Category Archives: Varroa

Midseason mite management

The Varroa mite and the potpourri of viruses it transmits are probably the greatest threat to our bees. The number of mites in the colony increases during the spring and summer, feeding and breeding on sealed brood.

Pupa (blue) and mite (red) numbers

In early/mid autumn mite levels reach their peak as the laying rate of the queen decreases. Consequently the number of mites per pupa increases significantly. The bees that are reared at this time of year are the overwintering workers, physiologically-adapted to get the colony through the winter.

The protection of these developing overwintering bees is critical and explains why an early autumn application of a suitable miticide is recommended … or usually essential.

And, although this might appear illogical, if you treat early enough to protect the winter bees you should also treat during a broodless period in midwinter. This is necessary because mite replication goes on into the autumn (while the colony continues to rear brood). If you omit the winter treatment the colony starts with a higher mite load the following season.

And you know what mites mean

Mites in midseason

Under certain circumstances mite levels can increase to dangerous levels 1 much earlier in the season than shown in the graph above.

What circumstances?

I can think of two major reasons 2. Firstly, if the colony starts the season with higher than desirable mite levels (this is why you treat midwinter). Secondly, if the mites are acquired by the colony from other colonies i.e. by infested bees drifting between colonies or by your bees robbing a mite infested colony.

Don’t underestimate the impact these events can have on mite levels. A strong colony robbing out a weak, heavily infested, collapsing colony can acquire dozens of mites a day.

The robbed colony may not be in your apiary. It could be a mile away across the fields in an apiary owned by a treatment-free 3 aficionado or from a pathogen-rich feral colony in the church tower.

How do you identify midseason mite problems?

You need to monitor mite levels, actively and/or passively. The latter includes periodic counts of mites that fall through an open mesh floor onto a Varroa board. The National Bee Unit has a handy – though not necessarily accurate – calculator to determine the total mite levels in the colony based on the Varroa drop.

Out, damn'd mite ...

Out, damn’d mite …

Don’t rely on the NBU calculator. A host of factors are likely to influence the natural Varroa drop. For example, if the laying rate of the queen is decreasing because there’s no nectar coming in there will be fewer larvae at the right stage to parasitise … consequently the natural drop (which originates from phoretic mites) will increase.

And vice versa.

Active monitoring includes uncapping drone brood or doing a sugar roll or alcohol wash to dislodge phoretic mites.

Overt disease

But in addition to looking for mites you should also keep a close eye on workers during routine inspections. If you see bees showing obvious signs of deformed wing virus (DWV) symptoms then you need to intervene to reduce mite levels.

High levels of DWV

High levels of DWV …

During our studies of DWV we have placed mite-free 4 colonies into a communal apiary. Infested drone cells were identified during routine uncapping within 2 weeks of our colony being introduced. Even more striking, symptomatic workers could be seen in the colony within 11 weeks.

Treatment options

Midseason mite management is more problematic than the late summer/early autumn and midwinter treatments.

Firstly, the colony will (or should) have good levels of sealed brood.

Secondly, there might be a nectar flow on and the colony is hopefully laden with supers.

The combination of these two factors is the issue.

If there is brood in the colony the majority (up to 90%) of mites will be hiding under the protective cappings feasting on sealed pupae.

Of course, exactly the same situation prevails in late summer/early autumn. This is why the majority of approved treatments – Apistan (don’t), Apivar, Apiguard etc. – need to be used for at least 4-6 weeks. This covers multiple brood cycles, so ensuring that the capped Varroa are released and (hopefully) slaughtered.

Which brings us to the second problem. All of those named treatments should not be used when there is a flow on or when there are supers on the hive. This is to avoid tainting (contaminating) the honey.

And, if you think about it, there’s unlikely to be a 4-6 week window between early May and late August during which there is not a nectar flow.

MAQS

The only high-efficacy miticide approved for use when supers are present is MAQS 5.

The active ingredient in MAQS is formic acid which is the only miticide capable of penetrating the cappings to kill Varroa in sealed brood 6. Because MAQS penetrates the cappings the treatment window is only 7 days long.

I have not used MAQS and so cannot comment on its use. The reason I’ve not used it is because of the problems many beekeepers have reported with queen losses or increased bee mortality. The Veterinary Medicines Directorate MAQS Summary of the product characteristics provides advice on how to avoid these problems.

Kill and cure isn’t the option I choose 😉 7

Of course, many beekeepers have used MAQS without problems.

So, what other strategies are available?

Oxalic acid Api-Bioxal

Many beekeepers these days – if you read the online forums – would recommend oxalic acid 8.

I’ve already discussed the oxalic acid-containing treatments extensively.

Importantly, these treatments only target phoretic mites, not those within capped cells.

Trickled oxalic acid is toxic to unsealed brood and so is a poor choice for a brood-rearing colony.

Varroa counts

In contrast, sublimated (vaporised) oxalic acid is tolerated well by the colony and does not harm open brood. Thomas Radetzki demonstrated it continued to be effective for about a week after administration, presumably due to its deposition on all internal surfaces of the hive. My daily mite counts of treated colonies support this conclusion.

Consequently beekeepers have empirically developed methods to treat brooding colonies multiple times with vaporised oxalic acid Api-Bioxal to kill mites released from capped cells.

The first method I’m aware of published for this was by Hivemaker on the Beekeeping Forum. There may well be earlier reports. Hivemaker recommended three or four doses at five day intervals if there is brood present.

This works well 9 but is it compatible with supers on the hive and a honey flow?

What do you mean by compatible?

The VMD Api-Bioxal Summary of product characteristics 10 specifically states “Don’t treat hives with super in position or during honey flow”.

That is about as definitive as possible.

Another one for the extractor ...

Another one for the extractor …

Some vapoholics (correctly) would argue that honey naturally contains oxalic acid. Untreated honey contains variable amounts of oxalic acid; 8-119 mg/kg in one study 11 or up to 400 mg/kg in a large sample of Italian honeys according to Franco Mutinelli 12.

It should be noted that these levels are significantly less than many vegetables.

In addition, Thomas Radetzki demonstrated that oxalic acid levels in spring honey from OA vaporised colonies (the previous autumn) were not different from those in untreated colonies. 

Therefore surely it’s OK to treat when the supers are present?

Absence of evidence is not evidence of absence

There are a few additional studies that have shown no marked rise in OA concentrations in honey post treatment. One of the problems with these studies is that the delay between treatment and honey testing is not clear and is often not stated 13.

Consider what the minimum potential delay between treatment and honey harvesting would be if it were allowed or recommended.

One day 14.

No one has (yet) tested OA concentrations in honey immediately following treatment, or the (presumable) decline in OA levels in the days, weeks and months after treatment. Is it linear over time? Does it flatline and then drop precipitously or does it drop precipitously and then remain at a very low (background) level?

Oxalic acid levels over time post treatment … it’s anyones guess

How does temperature influence this? What about colony strength and activity?

Frankly, without this information we’re just guessing.

Why risk it?

I try and produce the very best quality honey possible for friends, family and customers.

The last thing I would want to risk is inadvertently producing OA-contaminated honey.

Do I know what this tastes like? 15

No, and I’d prefer not to find out.

Formic acid and thymol have been shown to taint honey and my contention is that thorough studies to properly test this have yet to be conducted for oxalic acid.

Until they are – and unless they are statistically compelling – I will not treat colonies with supers present … and I think those that recommend you do are unwise.

What are the options?

Other than MAQS there are no treatments suitable for use when the honey supers are on. If there’s a good nectar flow and a mite-infested colony you have to make a judgement call.

Will the colony be seriously damaged if you delay treatment further?

Quite possibly.

Which is more valuable 16, the honey or the bees?

One option is to treat, hopefully save the colony and feed the honey back to the bees for winter (nothing wrong with this approach … make sure you label the supers clearly!).

Another approach might be to clear then remove the supers to another colony, then treat the original one.

However, if you choose to delay treatment consider the other colonies in your own or neighbouring apiaries. They are at risk as well.

Finally, prevention is better than cure. Timely application of an effective treatment in late summer and midwinter should be sufficient, particularly if all colonies in a geographic area are coordinately treated to minimise the impact of robbing and drifting.

I’ve got two more articles planned on midseason mite management for when the colony is broodless, or can be engineered to be broodless 17.


 

Bait hive guide

Spring this year is developing well. Even here on the chilly east coast of Scotland colonies are looking good and flying strongly when the sun is out. Large amounts of pollen are being taken in and there’s every sign that the hives are queenright and rearing lots of brood 1.

