Category Archives: Behaviour

Swarm prevention

Swarm prevention and control are distinct phases in the management of colonies during the next few weeks of the beekeeping season 1.

Not all beekeepers practice them and not all colonies need them.

But most should and will … respectively ūüėČ

Swarm prevention involves strategies to delay or stop the colony from initiating events that lead to swarming.

Swarm control strategies are more direct interventions that are used to prevent the loss of a swarm.

Why do colonies swarm?

Without swarming there would be no honey bees.

Swarming is honey bee colony reproduction. Without management (e.g. splitting colonies) colony numbers would remain static. And, since bees have only been managed for a few thousand years, they must have been successfully reproducing – by swarming – for millions of years before then.

So one of the major drivers of swarming is the innate need to reproduce.

Bees also swarm if their current environment is unable to accommodate further colony expansion. Therefore, another driver of swarming is overcrowding.

And, of course, there is some overlap in these two drivers of swarming.

You can therefore expect that strong, healthy, populous colonies will probably try to swarm on an annual basis.

The mechanics of swarming

When a colony swarms about 75% of the worker bees – of all age groups – leave with the queen. They set up a temporary bivouac near the original hive and subsequently relocate to a new nest site identified by the scout bees.

The original colony is left with all the brood (eggs, larvae and sealed brood), a significantly-depleted adult bee workforce and almost 2 all of the honey stores.

What they lack is a queen.

But what the swarm also leaves behind, amongst the brood, is one – or more often several – newly developing queens. These occupy specially enlarged cells that are located vertically on the edges or face of the comb.

Queen cells ...

Queen cells …

Queen cells look distinctive and their initial appearance – before the swarm leaves – is a clear indication that the time for swarm prevention has gone and swarm control is now urgently needed 3.

This is one of the reasons why regular colony inspections are essential, particularly during mid/late Spring and early summer which is the time of the season when swarming is most likely.

Colony fate and the risks of swarming

But back to the recently swarmed colony. In a few days the new queen(s) emerges. If there’s more than one they usually fight it out to leave just one. She goes on one or more mating flights and a few days later starts laying eggs.

This colony should survive and thrive. They have time to build up strength (and collect more stores) before the end of the season. Under natural conditions 87% of swarmed colonies overwinter successfully 4.

Alternatively, the swarmed colony may swarm again (and again), each with a virgin queen and each further depleting the worker population. Colonies can swarm themselves to destruction like this.

Swarms headed by virgin queens are termed casts. I’m not sure what determines whether a swarmed colony also produces one or more casts. Colony strength is a determinant, but clearly not the only one as some casts contain little more than a cup full of bees.

Under natural conditions swarming is a very risky business. Swarm survival is less than 25% 5 – many will not collect sufficient stores to overwinter – and the survival of casts will be even lower because of their size and the risks associated with queen mating.

But ‘our’ bees don’t live under natural conditions

For beekeeping the ‘risks’ associated with swarming are somewhat different.

When a colony swarms you lose the majority of the workforce. Therefore honey production will be significantly reduced. You’re unlikely to get a surplus from the swarmed colony.

Of course, honey might not interest you but propolis and wax production are also reduced, as is the strength of the colony to provide efficient ecosystem services (pollination).

Secondly, despite swarms being one of the most captivating sights in beekeeping, not everyone appreciates them. Non-beekeepers may be scared and – extraordinary as it may seem – resent the swarm establishing a new nest in the eaves of their house.

Incoming! from The Apiarist on Vimeo.

Inevitably some beekeepers will claim they’ve never met anyone scared of bees, or swarms are always welcomed in the gardens that abut their apiary.

Unfortunately, that does not alter the reality that – to many – swarms are a nuisance, a potential threat and (to a small number of people 6 ) a very real danger.

Therefore, as beekeepers, we have a responsibility to practice both swarm prevention and control. This prevents our hobby/obsession irritating other people and means we have more bees to make delicious honey for family, friends and customers.

Overcrowding

I’ve already defined the event that separates swarm prevention from swarm control. It is the appearance of queen cells during the weekly colony inspection.

Swarm prevention involves managing the colony to delay the appearance of queen cells. Once queen cells are produced, swarm control is required 7

I’ve also defined the two major drivers of swarming – overcrowding and the need to reproduce 8.

How does a colony determine that it is overcrowded? As beekeepers, how can we monitor and prevent overcrowding?

As a colony expands during the spring the queen lays concentric rings of eggs from the centre of the brood nest. Imagine this initially as a kiwi fruit-sized ball, then an orange, then a grapefruit, until it is the size of a large football.

Brood frame

Perhaps a slightly squashed football, but you get the general idea.

Running out of storage space

It takes bees to make bees. The initial brood reared helps feed subsequent larvae and keeps the maturing brood warm.

As the season develops more sources of nectar and pollen become available. These are collected in increasing amounts by the expanding numbers of foragers.

This all needs to be stored somewhere.

One possibility is that the stores are loaded into the cells recently vacated by emerging workers within the brood nest. This is often termed “backfilling”. Sometimes you find a frame in which the central concentric rings of brood have emerged and, before the queen has had a chance to re-lay the frame with new eggs, workers have backfilled the cells with nectar (or, less frequently, pollen).

But, at the same time as the space available for the queen to lay is reducing, the colony population is increasing. Very fast. There are larger numbers of unemployed young bees. Unemployed because there are reduced amounts of brood to rear because the queen is running out of space.

Pheromones

And the increased number of workers means that the pheromones produced by the queen, in particular the queen mandibular pheromone, are effectively diluted. Studies by Mark Winston and colleagues 9 investigated the relationship between queen mandibular pheromone (an inhibitor of queen cell production) and colony congestion. In it he concluded that overcrowding inhibits the transmission of this pheromone, so favouring queen cell production.

Play cup or queen cell?

Play cup or are they planning their escape …?

The distribution of other pheromones is also reduced in overcrowded colonies. Lensky and Slabezki 10 showed that the queen rarely visited the bottom edges of comb in overcrowded colonies. Consequently, the levels of queen footprint pheromone was reduced. This pheromone is an inhibitor of queen cup production, the very earliest stages of queen cell development.

So, overcrowded colonies start to prepare queen cells … and swarm control is needed.

Make space

If the colony is overcrowded then you have to provide more space for colony expansion.

Just piling supers on top may not be sufficient, though it may temporarily ease congestion and partially help. Leaving a colony with no supers during a strong nectar flow is a surefire way to fill the brood box with nectar and trigger swarm preparation.

If the colony is backfilling the brood nest with nectar then the addition of supers is likely to encourage them to move the stores up, providing more space for the queen.

It will additionally have the beneficial effect of moving some bees ‘up’, to store and process the nectar, again reducing congestion in the brood nest.

However, you probably also need to encourage the bees to expand the brood nest by providing frames for them to draw out as comb. Essentially you’re spreading the brood nest by inserting one or two empty frames within it.

Expanding or spreading the brood nest

I routinely do this by removing the outer frames, which often contain stores, and adding new foundationless frames on one or both sides of the centre of the brood nest. Usually I would place these about three to four frames apart 13.

