Tag Archives: Christina Grozinger

Diutinus bees

Diutinus is Latin for long-lasting.

Diutinus bees are therefore long-lasting bees. These are the bees that, in temperate regions, maintain the colony through the winter to the warmer days of spring.

I’ve discussed the importance of these bees recently., and I’ve regularly made the case that protecting these ‘long-lived’ bees from the ravages of Varroa-vectored viruses is critical to reduce overwintering colony losses.

Winter is coming …

In most cases the adjective diutinus is replaced with ‘winter’, as in winter bees; it’s a more familiar term and emphasises the time of year these bees are present in the hive. I’ll generally use the terms interchangeably in this post.

Diutinus does not mean winter

From a scientific standpoint, the key feature of these bees is that they can live for up to 8 months, in contrast to the ~30 days a worker bee lives in spring or summer. If you are interested in what induces the production of long-lived bees and the fate of these bees, then the important feature is their longevity … not the season.

Furthermore, a proper understanding of the environmental triggers that induce the production of long-lived bees might mean they could be produced at other times of the season … a point with no obvious practical beekeeping relevance, but one we’ll return to in passing.

It’s worth emphasising that diutinus bees are genetically similar to the spring/summer bees (which for convenience I’ll refer to as ‘summer bees’ for the rest of the post). Despite this similarity, they have unique physiological features that contribute to their ability to thermoregulate the winter cluster for months and to facilitate spring build-up as the season transitions to spring.

What induces the production of winter bees? Is it a single environmental trigger, or a combination of factors? Does summer bee production stop and winter bee production start? What happens at the end of the winter to the winter bees?

Segueing into winter bee production 

The graph below shows the numbers of bees of a particular age present in the hive between the end of August and early December.

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

Each distinct colour represents bees reared in a particular 12 day ‘window’. All bees present before the 31st of August are blue. The next 12 day cohort of bees are yellow etc. The area occupied by each colour indicates the number of bees of a particular age cohort.

Note that egg laying (black) is negligible between early October and late November when it restarts.

The graph shows that that there is no abrupt change from production of summer bees to production of winter bees.

For example, about 95% of the blue bees have disappeared by December 1. Of the yellow bees, which first appeared in mid-September, about 33% are present in December. Finally, the majority of the lime coloured bees, that first put in an appearance in early October, are present at the end of December.

The colony does not abruptly stop producing short-lived summer bees on a particular date and switch to generating long-lived ‘diutinus’ winter bees. Instead, as late summer segues into early autumn, fewer short lived bees and more long lived bees are produced. 

Note that each cohort emerge from eggs laid 24 days earlier. The orange cohort emerging from 24/09 to 05/10 were laid within the first two weeks of September. This emphasises the need to treat early to reduce mite levels sufficiently to protect the winter bees.

Winter bees are like nurse bees but different

Before we consider what triggers the production of diutinus bees we need to discuss how they differ from summer bees, both nurses and foragers.

Other than being long-lived what are their characteristics?

Interaction of key physiological factors in nurse (green), forager (red) and winter bees (blue). Colored disks indicate the relative abundance of each factor.

The four key physiological factors to be considered are the levels of juvenile hormone (JH), vitellogenin (Vg) and hemolymph proteins and the size of the hypopharyngeal gland (HPG).

As summer nurse bees transition to foragers the levels of JH increases and Vg decreases. This forms a negative feedback loop; as Vg levels decrease, JH levels increase. Nurse bees have high levels of hemolymph proteins and large HPG, the latter is involved in the production of brood food fed to larvae.

So if that describes the summer nurse bees and foragers, what about the winter bees?

Winter bees resemble nurse bees in having low JH levels, high levels of VG and hemolymph proteins and large HPG’s. 

Winter bees differ from nurse bees in being long lived. A nurse bee will mature into a forager after ~3 weeks. A winter bee will stay in a physiologically similar state for months.

There have also been time course studies of JH and Vg levels through the winter. In these, JH levels decrease rapidly through October and November and are at a minimum in mid-January, before rising steeply in February and March.

As JH levels rise, levels of Vg and hemolymph proteins decrease and the size of the HPG decreases i.e. as winter changes to early spring winter bees transition to foragers.

Now we know what to look for (JH, Vg levels etc) we can return to think about the environmental triggers that cause these changes.

No single trigger

In temperate regions what distinguishes winter from autumn or spring? 

Temperatures are lower in winter.

Daylength (photoperiod) is shorter in winter.