It’s too soon 2 to open the colonies but it’s not too soon to be thinking about the consequences of the inevitable continued expansion over the next few weeks.

Most healthy colonies will make preparations to swarm, often between late April and mid-June. The timing varies depending upon a host of factors including colony strength, climate, weather, forage, build up and beekeeper interventions.

Swarm prevention and control

You, like all responsible beekeepers, will use appropriate swarm prevention methods. Supers added early, ensure the brood box has space for laying etc.

In due course, once the colony gets bigger and stronger, you’ll notice queen cells and immediately deploy your chosen swarm control method e.g. the classic Pagden artificial swarm, the nucleus method I described last week, Demaree, vertical splits or – if you’re feeling ambitious – a Taranov board 3.

Which will of course be totally successful 😉

But just in case it isn’t …

… and just in case the beekeeper a couple of fields away is forgetful, unobservant, clumsy, on holiday, in prison or has some other half-baked excuse, be prepared for swarms.

As an aside, other than just walking around the fields, you can easily find hives near you by searching on Google maps and you can get an idea of the local beekeeper density 4 using the National Bee Unit’s Beebase.

You might think you know all the local beekeepers through your association, but it’s surprising the number who just ‘do their own thing’.

Swarms

This isn’t the place to discuss swarms in much detail. Here’s a quick reminder:

  1. The colony ‘decides’ to swarm and starts to make queen cells.
  2. Almost certainly, scout bees start to check out likely sites the swarm could occupy in the future 5.
  3. The swarm leaves the hive on the first calm, warm, sunny day, usually early in the afternoon, once the queen cells are capped. The prime swarm contains the mated, laying queen and about 75% of the worker bees 6.
  4. The swarm gathers around the queen and sets up a bivouac hanging from a convenient spot (tree, gatepost, bush, fence etc.) near to the hive. They rarely move more than 50 metres. It’s worth emphasising here that the spot they choose is convenient to the bees, but may be at the top of a 60 foot cypress. It may not be particularly convenient for the beekeeper 😉
  5. Scout bees continue to check out likely final sites to establish the new colony, returning to the swarm and ‘persuading’ other scouts (by doing a version of the waggle dance) so that, finally, a consensus is reached. This consensus is essentially based upon the suitability of the sites being surveyed.
  6. The scout bees lead the swarm to the new location, they move in and establish a new colony.

If you’re lucky you will be able to recapture the swarm if the spot they choose for their bivouac is within reach, not above a stream, in a huge thorny bush or on an electricity pylon.

A small swarm ...

A small swarm …

I say ‘recapture’ because, since the bivouac is usually near the issuing hive, it’s probably come from one of your own hives (unless you are snooping around your neighbouring apiaries 7).

But what if you miss the bivouacked swarm? Or if your neighbour misses it?

Those bees are going to look for a suitable location to set up home.

If you provide a suitable location, you can get them to hive themselves without the grief of falling off a ladder, toppling into a stream, getting lacerated with thorns or electrocution

This is where the bait hive comes in. Leave a couple in suitable locations and you can lure your own and other swarms to them.

Freebees 🙂

What do scouts look for?

The scout bees look for the following:

  1. A dark empty void with a volume of about 40 litres.
  2. Ideally located reasonably high up.
  3. A solid floor.
  4. A small entrance of about 10cm2, at the bottom of the void, ideally south facing.
  5. Something that ‘smells’ of bees.

What I’ve just described is … a used beehive 8.

More specifically, it’s a single National brood box (or two stacked supers) with a solid floor and a roof, containing one old dark frame of drawn comb pushed up against the back wall.

No stores, no pollen 9, just a manky old dark comb. The sort of thing you should be turning into firelighters.

That’s all you need.

However, you can improve things by giving the bees somewhere to start drawing comb and siting the hive in a location that makes your beekeeping easier.

Des Res

The first thing swarms do when they move in is start drawing comb. You can populate the bait hive with a few foundationless frames so they’ve got somewhere to start.

Bait hive ...

Bait hive …

In my view foundationless frames are much better than frames with foundation for bait hives. The scout bees measure the size of the void by flying around randomly inside 10. If you have sheets of foundation they’ll crash into it frequently, effectively giving them the impression that the void is smaller than it really is. And therefore making it less attractive to the scouts.

You can improve the smell of the hive by adding a little lemongrass oil to the top bar of one of the frames. Don’t overdo it. A drop or two every 7-10 days is more than ample.

If you do use foundationless frames make sure the hive is level. If you don’t the comb will be drawn at an angle to the frames which makes everything harder work later in the season. Your smartphone probably contains a spirit level function that makes levelling the bait hive very easy.

Location

But not if it’s above head height, or you’re teetering on top of a ladder …

It was Tom Seeley who worked out most things about scout bees and swarms (see his excellent book Honeybee Democracy). This included the observations that they favoured bait hives situated high up.

Believe me, it’s a whole lot easier if the bait hive is on a standard hive stand. It’s easier to level, it’s easier to check and it’s easier – in due course – to retrieve.

Bait hive

Bait hive

I’ve previously discussed how far swarms prefer to move from their original hive. Contrary to popular opinion (and perhaps illogically) they tend to prefer to move shorter distances i.e. 20m >> 200m >> 400m. However, there are also studies that show swarms moving a kilometre or more.

Don’t get hung up on this detail. Stick out a bait hive or two and, if there are swarming colonies in range, they’ll find it.

I always leave a bait hive in my apiaries and one or two in odd corners of the garden. In the last few years I’ve never failed to attract swarms to the bait hives, and know for certain that some have moved in from over a mile away as the bee flies (thanks Emma 😉 ).

Mites and swarms

Assuming you don’t have the luxury of living in Varroa-free areas of the UK (or anywhere in Australia) then the incoming swarm will contain mites. Studies have shown that ~35% of the mite population of a colony leaves with the swarm.

But, for about the first week after the swarm sets up home in your bait hive, what’s missing from the new arrivals is sealed brood. Therefore the mites are all phoretic.

Do not delay. Treat the swarm with an appropriate miticide to knock back the mite population by ~95%. An oxalic acid-containing treatment is ideal. Single dose, relatively inexpensive, easy to administer (trickled or vaporised) and well tolerated by the bees.

Varroa treatment ...

Varroa treatment …

You have eight days from the swarm arriving to there being sealed brood in the colony

Far better to slaughter the mites now. In a few months their numbers will have increased exponentially and the majority will be in capped cells and more difficult to treat.


 

Hanging around

I’ve recently discussed the misnamed ‘phoretic’ phase in the life cycle of Varroa destructor. Here we’ll briefly explore some features of this important, but non-reproductive, phase. It’s important because, as I’ll show, it influences subsequent mite reproduction.

I won’t rehash the life cycle of Varroa in detail as I’ve covered it previouslyVarroa is an ectoparasite of honey bees. It reproduces in capped cells, feeding on the developing pupa. A mated female mite enters the cell a few hours before capping and she and her incestuously mated daughters are released when the bee emerges.

The longer pupal development takes, the more progeny mites are produced, so Varroa has evolved to preferentially infest drone brood.

A smorgasbord of viruses

As an ectoparasite of honey bees, Varroa is responsible for the transmission of a smorgasbord of pathogenic viruses to the developing pupa. Subsequent virus replication, particularly by the aptly-named deformed wing virus, can result in developmental deformities.

Worker bee with DWV symptoms

Worker bee with DWV symptoms

Emerging workers with deformities are rapidly ejected from the hive. Other infested workers, with high viral levels, have reduced longevity. This is probably what accounts for the majority of overwintering colony losses. It may also explain so-called ‘isolation starvation‘.

The ‘phoretic’ phase

Mites outside capped cells are termed ‘phoretic’ mites. Recent studies have indicated that these mites are feeding on the workers to which they are attached. The same studies have shaken the long-held assumption that Varroa feeds on haemolymph 1 by rather neatly demonstrating that it is the fat body tissue of the bee that is the plat du jour.

The duration of the ‘phoretic’ phase is dependent upon the state of the colony. Since it is defined as the phase in which mites are not associated with developing pupae, in a broodless colony all the mites are ‘phoretic’. Under these circumstances mites remain ‘phoretic’ until either brood is produced, or they fall (or are groomed) off and drop through the open mesh floor.