You can routinely replace queens by purchasing new ones, by rearing your own, or through colony manipulation during swarm control e.g. by reuniting a vertical split.

Of these, I’d strongly recommend one of the last two approaches. It’s more interesting, it’s a whole lot more satisfying and it is a lot easier than many beekeepers realise.

Locally bred queen ...

Locally bred queen …

You have the additional advantage that the queens produced in your own apiary will Рby definition Рbe local and there is good evidence that local queens are better adapted to local conditions.

Robbing brood and making nucs

There are at least two additional, and related, ways of increasing the space available so helping swarm prevention in a rapidly expanding colony.

The first is stealing a frame of brood 14 and using it to boost a weaker colony.

Take care when doing this.

If the recipient colony is weak due to disease or a failing queen then you’re just wasting the donated brood. However, if the colony is healthy but small it can be a good investment of resources and may help delay swarming in the donor colony as well.

More drastically, it may be possible to remove a frame (or perhaps even two) of brood and adhering bees to make up a nucleus colony. In my experience, a strong donor colony can almost always be used to produce a nuc without compromising honey production, and with the added benefit of delaying swarm preparations.

I’m going to write about nuc production in more detail in a few weeks as it deserves a full post of its own. It’s worth noting here that the nuc should also be provided with sufficient bees and stores to survive and you will need a queen¬†for it (or at least a queen cell).

Do not just dump a couple of brood frames and bees into a box and expect them to rear a half-decent queen on their own.

However, if you have a queen (or mature queen cell) then splitting a nuc off a strong colony is usually a win-win solution for swarm prevention.


 

Who’s the daddy?

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

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

How do you measure polyandry?

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

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

Search and ye shall find – detecting rare patrilines

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

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

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

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

Half-sisters and super-sisters

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

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

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

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

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

Emergency queens and nepotism

What’s the¬†most important decision a colony makes?

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

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

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

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

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

Do worker bees exhibit nepotism when rearing emergency queens?

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

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

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

Royal patrilines

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

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

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

Experimental details

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

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

The nucs then started rearing new queens … again.

And again … and again.

This process was repeated until the nucs failed.

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

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

Cryptic “royal” subfamilies

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

Subfamily distribution per colony.

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

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

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

Frequency distribution of subfamilies.

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

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

Implications for our understanding of honey bee reproduction

Two points are immediately apparent:

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

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

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

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

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

Who’s the daddy?

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

Mechanistically, what accounts for this?

Are these larvae selected solely because they are rare?

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

Perhaps it is not even worker selection?

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

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

Or whether they’re the wrong questions entirely.

Swarming and supercedure

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

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

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

Play cups

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

Queen rearing

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

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

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

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

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

3 day old QCs ...

3 day old QCs …

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

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

Are these accepted less well by the nurse bees?

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

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

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

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


Colophon

Ray Winstone (as Carlin) 1979

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

Polyandry and colony fitness

Honey bees are polyandrous. The queen mates with multiple drones during her mating flight(s). Consequently, her daughters are of mixed paternity.

In naturally mated queens there is a relationship between the number of patrilines (genetically distinct offspring fathered by different drones) and the ‘fitness’ of a colony.

Colony fitness

A ‘fit’ colony is one that demonstrates one or more desirable traits (those that benefit the colony … and potentially the beekeeper) such as better population growth, weight gain, resistance to pathogens or survival.

If you analyse the molecular genotype of the worker offspring you can determine which patriline they belong to. If you genotype enough workers you start to see the same patrilines appearing again and again. The more patrilines, the more drones the queen mated with 1.

Shallow depth of field

One of many …

Naturally mated queens mate with ~13 drones. Depending upon the study a range from as low as 1 to as high as 40 (and exceptionally into the high 50’s) has been demonstrated, though different studies all tend to produce an average in the low- to mid-teens.

There is a well-established link between polyandry and colony fitness 2. Essentially, the more genetically diverse a colony i.e. the larger the number of patrilines, the fitter that colony is.

The benefits of polyandry

Why should colonies with increased genetic diversity be fitter?

There are a number of hypotheses that attempt to explain why intracolonial genetic diversity is beneficial. These include the increased behavioural repertoire of the worker bees, a reduced production of diploid drones (which would otherwise be produced due to the single-locus sex determination system) and an increased resistance to a wide range of parasites and pathogens 3.

Parasites and pathogens are an extremely effective evolutionary selective pressure. Several studies from David Tarpy and Thomas Seeley have shown that increased polyandry results in better resistance to chalkbrood and American foulbrood.

But what about¬†Varroa? It’s a new pathogen (evolutionarily speaking) to honey bees and there is evidence that the resistance mechanisms observed are genetically determined 4.

Does polyandry contribute to Varroa resistance? 

Would increased polyandry result in improved resistance to mites?

Limits of polyandry and natural resistance

Why is the average number of drone matings in the low teens?

If polyandry is beneficial – and there’s no doubt it is – then surely¬†more patrilines (hyperpolyandry) would be even more beneficial?

How could this be tested?

Naturally mated queens only very rarely exhibit 30+ drone matings. Not only are these colonies hard to find, but they are so rare that doing any sort of statistical analysis of the improved (or otherwise) fitness is probably a non-starter.

Perhaps there’s an alternative way to approach the question? Rather than look at individual colonies within a mixed population, why not study the overall level of polyandry within a population that demonstrates resistance?

For example, do queens that head colonies of untreated feral bees that exhibit a demonstrated enhanced resistance to Varroa, the most important pathogen of honey bees, exhibit higher levels of polyandry?

Two relatively recent scientific papers have tackled these questions. Both have produced clear answers.

Drones : if more is better, is lots more better still?

Yes.

Keith Delaplane and colleagues used instrumental insemination¬†(II) of virgin queens to produce queens ‘mated’ with 15, 30 or 60 drones. Sperm was collected from 1, 2 or 4 drones from 15 donor colonies, mixed thoroughly and used for queen insemination.

Full-sized colonies were requeened with the II queens and left for 6 weeks 5 after which sampling started. Over two seasons a total of 37 colonies (with 11, 13 and 13 colonies respectively headed by queens ‘mated’ with 15, 30 and 60 drones) were tested at approximately monthly intervals.

Testing involved visual analysis of colony strength 6 and comb construction. Mite levels were measured using standard alcohol wash of ~300 bees at mid- or late-summer timepoints.

Brood frame with a good laying pattern

The results of this study are commendably brief … just 8 lines of text and two tables. I’ll summarise them in just a couple of sentences.

Colonies headed by queens ‘mated’ with 30 or 60 drones produced significantly more brood than the colony headed by the queen ‘mated’ with only 15 drones. Conversely, significantly more colonies headed by queens mated with only 15 drones had a higher level of mite infestation 7.

Natural Varroa resistance and polyandry

One of the best studied populations of feral bees co-existing with Varroa are those in the Arnot Forest in New York State. These are the bees Thomas Seeley and colleagues study.

