. 

But it gets worse. I rummaged through Wikipedia and discovered that Kit Hain, the vocalist, subsequently had a very successful songwriting career with people like Roger Daltrey, Chaka Khan and Fleetwood Mac. Impressive. 

In contrast, Julian Marshall, the keyboard player became a member of the Flying Lizards who had a minor hit with a cover version of Barrett Strong’s 17 Money (That’s What I Want)

… and now I can’t get that out of my head 🙁

Cut your losses

The stats for winter losses in the UK, Europe and USA can make for rather sobering reading.

In the UK, losses over the last 12 years have fluctuated between 9% and 34%. This self-selecting survey includes responses from about 10% of the British Beekeepers Association membership (primarily England and Wales, despite the name). The average number of hives maintained by a BBKA member is about 5, meaning – all other things being equal 1 – that most beekeepers should expect to lose about 1 hive every winter.

BBKA winter losses survey

About 30 countries, mainly Northern hemisphere, contribute to the COLOSS survey which is significantly larger scale. The most recent 2 data published (for the ’16/’17 winter) had data from ~15,000 respondents 3 managing over 400,000 hives. Of these, ~21% were lost for a variety of reasons. COLOSS data is presented as an unwieldy table, rather than graphically. Further details, including recently published results, are linked from their website.

In the USA the Bee Informed Partnership surveys losses – both winter and summer – and claims to have results that cover ~10% of all the colonies in the country (so probably between 250,000 and 275,000 hives). Winter losses in the USA are rarely reported at less than 20% and were as high as 35% in the ’18/’19 winter 4.

Bee Informed Partnership annual colony losses

Are these figures to be trusted?

Who knows?

Each survey is accompanied by a variety of statistics. However, since they all appear to be based upon voluntary reporting by a subset of beekeepers, there are opportunities for all sorts of data to be included (and even more to be missed entirely). 

The problem with surveys

Is the successful beekeeper who managed to get all her colonies through the winter more likely to respond?

A form of ‘bragging rights’.

What about the beekeeper that lost all his colonies?

Does he respond out of a sense of responsibility?

Or does he keep quiet because he doesn’t want to be reminded of those cold, quiet, mouldy boxes opened on the first warm day of spring?

One and two year beekeepers

What about the high level of annual ‘churn’ amongst beekeepers? They buy a nuc in May, filled with enthusiasm about the jars of golden honey they’ll have for family and friends in late summer.

To say nothing of all the “saving the bees” they’ll be doing.

But by late summer the colony is queenless and has an unpleasant temperament

Beekeeping should be enjoyable ...

Beekeeping should be enjoyable …

Psychopathic you might say … if you were feeling uncharitable.

Consequently the Varroa treatment goes on far too late,. Or is quietly forgotten. The winter bees have high viral loads and ‘die like flies’ 5, resulting in the colony succumbing by the year end.

But this colony loss is never recorded on any surveys.

The once enthusiastic beekeeper has moved on and is now passionate about growing prize-winning vegetables or cheesemaking or keeping chickens. 

Beekeeping associations train lots of new beekeepers and – although membership numbers are increasing – it’s well below the rate they’re trained at.

Some may not be ‘joiners’ and go their own way.

Many just quietly stop after a year or two.

How many people have you met that say “Oh yes, I used to keep bees”

Did you ask them whether they ever completed a winter losses survey?

I’m not sure any of the surveys listed above do much ‘groundtruthing’ to establish whether the data they collect is truly representative of the population actually surveyed. With large numbers of respondents spread across a wide geographic and climatic range it’s not an easy thing to do.

So, treat these surveys with a healthy degree of scepticism.

Undoubtedly there are high levels of winter losses – at least sometimes – and the overall level of losses varies from year to year.

Losses and costs

The direct financial cost of these colony losses to beekeepers is very high.

Ignoring time invested and ‘consumables’ like food, miticides and foundation these costs in ’16/’17 for just Austria, the Czech Republic and Macedonia were estimated at €56 million 😯  

These figures simply reflect lost honey production and the value of the lost colonies. They do not include the indirect costs resulting from lost pollination.

But, for the small scale beekeeper, these economic losses are irrelevant.

Most of these beekeepers do not rely on bees for their income.

The real cost is emotional 🙁

It still saddens me when I lose a colony, particularly when I think that the loss was avoidable or due to my incompetence, carelessness or stupidity 6.

Little snow, big snow. Big snow, little snow.

Your hives should be quiet in winter, but it hurts when they are silent in spring.

Anatomy of a death

The COLOSS surveys give a breakdown of winter losses in three categories:

  • natural disasters
  • queen problems
  • dead colonies

Natural disasters are things like bears, honey badgers, flooding or falling trees.

We can probably safely ignore honey badgers in the UK, but climate change is increasing the weather extremes that causes flooding and falling trees.

Moving to higher ground ...

Moving to higher ground …

Don’t assume that poly hives are the answer to potential flooding.

They do float, though not necessarily the right way up 🙁

Queen problems cover a variety of issues ranging from reduced fecundity to poor mating (and consequent drone laying) to very early or late – and failed – supersedure 7.

Beekeepers with a lot more experience than me report that queen problems are increasing.

Drone laying queen ...