The duration of the ‘phoretic’ phase

In colonies with ample brood the ‘phoretic’ phase is, on average, 6 days in length. The range often quoted is 4 – 11 days. The absolute figure must depend upon a number of factors. These include the chance of a mite encountering a late-stage larva. This is presumably influenced by the amount of suitable-aged brood in the colony and – because there is a division of labour in the hive – the type of bee upon which the mite is riding around the colony on.

For the purpose of this post we’ll consider bees of three ages – newly emerged, nurse bees and foragers. Newly emerged bees (days 1-2 post emergence) clean cells and nurse bees (days 3-11) feed developing larvae. Older bees are involved in wax production (days 12-17) and foraging (>18 days until death).

Logic would dictate that mites would ‘choose’ 2 to associate with bees that bring them into contact with developing larvae of the right age to infest.

Do they?

Hanging around with nurses

Xie et al., (2016)3 assembled artificial colonies containing equal numbers of new bees, nurse bees and foragers, all suitably marked so their age was known. These colonies were provided with a queen, open brood and stores. At the start of the experiment the colonies had 1500 bees and low Varroa levels.

The scientists then introduced 200 ‘phoretic’ mites from another colony 4 and left the colonies for 48 hours. They then age-sorted the bees and harvested all the ‘phoretic’ mites by washing them off in alcohol.

'Phoretic' mites prefer nurse bees

‘Phoretic’ mites prefer nurse bees

On average, ~16% of the nurse bees had ‘phoretic’ mites attached. In contrast, only ~10% of the foragers and ~5% of the new bees had mites. These studies involved seven individual experiments, in two countries and two separate years, using a different source colony for the mites. The statistics are significant.

So mites prefer nurse bees.

Is this ‘simply’ 5 because the nurse bees are more likely to bring the mite into close proximity with a suitably-aged larvae?

Or does associating with, and presumably feeding on, nurse bees have other benefits for the mite?

Mite fecundity and fitness

Fecundity is the reproductive productiveness 6 of an organism.

We’ve obliquely tackled this subject recently. Using an in vitro artificial ‘feed packet’ system, Ramsey and colleagues demonstrated that mites fed on the fat body of bees laid more eggs i.e. they had higher fecundity.

Xie et al., also tested fecundity of ‘phoretic’ mites from newly emerged bees, nurse bees and foragers. They did this by manually harvesting mites after a 3 day ‘phoretic’ phase, adding them to a pre-pupa and then counting the number of progeny female mites 9 days later.

Mite fecundity and fitness

Mite fecundity and fitness

‘Phoretic’ mites from nurse bees exhibited higher fecundity (more female offspring), higher fitness (more mature female offspring) and lower infertility (female mites that did not generate offspring).

So evolution has elegantly resulted in ‘phoretic’ mites associating with the right type of bee to bring them close to developing larvae (upon which they reproduce) and made them better able to reproduce once they get there.

Why do Varroa mites prefer nurse bees?

This is the title of the Xie et al., paper.

Xie et al., sort of answer the question they posed in the title. What they don’t do is explain why ‘phoretic’ mites on nurse bees are more fecund. However, the recent Ramsey paper suggests that this may be because nurse bees have a larger fat body and higher levels of vitellogenin.

If they’re better fed perhaps they produce more viable offspring? 7.

Another known unknown (semiochemicals)

How do mites detect the differences between new, nurse and older bees?

Perhaps they ‘smell’ different?

Mites preferentially infest drone brood because it produces a range of methyl and ethyl esters of straight-chain fatty acids, in particular methyl palmitate.

Similarly, the preference for nurse bees might be explained by their production of another semiochemicals 8. If we could identify this semiochemical it might be possible to create a ‘sponge’ soaked in it that attracted all the mites in the colony.

A bit simplistic, but you get the idea.

In reality, it’s likely that nurse bees are identified by the relative strengths of a range of semiochemicals produced by bees of different ages.

In reality it’s also likely that dropping a ‘sponge’ soaked in eau de nurse bee into the colony could unbalance all sorts of other events in the hive … 🙁

I told you it wasn’t simple.


Rattus norvegicus

Rattus norvegicus

Hanging Around is the fifth track on Rattus norvegicus, the 1977 debut album of one of the finest rock/punk bands of all time, The Stranglers. The album also includes the incomparable Peaches and (Get a) Grip (On yourself), both of which were released as singles.

No more heroes from the same year, but a different album, is also a classic by the same band.

You had to be there … I was 😉

 

 

Pedantically not phoresy

The life cycle of the ectoparasitic mite Varroa destructor essentially consists of two stages. The first is within the capped cell, where reproduction takes place. The second occurs outside the capped cell when the recently-mated female progeny mites matures while riding around the colony attached to a nurse bee.

Almost without exception this second stage is termed the phoretic phase.

It isn’t.

Phoresy

Phoretic is an adjective of the word phoresy. Phoresy is derived from the French phorésie which, in turn, has its etymological origins in the Ancient Greek word φορησις.

And φορησις means being carried.

Which partly explains why the correct definition of the word phoresy is:

An association between two organisms in which one is carried on the body of the other, without being a parasite [OED]

Phoresy has been in use for about a century, with the word phoretic first being recorded in the Annals of the Entomological Society of America (25:79) in 1932:

It is possible, as suggested by Banks (1915), that such young mites are phoretic, being carried about from place to place on the host’s surfaces.

And, no, they weren’t discussing Varroa.

“Without being a parasite”

These are the critical words in the dictionary definition of phoresy which makes the use of the word phoretic incorrect when referring to mites on nurse bees.

Because mites on nurse bees are feeding – or at least a significant proportion 1 of them are.

They are therefore being parasitic and so shouldn’t be described as phoretic.

Om, nom, nom 2

Last week I discussed the recent Samual Ramsey paper presenting studies supporting the feasting of Varroa on the fat body of bees.

In the study they harvested bees from a heavily mite-infested hive and recorded the location on the bee to which the mite was attached.

The majority were attached to the left underside of the abdomen. More specifically, the mite was wedged underneath the third abdominal tergite 3.

What were they doing there? Hiding?

Yes … but let’s have a closer look.

Ramsey and colleagues removed some of the mites and used a scanning electron microscope to examine the attachment point on the bee. Underneath the tergite there is a soft membrane. The imprint of the body of the mite was clearly visible on the membrane.

Varroa feeding location on adult bee

Scanning EM of Varroa feeding location on adult bee

The footpads of the mite were left attached to the membrane (left image, white arrows), straddling an obvious wound where the mouthparts had pierced the membrane (black arrow). Between them, the inverted W shape is presumably the imprint of the lower carapace of the mite.

The close-up image on the right even shows grooves at the wound site consistent with the mouthparts of the mite.

These mites were feeding.

Extraoral digestion

Varroa belongs to the order (a level of classification) Mesostigmata. Most mesostigmatids feed using a process termed extraoral digestion.

Extraoral digestion has also been termed ‘solid-to-liquid’ feeding. It involves the injection of potent hydrolytic enzymes which digest solid tissue, converting it to a semi-solid that can be easily ingested. It can reduce the time needed to feed and it increases the nutrient density of the consumed food.

If Varroa fed on haemolymph it wouldn’t need to use extraoral digestion. Instead it would need all sorts of adaptations to a high volume, low nutrient diet. Varroa doesn’t have these. It has a simple tube-like gut parts of which lack enzymatic activity … implying that digestion occurs elsewhere.

A picture is worth a thousand words

Do the images of feeding mites support the use of extraoral digestion?

EM cross-section of Varroa feeding

EM cross-section of Varroa feeding

The image above 4 shows the cross-section of a Varroa (V), wedged under the tergite (Te), feeding through a hole (arrow in the enlargement on the right) in the membrane (M). The fat body (FB) is immediately underneath the membrane. The scale bar is incorrectly labelled 5.

A close-up of the wound site shows further evidence for extraoral digestion.

Feeding wound at higher magnification

Feeding wound at higher magnification

Beneath the wound site (C, arrow) are remnants of fat body cells (white arrow) and bacteria (black arrow; of two types, shown in D). A closer look still at the remnants of the fat body (E and F) shows cell nuclear debris (blue arrows) and lipid droplets (red arrows).

These images are entirely consistent with extraoral digestion of fat body tissue by feeding Varroa. The presence of bacteria near the wound suggests that bacterial infection may result from Varroa feeding, possibly further contributing to disease in bees.