These colonies live in natural holes in trees at low density through the forest. The colonies are small and they swarm frequently. Their spatial distribution, size and swarminess (is that a word?) are all evolutionary traits that enable resistance, or at least tolerance, to Varroa and the pathogenic viruses the mite transmits.

I’ve discussed Seeley’s studies of the importance of colony size and swarming previously. I don’t think I’ve discussed his work on spatial separation of colonies, but I have described related studies by Delaplane and colleagues.

Essentially, by being well-separated, mite transmission between colonies (e.g. during robbing) is minimised. Similarly, by existing as small colonies that swarm frequently Iwith concomitant brood breaks) the mite population is maintained at a manageable level.

Marked queen surrounded by a retinue of workers.

Her majesty …

Do the Arnot Forest Varroa-resistant 8 bees exhibit especially high levels of polyandry, suggesting that this contributes their survival?

No.

Seeley and colleagues determined the number of patrilines in 10 Arnot Forest colonies using the same type of genotyping analysis described earlier. They compared these results to a similar analysis of 20 managed honey bee colonies located nearby.

On average, Arnot Forest queens had mated with ~18 drones (17.8 ¬Ī 9.8) each. In contrast, queens in managed colonies in two nearby apiaries had mated with ~16 and ~21 drones. These figures are not statistically different from each other or from the natural mating frequencies reported for honey bees in other studies.

Hyperpolyandry and colony fitness

The first of the studies confirms and extends earlier work demonstrating the polyandry (and in this instance hyperpolyandry i.e. at an even greater level than seen normally) increases colony fitness Рat least in terms of colony strength and Varroa resistance.

Delaplane and colleagues hypothesise that the increased mite resistance in hyperpolyandrous (30 or 60 drones) colonies may be explained by either:

  • the importance of extremely rare alleles (gene variants), which would only be present in colonies in which the queen had mated with a very large number of drones.
  • the presence of beneficial non-additive interactions between genetically-determined traits¬†e.g. grooming and hygienic behaviour and reduced mite reproduction.

Neither of which are mutually exclusive and both fit at least some of the extant data on natural mite resistance. Discriminating between these two hypotheses and teasing apart the variables will not be straightforward.

Absence of hyperpolyandry in naturally mite-resistant colonies

At first glance, the absence of the hyperpolyandry in the mite-resistant Arnot Forest bees studied by Thomas Seeley and colleagues appears to contradict the studies using the instrumentally inseminated queens.

The Arnot Forest bees exhibit the same level of polyandry as nearby managed colonies, and for that matter, as colonies studied elsewhere. They are mite-resistant but the queen has not mated with an increased number of drones.

In other studies 9, naturally mated colonies exhibiting different levels of polyandry (within the normal range) showed no correlation between Varroa levels and queen mating frequency.

Perhaps it’s surprising that the Arnot Forest queens hadn’t mated with fewer drones considering the extreme separation of the colonies (when compared with managed colonies). The colony density within the Forest is approximately one per square kilometre.

However, at least during the peak swarming and mating period in the season, drone availability is rarely limiting.

This is because drones are not evenly spread in the environment. Instead, they accumulate in drone congregation areas (DCA) to which the queen flies for mating.

What limits polyandry?

Polyandry is beneficial and, apparently, hyperpolyandry is more beneficial. However, queens mate with 10 – 20 drones, rather than 50 or more. Why is this?

Queen mating is a risky business. The queen has to fly to the DCA, mate with multiple drones and then return to the hive. She may make one or several mating flights.

I’ve discussed how far drones and queens fly to reach the DCA previously. Most drones fly less than 3 miles and 90% of matings occur within about 5 miles of the virgin queen’s hive. The queen probably flies further to the DCA.

All the time she is travelling to and from the DCA, and all the time she is present within it mating, she’s potentially at risk from hungry house martins, swallows, bee eaters (!) or from thunderstorms.

Or simply from getting lost.

Additionally, a number of honey bee pathogens are transmitted between drones and queens during mating. Hyperpolyandrous queens 10 are therefore at risk from these sexually transmitted diseases 11.

It’s therefore likely that the level of polyandry observed in honey bees has evolved as a consequence of the beneficial pressures polyandry brings balanced by the risks associated with mating multiple times.

Practical beekeeping

Although the two studies described here don’t have an immediate relevance to day-to-day practical beekeeping, it’s worth remembering that poor queen mating is regularly blamed for queen failures e.g.¬†queens that develop into drone layers during the winter.

I’m going to write about drones later this year so for the moment will just make these points:

  • drone production is maximised to generate sexually mature drones for the swarming season
  • after eclosion, drones need to mature before being able to mate
  • drones live about 30 days and their sperm volume, though not necessarily viability, decreases as they age

Together this means that late in the season – perhaps late July or early August (though this will vary depending upon location) – the number of drones will decrease.

More significantly, the drones will be ageing.

In turn this means that late-mated queens may not mate with as many drones, or that the matings may not result in insemination.

Most beekeepers will be aware of queens that apparently ‘run out of sperm’ and become drone layers.

However, there may be less obvious problems with late-mated queens. I’m not aware of any studies on seasonality of queen mating and polyandry. However,¬†I would not be at all surprised if they exhibited a reduced level of polyandry.

And, as described above, these colonies are likely to exhibit reduced fitness.

Something else to consider when deciding whether to unite a colony late in the season or hope the last of your virgin queens mates successfully …


 

Bee bombs

The last couple of posts on overwintering survival and local bees have been heavy going 1. So, rather than more of the same, here’s something that is both informative and entertaining 2.

Though it maybe wasn’t at the time.

Six-legged soldiers

I’m currently enjoying reading¬†Six-legged soldiers by Jeffrey A. Lockwood. This is an account of the many and devious applications man has found for employing insects in warfare. Whilst the topic certainly isn’t ‘laugh out loud’ entertaining, the book is written in an engaging style with plenty of graphic descriptions, ample Biblical and historical references, and enough wriggly, stinging, aggressive insects to make¬†“I’m a celebrity, get me out of here!”¬†3 appear like a walk in the park.

As a beekeeper I’m pleased to see that bees feature significantly in the book.

And as a beekeeper who appreciates the importance of the integrity of the colony to bee survival I also found it a little distressing.

But as a source of all sorts of stories for friends and families over the forthcoming holiday season it probably cannot be beaten.

It’s pretty good on mosquitos as well …

Bombus away

The genus Bombus includes lots of the well known bumble bees e.g. Bombus terrestris (Buff tailed), B. pascuorum (Common carder) and B. hypnorum (Tree) . The generic name Bombus is derived etymologically from the Latin word bombus which means buzzing i.e. the noise a bee makes when it flies.

Bombus lucorum

Etymologically, the word bomb, has a similar origin Рvia bombe in French, bomba in Spanish, bombo and then the Latin bombus.

Boom also has a similar origin.

But that’s not the only link between bees and bombs.

Mushroom shaped clouds

Have you ever seen anyone drop a full brood box?

It is an amazing sight and one best appreciated from a distance and when wearing a full beesuit.

Bees do not appreciate being knocked, shocked or jarred. When I transport hives between apiaries I always give them several minutes to settle before removing the entrance block. If you don’t they tend to boil out the front spoiling for a fight.

