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The Lives of Bees

The Lives of Bees

The untold story of the honey bee in the wild by Thomas D. Seeley, Professor of Biology at Cornell University.

Well, not quite untold, but this is a highly informative and entertaining book about the biology of honey bees living wild, primarily in the Arnot Forest, near Ithaca in the Finger Lakes region of New York.

Thomas Seeley conducts simple, elegant experiments to address interesting or important questions about bees. He then presents the studies and the conclusions in an easily understandable form, unencumbered by statistical mumbo-jumbo or extensive caveats and qualifications. 

This makes the work very accessible, even for those with no scientific training. You don’t even need extensive knowledge of honey bees; he explains the background to the experiments in sufficient detail that they are comprehensible without lots of prior knowledge.

For this reason, this is an ideal book to introduce a new beekeeper to the biology of bees.

However, for reasons to be covered separately, I think the suggestions it makes on practical beekeeping is very poor advice for the new beekeeper 🙁

A three part story

Essentially the book is in three parts, divided into eleven chapters.

After a general introduction there are three chapters that provide a historical perspective to the bees in the Arnot Forest and, more generally, to beekeeping. Not the practical aspects of beekeeping, but the interaction of humans and bees over tens of thousands of years.

The Beekeepers and the Birdnester by Pieter Bruegel (c. 1568)

Chapters 5 to 10 cover key aspects of the biology of the colony. These are:

  • the features that influence selection of a nest site
  • an overview of the annual cycle; spring build up, overwintering etc.
  • colony reproduction i.e. swarming
  • thermoregulation of the colony
  • collection of pollen, nectar and water – the food and stores needed for survival
  • defence of the colony – from microscopic viruses to (distinctly) macroscopic black bears

The final chapter – Darwinian beekeeping – contains Seeley’s suggestions for changes to beekeeping practice, informed by the observations presented in the six preceding chapters.

I’ll discuss Darwinian beekeeping another time as it deserves a post of its own.

Something for everyone

Each chapter is accompanied by a couple of pages of explanatory notes and there is a 19 page bibliography should the reader want to consult the primary sources.

An interested lay person could spend hours enjoyably reading about the biology of wild-living honey bees without ever consulting the notes or references. These don’t litter the text, making the book very much more accessible to those unused to the sort of cite-every-statement-to-avoid-offending-the-peer-reviewers style of writing that plagues most reviews (Bloggs et al., 1929b).

Alternatively, if you really do want to find out the original source you usually can, by consulting the notes and the references. Inevitably some things are missed, but that’s the nature of an eminently readable tome covering about a million years of Apis mellifera biology, 4500 years of beekeeping and at least 300 years of scientific observations about bees.

One of the great aspects of Seeley’s writing is that things are often presented with reference to some long-lost study which would otherwise have been forgotten.

A couple of weeks ago I discussed the importance of checking hive weights at this time of the year. The rate of stores usage increases significantly as more brood is reared. How do we know this increased rate of stores usage is due to increased brood rearing, rather than just correlating with it?

Seeley presents his data on colony weight changes but does so with reference to Clayton Farrar’s study of brood rearing by colonies lacking pollen in the 1930’s. These used only half as much of their stores because brood rearing needs pollen. Farrar’s study was published in the American Bee Journal in 1936.

There are several examples in the book where modern molecular studies are juxtaposed with some of the great observational science of the first part of the 20th Century. As someone involved daily at the gene-jockey end of science, this historical perspective alone makes the book worth purchasing.

Wild vs. domesticated bees

Throughout the book Seeley focuses on bees living in the wild i.e. without help intervention from beekeepers. His contention is that it is only by studying bees in their natural habitat that we’ll be able to properly understand what they need to survive and thrive when managed.

Seeley has studied bees in the Arnot forest for at least 40 years. He can therefore provide a ‘before and after’ view of the impact of the introduction of Varroa which probably occurred in the early 1990’s. Surprisingly, the overall number and density of colonies living in the forest in the 1970’s is about the same as it is now. This is discussed in several places in the book.

How can wild bees cope with the mites that, uncontrolled, generally destroy a hived colony within a year or two? His explanations of this is the underlying thread running through much of the book and the primary topic of the final chapter.

Are bees domesticated? This topic gets an entire chapter of its own. The genetic changes that species undergo during domestication 1 are not seen in honey bees.

Although perhaps not ‘domesticated’, through environmental manipulation we have significantly changed our relationship with bees. We now determine the size of the colony (or at least the space it has). By moving or manipulating the hive we influence what it produces (e.g. propolis, Royal Jelly, heather honey). We also control whether or not it reproduces. Indeed, most beekeepers try to stop their colonies reproducing (swarming) as it results in the loss of bees, and honey.

Throughout the book comparisons are made between the choices ‘wild’ bees make and the choices made for them by beekeepers. For example, the thermal conductivity of the hives used by beekeepers compared with a nest in a tree trunk.


Not really.