So, pedantically it’s not phoresy

So-called phoretic mites, unless they’re on the thorax or head of the bee, are not really phoretic. They are being carried about, but they are also likely feeding. By definition that excludes them from being phoretic.

Instead they are ectoparasites of adult bees.

What are the chances that beekeepers will stop using the term phoretic?

Slim to none I’d predict 6.

And, of course, it doesn’t really matter what the correct term for them is.

What’s more important is that beekeepers remember that it’s at this stage that mites are susceptible to all miticides.

The June gap

But it’s also worth thinking about the potential impact of brood breaks.

During brood breaks all the mites in the colony must be ‘phoretic’.

Generally, the majority of the mites in a hive are in capped cells. Depending upon the stage of the season, the egg-laying rate of the queen and other factors, up to 90% of the mites are associated with developing pupae.

But as the laying rate dwindles more and more mites are released from cells and become ‘phoretic’, unable to find a suitable late-stage larva to infest.

And which bees do the mites associate with?

Nurse bees primarily, for reasons I’ll discuss in the future. But – spoiler alert – one of the reasons is likely to be that they have a larger fat body.

So, a mid-season brood break (e.g. the ‘June gap’) is likely to result in lots more nurse bees becoming both the carriers and the dinner of the mite population.

Some or many of the nurse bee cohort may perish, perhaps from damage to the fat body or from the viruses acquired from the mite. However, bees exhibit phenotypic plasticity, meaning that older bees can revert to being nurse bees when the queen starts laying again.

Late season brood breaks

In late summer mite levels are usually at their highest in the hive. A brood break occurring now will release a very large number of mites to parasitise the adult bee population.

Presumably these mites select the bees best able to support them 7.

And which bees are these? The nurse bees of course. But it’s also worth remembering that there are key physiological similarities between nurse bees and winter bees. Both have low levels of juvenile hormone and high levels of vitellogenin (stored in the fat body).

So I’d bet that the ‘phoretic’ mites during a late season brood break would also preferentially associate with any early-produced winter bees.

Furthermore, once the queen starts laying again – perhaps in early/mid-autumn – the winter bees being produced would be subjected to the double-whammy of high levels of mite infestation and potential damage from ‘phoretic’ mites.

Practical considerations

More work is required to model or actually measure the impact of late season brood breaks, high levels of ‘phoretic’ mites, nurse bee numbers and winter bee development.

Compare two colonies of a similar size with a similar mite load, treated at the same time in early autumn with an appropriate miticide. If one of them experienced a late summer brood break (pre-treatment) and consequent high levels of ‘phoretic’ mites, does this reduce the chances of the colony surviving overwinter?

Who knows? Lots and lots of variables …

Fundamentally, it remains important to treat colonies early enough to protect the winter bee population. You’ve heard this from me before and you’ll hear it again.

However, it’s something to think about and I can see ways in which it might influence the strategy and timing of mite control used. I’ll return to this sometime in the future.


 

Chewin’ the fat

A little over a year ago reports started to circulate of a study showing that Varroa feed on the fat body of bees rather than on haemolymph.

Having worked in Glasgow through the early noughties the title of this post was a no-brainer and an outline draft was written in December 2017. However, the peer-reviewed paper wasn’t published until last month, so it’s only now we’ve got the chance to judge the study and consider its implications.

Varroa feed on hameolymph, right?

Historically this was the accepted dogma. However, the experimental data supporting this conclusion – based upon labelling bees with radioactive isotopes and seeing what the mites acquired after feeding – was really not definitive. The experiments had been done in the 1970’s and the specificity of the labelling was a bit dubious. In addition, during the intervening period scientists had determined that, unlike vertebrate blood which is rich in cells and nutrients 1, haemolymph has little of either and is actually a pretty lousy food source.

In addition, and somewhat more circumstantially, Varroa control using chemotherapeutics fed to bees (and subsequently taken up by the mite during feeding) had been relatively disappointing.

Perhaps these chemicals weren’t getting to the right tissues of the bee?

Perhaps Varroa don’t feed on haemolymph after all?

The Ramsey study

This new study reports three independent experiments that, together, indicate that Varroa actually feed on the fat body of bees, rather than on haemolymph. The paper is so-called ‘open access’, so anyone can access it and therefore I’ll just provide a synopsis of the important bits.

The questions Samual Ramsey and colleagues attempted to answer were:

  1. Where on the bee do mites feed? Is it primarily or exclusively near the fat body?
  2. When Varroa feeds, what host tissues are ingested?
  3. What sort of diet is required to maintain Varroa and allow their reproduction in vitro2.

Location, location, location

The authors counted phoretic mites on 104 bees. Over 95% of them were located on the underside of the body, predominantly on the left side of the bee, under the tergite or sternite3 on the third metasomal segment (i.e. the second visible segment of the abdomen).

Mite location on nurse bees

Mite location on nurse bees

This position is consistent with feeding on the fat body tissues which are most abundant under the inner ventral surface of the metasoma.

Seeing red

Bees were fed with Nile red, a lipophilic fluorescent stain that preferentially accumulates in the fat body. They co-fed bees with uranine, a differently coloured fluorophore that accumulates in the haemolymph. They then allowed mites to feed on the fluorescently labelled bees and subsequently photographed the mites under fluorescent light.

The rationale here was straightforward. If the mites fed on the fat body they would stain red due to taking up the Nile red stain.

Mites visualised after feeding on fluorescently labelled bees

Mites visualised after feeding on fluorescently labelled bees

Which they did.

It was notable that the red stain predominantly accumulated in the rectum and gut of the mite (image O above). The authors conducted all sorts of controls to confirm that the stains actually stained what they were supposed to – you can view these in the paper.

Babies!

In the final part of the study the authors maintained mites in vitro (in an incubator), feeding them on a diet containing increasing amounts of fat body or haemolymph. These are tricky experiments and in some way the least satisfactory part of the study.

Two results suggest that fat body was beneficial or essential to the mites. Firstly, only mites that had 50% or more fat body in the diet survived for 7 days. Secondly, there was a dose response to the amount of fat body in the diet and fecundity. Mites on a 100% fat body diet exhibited 40% fecundity, the highest level observed in the study.

What can we conclude from the Ramsey study

Of the three experiments presented, the Nile red fat body stain uptake by mites is reasonably compelling.

The feeding position study is essentially correlative, but there could be other interpretations of the data. For example, that location on the bee might be the least accessible to a ‘grooming’ bee. Perhaps it’s a survival mechanism?

Survival and fecundity in in vitro studies wasn’t great. However, in defence of the authors, fecundity of mites under natural conditions can be as low as 40% and is not higher than 80%. Not all mites have baby mites. Thankfully.

Only 20% of the mites survived one week under in vitro conditions, even on a 100% fat body diet. In contrast, mites fed haemolymph alone died within 48 hours. This poor level of survival was surprising and suggests other essential components of the diet were probably missing.

Other published studies have shown reasonable survival of Varroa for at least 3 days, with at least one report of mites surviving on flowers for up to 7 days. I’m also aware that other laboratories can maintain mites in vitro for longer than 7 days without using any honey bee-derived components in the diet.

Hang on … what is the fat body anyway?

The fat body is multi-functional. It has been compared to the vertebrate liver and adipose tissue. It acts as a major organ for nutrient storage, energy metabolism and detoxification of things like pesticides.

Vitellogenin made by and stored in the fat body reduces oxidative stress and is associated with extending the longevity of overwintering bees. The fat body also has critical roles in metamorphosis.

So, not only multi-functional, but also very important.

Significance of the results … is this a game changer?

This paper has been discussed online as a ‘game changer’. That’s probably a bit strong. Whilst the fluorescent stain uptake study is reasonably convincing it must be remembered that it was conducted on adult bees.

Do mites on pupae also feast on the fat body?

This will have to be determined in the future. It’s a more difficult experiment of course.

The other two studies, and a number of additional small observations I’ve not discussed here, are certainly supportive, but not alone hugely convincing. The in vitro study in particular will be interesting to compare with (currently unpublished) studies from other laboratories that do not use honey bee fat bodies in their mite feeding and maintenance diet.

Practical matters

Does it matter what part of the bee the mite feeds on?

Clearly it does for the mite, but what about the beekeeper?

I think this study is significant for the beekeeper for two reasons – the first will only be relevant if and when lipophilic miticides are developed, the second matters right now.