So you can imagine that dropping a brood box from waist height achieves Рsimultaneously Рthe sudden jarring of the colony and the release of the bees.

Not so much ‘shock and awe‘ as shock and aargh!

The mushroom-shaped cloud of bees that are released are distinctly agitated.

In the absence of a beesuit you’re likely to get hammered.

Even with a beesuit there can be some uncomfortable moments.

And, since soldiers don’t routinely go into battle wearing camouflage BBwear battledress with an inbuilt fencing-style veil, this¬†neatly¬†brings us to using bees as weapons.

Package bees

These days a ‘package’ is one way to buy bees to start a colony.

But as a weapon, a colony of bees isn’t much use until it’s actually in something.

How do you carry them? How do you use them as a projectile?

Well, man is nothing if not ingenious when it comes to weapons development.

The Tiv people of Nigeria used a specially shaped horn, loaded with angry bees (presumably not so much Africanized as African ūüėĮ ). In the heat of battle these would be fired at the opposition, with the horn-shape ensuring the bees both reached the enemy and were kept at a distance from friendly forces 4.

But then, 9000 years ago, pottery containers started to be used for beekeeping … and it got a whole lot easier to move the bee bombs to the front line and drop them on the opposition.

A big, beautiful wall

A wall seems like an obvious way to defend yourself.

The enemy have to knock the wall down, or go over or under the wall.

And if they choose to tunnel under the wall then they’re going to be less than enthusiastic if the tunnel is filled with bees.

Which is what happened in 908 when the Scandinavians laid siege to Chester. The city’s fortifications were impenetrable, so they tunnelled underneath them. The siege was ended when all the city’s beehives were dumped into the tunnel.

Chester City walls. Originally built in ~100 AD by the Romans.

The Scandinavians appear not to have learnt their lesson as they were again repelled by bees while storming the walled city of Kissingen (Germany) during the Thirty Years War (1618-48). In this instance the bees were dropped from a height onto the Swedish forces.

The troops were heavily armed and armoured, and were unfazed.

Their horses were not.

The siege collapsed as the cavalry mounts were driven into a frenzy by the bees. Even now, most beekeepers are aware that bees and horses don’t mix well.

Again in the 1600’s, the besieged nuns of Wuppertal (Germany) knocked over all the hives in their apiaries before – wisely – hiding indoors. The maelstrom of bees drove the marauding soldiers away and the town was subsequently renamed Beyenberg (‘bee town’).

Bee boles

A bee bole is a recess in a wall 5 designed to house a hive of bees, which Рin the days when they were constructed Рwas likely a skep. Many castles and fortified town walls have bee boles built into them.

How convenient.

What could be easier than to drop these on the marauding troops trying to scale the ramparts or storm the drawbridge?

Bee boles in Kellie Castle, Fife, Scotland

As an aside, IBRA (The International Bee Research Association) maintain the comprehensive Bee Boles Register which is well worth searching if you are interested in historical beekeeping (or early bomb design).

Avoiding friendly fire

You’ll notice that a lot of these bees were being used in relatively close combat situations.

Having witnessed a brood box being dropped, I can assure you that bees are rather indiscriminate after a¬†“dropped from a great height onto a hard surface” experience.

Far better to use the container housing the bees as a projectile, launching them at the opposition from a safe distance.

Safe in terms of contact with the enemy … and the bees ūüėČ

The Greeks and subsequently the Romans developed and perfected the siege engine, capable of launching all sorts of things up and over defensive walls.

Including beehives.

Illustration of a ballista being loaded and drawn – note BBwear ‘Corinthian helmet’ style beesuit and veil.

The Greek ballista and the Roman onager were torsion powered siege engines developed between 400 BC and 350 AD. Both were capable of firing stones, often wrapped in combustible material set alight, with smaller later models also used as battlefield weapons firing projectiles 500 Р1000 yards.

They’d have barely broken a sweat firing one or more skeps at the enemy.

The Romans were so keen on bee bombs that there was a documented decline in hive numbers during the late Roman Empire.

And this enthusiasm continued … as did the demand for hives to hurl.

By the 14th Century those dastardly weapons designers had developed a windmill-like device capable of launching hive after hive from the end of its rapidly rotating arms.

Bees will not fly over water

But they will in a skep catapulted from a ship.

As the army developed entomological weaponry the navy exploited it.

As early as 330 BC pottery hives were being thrown at enemy ships during naval battles. Cannons and cannonballs eventually superseded 20,000¬†A. mellifera ligustica in a skep, but there is well-documented use of bees in naval warfare until at least the 1600’s.

Bees would therefore have been carried by warships for hundreds of years. It’s not documented how the colonies were managed or maintained. Perhaps they only fought local battles? However, since that rather defeats the purpose of a highly mobile navy it can be assumed that bees were probably transported long distances by sea … bringing a whole new meaning to the term migratory beekeeping.

Gunpowder and bees

Eventually the development of modern weaponry overtook the use of bees and beehives. Fortunately we don’t have to discuss the aerodynamic benefits of cedar¬†vs. poly hives 6.

Gunpowder and explosives made the Gatling gun-like skep-launching windmill catapult a relic of the good old days of warfare, when the infantry hankered after really cold days when the bees would be torpid and much less aggressive.

But, as a couple of masochists have already demonstrated, Apis mellifera is pretty tame where stinging is concerned.

Apis dorsata, the giant honey bee of South East Asia, is much bigger than our honey bee, and is reputed to pack more of a punch when stinging 7.

These bees build large exposed nests and the colony may have up to 100,000 bees in it.

Apis dorsata nest, a single exposed comb which may be a metre wide.

Which doesn’t mix too well with gunpowder or, more specifically, a firecracker containing gunpowder.

During the Vietnam War the Viet Cong would attach firecrackers to dorsata nests relocated to the jungle trails used by the enemy. As a patrol passed by they would ‘light the blue touch paper’ and set off the firecracker.

And then stand well back.

And, at about the same time (1960’s), the Americans were developing chemical warfare approaches using isopentyl acetate, the alarm pheromone, with the intention of spraying it onto enemy troops and redirecting the bees to attack them instead.

Six-legged soldiers

There’s lots more in¬†Six-Legged Soldiers … get a copy and enjoy reading it over the Christmas vacation. Jeffrey Lockwood is an entomologist and University of Wyoming Professor. The Sunday Times 2009 review of the book criticised it as ‘scarcely scholarly’, being a mix of myth, legend and historical facts.

I cannot imagine a better review and it probably explains why it is so entertaining to read ūüôā

Mite bombs

These are something altogether different to bee bombs … and for regular beekeepers, much more relevant.

A mite bomb is a heavily mite infested and collapsing colony that liberally spreads Varroa mites around the neighbourhood. Recent evidence suggests that this occurs primarily during late-season robbing of weak (mite infested) colonies by strong colonies.

This is the primary reason late summer miticide treatment should be coordinated over a wide geographic area. What’s the point of treating your strong colonies if they’re going to load up on mites when robbing weak colonies in the adjacent fields?