Drone laying queen …

Perhaps the issues with fecundity and drone laying are related to toxic levels of miticides in commercial foundation? It’s certainly known that these residues reduce drone sperm fertility significantly. I intend to return to this topic sometime during the approaching winter … perhaps in time to encourage the use of some foundationless frames for (fertile) drone production 😉

In the ’16/’17 COLOSS data, natural disasters accounted for 1.6% of all overwintered colonies (so ~7.5% of losses), queen problems resulted in the loss of 5.1% of colonies (i.e. ~24% of losses) and the remainder (14.1% of colonies, ~68% of losses) just died.

Just died?

We’ll return to natural disasters (but not bears or honey badgers) and queen problems shortly. What about the majority of losses in which the colony ‘just died’?

If you discuss colony post-mortems with beekeepers they sometimes divide the ‘just died’ category (i.e. those not readily attributable to failed queens, marauding grizzlies or tsunamis) into four groups:

  • disease
  • isolation starvation
  • starvation
  • don’t know 

The most important disease associated with overwintering colony losses is high levels of Deformed wing virus (DWV). This results from uncontrolled or inadequately controlled Varroa infestation. For any new readers of this site, please refer back to many of the articles I’ve already written on Varroa management 8.

I strongly suspect that a significant proportion of the reported isolation starvation is actually also due to disease, rather than isolation per se.

A consequence of high levels of DWV is that winter bees die prematurely. Consequently, the colony shrinks faster than it would otherwise do. It starts the size of a basketball but (too) rapidly ends up the size of a grapefruit … or an orange.

Isolation starvation and disease

The small cluster is then unable to remain in contact with stores, and so starves. 

Yes, the colony died from ‘isolation starvation’, but the cause was the high levels of Varroa and the viruses it transmits.

Isolation starvation ...

Isolation starvation …

What about regular starvation?

Not because the cluster became isolated from the stores, but simply because they had insufficient stores to get through the winter.

Whose fault was that?

And the last category, the “don’t knows”?

I bet most of these are due to high levels of Varroa and DWV as well 🙁

Yes, there will be other reasons … but probably not a huge number. 

What’s more … if you don’t know the reason for the colony loss there’s very little you can do to mitigate against it in future seasons.

And, other than wild and increasingly vague speculation, there’s little I can write about if the reason for the loss remains unknown 9.

Avoiding winter losses

So, let’s rationalise those earlier lists into the probable (known) major causes of overwintering colony losses:

  • natural disasters
  • queen problems
  • starvation
  • disease (but probably mainly DWV and Varroa

As the long, hot days of summer gradually shorten and cool as early autumn approaches, you should be thinking about each of these potential causes of overwintering colony loss … and doing what you can to ensure it doesn’t happen to you (or, more correctly, your bees).

Ardnamurchan autumn

Ardnamurchan autumn

Some are easier to deal with than others.

Here’s a whistle-stop tour of some more specific problems and some practical solutions 10. Some, all or none may apply to your bees – it depends upon your location, your climate, your experience and future plans as a beekeeper. 

Natural disasters

These fall into two broad groups:

  • things you can do almost nothing about (but might be able to avoid)
  • things you can relatively easily solve

Flooding, falling trees, lightning, landslides, earthquakes, volcanoes, meteor strikes etc. all fall into the first group.

If you can avoid them, do. 

Your local council will have information on areas at risk from flooding. There are also searchable maps available from SEPA. Do not underestimate the severity of some of the recent flooding. Some parts of Scotland and Northern England had 600 mm of rain in two days in 2015.

You might be surprised (and from an insurance aspect, devastated) at the classification of some areas now ‘at risk’. 

Where did Noah keep his bees? In his Ark hive.

Where did Noah keep his bees? In his Ark hive.

Consider moving hives to higher ground before the winter rains start. One consequence of climate change is that heavy rainfall is now ~20% heavier than it was a few decades ago. This means that floods occur more frequently, are more extensive and the water levels rise faster. You might not have a chance to move the hives if flooding does occur,

More rain and stronger winds (particularly before leaf fall) mean more trees will come down. You might be able to identify trees potentially at risk from falling. It makes sense to remove them (or site your hives elsewhere). 

No risk of this larch tree falling on my hives

Lightning, earthquakes, volcanoes, meteor strikes … all a possibility though I would 11 probably worry about Varroa and woodpeckers first 😉

Solvable natural disasters

The ‘solvable’ natural disasters include preventing your colonies being robbed by other bees or wasps. Or ransacked by mice or woodpeckers after the first hard frosts start. A solution to many of these are ‘reduced size entrances’ which either enable the colony to better defend itself, or physically restricts access to critters.

The L-shaped ‘kewl floors‘ I use prevent mice from accessing the brood box. They are also easier for the colony to defend from bees/wasps, but can also easily be reduced in size with a narrow piece of hardwood. If you don’t use these types of floor you should probably use a mouseguard.

Polyhives and polythene

Polyhives and polythene …

Woodpeckers 12 need to cling onto the outside of the hive to hammer their way through the side. You can either place a wire mesh cage around the hive, or wrap the box in something like damp proof membrane (or polythene) to prevent them gaining purchase on the side walls.

Keep off Woody

Keep off Woody

Doing both is probably overkill though 😉

Strong colonies

Before we move onto queen problems – though it is related – it’s worth emphasising that an even better solution to prevent robbing by bees or wasps is to maintain really strong colonies.

Strong colonies with a well balanced population of bees can almost always defend themselves successfully against wasps and robbing bees.

Nucs, that are both weaker and – at least shortly after being made up – unbalanced, are far less able to defend themselves and need some sort of access restriction.