  1. Strategies are being developed to add highly specific miticides to the diet of bees which are then delivered to Varroa when the mite feeds. To date, these have been rather underwhelming in their performance. If Ramsey is right, modification of these miticides to make them lipophilic (like the Nile red fluorphore) will concentrate them in precisely the right place to ensure the mites get a lethal dose.
  2. A key product of the fat body is vitellogenin. The long-lived overwintering bees have high levels of vitellogenin. Mites feeding on, and depleting, the fat body would be expected to result in reduced vitellogenin levels in the bee 4. This would explain why high Varroa levels are associated with reduced longevity of winter bees and consequently increased overwintering colony losses.

The most important take home message

To prevent mites that feed on fat bodies from damaging vitellogenin production miticides have to be used early enough to protect the winter bees.

In the paper Ramsey makes the statement:

Simple reduction of mite loads late in the season to decrease the overwinter parasite load may not be enough, as it still allows for the mites to damage tissue critical to the process of overwintering …

Instead …

A treatment schedule that includes treatment in late summer or early fall before mites can significantly damage fat body in developing winter bees would likely be more effective.

Which is precisely the point I’ve made previously about treating early enough to protect winter bees.

What the Ramsey paper adds is the piece of the jigsaw possibly explaining why late summer treatment is so important.


Colophon

Chewin’ the Fat was a four-series Scottish comedy sketch show. It was broadcast from 1999 to 2002, with further Hogmanay specials until 2005. The show had a recurring cast of characters and sketches including The Big ManThe Banter BoysThe Lighthouse KeepersBallistic Bob and Taysiders in Space.

Gonna no' dae that

Gonna no’ dae that – The Lighthouse Keepers

Chewin’ the Fat was filmed in and around Glasgow (where I worked at the time) and the characters parodied a range of local ‘types’ … pretentious Kelvinsiders, Glaswegian gangsters, narcissistic golfers, The man from Kilmacolm, and shellsuit-wearing, chain-smoking, hard-drinking Glaswegian neds.

It was a bit rude and definitely an acquired taste. Without subtitles, some of the scenes would probably have been unintelligible south of the border.

Mites equal viruses

Healthy bees are happy bees 🙂

Sounds good doesn’t it?

Actually, there’s no evidence that bees display or perceive most of the emotions often attributed to them 1.

Happy? Who knows? But certainly not healthy ...

Happy? Who knows? But certainly not healthy …

A more accurate statement might be “Healthy bees are more productive, they are less likely to die overwinter, less likely to be robbed out by wasps or neighbouring strong colonies and their parasites and pathogens cannot threaten the health of other honey bee colonies or, through so-called-pathogen overspill, the health of other pollinators.”

More accurate?

Yes … but it doesn’t exactly trip off the tongue 😉

Whether it makes the bees happy or not, beekeepers have a responsibility to look after the health of their livestock. This includes controlling Varroa numbers to reduce the levels of pathogenic viruses in the hive.

How well are virus levels controlled if mite levels are reduced?

I’ll get to that in due course …

Midwinter mite massacre

The 2018 autumn was relatively mild through until mid/late November. In the absence of very early frosts colonies continued rearing brood.

We opened colonies in mid-November (for work) and found sealed brood, though it was clear that the laying rate of the queen was much-reduced.

These are ideal conditions for residual mite replication. Any mites that escaped the late summer/early autumn treatment (the ideal time to treat to protect the overwintering bees) continue to replicate, resulting in the colony starting the following season with a disappointingly high level of mites.

I’ve noted before that midwinter mite levels are paradoxically higher if you treat early enough in the autumn to protect the all-important winter bees.

Consequently, to start the year with minimal mite levels, I treat in midwinter with a trickled or vaporised oxalic acid-containing (OA) treatment.

A combination of colder weather (hard frosts in late November) and brood temperature measurements 2 indicated mid-December was a good time to treat.

Midwinter mite massacre

Midwinter mite massacre

18th December

In one of my apiaries ten colonies were treated. Some were definitely broodless (based upon Arnia hive monitoring). Others may have had brood, but colonies were not routinely checked.

Over the four day period after vaporising these ten colonies dropped a total of 92 mites. More than 50% of these were from just one double-brooded colony. Overwintering nucs 3 dropped no mites at all in the 12 days following treatment.

This was very encouraging. These are lower midwinter mite levels than I’ve seen since returning to Scotland in 2015.

The one colony with ‘high’ mite levels received two further treatments (on the 22nd and 27th) in an attempt to minimise the mite levels for the start of the season. Going by the strength of the colony and the debris on the Varroa tray it was presumed that this colony was still rearing brood.

Mite drop following the third treatment was negligible 4.

Why are mite levels so low?

I think it’s a combination of:

  • Luck
  • Use of natural, organic, bee-centric and biodynamic beekeeping methods
  • Varroa-resistant bees
  • Very tight control of mite numbers in the 2017/18 season, primarily by correctly timing the winter and the late-season autumn treatments. This is simply good colony management. Anyone can achieve this.
  • A brood break midseason and/or a broodless period when splitting colonies (both give opportunities for more phoretic mites to be lost through grooming). Undoubtedly beneficial but season-dependent. I’ll be discussing ways to exploit these events in posts next year.
  • A low density of beekeepers in Fife, so relatively little drifting or robbing of poorly managed colonies from neighbouring apiaries. Geography-dependent. Much easier in Fife than Warwickshire … and easier still in Lochaber.

And what do less mites mean?

Varroa is a threat to bee health because it transmits pathogenic viruses when feeding on developing pupae.

The most important of these viruses is deformed wing virus (DWV).

Generally, the higher the level of infestation with mites, the higher the viral load 5. This has been repeatedly demonstrated by studies from researchers working in the UK, Europe and the USA.

It is well-established that colonies with high viral loads have an increased chance of dying overwinter, due to the decreased longevity of bees infected with high levels of virus.

DWV symptoms

DWV symptoms

In our work apiaries we regularly measure DWV levels. For routine screening our limit of detection is around 1,000 viruses per bee.

We don’t actually count the viruses. They’re too small to see without an electron microscope 6.

Instead, we quantify the amount of the virus genetic material present 7, compare it to a set of standards and express it as ‘genome equivalents (GE)’.

Many of the bees tested this year contained ~103 (i.e. 1000) GE, which is extremely low. Bees from Varroa-free regions (e.g. Colonsay) carry similar levels of DWV.

Most of our colonies were at or close to this level of virus much of the 2018 season. This is 100-1,000 times lower than we often see even in apparently perfectly healthy colonies in other years or other apiaries.

For comparison, using the same assay we usually detect about 1010 (ten billion) DWV GE per bee in symptomatic adult bees from heavily mite-infested colonies.

So, less mites means less viruses which means healthier bees 🙂

And they might even be happier bees 😉

And your point is?

It’s worth remembering that the purpose of treating a colony with miticides is to reduce the transmission of viruses between bees. This transmission results in the amplification of DWV. This is why the timing of treatments is so important.

Yes, it’s always good to slaughter a few (or a few thousand 🙂 ) mites. However, far better massacre them when you need to protect particular populations of bees.

This includes the overwintering bees, raised in September, that get the colony through to the Spring.

Remember also that it ‘takes bees to make bees’ i.e. the rearing of new brood requires bees. Therefore strong colony build-up in Spring requires healthy workers rearing healthy brood.

This is why it’s important to minimise mite levels in midwinter when colonies are broodless.

What do most beekeepers do?

Fifteen months ago I published a post on the preparation of oxalic acid solutions for trickling colonies in midwinter.

Whatever the vapoholics on the online forums claim, trickling remains the easiest, quickest and least expensive way to treat colonies in midwinter 8.

The best time to treat in the winter is when the colony is broodless. Here in Fife, and often elsewhere, I believe that this usually occurs earlier in the winter than many beekeepers treat (if it happens at all … or if they treat at all).

I usually treat between the end of the third week in November and mid-December, at the end of the first extended cold period.

Oxalic acid preparation recipe page views

Oxalic acid preparation recipe page views

Looking at the page views for these oxalic acid recipes it looks as though many beekeepers treat after Christmas 9 … which may be suboptimal if colonies had a broodless period and now started rearing brood again.

Mine have.

This winter has been quite mild (at least at the time of writing) so there may yet be opportunities to treat really effectively during a broodless period.