Which reminds me, and should remind you, that winter mite treatments will be needed in the next few weeks to ensure your bees get the best possible start to the new season.

We’ve had a protracted cold period here in Fife and my colonies will probably be treated in the next 5-7 days before there’s a chance they start brood rearing again.


 

Downstairs? Upstairs?

Colony inspections usually concentrate on the brood box. This is where all the action is. This is where the queen is and where there needs to be sufficient space for the colony to expand.

Or, if times are lean, sufficient stores and pollen to survive.

In contrast, the honey supers get no more than a cursory glance. There’s little of interest going on up there until it’s time to harvest the honey for extraction.

If the supers are light there’s nothing more to do other than hope for a good nectar flow in the future. In contrast, if they’re really heavy they might be ready to remove for extraction. If the frames are all capped the honey is ready.

Usually the supers are not heavy enough (a full super weighs something like 25kg) and they often don’t even get a glance, instead being bodily lifted off and left in a pile while the brood box is inspected.

Checking supers

Nectar has a high water content which the bees evaporate off during the production of honey. If they didn’t get rid of the water the stores would ferment.¬†Since honey is hygroscopic they then add a wax ‘cap’ to the honey-filled cell to protect their stores for the winter.

Nectar is generally stored in the supers, starting in the middle of the middle frames and moving towards the periphery. This is the warmest part of the hive and presumably the easiest to evaporate water from. Therefore, the central frames in the super are most likely to contain capped honey stores.

Ready to extract

Ready to extract …

All I do when checking a heavy super is to first briefly look at the central frame to see if the stores are capped. If they are not then there’s no point in looking anywhere else in the super.

If the central frame is capped then it’s worth looking to see if the outside frames are as well. If so a clearer board can be placed below the super and you can take the honey for extraction.

Actually, there’s a bit more complexity as sometimes the honey is ready to extract, but isn’t capped. I’ll deal with that another time. The point I’m (slowly) trying to make is that supers are rarely checked in any detail … until they’re full.

It’s therefore interesting what turns up when you do remove them for extraction.

Pollen and stores-free area

With a strong colony, the bottom super i.e. the one immediately above the queen excluder, often has no honey stored in a semi-circular area immediately above the brood nest. Sometimes the edge of this clear area, adjacent to the honey, contains a band of stored pollen.

This clear area indicates that the colony need more space. The workers are keeping it clear for the queen to lay, but the queen excluder prevents her from accessing it. Sometimes you can get the bees to backfill this area by switching the super with one higher in the stack.

“Billy no mates” brood

It’s not unusual to find a very few scattered capped pupae in a stack of supers. These are almost invariably drone pupae, irrespective of whether the drawn super comb is on worker or drone foundation. In ~24 supers I extracted last weekend I saw three or four.

Billy no-mates ...

Billy no-mates …

I’ve always assumed that these were due to laying worker activity. There are always a few laying workers in a colony, but their numbers are suppressed by a pheromone produced by unsealed brood. Laying workers can be a significant problem in queenless and broodless colonies.

Since workers are unmated, the eggs that laying workers produce are unfertilised and so develop as drones 1.

There may be other explanations for these singleton pupae¬†e.g. workers moving eggs up from the brood box. However, this doesn’t explain why they are almost always drones 2.

Clustered brood

Sometimes you’ll find a super packed with brood in all stages … wall to wall eggs, open and sealed brood. This happens when the queen has somehow sneaked above the queen excluder.

When this has happened to me I usually put it down to a lack of attentiveness in checking the underside of the queen excluder when opening the box. If the queen was on the underside and the QE is leant against the hive stand she can easily wander round to the other side, thereby giving her access to the supers.

Spot the queen

While checking supers for extraction last month I found one box – the lowest super of a stack of three – contained two or three frames with small amounts of clustered brood 3.

Another example of inattentiveness? Possibly, but there were some oddities about this colony.

Eggs and sealed brood ...

Eggs and sealed brood …

Firstly, there was no open brood … just eggs and sealed brood. I uncapped a few cells and the pupae were all just at the purple eyed stage. This is day 15 for workers and day 16 for drones. Since eggs hatch after 3 days this means that there had been a gap of at least 12 days when the queen wasn’t laying.

Half-sisters of the same age ...

Half-sisters of the same age …

Secondly, there was both worker and drone sealed brood present, but it was on separate frames. There was no drone brood in worker cells, which have characteristically domed caps 4.

Finally, I checked the brood box. There was plenty of brood in all stages – eggs, larvae and sealed pupae – in a busy hive. However, I didn’t see the queen (who was nominally marked and clipped) but by this time I was in a bit of a rush.

A partial solution

Some of these apparent oddities have a straightforward explanation.

The separation of drone and worker brood is because I use a range of different frames in my supers – worker foundation, drone foundation and foundationless. They start as matched boxes, but over the years have got completely mixed up.

All the drone brood was in a super frame originally drawn from drone foundation.

That was easy ūüėČ

However, why was there brood at all in the super if the brood box contained the laying queen?

Or should that read a laying queen?

Perhaps there was another queen in the super?

Aside from speculating about how she got there, or – if she was the original queen in the box – where the one ‘downstairs’ came from, there’s also the puzzle about why she’d taken a 12 day holiday from egg laying.

And where the hell was she now?

She’d been in the top box sometime in the last 3 days (because there were eggs present). However, although I’m reasonably good at finding queens, I searched in vain in this super (and the two above) and couldn’t find her.

Time to be pragmatic

Carefully looking through ~30 super frames takes time and I was running out of both time and patience. These three supers were ready for extraction and I still had half a dozen colonies to check.

I could continue looking and eventually find her … if she was there at all.

If she wasn’t, I’d obviously never find her.

What did I do?

I shook all the bees off the super frames – directly over the brood box5 – and took them away for extraction.

I’m a great believer in Occam’s Razor. The simplest explanation is probably the correct one.

I reasoned that there was probably one queen in the box. Any other explanation was going to get convoluted.

If there was only one queen she was either in the brood box or the supers.

If she was in the brood box then all was well.

If she was in the supers she’d hopefully end up in the brood box.

There was little point in using a clearer board if the queen was in the supers. Firstly, with brood present many bees would probably remain. Secondly, if the queen was present in the supers, they’d definitely not clear.

Super frames with brood ...

Super frames with brood …

And … what happened?

I got well over 60 lb of honey from the colony ūüôā

There was a blue marked and clipped queen in the bottom box when I checked the colony a few days later.

She was (still) laying well.

Unsatisfactory explanation

I suspect that the queen excluder was faulty or damaged. It was a wooden-framed wire one. If the wires were prised apart during cleaning or through carelessness the queen could get up into the super.

She could also therefore return to the brood box.

The 12 day gap in laying was probably explained by the queen returning to the brood box during this period.

The two short stints when she’d been ‘upstairs’ hadn’t noticeably left gaps in the brood pattern in the brood box – she might have only nipped up for a few hours or so. There were only a few hundred cells with eggs or pupae in the super.