There is less pollen and nectar (forage) available in winter.

Under experimental conditions it’s quite difficult to change one of these variables without altering others. For example, shifting a colony to a cold room (i.e. lowering the ambient temperature to <10°C) leads to a rapid decrease in JH levels (more winter bee-like). However, the cold room was dark, so perhaps it was daylength that induced the change? Alternatively, a secondary consequence of moving the colony is that external forage was no longer available, which could account for the changes observed.

And forage availability will, inevitably, influence brood rearing.


Reducing photoperiod alone does induce some winter bee-like characteristics, such as increases in the protein and lipid content of the fat bodies. It also increases resistance to cold and starvation. It can even cause clustering at elevated (~19°C) temperatures. However, critically, a reduced photoperiod alone does not appear to make the bees long lived. 

Remember also that a reduced photoperiod will limit foraging, so reducing the nutritional status of the colony. This is not insignificant; pollen trapping 2 in the autumn accelerates the production of winter bees.

But again, this may be an indirect effect. Reduced pollen input will lead to a reduction in brood rearing. Feeding pollen to broodless winter colonies induces egg-laying by the queen.

Brood, brood pheromones and ethyl oleate

One of the strongest clues about what factor(s) induces winter bee production comes from studies of free-flying summer colonies from which the brood is removed. In these, the workers rapidly change to physiologically resemble winter bees 3.

How does the presence of brood prevent the generation of diutinus bees?

There are some studies which demonstrate that the micro-climate generated in the colony by the presence of brood – elevated temperature (35°C) and 1.5% CO2 – can influence JH levels. 

However, brood also produces pheromones – imaginately termed brood pheromone – which does all sorts of things in the colony. I’ve discussed brood pheromone previously in the context of laying workers. The brood pheromone inhibits egg laying by worker bees.

Brood pheromone also contributes to a enhancement loop; it induces foraging which results in increased brood rearing and, consequently, the production of more brood pheromone.

One way brood pheromone induces foraging is by speeding the maturation of middle-aged hive bees into foragers. Conversely, when raised in the absence of brood, bees have higher Vg levels, start foraging later and live longer.

But it’s not only brood that produces pheromones.

Workers also produce ethyl oleate, a pheromone that slows the maturation of nurse bees, so reducing their transition to foragers.

A picture is worth a thousand words

All of the above is quite complicated.

Individual factors, both environmental and in the hive, have direct and indirect effects. Experimentally it is difficult to disentangle these. However, Christina Grozinger and colleagues have produced a model which encapsulates much of the above and suggests how the production of winter bees is regulated. 

Proposed model for regulation of production of winter bees.

During autumn there is a reduction in forage available coupled with a reduced daylength and lower environmental temperatures. Consequently, there is less foraging by the colony. 

Since more foragers are present within the hive, the nurse bees are exposed to higher levels of ethyl oleate, so slowing their maturation.

There’s less pollen being brought into the colony (reduced nutrition), so brood production decreases and so does the level of brood pheromone. The reduced levels of brood pheromone also reduces nurse bee maturation.

As shown in the diagram, all of these events are in a feedback loop. The reduction in levels of brood pheromone further reduces the level of foraging … meaning more foragers are ‘at home’, so increasing the effects of ethyl oleate.

All of these events have the effect of retarding worker bee maturation. The workers remain as ‘nurse-like’ long-lived winter bees.

Is that all?

The difference between nurse bees and winter bees is their longevity … or is it?

In the description above, and in most of the experiments conducted to date, the key markers of the levels of JH, Vg and hemolymph proteins, and the size of the HPG, are what has been studied. 

I’d be astounded if there are not many additional changes. 

Comparison of workers and queen bees have shown a large range of epigenetic changes induced by the differences in the diet of young larvae 4. Epigenetic changes are modifications to the genetic material that change gene expression.

I would not be surprised if there were epigenetic changes in winter bees, perhaps induced by alteration of the protein content of their diet as larvae, that influence gene expression and subsequent longevity. Two recent papers suggest that this may indeed happen; the expression of the DNA methyltransferases (the enzymes that cause the epigenetic modifications) differs depending upon the demography of the colony 5 and there are epigenetic changes between the HPG in winter bees and bees in spring 6.

Clearly there is a lot more work required to fully understand the characteristics of winter bees and how they are determined.

Don’t forget …

It’s worth emphasising that the local environment (forage and weather in particular) and the strain of the bees 7 will have an influence on the timing of winter bee production.