Or the chance may have gone …


 

Responsibilities

In draughty church halls the length and breadth of the country potential apiarists are just starting their “Beginning beekeeping” courses run by local associations. The content of these courses varies a bit but usually contains (in no particular order):

  • The Beekeeping Year
  • The hive and/or beekeeping equipment
  • The life cycle of the honey bee
  • Colony inspections
  • Pests and diseases
  • Swarm prevention and control
  • Products of the hive

I’ve seen these courses from both sides. I took one before I started beekeeping and I’ve subsequently taught on them.

Although I’m not convinced the seven topics above are the optimal way to cover the basics of beekeeping (perhaps that’s something for a future post?), I am a strong supporter of the need to educate new beekeepers.

Theory and practice

You can learn some of the theoretical aspects of beekeeping on dark winter evenings. In my experience a liberal supply of tea and digestives hugely helps this learning process 😉

However, beekeeping is essentially a practical subject and any responsible association will offer apiary-based training sessions once the season starts. A good association will run these throughout the season, enabling beginners to experience all aspects of the beekeeping year.

Trainee beekeepers

Trainee beekeepers

If they don’t, they should (both run them and run them through the season).

The reason is simple … ‘hands on’ with the bees is a much better way of appreciating some of the most important characteristics of the colony. It’s strength and temperament, the rate at which it’s developing, the levels of stores etc.

But all this takes time. A couple of early-season apiary sessions might be held on cool evenings in failing light, or dodging Spring weekend showers. This means that ‘hive time’ is often restricted and beginners only get a small snapshot of the beekeeping season.

Curb your enthusiasm

Inevitably, many new beekeepers are desperate to get their own bees as soon as possible. After all, the season has started and there are kilograms of nectar out there waiting to be collected and converted into delicious honey for friends and family.

Demand for overwintered nucs is very high (usually significantly outstripping supply, meaning a considerable price premium) and a purchased colony, which should be strong and building up fast, becomes the property of someone who potentially has yet to see an open hive.

The seasonal nature of the hobby and the way we train beginners creates a very steep learning curve for new beekeepers 1. Almost as soon as they’re out of the classroom (or draughty church hall) they’re faced with the start of their first swarm season.

Queen cells ...

Queen cells …

Their inevitable – and completely understandable – enthusiasm to start practical beekeeping reaches a crescendo at a time when they are singularly poorly equipped to manage the colony 2.

What’s missing?

The emphasis on the theory and practical aspects of beekeeping is understandable. There’s a lot to learn in a relatively short time.

However, this focus on the practicalities often overlooks emphasising the responsibilities of beekeepers.

In the frenetic early-season enthusiasm to ‘become a beekeeper’ these might seem unimportant, superfluous or entirely obvious.

But they’re not.

Oil seed rape (OSR) ...

Oil seed rape (OSR) …

Later in the season the colony can become bad tempered, unmanageably large or ignored. Some or all of these happen with new (and not-so-new) beekeepers. The OSR goes over and colonies get stroppy, April’s 5-frame nuc “explodes” to occupy a towering double brood monstrosity or a new-found enthusiasm for dahlias or crown green bowls becomes all-consuming.

Bees? What bees? Have you seen my dahlias?

Bees? What bees? Have you seen my dahlias?

This is when the responsibilities of beekeepers become really important.

What are the responsibilities of beekeepers?

As I see it, as beekeepers we have responsibilities to:

  • The general public
  • Other beekeepers
  • The bees 3

As I stated above, these might seem entirely obvious. However, every year new beekeepers start with the best of intentions but some have a near-total lack of awareness of what these responsibilities are (or mean).

The general public

The combination of calm bees, careful handling and appropriate protective clothing means that bees essentially pose no risk to the beekeeper.

However, strange as it may seem to a beekeeper, some people are terrified of bees (mellisophobics). Others, due to adverse allergic reactions (anaphylactic shock), may have their lives endangered by bee stings. Finally – and thankfully by far the largest group – are the remainder of the public who should never feel bothered or threatened by our bees, whether we consider this a rational response or not.

What does this mean in terms of practical beekeeping? I think it can be distilled to just three points:

  1. Keep calm bees
  2. Keep bees and the public well-separated
  3. Restrict beekeeping activities to times when the public are not inconvenienced

The first point is sensible, whether or not there’s anyone else around. It makes beekeeping a much more relaxing and rewarding experience.

The second point involves either keeping bees in unfrequented locations (infinitely preferable) or ensuring that bees are forced to fly up and away from the hives (by suitable screening) and well-away from passers-by.

The final point is the most inconvenient, but also the most important. If there are members of the public around who might be bothered by your bees – walkers strolling across the field towards your apiary, kids playing in the garden next door – don’t open the hives.

My apiaries have generally been in large rural gardens, private farmland and very well screened. I’ve also kept bees in urban environments, with no problems from the neighbours. However, I have always maintained out apiaries to move my bees to should they exhibit poor temper. Additionally, I’d only conduct inspections when the adjacent gardens were empty … meaning inspections were often carried out in sub-optimal weather or late in the evening.

Finally, while many beekeepers consider the sight of a swarm is one of the truly great sights of beekeeping, this isn’t a sentiment shared by most non-beekeepers.

Swarm on a swing ... not ideal if it's in the next door garden

Swarm on a swing … not ideal if it’s in the next door garden

Keep non-swarmy bees, clip the queen and keep a bait hive prepared to lure any swarms that do emerge.

Other beekeepers

The responsibilities beekeepers have to other beekeepers are probably restricted to:

  1. Courtesy
  2. Disease

The first is straightforward. Don’t do things that negatively impact other beekeepers 4. For example, don’t plonk two dozen hives over the fence from an established apiary, unless you’ve first discussed it with the beekeeper and you’re both happy that the local forage is sufficient.

And, of course, don’t steal hives or colonies 5.

Disease is perhaps less obvious and more insidious. The health of your bees influences the health of other colonies in the area. Over short distances bees drift from one hive to another. Over much longer distances strong colonies can rob weaker colonies.

All these bee exchanges also move the parasites and diseases they carry between hives. This includes VarroaNosema, a panoply of pathogenic viruses and European and American foulbrood.

Of these, the foulbroods are statutory notifiable diseases and beekeepers are legally required to report suspected diseased colonies under the Bee Diseases and Pests Control Order 2006 (and amendments). Responsible beekeepers will register their apiaries on the National Bee Unit’s Beebase so they are notified of local outbreaks, and so the bee inspectors can check their colonies if there is a nearby outbreak.

National Bee Unit Beebase

National Bee Unit Beebase

Whilst not notifiable, the remaining parasites and pathogens are also best avoided … and certainly should not be foisted upon other local beekeepers.

If your colony is weak, disease-riddled and poorly managed it may get robbed-out by other local strong colonies. In doing so, your bees will transfer (some of) the pathogen load to the stronger colony.

That is irresponsible beekeeping.

US beekeepers use the term ‘mite bomb’ to refer to an unmanaged, Varroa-riddled, collapsing colony that introduces significantly higher mite levels to local strong colonies as it’s robbed. This is more extreme, but not dissimilar, to beekeepers that treat with miticides far too late in the season. Their colonies retain high mite levels and can spread them to nearby hives. One way to avoid this is to coordinately treat mites in the same geographic area.

The bees

Bees may or may not be classified as livestock. The standard definition 6 of “domestic animals kept on a farm for use or profit; esp. cattle, sheep, and pigs” is perhaps a little restrictive 7 so lets accept for the moment that they are livestock.

If you keep livestock you usually need to register them and vaccinate them, and you always need to look after their health, feed and transport them properly and generally take responsibility for them.

If you don’t look after their welfare you may be prosecuted.

Of course, bees are invertebrates, not mammals or animals with backbones. Legally invertebrates are not usually considered as animals in the Animal Welfare Act 2006 8 which defines the law on animal welfare.

But all these definitions are a distraction.

In my view, if you keep bees you have a responsibility to look after them properly.

Even if this isn’t a legal requirement, its a moral responsibility.

This responsibility to your bees includes – but is not restricted to – preventing and treating them for disease when appropriate and ensuring they have sufficient stores going into winter (and during periods with no nectar).

If you can’t do this perhaps take up crown green bowls instead.

Blimey, this is all getting a bit heavy isn’t it?

Bees are not ‘fit and forget’.

Actually, they’re quite the opposite.

Proper management means that there are certain things that must be done at a particular time. This includes treating for mites at the end of the summer honey season, feeding the colony up for winter and swarm prevention and control.

If you work abroad for April and May or if you holiday on the Maldives for six weeks every autumn you’re unlikely to become a successful beekeeper.