And the most unsatisfactory thing of all … I thoughtlessly stacked the queen excluder with five others from the same apiary and so now need to carefully inspect all of them for damage ūüôĀ


 

Robbery

Robber

Robber

Another apiculture-flavoured tale of daylight robbery, literally, to follow the post on hive and bee thefts last week.

However,¬†this time¬†it’s not dodgy bee-suited perps with badly inked prison tats offering cheap nucs down the Dog and Duck.

Like other offenders, the robbers this week wear striped apparel, but this time it’s dark brown and tan, or brown and yellow or black and yellow.

I am of course referring to honey bees and wasps (Vespa vulgaris and V. germanica), both of which can cause major problems at this time of year by robbing weak colonies.

Carb loading

The season here – other than for those who have taken colonies to the heather – is drawing to a close. The main nectar sources have more or less dried up in the last fortnight. There’s a bit of rosebay willow herb and bramble in the hedgerows and some himalayan balsam in the river valleys, but that’s about it.

Colonies are strong, or should be. With the dearth of nectar in the fields, the foragers turn their attention to other colonies as a potential source of carbohydrates. Colonies need large amounts of stores to get through the winter and evolution has selected a behavioural strategy – robbing of weaker colonies – to get as much carbohydrate from the easiest possible sources.

Like the nucs you carefully prepared for overwintering ūüôĀ

At the same time, wasps are also wanting to pile in the carbs before winter 1. In the last fortnight the wasp numbers in my apiaries and equipment stores have increased significantly.

Jekyll and Hyde

Within a few days in late summer/early autumn the mood and attitude of colonies in the apiary changes completely.

During a strong nectar flow the bees single-mindedly¬†pile in the stores. They alight, tail-heavy, on the landing board, enter the hive, unload and set out again. There’s a glut and they ignore almost anything other than bingeing on it. Inspections are easy. Most bees are out foraging and they are – or should be – well-tempered and forgiving.¬†

Laden foragers returning ...

Laden foragers returning …

But then the nectar flow, almost overnight, stops.

Colonies become markedly more defensive.¬†They are packed with bees and they’re tetchy. There’s nothing to distract them, they resent the intrusion and they want to protect their hard-won stores 2.

At the same time, they quickly become more inquisitive, investigating any potential new source of sugar. If you shake the bees off a frame and leave it standing against the leg of the hive stand there will be dozens of foragers – many from nearby colonies – gorging themselves on the nectar.

If you spill unripened nectar from a frame they’re all over it, quickly forming a frenzied mass – probably from several different hives – scrabbling to ‘fill their boots’.

They also closely investigate anything that smells of nectar or honey. Stacks of equipment, empty supers, hive tools, the smoker bellows … anything.

Robbing

And it’s this behaviour that can quickly turn into robbing.

The foragers investigate a small, dark entrance that smells of honey … like a nuc in the corner of the apiary. They enter unchallenged or after a little argy-bargy 3, find the stores, stuff themselves, go back to their colony and then return mob-handed.

Before long, the nuc entrance had a writhing mass of bees trying to get in, any guards present are soon overwhelmed and, in just a few hours, it’s robbed out and probably doomed.

This is the most obvious – and rather distressing – form of robbing. Wasps can do almost exactly the same thing, with similarly devastating consequences.

Prevention is better than cure

Once started (and obvious), robbing is difficult to stop. About the only option is to seal the target hive and remove it to another apiary a good distance away.

Far better to prevent it happening in the first place.

The best way of preventing robbing is to maintain large, strong and healthy colonies. With ample bees there are ample guards and the colony will be able to defend itself from both bees and wasps. Strong colonies are much more likely to be the robbers than the robbed.

For smaller colonies in a full-sized hive, or nucleus colonies or – and these are the most difficult of all to defend – mini-nucs used for queen mating, it’s imperative to make the hive easy to defend¬†and¬†minimise attracting robbers to the apiary in the first place.

The underfloor entrances on kewl floors are much easier to defend than a standard entrance and small entrances are easier to defend than large ones. ‘Small’ might mean as little as one bee-width …¬†i.e. only traversable by a single bee at a time.

Smaller is better ...

Smaller is better …

You can even combine the two; insert a 9mm thick piece of stripwood into the Kewl floor entrance to reduce the space to be defended to a centimetre or two. If – as happened tonight when returning wet supers to the hives – I don’t have a suitable piece of stripwood in the apiary I use a strip of gaffer tape to reduce the entrance 4.

Gaffer tape is also essential to maintain the integrity of the hive if some of the supers are a bit warped. Wasps can squeeze through smaller holes than bees and the quick application of a half metre along the junction between boxes can save the day 5.

The poly nucs I favour have a ridiculously large entrance which I reduce by 90% using foam blocks, dried grass, gaffer tape, wire mesh or Correx.

Correx, the beekeepers friend ...

Correx, the beekeepers friend …

Don’t tempt them

Finally, reduce the inducement robbers – whether bees or wasps – have to investigate everything in the apiary by not leaving open sources of nectar, not spilling honey or syrup, clearing up brace comb and ensuring any stored equipment is ‘bee proof’.

You don’t need to inspect as frequently at this time of the season. The queen will have reduced her laying rate and colonies are no longer expanding. With no nectar coming in they should have sufficient space in the brood nest. There’s little chance they will swarm.

If you don’t need to inspect, then don’t. The ability to judge this comes with experience.

If you do have to inspect (to find, mark and clip a late-season mated queen for example 6 do not leave the colony open for longer than necessary. Any supers that are temporarily removed should be secured so bees and wasps cannot access them.

Wet supers

If you’re returning wet supers after extraction, do it with the minimum disruption late in the evening. These supers absolutely reek of honey and attract robbers from far and wide. Keep the supers covered – top and bottom – gently lift the crownboard, give them a tiny puff of smoke, place the supers on top, replace the roof and leave them be.

Returning wet supers

Returning wet supers …

In my experience wet supers are the most likely thing to trigger a robbing frenzy. I usually reduce the entrance at the same time I put the wet supers back and try to add wet supers to all the colonies in the apiary on the same evening 7.

I generally don’t inspect colonies until the supers are cleaned out and ready for storage.


 

Sphere of influence

How far do honey bees fly? An easy enough question, but one that is not straightforward to answer.

The flight range of the honeybee ...

The flight range of the honeybee …

Does the question mean any honey bee i.e. workers, drones or the queen? As individuals, or as a swarm?

Is the question how far can they fly? Or how far do they usually fly?

Why does any of this matter anyway?

Ladies first …

Workers

The first definitive experiments were done by John Eckert in the 1930’s. He located apiaries in the Wyoming badlands at increasing distances from natural or artificial forage 1. Essentially the bees were forced to fly over a moonscape of rocks, sand, sagebrush and cacti to reach an irrigated area with good forage. He then recorded weight gain or loss of the hives located at various distances from the forage.

Wyoming badlands

Wyoming badlands …

The original paper can be found online here (PDF). The experiments are thorough, explained well and make entertaining reading. They involved multiple colonies and were conducted in three successive years.

Surprisingly, Eckert showed that bees would forage up to 8.5 miles from the colony. This means they’d be making a round trip of at least 17 miles – and probably significantly more – to collect pollen and nectar.