Last week I discussed a colony in my bee shed that had very little brood on the 2nd of October (less than one side of one frame). When I checked the colonies last weekend (11th) there were almost no bees flying and no pollen coming in. A colleague checked an adjacent colony on Monday (13th) and reported it was completely broodless. These bees are ‘local mongrels’, selected over several years to suit my beekeeping.

Early autumn colonies

In contrast, my colonies on the west coast are still busy. These are native black bees. On the 14th they were still collecting pollen and were still rearing brood. 

The calendar dates in the second figure (above) are therefore largely irrelevant.

The transition from summer bees to the diutinus winter bees will be happening in your colonies, sooner or later. I suspect it’s already completed in my Fife bees.

Whether genetics or environment has a greater influence on winter bee production remains to be determined. However, I have previously described the good evidence that local bees are better adapted to overwintering colony survival.

To me, this suggests that the two are inextricably linked; locally selected bees are better able to exploit the environment in a timely manner to ensure the colony has the winter bees needed to get the colony through to spring.


Midwinter, no; mites, yes

There’s a certain irony that the more conscientious you are in protecting your winter bees from the ravages of Varroa in late summer, the more necessary it is to apply a miticide in the winter.

Winter bees are the ones that are in your hives now 1.

They have a very different physiology to the midsummer foragers that fill your supers with nectar. Winter bees have low levels of juvenile hormone and high levels of vitellogenin. They are long-lived – up to 8 months – and they form an efficient thermoregulating cluster when the external temperature plummets.

Winter bees production

In the temperate northern hemisphere, winter bees are reared from late summer/early autumn onwards. The combination of reductions in the photoperiod (day length), temperature and forage availability triggers changes in brood and forager pheromones.

Factors that influence winter bee production

Together these induce the production of winter bees.

For more details see Overwintering honey bees: biology and management by Döke et al., (2010).

Day length reduces predictably as summer changes to autumn. In contrast, temperature and forage availability (which itself is influenced by temperature and rainfall … and day length) are much more variable (so less predictable).

All of which means that you cannot be sure when the winter bees are produced.

If there’s an “Indian summer“, with warm temperatures stretching into late October, the bees will be out working the ivy and rearing good amounts of brood late into the year. The busy foragers and high(er) levels of brood pheromone will then delay the production of winter bees.

Conversely, low temperatures and early frosts reduce foraging and brood production, so bringing forward winter bee production.

It’s an inexact science.

You cannot be sure when the winter bees will be produced, but you can be sure that they will be reared.

Protect your winter bees

And if they are being reared, you must protect them from Varroa and the viral payload it delivers to developing pupae. Most important of these viruses is deformed wing virus (DWV).

Worker bee with DWV symptoms

Worker bee with DWV symptoms

Aside from “doing what it says on the tin” i.e. causing wing deformities and other developmental defects in some brood, DWV also reduces the longevity of winter bees.

And that’s a problem.

If they die sooner than they should they cannot help in thermoregulating the winter cluster.

And that results in the cluster having to work harder to keep warm as it gets smaller … and smaller … and smaller …

Until it’s so small it cannot reach its food reserves (isolation starvation) or freezes to death 2.

So, to protect your winter bees, you need to treat with an appropriate miticide in late summer. This reduces the mite load in the hive by up to 95% and so gives the winter bees a very good chance of leading a long and happy life 😉

Time of treatment and mite numbers

Time of treatment and mite numbers

I discussed this in excruciating detail in 2016 in a post titled When to treat?.

The figure above was taken from that post and is described more fully there. The arrow indicates when winter bees are produced and the variously coloured solid lines indicate mite numbers when treated in mid-July to mid-November.

The earlier you treat (indicated by the sudden drop in the mite count) the lower the peak mite numbers when the winter bees are being reared.

Note that the mite numbers indicated on the right hand vertical axis are not ‘real’ figures. They depend on the number present at the start of the year. In the figure above I “primed” the in silico modelled colony with just 20 mites. This will become very important in a few paragraphs.

Late season brood rearing

Compare the blue line (mid-August treatment) with the cyan line 3 (mid-October treatment) in the figure above.

The mid-October treatment really hammers the mite number down and they remain low until the end of the year 4.

The reason the mite numbers remain low after a mid-October treatment is that there is little or no brood being reared in the colony during this period.

Mites need brood, and specifically sealed brood, to reproduce on.