Powder blue surgeonfish, Maldives

Bees? What bees? They’ll be OK …

And you’re certainly unlikely to be a responsible beekeeper.

You might start with bees, but you’re unlikely to keep them …

What prompted this post? A combination of things … cabin fever and online discussion forum posts from beekeepers puzzling why their colonies all died (no mite treatment, ever) or starved (no feeding before winter) or hadn’t been inspected in the last 15 months (“I’ve been busy”).

It’s going to be a long winter … 9


 

Know your enemy

What less appropriate time is there, as we enter the festive season of goodwill, to provide a brief account of the incestuous and disease-riddled life cycle of the Varroa mite?

Happy Christmas 🙂

Scanning electron micrograph of Varroa destructor

Scanning electron micrograph of Varroa destructor

Varroa is the biggest enemy of bees, beekeepers and beekeeping. During the replication cycle the mite transfers a smorgasbord of viruses to developing pupae. One of these viruses, deformed wing virus (DWV), although well-tolerated in the absence of Varroa 1replicates to devastatingly high levels and is pathogenic when transferred by the mite.

Without colony management methods to control Varroa, mite and virus replication will eventually kill the colony.

I’ve written extensively on ways to control Varroa. Most of these have focused on early autumn and midwinter treatment regimes. However, next season I’m hoping to discuss some alternative strategies and will need to reference aspects of the life cycle of Varroa … hence this post.

What is Varroa?

Varroa destructor is a distant relative of spiders, both being members of the class Arachnida … the joint-legged invertebrates (arthropods). It was originally (and remains) an external parasite (ectoparasite) of Apis cerana (the Eastern honey bee) and – following cross-species transfer a century or so ago – Apis mellifera, ‘our’ Western honey bee.

Apis cerana, having co-evolved with Varroa, has a number of strategies to minimise the detrimental consequences of being parasitised by the mite.

Apis mellifera doesn’t. Simple as that 2.

One hundred years is the blink of an eye in evolutionary terms and, whilst there are bees that have partial solutions – largely behavioural (small colonies and very swarmy) – they’re probably unable to collect meaningful amounts of honey 3.

Varroa-resistant honey bees will probably evolve (as much as anything is predictable in evolution) but not in my time as a beekeeper … or possibly not until Voyager 2 leaves the Oort Cloud 4.

And there’s no guarantee they’ll be any use whatsoever for beekeeping …

The replication cycle of Varroa

Varroa has no free-living stage during the life-cycle. The adult mated female mite exhibits two distinct phases during the life-cycle. It has a phoretic phase on adult bees and a reproductive phase within sealed (‘capped’) worker and drone brood cells. Male mites only ever exist within sealed brood cells.

I’m going to discuss phoretic mites in a separate post. I’ll concentrate here on the replication cycle.

The mated female mite enters a cell 15-50 hours before brood capping. Drone brood is chosen preferentially (at ~10-fold greater rates than worker brood) and entered earlier. Depending upon the time of the season and the levels of mites and brood, up to 70-90% of mites in the colony occupy capped cells.

The first egg is laid ~70 hours after cell capping. This egg is unfertilized and develops into a haploid male mite. Subsequent eggs are fertilised, diploid, and so develop into female mites. These are laid at ~30 hour intervals.

The replication cycle of Varroa

The replication cycle of Varroa

Worker and drone brood take different times to develop. Therefore a typical reproductive cycle involves five eggs being laid in worker brood and six in drone brood. Not all of these eggs mature, their development being curtailed by the bee emerging as an adult.

There are all sorts of developmental stages involved in getting from an egg to a mature unfertilised mite, but these are not important in terms of the overall outcome. Mite-geeks love this sort of detail 5, but we need to cut to the chase …

Keeping it in the family

The foundress ‘mother’ mite and her progeny all share a single feeding hole through the cuticle of the developing pupa.

What a lovely scene of family ‘togetherness’. 

Male and female mites take 6.6 and 5.8 days respectively to develop to sexual maturity. Therefore the male mite reaches sexual maturity before the first of his sisters.

He then lurks around the attractive-sounding “faecal accumulation site” and mates with each of the (sister) females in turn.

What a little charmer 😉

Male mites are short lived and the eclosion of the adult worker or drone curtails further mating activity, releasing the foundress mite and the mated mature daughters 6.

Reproductive rate (mites per cell)

The three day difference in the duration of worker and drone development means that more mites are produced from drone cells than worker cells. Depending on conditions the reproductive rate is 1.3 – 1.45 in worker brood and 2.2 – 2.6 in drone brood.

Remember that the foundress is also released from the cell. She can go on to initiate one or two further reproductive cycles (or up to 7 in vitro). Consequently, the average yield of mature, mated female mites from worker and drone cells is a fraction over 2 and 3 respectively.

Before entering a fresh cell containing a late stage (5th instar) larva the newly-mated mites need to mature. They do this during the phoretic phase which lasts 5-11 days. Therefore the full replication cycle of the mite probably takes a minimum of about 17 days.

Exponential growth

Two to three mites per infested cell doesn’t sound very much. However, under ideal conditions this leads to exponential growth of the mite population in the colony. Assuming 10 reproductive cycles in 6 months, a single mite would generate a population of >1,000 in worker brood and >59,000 in drone brood 7.

Fortunately (for our bees, not for the mites), ideal conditions don’t actually occur in reality.

Lots of things contribute to the reduction in reproductive potential. For example, only 60% of male mites achieve sexual maturity due to developmental mortality, drone brood is only available at certain times in the season, brood breaks interrupt the availability of any suitable brood and grooming helps rid adult bees of phoretic mites.

Out, damn'd mite ...

Out, damn’d mite …

However, these reductions aren’t enough. Without proper management mite levels still reach dangerously high levels, threatening the long-term viability of the colony.

In the next few months I will discuss some additional opportunities for reducing the mite population.

In the meantime, as we reach the winter solstice, colonies in temperate regions may well be broodless and – as emphasised last week – this is an ideal time to apply a midwinter oxalic acid-containing treatment. This will effectively reduce mite levels for the start of the coming season.

Happy Christmas … unless you’re a mite 😉


Colophon

Today is the winter solstice in the Northern hemisphere. This is actually the precise time when the Earth’s Northern pole has its maximum tilt away from the Sun. However, the term is usually used for the day with the shortest period of daylight and the longest period of night. In Fife, sunrise is at 08.44 and sunset at 15.37, meaning the day length is 6 hours and 53 minutes long.

With increasing day length queens will start laying again … but there’s a long way to go until winter is over.

 

Convenience or laziness?

It’s cold and dark and all is quiet in the apiary. Hives appear somnolent. Colonies are clustered 1 and, other than the odd corpse or two on the landing board, I’ve not seen a bee for at least a fortnight.

The apiary in winter ...

The apiary in winter …

Based upon previous experience I suspect colonies are – or very soon will be – broodless. I usually reckon that the first extended (2-3 weeks) period of cold weather 2 in the winter is the most likely time for the colony to be broodless.

In 2016/17 this was the first week in December.

In 2017/18 it was just a day or two later.

In both instances, when the hives were checked, they had no brood.

What’s all this about being broodless?

If a colony is broodless there are no capped cells in which the Varroa mite can ‘hide’. As a consequence it’s an ideal time to apply a miticide like a trickled solution of Api-Bioxal 3.

There are very good reasons why a midwinter OA treatment is necessary, particularly if you treated early enough in the autumn to protect the overwintering workers from the ravages of Deformed Wing Virus (DWV). High DWV levels reduce the lifespan of bees and contribute to many (possibly most) winter colony losses. I’ve even suggested here that “isolation starvation” might actually be due to Varroa-transmitted viral disease.

Time of treatment and mite numbers

Time of treatment and mite numbers

Early autumn treatment protects the winter bees but also leaves the long autumn for the residual mites to continue replicating.

And there will be residual mites. No treatment is 100% effective.

So, paradoxically, if you treated early enough in the autumn to really help protect the winter bees, your mite levels will be higher at the end of the year.

Which also means they’ll be higher at the beginning of next year.

Not a good start to the 2019 season 🙁

Convenience or laziness?

Many beekeepers, for convenience, laziness or historical precedent, choose to apply the winter OA treatment between Christmas and New Year. I suspect that this is often too late. If the queen starts laying again around the winter solstice there will be sealed brood – and therefore unreachable Varroa – by the end of the month.