However, although colonies situated within 2 miles of the nectar source gained weight, those situated more than 5 miles away lost weight during the experiments.

Gain or loss in hive weight ...

Gain or loss in hive weight …

Therefore, bees can forage over surprisingly long distances, but in doing so they use more resources than they gain.

John Eckert was the co-author (with Harry Laidlaw) of one of the classic books on queen rearing 2. His studies were probably the first thorough analysis of the abilities of worker bees to forage over long distances. Much more recently, Beekman and Ratnieks interpreted the waggle dance (PDF) of bees to calculate foraging distances to heather. In these studies, only 10% of the bees foraged ~6 miles from the hive, although over 50% travelled over 3.5 miles.

Queens

Queens don’t get to do a lot of flying. They go on one or two matings flights, perhaps preceded by shorter orientation flights, and they might swarm.

Heading for a DCA near you ...

Heading for a DCA near you …

I’ll deal with swarms separately. I’ll also assume that the orientation flights are no greater than those of workers (I don’t think there’s any data on queen orientation flight distance or duration) at no more than ~300 metres 3.

On mating flights the queen flies to a drone congregation area (DCA), mates with multiple drones and returns to the colony. DCA’s justify a complete post of their own, but are geographically-defined features, often used year after year.

There are a number of studies on queen mating range using genetically-distinguishable virgin queens and drones in isolated or semi-isolated locations. They¬†‚Äėdo what they say on the tin‚Äô, drone congregate there and wait for a virgin queen

In the 1930’s Klatt conducted studies using colonies on an isolated peninsula and observed successful mating at distances up to 6.3 miles

Studies in the 1950’s by Peer demonstrated that matings could occur between queens and drones originally separated by 10.1 miles 4. These studies showed an inverse relationship between distance and successful mating.

More recently, Jensen et al., produced data that was in agreement with this, with drone and queen colonies separated by 9.3 miles still successfully mating 5.

However, this more recent study also demonstrated that more than 50% of matings occurred within 1.5 miles and 90% occurring within 4.6 miles.

Just because they can, doesn’t mean they do ūüôā

Drones … it takes 17 to tango …

Seventeen of course, because that’s one queen and an average of 16 drones ūüėČ

There’s a problem with the queen mating flight distances listed above. Did the queen fly 9 miles and the drone fly just a short distance to the DCA?

Or vice versa?

10 miles ... you must be joking!

10 miles … you must be joking!

Or do they meet in the middle?

Do queens choose 6 to fly shorter distances because it minimises the risk of predation and because they are less muscle-bound and presumably less strong flyers than drones?

Alternatively, perhaps drones have evolved to visit local DCAs to maximise the time they have aloft without exhausting themselves flying miles first?

Or getting eaten.

It turns out that – at least in these long-distance liaisons – it’s the queen that probably flies further. Drones do prefer local DCAs 7 and most DCAs are located less than 3 miles from the ‘drone’ apiary 8.

Swarms

I’ve discussed the relocation of swarms recently. Perhaps surprisingly (at least in terms of forage competition), swarms prefer to relocate relatively near the originating hive. Metres rather than miles.

The sphere of influence

Effective foraging – in terms of honey production (or, for that matter, brood rearing) – occurs within 2-3 miles of the hive. This distance is also the furthest that drones usually fly to occupy DCAs for mating.

Queens can fly further, but it’s the law of diminishing returns.¬†Literally.¬†The vast majority of matings occur within 5 miles of the hive.

In fact, other than under exceptional circumstances, a radius of 5 miles from a colony probably represents its ‘sphere of influence’ … either things that can influence the colony, or that the colony can influence.

Why does this matter?

Worker flight distances are relevant if you want to know the nectar sources your bees are able to exploit, or the pollination services they can provide. In both cases, closer is better. It used to also be relevant in trying to track down the source of pesticide kills, though fortunately these are very much rarer these days.

Closer is better ...

Closer is better …

Workers not only fly to forage on plants and trees. They also fly to rob other colonies. I don’t think there are any studies on the distances over which robbing can occur, but I’ve followed bees the best part of a mile across fields from my apiary to find the source of the robbing 9.

All of these movements can also transport diseases about, either in the form of phoretic Varroa mites piggybacking and carrying a toxic viral payload, or as spores from the foulbroods.

Drone and queen flight distances are important if you’re interested in establishing isolated mating sites to maintain particular strains of bees. My friends in the Scottish Native Honey Bee Society have recently described their efforts to establish an isolated queen mating site in the Ochil Hills.

And I’m interested as I now have access to a site over 6 miles from the nearest honey bees in an area largely free of Varroa.

It’s not the Wyoming badlands, but it’s very remote ūüôā


 

Stroppiness

Perhaps surprisingly this isn’t about some of the contributors to online beekeeping discussion forums … ūüėČ I’ll discuss those next winter when their “shack nasties” – and associated rantings – get really bad.

What beekeeping is

Beekeeping should be an enjoyable pastime. It’s a great way to work with nature, to learn and continue learning, to understand and interact with the environment … and to make delicious honey.

Of course, it’s lots of other things as well. It can be hard and hot physical work at times. It can be infuriating when the weather and the bees and a thousand other things conspire to frustrate your plans.

And in our long winters it can require a significant level of patience.

What beekeeping isn’t

What it isn’t, or at least what it shouldn’t be, is something that fills you with dread, that hurts like hell or that threatens, frightens or – even worse – harms other people.

All of these things can be the result of having aggressive bees.

You can and should do something to ‘cure’ the colony of its aggression.

Bees should not naturally be aggressive. When not threatened they go about their daily business in a workmanlike 1 way, collecting pollen or nectar or water. Unless inadvertently trapped in clothing or hair they almost never sting; when they do it’s because the bee is trying to¬†defend itself.

Defensive bees can behave similarly to aggressive bees but¬†they are not the same thing at all. In this case the ‘cure’ is very different and probably involves the beekeeper rather than the bees.

Colony management, aggression and defensiveness

Beekeeping involves managing the colony 2. This necessitates regular inspections during the season.

It’s during inspections that both the nature of the colony¬†and the abilities of the beekeeper are tested.¬†It’s during these inspections that the beekeeper should try and distinguish between aggressive and defensive bees.

Aggression in bees is an unpleasant characteristic with predominantly genetic causes.

Defensive bees are reacting to a perceived threat and need to be treated more appropriately.

An aggressive colony

With little or no provocation, aggressive bees are out to get you. They buzz you a couple of times from yards away as you approach the hive, they ‘boil’ out of from under the crownboard when you gently prise it up, they bounce off your veil repeatedly or cling on tightly with the abdomen curled under them trying to sting.

Beekeeping should be enjoyable ...

Beekeeping should be enjoyable …

They burrow into the folds in your beesuit, they worm their way under the cuffs of your gloves, they attack your hands and the hive tool as it is used to lift the frame.

And they don’t stop when you close everything up and thankfully retire. They follow you across the apiary and continue to bombard your¬†(hopefully still veiled)¬†head.