By ‘balanced’ I mean that the numbers and proportions of bees fulfilling the various roles in the nucleus colony are reflective of a full hive e.g. nurse bees, foragers, guard bees. 

Reduced entrance ...

Reduced entrance …

But the benefits of strong colonies are far greater than just being able to prevent wasps or robbing bees. There is compelling scientific evidence that strong colonies overwinter better

I don’t mean strong summer colonies, I mean colonies that are strong in the late autumn when they are fully populated with the winter bees.

Almost the entire complement of bees in the hive are replaced between late summer and late autumn. Remember that a really strong summer colony may not be strong in the winter if Varroa and virus levels have not been controlled.

How do you ensure your colonies are strong?

  1. Minimise disease by controlling Varroa levels in early autumn to guarantee the all-important winter bees are reared without being exposed to high levels of DWV.
  2. Try and use a miticide treatment that does not reduce the laying rate of the queen.
  3. Avoid blocking the brood nest with stores where the queen should be laying eggs.
  4. Requeen your colonies regularly. Young queens lay more eggs later into the autumn. As a consequence the colonies have increased populations of winter bees.
  5. Unite weak colonies (assuming they are disease-free) with stronger colonies. The former may well not survive anyway, and the latter will have a better chance of surviving if it is even stronger – see below. 
  6. Use local bees. There’s good evidence that local bees (i.e. reared locally, not imported from elsewhere) overwinter better, not least because they produce stronger colonies.

Uniting – take your losses in the autumn

My regular colony inspections every 7-10 days during May and June are pretty much abandoned by July. The risk of swarming is very much reduced after the ‘June gap’ in my experience. 

I still check the colonies periodically and I’m usually still rearing queens. However, the rigour with which I check for queen cells is much reduced. By July my colonies are usually committed to single-mindedly filling the supers with summer nectar.

They are already making their own preparations for the long winter ahead.

Although the inspections are less rigorous, I do keep a careful watch on the strength of each colony. Often this is directly related to the number of supers I’ve had to pile on top.

Colonies that are underperforming, and – more specifically – understrength are almost always united with a stronger colony.

An Abelo/Swienty hybrid hive ...

An Abelo/Swienty hybrid hive …

Experience has taught me that an understrength colony is usually more trouble than it’s worth. If it’s disease-free it may well overwinter reasonably well. However, it’s likely to start brood rearing more slowly and build up less well. It may also need more mollycoddling 13 in the autumn e.g. protection from wasps or robbing bees.

However, a colony that is not flourishing in the summer is much more likely to struggle and fail during the winter. Perhaps the queen is not quite ‘firing on all cylinders’ and laying at a really good rate, or she might be poorly mated.

Far better that the workforce contributes to strengthening another hive, rather than collect an underwhelming amount of honey before entering the winter and eventually becoming a statistic.

My winter losses are low and, over the last decade, reducing.

That’s partly because my Varroa management is reasonably thorough.

However, it’s probably mainly due to ensuring only strong colonies go into the winter in the first place.

Newspaper

I’ve dealt with uniting in several previous posts.

It’s a two minute job. 

You remove the queen from the weak colony, stack one brood box over the other separated by a sheet or two of newspaper with a very small (~3mm) hole in the middle. Add the roof and leave them to get on with things.

I don’t think it makes any difference whether the strong colony goes on the top or the bottom.

I place the colony I’m moving above the box I’m uniting it with. My – wildly unscientific – rationale being that the bees in the top box will have to negotiate the route to the hive entrance and, in doing so, will help them orientate to the new location faster 14.

If you unite colonies early or late in the day most foragers will be ‘at home’ so not too many bees will return to find their hive missing.

If there are supers on one or both hives you can separate them with newspaper as well. Alternatively, use a clearer the day before to empty the supers prior to uniting the colonies. You can then add back the supers you want and redistribute the remainder to other hives in the apiary.

Successful uniting ...

Successful uniting …

Don’t be in too much of a hurry to check for successful uniting.

Leave them a week. The last thing you want is for the queen to get killed in an unseemly melee caused by you disturbing them before they have properly settled.

Done properly, uniting is almost foolproof. I reckon over 95% of colonies I unite are successful.

That’s all folks … more on ‘Cutting your losses’ next week 🙂


Notes

At just over 3000 words this post got a bit out of control … I’ll deal with more significant queen problems, feeding colonies, the weather and some miscellaneous ‘odds and sods’ next week.

Irrational Varroa control

About a month ago I wrote a post on rational Varroa control. I define this as choosing a miticide appropriate for your colony 1 and environment, administering it properly and at the right time so providing the maximum benefit to your bees. The long term goal of rational Varroa control is reduced colony losses due to mite-transmitted viruses.

Primarily this means reduced overwintering losses due to Deformed wing virus.

I regularly give talks on this topic. In these I provide a few brief examples of the misuse of miticides. Some of these examples originate – anonymised – in questions I’ve received in previous talks or by email. Some are from surveys of miticide usage.

Winter is coming – miticide treatments and fondant on

A few (a very few) are from direct personal experience 🙁

I’m aware that these examples qualify as ‘misuse’ because I’ve read up about the active ingredient in miticides and have a reasonable understanding of how, when and why they work.

Misuse could probably be broken down into three divisions:

  • Incorrect usage that does not reduce the efficacy of the treatment. 
  • Using the miticide in a manner that significantly reduces its efficacy, but otherwise does little or no harm.
  • Incorrect usage that has no impact upon the mite population and/or that significantly harms the colony and/or spoils the honey.