I’d prefer to have a cold and damp afternoon in the apiary slaughtering Varroa now than the convenience of treating them less effectively during the Christmas holiday period.

The latter might be more convenient … the office will be closed, I’ll be replete with turkey and sprouts and it will be a good excuse to ‘escape’ visiting relatives and yet more mince pies 4.

But is it the best time for your bees?

We have the technology

We have a couple of hives with Arnia hive monitors fitted 5. These have a temperature probe inserted into the brood nest. Brood rearing temperature is around 34°C. Here is a trace of one colony over the last month.

Arnia hive monitor temperature

Arnia hive monitor temperature

The colony temperature was pretty stable (around 33-35°C) until about the 19th of November and has dropped about 10°C since then. Although I’ve not opened the colony I think that this is additional evidence that the colony is broodless 6.

Beekeeping by numbers

Keeping bees properly involves being aware of the seasons, the available forage and the state of the colony. This varies from month to month and year to year 7.

You can’t mechanically (‘by the numbers’) add supers on the 5th of May and harvest honey on the 15th of June. Sure, it might work some years, but is it the best time to do it?

Similarly, you can’t optimally treat a colony for Varroa on the 30th of December unless the climatic conditions and state of the colony coincide to make that the best time to treat.

It might be, but I suspect that generally it’s a bit late if there is a brood break.

If you’re going to the trouble of preparing the OA treatment, donning the beesuit and disturbing the colony you might as well do it at the right time for the bees.

I’ll be treating in between the predicted sleet showers and sunny periods this weekend.

Time to treat

Time to treat

Isn’t evolution a wonderful thing? This post started with a working title of Know your enemy” and was on a different topic altogether. I’ll save that for next week.


STOP PRESS

The above was written at the beginning of the week. Now the weekend is closer it’s clear the weather is going to be cold with heavy snow predicted. Unless the forecast is wrong (and how often does that happen?!) I’ll hold off treating until a) it’s over 5°C, and b) the roads are safe.

Resistance is not futile

Apivar ...

Apivar …

Amitraz-containing miticides are sold in the UK as Apivar and Apitraz.

Until recently they were only available with a veterinary prescription. I expect – though I have not yet seen data to support this – that their usage in the UK will increase now they are off-prescription. These miticides are now widely available and so there is greater opportunity to use – and misuse – them.

If you’re using Apivar 1 for the first time this year you will soon have to remove the strips from the hive.

That’s assuming you started treating early enough to protect the all-important winter bees from Varroa and its deadly viral payload.

This post is a reminder to remove the strips at the right time. The alternative – leaving them in place until the first Spring inspections – risks help the development of resistance to amitraz, so further reducing our opportunity to control mites effectively.

Leave and let die

Without careful integrated pest management (IPM) 2 Varroa levels build up in the hive. Varroa transmits viruses – most important of which is deformed wing virus (DWV) – to developing pupae. High levels of DWV either kills the pupa or results in emergence with or without significant developmental defects. Even those bees that are apparently normally developed have a reduced lifespan 3.

Winter bees with a reduced lifespan prevent the colony from surviving through the winter until the queen starts laying again. I’ve also proposed recently that high levels of DWV, and the resulting increased rate of winter bee die-offs, probably accounts for some cases of isolation starvation.

So … intervention is needed to reduce mite levels, protect your bees and save your colonies.

Follow the instructions!

Apivar is one solution to reduce mite levels. It is an easy-to-apply chemical treatment that is very effective in reducing the Varroa load by ~95%. For a National hive it is applied as two polymer strips, each containing 500mg of slow-release Amitraz. Strips are hung between brood frames for 6-10 weeks and – for maximum efficacy – should be scratched with a hive tool and repositioned half way through the treatment period.

Amitraz

Amitraz …

Unlike some other miticides (e.g. Apiguard and MAQS) there are no temperature restrictions for Apivar usage. The colony does not need to be broodless (a requirement for trickled oxalic acid-based treatments) as the treatment period covers multiple brood cycles.

Other than not using it with supers present the only contraindication for Apivar is to not use it if Amitraz-resistant mites are present.

How does resistance develop?

When discussing parasites and pathogens, resistance 4 is a consequence of two things:

  1. A selective pressure that kills the pathogen
  2. A population which exhibits genetic diversity

The selective pressure could be anything … heat for example, antibiotics prescribed by your GP, an antiviral against HIV or – of relevance here – Apivar against Varroa.

Killing – at the population level – is not absolute. Some individuals within the population survive longer than others. They could be exposed to a slightly lower dose, or be located in a protected niche for example. However, treat for long enough and the majority will be killed.

But there’s more …

Pathogen populations are not genetically invariant. Actually, many are quite diverse and have replication cycles that – deliberately 5 – generate diversity.

Therefore some pathogens are genetically slightly less resistant and some are genetically slightly more resistant to a selective pressure. We can ignore the former as they’ll rapidly be killed off … but we must be concerned about the more resistant ones.

Keep taking the pills

All of this is a ‘numbers game’, better represented with graphs and equations. However, the take-home message is simple … to effectively control a pathogen you need to treat for long enough and with a high enough dose to kill the vast majority of the population.

That’s why you’re encouraged to “complete the course” of antibiotics … or to remove the Apivar strips after 10 weeks and not leave them in over the winter.

Because both courses of action result in selection of more resistant pathogens.

If you stop taking antibiotics too soon, you won’t have treated for long enough and with a high enough dose. You end up selecting for the more genetically resistant pathogens.

Similarly, if you leave Apivar strips in overwinter you’ll be “treating” the remaining mites 6 with a lower dose of the miticide, which is an ideal situation to favour the growth of the slightly more genetically resistant mites.

How does Amitraz resistance develop?

Resistance to Amitraz in Varroa is well documented. It’s been described in a number of countries including the USA and Europe, Mexico and Argentina 7. Generally resistance is defined in terms of a reduced level of mite killing, or – in laboratory experiments – an increased dose required to kill a certain proportion of mites.

However, I’m unaware of any studies defining the genetic basis of Amitraz resistance in Varroa.

But Amitraz is a widely-used acaricide 8 and the genetic basis of resistance in cattle ticks is well understood. In these, ticks resistant to Amitraz carry a mutation in the RMβAOR gene 9.

What 10 is the RMβAOR gene?

I’m glad you asked 😉

This gene encodes the β-adrenergic octopamine receptor protein and readers with good memories will recall that this is one of the targets that Amitraz binds to and inactivates 11.

If the protein carries a mutation the Amitraz cannot bind to it and so the mite – or more correctly the tick as it’s yet to be formally demonstrated in mites – is therefore resistant.

(Bad) practical beekeeping

What does all this mean in terms of practical beekeeping?

It means use the correct number of Apivar strips for the colony and leave them in for the right length of time.

Do not …

  • Use one strip on a full colony mid-season to ‘knock back the mites a bit’ 
  • Re-use the strips in another colony (yes really!)
  • Use improperly stored strips (or out of date strips) in which the effective Amitraz dose is reduced

I’ve heard examples of these types of misuse this season. All increase the chance of selecting for Amitraz-resitant mites.

And (the real reason for posting this at this time of year) …

  • Do not leave the strips you added in late summer in the colony throughout the winter

Removing the strips takes seconds. Prize off the crownboard, grab the tab projecting above the top bars, gently withdraw the strip and close the hive up again.

Finally, because of the incestuous lifestyle 12 of Varroa the genetic diversity (and therefore potential presence of more resistant mites) in the population is likely to be increased by the high mite levels that prevail late in the season.

All the more reason to use the effective treatments we currently have in a way that helps ensure they remain effective.


Colophon

Resistance is futile

Resistance is futile

Resistance is futile is the title of a 2018 album by the Welsh rock band the Manic Street Preachers.

More specifically, in the context of this post, it was the phrase routinely used by the Borg – the alien cyborgs sharing a collective mind – in the Star Trek franchise. Borgs rarely speak, but when they do they usually include this phrase. For example “We are the Borg. Lower your shields and surrender your ships. We will add your biological and technological distinctiveness to our own. Your culture will adapt to service us. Resistance is futile.” The warning about resistance being futile was usually accompanied by the threat that the target would be assimilated”.

I’d started writing this post using the title ‘Resistance is futile’ but realised late on that – as far as Varroa are concerned – resistance is anything but futile 13.

Resistance – to miticides – gives Varroa a reason to live. Literally.

Let’s not help them 🙂