Truly psychotic bees follow you up the field back to the car. You have to hang around until they lose interest or drive off still wearing the veil 3.

Before, during and/or after the inspection you’re getting stung. Depending upon the thickness of your gloves, your beesuit or your skin this might not hurt … but the build up of sting pheromone incites them even more. At worst, you’re forced to retreat from the onslaught.

It’s bad enough for the beekeeper. It’s much worse for anyone else inadvertently going near the colony, particularly after an inspection.

A defensive colony

A defensive colony is reactive rather than proactive. They react – in some of the ways described above – to rough treatment, to poorly timed interventions or to other perceived threats. They can and do sting, but if treated properly (i.e. better) they¬†don’t.

A well behaved colony can become defensive if it is jarred, jolted or – and this has to be seen to be believed – dropped. Bees that should be perfectly calm and well tempered can ‘go postal’ if maltreated.

The significant difference here is that they’re being badly or poorly treated. This is where calmness, confidence and experience – or ideally, all three characteristics – shown by the beekeeper is the major influence on the behaviour of the colony.

An ideal place to observe this is in the training apiary of a large beekeeping association. With an experienced beekeeper the colony can be wonderfully well-tempered, barely stirring from the frames.

With an almost complete novice – who inevitably works slowly and remembers all of the good advice they were told in the theory lessons – the colony is also OK, though if the hive is open too long they can become a little tetchy.

But with the intermediate (in experience and ability) beekeeper, who knows just enough to be dangerous but who thinks they know it all, who squashes a few bees every time they drop a frame back into the hive, who crushes a few more as they lever the next frame up, who waves their hands to and fro over the top bars and who smokes the colony too heavily … to this beekeeper the colony can appear aggressive.

But they’re actually being defensive … because they’re being mistreated.

Going postal

Sometimes even the most experienced and careful beekeeper can have a¬†D’oh! moment, instantaneously converting the calmest of colonies into a mushroom cloud-shaped maelstrom of psychotic bees.

Trainee beekeepers

Trainee beekeepers

I’ve seen an experienced beekeeper in a full training apiary inadvertently lift both a super¬†and the brood box to which it was propolised off the floor. Once in mid-air the propolis gave way, dropping a full brood box onto the ground.

Kaboom!

I doubt there was a single beekeeper in the apiary over a 20 yard radius who wasn’t stung.

But these weren’t aggressive bees. They’d done nothing when the crownboard was removed. They were simply being defensive – understandably – once their home was dropped from a great height onto the ground.

The grey area between attack and defence …

I’ve been reasonably clear cut about the differences between aggressive and defensive bees. Overly so. There’s a grey area when an otherwise calm colony, almost irrespective of how well treated it is, can appear aggressive.

A number of “environmental factors” can influence the behaviour of the colony. The most important of these are forage, weather and queenlessness.

Double trouble ...

Double trouble …

Bees are usually really well behaved when there’s a good flow of nectar. Open a colony when the OSR or lime is at it’s peak and you can do no wrong. Well, almost. However, open a colony when the OSR has recently¬†gone over or the lime has stopped yielding and the bees can be a bit tetchy.

Short tempered perhaps, not truly aggressive.

Similarly, open a colony – or certain colonies – as the barometer plummets or there’s thunder rumbling in the near distance and they can also be rather short tempered.

In my experience most colonies get a bit stroppy when a strong nectar flow dries up. In contrast, only some overreact to poor weather. I have opened colonies during a thunderstorm Рa long story, but it was to do with the day job long before we had the bee shed Рand they were fine. In fact, they appeared to welcome the shelter provided from the rain as I stooped over the open box rummaging around for 2-3 day old larvae.

Finally, a queenless colony is usually more aggressive … or, perhaps more accurately, defensive. If the queenless colony does not rear a new queen it will fail.

Curing aggression

I don’t think aggressive bees should be tolerated. They make beekeeping a chore. Worse, they frighten passers by and terrify the mellisophobic 4.

More worrying still is that aggressive bees might, either unprovoked or before calming down after an inspection, sting a passer-by who then goes into anaphylactic shock.

I don’t believe that aggressive bees are better at collecting honey, though many do.

Aggression is a genetic trait 5. The only cure is therefore to change the genetics of the colony. This means culling the old queen and replacing her with a new one. If you haven’t got immediate access to a replacement queen I’d suggest culling the old queen and uniting the colony with a strong, well behaved, colony.

Often the behaviour improves quickly – presumably due to the different pheromones at work – but it’s worth remembering that it will be 6-9 weeks until all of the brood and workers originating from the old queen are replaced.

Curing defensiveness

Physician, heal thyself¬†6 … or, more correctly, Beekeeper, heal thyself. Since defensiveness is a reactive response to poor handling the best solution is to improve the quality and care of inspections. And possibly improve their timing as well.

Don’t inspect when the weather is poor and be particularly careful when a strong flow has recently stopped. Treat the hive and the colony gently. Use as little smoke as possible. Carefully remove one frame and set it aside. Break the propolis seal on the remaining frames, one side at a time, gently and without waving your hands over the box.

Remove and replace each frame without crushing bees under the frame lugs. Don’t crush bees between the side bars when pushing frames together. Don’t shake bees off the frames unless necessary.

Work reasonably quickly, carefully and confidently.

The bees will appreciate it.

Record keeping

When I inspect colonies, in addition to things like space, stores and queen cells, I’m also observing the behaviour of the colony. I record behavioural traits (temper, following and running on the frames) in my notes. Any colony consistently performing badly on these criteria is sooner or later requeened.

I can excuse one bad day. I can just about accept a second. But three weeks of poor temper – particularly if the other hives in the apiary are fine – and the monarch will be replaced 7.

My notes from late last season showed that one very strong colony was developing aggressive tendencies. I couldn’t really face going through a double-brood box on a cool autumn day to find the old queen and unite the colony (and had no spare queens anyway).

The first inspection of the year has demonstrated they’re still a bit surly and, whilst not awful (no stings and no following), they’re probably only going to get worse as the colony builds up this spring.

If this shows any signs of happening I’ll unite the colony with another one – it’s too early in the season to have new queens and I’m not going to put up with bad behaviour.

Or impose it on passers by.


Colophon

Stroppy¬†(and hence stroppiness) is probably derived from obstreperous¬†and means bad-tempered, rebellious, awkward, or unruly’. It’s a word that’s been in use since the early 1950’s.¬†

Going postal¬†is a phrase that more specifically means stress-induced extreme violence. If you use Google’s ngram viewer to look the term up you’ll see that, other than a brief blip in the 1880’s, it’s a phrase only found in recent (>1990’s) English books.

Going postal ...

Going postal …

Going postal¬†has tragic origins as it refers to a spate of shootings in US post offices, by post office workers, in the late 80’s and early 90’s.

This post was timetabled to appear last week … major access issues with the website (repeated timeouts with visitor number reduced by about 40%) were repeatedly denied by my hosting provider and¬†took them ~4 days to resolve, by which time I decided it was better to postpone posting. I got stroppy but didn‚Äôt need to go postal ūüėČ