The strapline on the front of the book indicates that this is the untold story of the honey bee in the wild.

In reality it’s not.

More accurately it’s a very readable compendium of studies published by Seeley and others over the last century or so.

But that’s hardly going to make copies of this £25 book fly off the shelves, so ‘untold’ it is.

In fact, several aspects of the biology of the wild-living honey bee will be familiar to readers of this site. I’ve covered studies by Seeley in discussion of bait hives, drifting, robbing, polyandry and mites in swarms. A quick search turns up ~25 posts in which he gets a mention.

In addition, anyone who is fortunate enough to have already read Honeybee Democracy will be familiar with many bits in the chapter that cover nest site selection. Similarly, the bee lining methods used to locate nests in the Arnot forest have been described in exhaustive detail in his previous book Following the Wild Bees.

Don’t let this put you off.

Honeybee Democracy takes ~250 pages to describe in exhaustive (but still entertaining) detail how swarms choose new nest sites. This topic, together with all sorts of fascinating stuff on comb building and propolis, takes just part of the 40 page ‘Nest’ chapter of The Lives of Bees.

Bee·lining box, in cutaway view to show construction detail.

Similarly, the mechanics of bee lining don’t really get described in the new book, but the wild-living nests discovered using this method feature throughout.


Absolutely. It’s an excellent book.

But be aware that, in addition to a comprehensive account of how bees live in the wild, there’s an agenda here as well.

The sleeve notes (does anyone really read these?) include the words ” … and how wild honey bees may hold the key to reversing the alarming die-off of the planet’s managed honey bee population”.

Global beehive numbers 1968 – 2018

What alarming die off?

The graph above is of the global total of beehive numbers over the last 50 years or so. During this period the number has increased by ~1.7 times.

Of course, there are more beekeepers over the last 50 years (and the global population has more than doubled). This increased number of beekeepers are having to work harder to maintain (and increase) the stocks they manage.

But increasing they are … 2

It is therefore both inaccurate and an oversimplification to claim that there’s an alarming die-off in honey bee colonies.

Perhaps the sleeve notes are just to help boost sales?

Something a bit spicy to entice the browser to think that the book they are holding contains the ‘untold’ secrets to ‘saving the bees’?

Save the bees … save humanity

It’s not the first time ‘Save the bees … save humanity’ has been used as a marketing ploy 3. Here’s a graphic I regularly use to introduce my talks on rational Varroa control.

Save the bees ...

Save the bees …

Pity the image is of a wasp 🙂

Again, don’t let these minor errors in the sleeve blurb put you off.

Whatever the relevance to practical beekeeping (or reversing the “alarming die-off”), the first ten chapters provide the best overview of the lives of wild-living honey bees written by an acknowledged master of science communication.

I read a lot of stuff about honey bees, for work and pleasure.

The Lives of Bees had a wealth of information I was unaware of.

Buy it, or borrow it from your library … you won’t be disappointed.


Scouting for girls

A swarm of bees is a wonderful sight … if it’s arriving in your bait hive. It’s still dramatic, but perhaps slightly less wonderful, if it’s disappearing over the fence from your apiary 😉

Although some will disagree, I think beekeepers have a responsibility to both control swarming of their own stock, and capture – or attract – swarms lost by others. Although perhaps incomprehensible to us, some don’t share our passion for bees. Many are frightened and a large swarm is an intimidating sight for the melissophobic.

A small swarm ...

A small swarm …

Aside from frightening people, if they move into the church tower or an old hollow tree, they’re likely to develop high levels of Varroa and the pathogenic viruses the mite transmits. As a consequence, they can act as a source of disease to bees in local apiaries, until they’re killed off in the winter. Which they almost certainly will be.

I therefore always put out bait hives in late Spring, well ahead of the expected start of the swarming season (which often coincides with the oil seed rape finishing). I’ve described the basics of bait hives previously – a National-sized, bee-smelling box containing one frame of old, dark comb and half a dozen foundationless frames. I often use stacked supers from the, otherwise-awful, Paradise poly hives for this purpose.

Dyb dyb dyb

One of the greats sights of the swarming season is the appearance of the first scout bees at the bait hive. First one or two, then a dozen and, within hours or days, hundreds. They check out the entrance and the inside the bait hive. They fly all around the perimeter. They’re unaggressive and you can get up close to watch them at work. If you listen carefully you can hear them pinging into the sidewalls and floor of the bait hive as they move about inside.

They actually probably measure the volume by a combination of walking around the inner walls and determining the mean free path length – the average length of all straight lines from wall-to-wall in the hive – in short flights. For an interesting and easily readable discussion of the physics behind this I recommend the short paper by Nigel Franks and Anna Dornhaus (PDF) How might individual honeybees measure massive volumes?

In my view, this alone is a good reason to use foundationless frames in a bait hive.

Scouts often arrive early at the bait hive and leave late. Their numbers will fluctuate with weather and temperature – they’ll disappear altogether in the rain, but reappear in force once a shower has passed.