After a short 2 preamble, I’ll outline some of the more glaring examples. Keep these in mind when planning your mite control strategy and you should see improved winter survival, better spring build up, stronger summer colonies … and more honey 🙂

What can I use?

The Veterinary Medicines Directorate maintain a list of miticides approved for use on honey bees (and other animals). It’s a large database and I find the easiest way to see the current product list – which changes quite regularly as things are added and removed – is to use the search facility.

VMD database search … bees, budgies and bearded dragons

Simply select ‘Bees’ from species list (they’re between ‘Bearded dragons’ and ‘Budgies’!) and then hit the big Run search button at the bottom.

There are currently 31 approved products though this contains a number of duplicates as Great Britain and Northern Ireland are listed as separate territories.

In my view, the two most important useful columns in the returned table are the ‘Active substance‘ and the ‘Aligned product‘. 

The active substance is the chemical in the miticide that is responsible for killing mites. 

Look down the list. There are relatively few different active substances, present alone or in combination.

I currently count just five.

And then there were five … VMD approved miticides classified by active ingredient

Thymol, amitraz, pyrethroids, formic acid and oxalic acid.

I’m ignoring the minor components like eucalyptus oil that, individually, do not have high levels of miticidal activity.

Flumethrin and tau-fluvalinate (the active ingredients of Bayvarol and Apistan respectively) are both pyrethroids and have the same mode of action (and, more importantly, resistance to one usually confers resistance to the other).

What about management methods to control mites?

The VMD database lists the miticidal products approved for treating honey bees. 

They don’t list management techniques like drone brood uncapping or small cell foundation, or the application of non-toxic compounds like dusting with icing sugar.

There’s nothing to stop you using these approaches.

However, first conduct some sort of cost-benefit analysis to determine if they are worthwhile.

Is the disruption to the colony, or the potential tainting of honey stores, justified by the reduction of Varroa numbers?

For example, Randy Oliver has done some analysis of the impact of sugar dusting and shows that – at best and with weekly applications – it might be able to hold Varroa levels steady. You can download his Excel calculations and play with some of the assumptions.

If you increase the percentage of mites capped in cells from 60% (which is probably rather conservative) and decrease the percentage of mites dislodged by sugar dusting from 40% (which is probably rather aggressive) then mite replication rapidly outruns the control method applied.

That’s a questionable ‘benefit’ in return for the disruption of blowing 120 g of icing sugar into the hive every 7 days 3 … but there’s nothing to stop you using it as a control method.

Though it might not do much controlling … 🙁

I choose miticides that kill at least 90% of mites when used properly. I prefer to use miticides once or twice during the season, rather than dabbling every other week or month.

What can’t or shouldn’t I use?

You cannot use things like fenpyroximate, spirotetramat or spirodiclofen.

What?

These are used for mite or tick treatment of other animals or plants 4. They have been shown to be effective against Varroa (though at high levels they may also kill bees) but are not approved for use. 

Remember also that VMD approval involves both the compound and the mode by which it is administered.

Amitraz is approved for use as the active ingredient in Apivar strips. However, in Great Britain and Northern Ireland you cannot fumigate colonies with amitraz … which, as Apiwarol, is a popular treatment method in Poland.

There are some (perhaps surprising) ‘restrictions’ in terms of approved usage. Api-Bioxal can be used to trickle treat twice per season, but only used once for vaporisation.

Why?

This is one of the many oddities buried in the depths of the VMD database.

Api-Bioxal is not approved for spray administration, but Oxuvar is. Both have the same active ingredient. 

No wonder beekeepers find this confusing … 🙁

Read the instructions and other documentation

Miticides approved by the VMD come with documentation. This takes the form of the instructions written (often in really tiny print) on the packet. The other thing written on the packet is the ‘use by’ date. 

Apivar instructions – duration of treatment

Keep the packaging 🙂

Read the instructions for use.

It may seem like an obvious thing to suggest but at least a third of the questions I get asked are answered in the documentation.

Part of the problem is the wording that’s often used or the apparent (and sometimes real) contradictions between the instructions provided for two miticides that have the same active ingredient and formulation 5

Api-Bioxal additional documentation

For more legible documentation you could refer again to the VMD database of approved miticides. If you following the Aligned product’ link you will have access to several additional pieces of information, most useful of which are the Summary of product characteristics and the Product literature. The latter is a copy of the literature in a very easy-to-read format, without any fancy colours or tiny fonts.

Keep records

You should keep records of when you treat and what you treat with, including batch numbers. I simply keep the packet the miticide was supplied in. I know when I treated as my copious 6 notes record the dates … and allow me to work out when the period of treatment is finished.

Apivar (but the batch number is on the back of the packet)

Even easier … just take a photograph of the empty packet, but make sure you include the batch number in the shot.

Examples of miticide misuse

It’s not possible to provide a comprehensive set of examples of miticide misuse (or Irrational Varroa control) … at least not in ~2000 words 😉 – after all, there might be one or two ways to use a miticide properly, but thousands of ways it could be used improperly.

Here is a non-exclusive and far-from-comprehensive list of miticide misuse or bad practice. 

Don’t do this at home … or in the out apiary 😉

Too little

Use the correct dose. Using less than the recommended dose ensures that some mites will survive. This may lead to the development of mite resistance.

Some beekeepers have been known to add a single strip of Apivar to colonies in preparation for going to the heather moors.

This is bad practice.