In the absence of brood the mites ‘colony surf‘, riding around as phoretic mites on nurse bees (or any bees if there aren’t the nurse bees they prefer).

And that late season brood rearing is the reason the end-of-year mite number for the colony treated in mid-August (the blue line) remains significantly higher.

Mites that survive the miticide in August simply carry on with their sordid little destructive lives, infesting the ample brood available (which could even include some highly mite-attractive and productive drone brood) and reproducing busily.

So, the earlier you treat, the more mites remain in the hive at the end of the year.

Weird, but true.

Early season brood rearing

The winter bees don’t ‘just’ get the colony through the winter.

As the day length increases and the temperature rises the colony starts rearing brood again. Depending upon your latitude it might never stop, but the rate at which it rears brood certainly increases in early spring.

Or, more correctly, in mid- to late-winter.

And it’s the winter bees that do this brood rearing. As Grozinger and colleagues state Once brood rearing re-initiates in late winter/early spring, the division of labor resumes among overwintered worker bees.”

Some winter bees revert to nurse bee activity, to rear the next generation of bees.

And this is another reason why strong colonies overwinter better … not because they (also) survive the cold better 5, but because there are more bees available to take on these brood rearing activities.

Strong, healthy colonies build up better in early spring.

Colonies that are weak in spring and stagger through the first few months of the year, never getting close to swarming, are of little use for honey production, more likely to get robbed out and may not build up enough for the following winter.

Midwinter mite treatments

Which brings us back to the need for miticide treatment in midwinter.

The BEEHAVE modelled colony shown in the graph above was ‘primed’ at the beginning of the season with 20 mites. These reproduced and generated almost 800 mites over the next 10-11 months.

What do you think would happen if you start the year with 200 mites, rather than 20?

Like the 200 remaining at the year end when you treat in mid-August?

Lots of mites … probably approaching 8000 … that’s almost as many mites as bees by the end of the season.

So, one reason to treat in the middle of winter is to reduce mite levels later in the season. The smaller the number you start with, the less you have later.

Vapour leaks out ...

Vaporisation … oxalic acid vapour leaks out …

But at the beginning of the season these elevated levels of mites could cause problems. High levels of mites and low levels of brood is not a good mix.

There’s the potential for those tiny patches of brood to become mite-infested very early in the season … this helps the mites but hinders the bees.

Logically, the more mites present at the start of brood rearing, the more likely it is that colony build up will be retarded.

So that’s two reasons to treat with miticides – usually an oxalic-acid containing treatment – in midwinter.

Midwinter? Or earlier?

When does the colony start brood rearing again in earnest?

This is important as the ‘midwinter’ treatment should be timed for a period before this when the colony is broodless. This is to ensure that all the mites are phoretic and ‘easy to reach’ with a well-timed dribble of Api-Bioxal.

In studies over 30 years ago Seeley and Visscher demonstrated that colonies have to start brood rearing in midwinter to build up enough to have the opportunity to swarm in late spring. These were colonies in cold climates, but the conditions – and season length – aren’t dramatically different to much of the UK.

Low temperatures regularly extend into January or February. The temperature is also variable year on year. It therefore seems (to me) that the most likely trigger for new brood rearing is increasing day length 6.

The apiary in winter ...

The apiary in winter …

I therefore assume that colonies may well be rearing brood very soon after the winter solstice.

I’m also aware that my colonies are almost always broodless earlier in the winter … or even what is still technically late autumn.

This is from experience of both direct (opening hives) or indirect (fresh brood mappings on the Varroa tray) observation.

Hence the “Midwinter, no” title of this post.

Don’t delay

I therefore treat with a dribbled or vaporised oxalic acid-containing miticide in late November or early December. In 2016 and 2017 it was the first week in December. Last year it was a week  later because we had heavy snow.

This year it was today … the 28th of November. With another apiary destined for treatment this weekend.

If colonies are broodless there is nothing to be gained by delaying treatment until later in the winter.

Most beekeepers treat between Christmas and New Year. It’s convenient. They’re probably on holiday and it is a good excuse to escape the family/mince pies/rubbish on the TV (delete as appropriate).

But it might be too late … don’t delay.

If colonies are broodless treat them now.

If you don’t and they start rearing brood the mites will hide away and be unreachable … but their daughters and granddaughters will cause you and your bees problems later in the season.

Finally, it’s worth noting that there’s no need to coordinate winter treatments. The bees aren’t flying and the possibility of mites being transferred – through robbing or drifting – from treated to untreated colonies is minimal.