Even worse … some have been found leaving the strip in the hive while they were at the heather. Is that forgetfulness or just reckless? The honey will be tainted  and the long-term exposure to low levels of amitraz may contribute to the development of resistance 7.

Too late

Remember that the goal of the late summer treatment is to prevent the developing winter bees from being exposed to Varroa and the viruses that the mite transmits.

If you treat too late in the season you may well kill lots of mites, but the winter bees will already have been exposed.

Mite levels (solid lines) when treated July to November and timing of winter bee production.

In the graph above, treating in mid-October will kill significantly more mites than treating in mid-August.

But that’s not really the goal.

An October treatment will kill more mites because they’ve been breeding like rabbits throughout September.

And, what have they been reproducing on? Your developing winter bee pupae 🙁

Winter bee production is not an all or nothing event. The colony does not switch from producing summer bees to winter bees on a particular date. As late summer segues into early autumn an increasing proportion of the developing brood will be winter bees.

It’s your responsibility to ensure that enough of them are protected from Varroa so that they can lead a long and protective life, getting the colony through until February or March next year.

Too much

Many miticides are reasonably well tolerated by bees. Nevertheless, overdosing is also to be avoided. If you read the product characteristics for Api-Bioxal for example it states that:

Significantly higher bee mortality was observed in hives that received double (by sublimation) or triple (by trickling) dosages of product. In addition, when overdosed, the over-wintering  capacity of colonies was diminished and there may be detrimental effects on colony development in the future 8.

Remember that nucleus colonies are smaller than full sized colonies. It’s not unusual for a  beekeeper to administer a full dose to what is essentially a half-sized colony. 

In the case of Amitraz, ‘overdosing’ is well tolerated and two strips in a nucleus colony is unlikely to do the bees any harm.

However, the same cannot be said of MAQS. The product characteristics for MAQS specifically state that it should not be used for colonies with less than 6 frames of brood.

Too high

The formic acid-containing MAQS is poorly tolerated by the colony at high ambient temperatures. The literature suggests it should not be used when the peak daily temperature might exceed 29.5°C.

I’ve never used MAQS. I’m told by other beekeepers that they’ve had problems with queen losses at temperatures as ‘low’ as 25°C.

Remember, when you add the MAQS strips they need to be in the hive for 7 days. You therefore need to check the forecast for the week ahead before starting treatment.

Too low

Alison Gray and Magnus Peterson (both at Strathclyde University) have conducted surveys of Scottish beekeepers for about the last 15 years, summaries of which appear annually in The Scottish Beekeeper. Results are also collated into the COLOSS analysis. 

The surveys have evolved over the years, but have included questions on the type and timing of treatment.

I read some of these carefully when I returned to live in Scotland (in 2015). I was interested to see what other beekeepers were using. One thing that surprised me was the amount and timing of thymol (Apiguard) treatment.

For example, in the 2014 report (PDF), ~25% of treatments used were thymol, with 60% of these being applied in September, October and November.

A quick check of the Apiguard Product characteristics turns up the following statement:

Do not use the product when the maximum daily temperature expected during the treatment is lower than 15°C or when the colony activity is very low or when temperature is above 40°C.

I don’t know anywhere in Scotland in where the maximum daily temperature exceeds 15°C for four weeks (the duration of treatment) between September and November. 

I checked a personal weather station a couple of miles from my apiaries in Fife … in October 2013 the average temperature was 11.5°C, but the maximum failed to reach 15°C on 10 days through the month.

Inevitably, the efficacy of treatment would be reduced.

West coast temperatures 26/7/21 to 26/8/21 … 7 days fail to reach 15°C making Apiguard a very poor choice here.

You need good long range weather forecasting skills. Prior experience of what might be expected can really help your planning in these circumstances. 

Too long

Do not leave Apivar strips in the colony longer than stated in the instructions. 

How long is that?

Again, from the Product characteristics documentation:

If brood is not present or at its lowest level, the strips can be removed after 6 weeks of treatment. If brood is present, leave the strips in place for 10 weeks and remove the strips at the end of treatment.

Strips added to a colony with a young queen (and therefore laying well, and late into the year) on the day this article appears 9 should probably be removed in the first week of November.

Used and removed Apivar strips

That’s not a great time of year to be lifting roofs and prising up crownboards.

However, the alternative is worse. If you leave the strips in situ you ensure that any surviving mites (and there will be some) are continuously exposed to a low level of amitraz … perfect conditions to help select for resistance 🙁

Too short

This is much the same as too little (see above). If you remove the miticide before the correct period of time has elapsed then some mites will escape treatment.

No miticide is 100% effective. However, why store up problems for the future by unnecessarily reducing the efficacy of the treatment?

Remember … the only good mite is a dead mite

Too old

Miticides are not inexpensive. It is therefore very tempting to save and re-use opened packets. 

For example, beekeepers with just a couple of hives still have to purchase a packet of Apivar 10 for an eye-watering £31. It must be very tempting to tuck the unused portion away in the shed for the next time.

And, when they do, how many do a before and after count of phoretic mites to confirm that the treatment worked?

I suspect very few. 

If they did they might be in for a bit of a disappointment.

Amitraz, the active ingredient in Apivar strips, is quite unstable and rapidly degrades. I’ve used strips from previously opened packets and observed that they were significantly less effective 11.

What I should have done was read the Product characteristics documentation where it clearly states:

Shelf life after first opening the immediate packaging: use immediately and discard
any unused product.

Similar problems (short or non-existent shelf life) apply to some of the oxalic acid-containing solutions, in this case because they degrade to product hydroxymethylfurfural which is toxic to bees at high concentrations (see the notes on this at the end of the post of preparing Api-Bioxal for trickle treating).

One and two hive owners should coordinate their purchases and treatment of colonies to avoid wasting money.

Coordinated treatment has additional benefits, which neatly takes me to the final topic … 

Too few

OK, these titles are getting a bit contrived now, but this is the last one 😉

Treat all the colonies in the apiary simultaneously. I’ve written extensively about drifting and robbing. Both activities redistribute adult bees and the mites piggybacking on them around the apiary (or the wider environment).

If you treat just one colony it will soon 12 become reinfested with mites from neighbouring colonies.

If you don’t treat one colony and it develops high mite levels in autumn, perhaps due to a late surge in brood rearing, it will shed mites (hitching a ride on young bees going on orientation flights) to your adjacent treated colonies.

A final note on Apistan and pyrethroid-containing miticides

Resistance to Apistan is widespread. It’s so widespread that the National Bee Unit apparently stopped keeping records a decade or so ago.

Apistan is a very effective miticide … against mites that are sensitive. 

If you intend to use Apistan (or any of the approved pyrethroid-containing miticides) I would strongly suggest determining the level of infestation before treatment, and confirming a 90+% reduction in mite levels after treatment.

Unfortunately, just counting the dead mites on the Varroa tray is not evidence that the treatment has been effective.

If you find 1000 dead mites on the tray it just tells you is that you have 1000 mites less in the colony.

There may still be 9000 left if the treatment was only 10% effective … 

That’s not going to end well 🙁


 

Rational Varroa control

It’s the end of July … in the next two to three weeks the first eggs will be laid that will develop into the winter bees that get your colonies through to next spring. Protecting these winter bees is necessary to prevent overwintering colony losses.

I’ve written and lectured extensively on Varroa control and related topics for at least 5 years. The following article is published in August’s BBKA Newsletter and The Scottish Beekeeper. It provides an overview of what I term rational Varroa control.

I define this as effective mite management based upon our current understanding of the biology of bees and Varroa. The goal of this control is to minimise winter losses due to Varroa and viruses.

It is not a recipe with easy to follow if this, then that instructions. Neither does it provide a calendar-based guide of what to do and when to do it.

It does not even tell you what you should use for mite control.

Instead it focuses on the principles … understanding these will enable you to implement control strategies that help your bees, in your environment, survive.

This version is hyperlinked to additional, more expansive, posts on particular topics, is slightly better illustrated than those that appeared in print and contains some additional footnotes with caveats and exceptions.


Introduction

Despite almost 30 years experience of Varroa in the UK, this ectoparasitic mite of honey bees remains the greatest threat to bees and beekeeping. With the exception of those fortunate to live in mite-free regions, all beekeepers must manage the mite population in their hives or risk losing the colony to the viruses transmitted when Varroa feeds on developing pupae. 

Fortunately, Varroa control is relatively straightforward; there are a range of approved and effective miticides that – used appropriately – reduce mite infestation levels significantly. The key words in that last sentence are ‘approved’, ‘effective’ and ‘used appropriately’. In reality annual colony losses, primarily occurring in the winter, often exceed 20% (Figure 1) and may be significantly higher in long or harsh winters 1. Many of these losses are attributable to Varroa and viruses. It is therefore clear that many beekeepers are not successful in managing Varroa; either they are not treating at all, or they are treating inappropriately.

Figure 1. BBKA winter survival survey – larger studies (COLOSS and BIP) often show much higher losses

This article is primarily aimed at relatively inexperienced beekeepers, but may also help the more experienced who still suffer with high levels of winter losses. It emphasises the importance of two, correctly timed, appropriate miticide treatments per season that should ensure colony survival. It is not going to deal with treatments of questionable or minor efficacy. These include the use of small cell foundation, drone brood culling or sugar dusting. These may reduce mite levels, but insufficiently to benefit colony health. Nor will it discuss the use of any miticides (or application methods) that are not approved by the Veterinary Medicines Directorate. I will also not discuss treatment-free beekeeping, selection of mite-resistant bees or advanced colony manipulations like queen trapping. In my view any or all of these could or should be tried … but only once a beekeeper can routinely successfully overwinter colonies using strategies similar to those described here.

The problem

Varroa is an ectoparasitic mite that feeds on developing honey bee pupae. During feeding it transmits a range of honey bee viruses, the most important of which is deformed wing virus (DWV). DWV is present in honey bees in the absence of Varroa. In our studies, using sensitive PCR-based detection methods, we never detect bees – even those from mite-free regions of Scotland – without DWV. The virus is transmitted horizontally between bees during trophallaxis, and vertically from drones or the queen through sperm or eggs. These routes of transmission are rarely if ever associated with any significant levels of disease and virus only replicates to modest levels (perhaps 1-10 thousand viruses per bee). However, when Varroa transmits DWV the virus bypasses the bee’s natural defence mechanisms and replicates to very high levels in recipient pupae (billions per pupa, 1 – 10 million times higher than in unparasitised pupae). Studies from our laboratory have shown that ~75% of pupae with these high virus loads either do not emerge, or emerge exhibiting the characteristic “deformed wings” that give the virus its name (Figure 2; Gusachenko et al., Viruses 2020, 12, 532; doi:10.3390/v12050532). The ~25% of bees that do emerge and appear ‘normal’ exhibit a range of symptoms including reduced fitness, impaired learning and reduced foraging. However, most importantly they also exhibit reduced longevity. During the summer this is probably not critical; the lifespan of a worker is only ~6 weeks and, assuming the queen is laying well, there are thousands of half-sisters around with more being produced every day.

DWV symptoms

Figure 2. DWV symptoms

But during the winter, brood rearing either stops completely or drops to a very low level. The bees reared from late summer onwards are physiologically very different. These are the ‘winter bees’, also termed the diutinus bees (from the Latin meaning long-lived). Physiologically these bees resemble juvenile workers and they can survive for many months. And they need to … it is these bees that get the colony through the autumn, winter and into the following spring. They protect the queen, they thermoregulate the hive and, usually around the winter solstice, they start to rear small amounts of new brood for the season ahead.

The longevity of the bees in the hive in winter is critical to colony survival. If the winter bees have high DWV levels their longevity is reduced (in addition to the reduced numbers due to overt disease or non-viability). This means that the winter cluster shrinks in size faster than it would do otherwise. With reduced numbers of bees it cannot keep brood warm enough and so the colony fails to expand early the following season. In cold spells it may be unable to reach the food stores resulting in the colony perishing from ‘isolation starvation’. It may not be able to maintain sufficient warmth to protect the queen, or may simply freeze to death.

The goal of rational Varroa control

Successful overwintering requires lots of winter bees. The size of the winter cluster is directly related to its survival chances. Therefore the goal of rational Varroa control is to prevent the winter bees from being exposed to mites and mite-transmitted viruses during their development. Winter bee production is induced by a range of factors including photoperiod, nectar and pollen availability, brood and forager pheromones. Together these induce slowed behavioural maturation of the winter bees. This is not like flicking a switch, instead it is a seamless transition occurring as late summer segues into early autumn (Figure 3). Winter bee production is also influenced by the queen. Young queens lay later into the autumn, so increasing the numbers of winter bees. 

Figure 3. Colony age structure from August to December.

It is important to note that these events are environment-driven, not calendar-driven. It will not happen at precisely the same time each year, or at the same time in different locations (or latitudes) each year.

To protect these winter bees the colony needs to be treated with an effective miticide before the majority of the winter bees are produced. This ensures that the developing winter bee pupae are not parasitised by virus-laden mites and so do not suffer from reduced longevity. 

When are winter bees produced in the UK? 

Unfortunately, I’m not aware of any direct studies of this. Scientists in Bern (49.9°N) in 2007/08, where the average temperatures in November and December were ~3°C, showed that the Varroa- and virus-reduced longevity of bees was first measurable in mid-November, 50 days after emergence. By extrapolation, the eggs must have been laid in the first week of September. 

Doing large scale experiments of Varroa control is time-consuming and subject to the vagaries of the climate (and, as a molecular virologist, beyond me in terms of the resources needed). I have therefore used the well-established BEEHAVE program of colony development (from scientists in the University of Exeter; https://beehave-model.net/) to model the numbers of developing and adult bees, and the mite numbers in a colony. BEEHAVE by default uses environmental parameters (climate and forage) based upon data from Rothamsted (51.8°N). Using results from this model system, the bees present in the hive at the end of December – by definition the diutinus winter bees – were produced from eggs laid from early/mid August (Figure 4).

Whatever the precise date – and it will vary from season to season as indicated above – at some point in September the adult bee population starts to be entirely replaced with winter bees. Large numbers of these need to live until the following February or March to ensure the colony survives and is able to build up again once the queen starts laying.

When to treat – late summer

The numbers of pupae and adult bees present in the colony are plotted in Figure 4 using dashed lines. Adult bee number decrease in early spring until new brood is reared. The influence of the ‘June gap’ on pupal numbers is obvious. Brood rearing gradually tails off from early July and stops altogether sometime in late October or early November. The shaded area represents the period of winter bee production – from early/mid August until brood rearing stops. 

Figure 4. Winter bee production and mite levels – see key and text for further details

Mite levels are indicated using solid lines. The impact on the mite population of treating in the middle of each month from July to November is shown (arrowed and labelled J, A, S, O and N) using the colours green, blue, red, cyan and black respectively. The absolute numbers of bees or mites is irrelevant, but bees (pupae and adults) are plotted on the left, and mites on the right hand axis, so they cannot be directly compared. The miticide treatment modelled was ‘applied’ for one month and was 95% effective, reproducing many licensed and approved products.

Mite levels peak in the colony in late September to October. If treatment does not occur until this time of the season then the majority of winter bees will have been reared in the presence of large amounts of mites. Unsurprisingly, the earlier the treatment is applied, the lower the mite levels during the period of winter bee production. 

Rational Varroa control therefore involves treatment soon after the summer blossom honey is removed from the hive, so maximising the winter bees produced in the presence of low mite numbers. If you leave treatment until mid-September, you risk exposing the majority of winter bees to high levels of Varroa in the hive. If your primary crop is heather honey, which is not harvested until September, you may need to consider treating earlier in the summer – for example during the brood break when requeening or during swarm control.

Why treat in midwinter?

A key point to notice from Figure 4 is that, paradoxically, the earlier the miticide is applied, the higher the mite levels are at the end of the year. Compare the August (blue) and October (cyan) lines at year end for example. This is because mites that survive treatment – and some always do – subsequently reproduce in the small amount of brood reared late in the season. This is what necessitates a ‘midwinter’ treatment. Without it, mite levels increase inexorably year upon year, and cannot be controlled by a single late-summer treatment. Beekeepers bragging on social media that their mite drop after the winter treatment was zero probably applied the summer treatment too late to effectively protect their winter bees.

And when is midwinter?

Historically beekeepers apply the ‘midwinter’ treatment between Christmas and New Year. This is probably too late. The usual miticide used at this time is oxalic acid, a ‘one shot’ treatment that is ineffective against mites in capped cells. For maximum efficacy this must be applied when the colony is broodless. Brood rearing usually starts (if it ends at all, again this is climate-dependent) around the winter solstice. By delaying treatment until a lull in the Christmas festivities or even early January some mites will already be inaccessible in capped cells. 

Figure 5. Biscuit coloured (or a bit darker) cappings indicating brood rearing in this colony

I check my colonies for brood – either by looking for biscuit-coloured cappings on the Varroa tray (Figure 5) or by quickly inspecting frames in the centre of the cluster – and usually treat in November or very early December. If I cannot check visually I apply the treatment during the first extended cold spell of the winter. By treating when the colony is broodless I can be certain my intervention will have maximal effect.

What to treat with?

I have deliberately avoided – other than mentioning oxalic acid – specific miticides. Rational Varroa control involves the choice of an appropriate miticide and its correct application. Examples of incorrect or inappropriate miticide choice include; use of Apistan when resistance is known to be very widespread, use of Apiguard when the average ambient temperature is below 15°C (which makes Apiguard of little use for effective control in much of Scotland) or the use of Api-Bioxal when there is capped brood present. In addition, use of a half-dose or a reduced period of application will both reduce efficacy and potentially lead to the selection of resistance in the mite population. Used correctly – the right dose at the right time and for the right duration – the majority of the currently licensed miticides are be capable of reducing mite levels by over 90%. If they do not, use one that does. Miticide choice should be dictated by your environment and the state of the colony.

All together now

Most beekeepers grossly underestimate the movement of bees (and their phoretic mites) between colonies. Numerous studies have shown that drifting and (to an even greater extent) robbing can result in the transfer of large numbers of mites from adjacent and, in the case of robbing, more distant colonies. 

Gaffer tape apiary

Figure 6. Gaffer tape apiary …

Rational Varroa control therefore involves treating all colonies within an apiary, and ideally the wider landscape, in a coordinated manner. In communal association apiaries (Figure 6), where beekeeping experience and therefore colony management and health can vary significantly, this is particularly important. Coordinated treatment is only relevant in late summer when bees are freely flying.

Swarms

Swarms originating from unmanaged or poorly managed colonies will have high mite levels. The bee population in a swarm is biased towards younger bees; these are the bees that phoretic mites preferentially associate with. Studies have shown that ~35% of the mite population of a colony leaves with the swarm.

Figure 7. Varroa treatment of a new swarm in a bait hive…

Since swarms contain no sealed brood until ~9 days after they are hived oxalic acid is the most appropriate treatment. I usually treat them using vaporised oxalic acid late in the evening soon after they are hived (Figure 7). Even casts get this treatment and I have not experienced any issues with the queen not subsequently mating successfully. I’d prefer to have a queenless low-mite colony than a queenright one potentially riddled with Varroa.

Midseason mite treatment

The text above describes the mite management strategies I have used for several years. I apply Apivar immediately the summer honey is removed and treat with oxalic acid when broodless before the end of the year. Doing this has almost never required any additional midseason treatments; if mite levels are sufficiently low at the beginning of the season they cannot rise to dangerous levels before the late summer treatment. I still get winter colony losses, but they are almost always due to poor queen mating and rarely due to Varroa and viruses.

Figure 8. Queenright splits and the window(s) of opportunity

However, if midseason treatments are required – either because there are signs of overt infestation, because regular mite counts have shown there is a problem, or to have low mite colonies after the heather honey is collected – then there are two choices. Treat with MAQS which is approved for use when there are supers on the hive and, more importantly, is effective against mites in capped cells 2. Alternatively, treat during swarm control. With care, the majority of splits (e.g. the Pagden artificial swarm or the nucleus method) can be performed to give a broodless period for both the queenright (Figure 8) and queenless partitions. That being the case, a single application of an oxalic acid-containing miticide can be very effective in controlling the mite population.

Costs

Many beekeepers complain about the cost of licensed and approved miticides. However, some perspective is needed. A colony with low levels of mites will be more likely to survive overwinter, so reducing the costs of replacement bees. In addition, a healthy colony will be a stronger colony, and therefore much more likely to produce a good crop of honey (and potentially an additional nuc). Over the last 5-6 years my miticide costs are equivalent to one jar of honey per colony per year. This is an insignificant amount to pay for healthy colonies.

Summary

Rational Varroa control requires an understanding of the goals of treatment – protecting the winter bees and minimising mite levels for the beginning of the following season – and an appreciation of how this can best be achieved using miticides appropriate for the environment and the state of the colony. Like so much of beekeeping, it involves judgement of the colony and will vary from season to season and your location. I’ve applied my midwinter treatment as early as the end of October or as late as mid-December, reflecting variation in timing of the broodless period. Rational Varroa control also involves an understanding of the biology of bees and an awareness of the influence of beekeeping (e.g. crowding colonies in apiaries which increases mite and disease transmission) on our bees. However, none of this is difficult, expensive or time consuming … and the benefits in terms of strong, healthy, productive colonies are considerable.


 

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 😉