Scouting around

This short video was taken about 9am, two or three days before a large prime swarm occupied the bait hive. The first scout bees I’d seen had been almost two weeks previously. By midday there were hundreds of bees checking the hive.

However, if you look closely, their behaviour is distinctively different from a colony ‘in residence’. They’re much more hesitant in entering the hive and they tend to check the immediate environment much more closely. In contrast, foragers returning to a colony don’t bother doing a couple of laps of the hive … they approach directly and enter with minimal delay.

Seeley’s swarms

The definitive guide to how scout bees choose suitable locations and then ‘persuade’ the swarm to relocate is Honeybee Democracy by Tom Seeley. This is an outstanding book, beautifully written and illustrated.

Swarm of bees

Swarm of bees

Swarming is a two-stage process. The queen and flying bees leave the hive and settle nearby – on a branch, a fence post or (irritatingly) the top of a nearby conifer – creating the classic ‘beard-shaped’ cluster of bees. If you’re lucky with your timing and their location you can knock these into a box and, voila, you have a new colony.

Tom Seeley describes his own studies (based on the equally elegant work of Martin Lindauer in the 1950’s) that determine how scout bees convince the swarm to move from this temporary staging post to a new nesting location – a tree cavity, the church tower or your bait hive. The scout bees use a variation of the classic waggle dance – on the surface of the swarm hanging in the tree – to ‘persuade’ other scouts to check out the location they’ve found. Through repeated cycles of recruitment and reinforcement a consensus is reached and the scouts then lead the swarm to their new home.

That’s the abridged version. Read the book. There are subtleties and anecdotes throughout Honeybee Democracy that mean it’s the sort of book you can go back to time and time again, learning something new each time.

Early scouts

I was puzzled by the swarm that arrived in my bait hive. The first scouts appeared early in the first week of May. I was abroad from the 7th to the 14th and confidently expected the swarm to be waiting for me when I got back. However, it wasn’t until at least another week had elapsed – during which scout bee interest continued unabated – that the swarm arrived.

Honeybee democracy

Honeybee democracy

I went back to Honeybee Democracy and re-read the second chapter (‘Life in a honeybee colony’) and learnt – or was reminded – that there are early scout bees that are able to judge both nest site quality and the state of the colony preparing to swarm. These scouts are at work before the colony swarms. Uniquely these bees are judging both the availability and suitability of new homes and the readiness of the colony to swarm.

They can also tell whether it’s a nice day. The coincidence of these factors – good weather, readiness of the colony to swarm (i.e. sealed queen cells) and potential nest sites – initiate a behavioural change in these scouts that leads to the colony swarming.

Are these scouts the earliest sign of swarm preparation?

What Seeley doesn’t say is just how early in the swarming cycle these scout bees start their initial explorations.

Queens take 16 days to develop from new-laid eggs to eclosion, and just nine days to the sealing of the queen cell. If we assume that the first scouts I saw were from the same hive that subsequently swarmed (and delivered itself to my bait hive) then these scouts were out and about well-before queen cells were even started.

Of course, I have no way of telling whether the first scouts I saw were from the same colony that finally swarmed and arrived. Nevertheless, it’s an interesting thought. Perhaps scout bee interest in a bait hive pre-dates the first definitive swarm preparation signs beekeepers can usually recognise – the appearance of charged queen cells?

Considering the density of beekeepers (by which I mean apiaries 😉 ) in the UK it’s not easy to see how this would be useful … unless you’re the only beekeeper on an isolated island.

However, if you see do scout bee activity at your bait hives it might be worth being more assiduous than usual when checking your own colonies in the neighbourhood.


Dyb dyb dyb is an abbreviation for ‘do your best‘. This was part of a cub (not scout) ceremony and was followed by Dob dob dob (‘do our best‘). It was abandoned in the late sixties, but lives on in tricky questions on Qi.


Scouting for Boys

Scouting for Boys …

The title of this post is a play on ‘Scouting for Boys‘, the book on Boy Scout training, written and illustrated (originally) by Robert Baden-Powell and published in 1908. The book contains sections on scoutcraft, woodcraft, tracking, camp life, endurance, chivalry, life saving and patriotism. It was the inspiration for the scout movement and Baden-Powell was the founder and first Chief Scout of the Boy Scouts Association (and the founder of the Girl Guides). It is estimated that 4 million copies sold in the UK alone, with global sales in the 20th Century exceeding 100 million.

The book even contains reference to honey bees with the statement that bees form a ‘model community, for they respect their Queen and kill their unemployed’.

The Boy Scouts of America used to offer merit badges in Beefarming (1915-1955) and Beekeeping (from 1955). The Beekeeping merit badge was discontinued in 1995.

Scouting for Girls is an English pop rock band. They have recently announced their 10th Anniversary Tour (Oct/Nov 2017) which means they’re much too new for me to know any of their music 😉


The Drifters cont.

The Drifters ...

The Drifters …

Not the legendary American doo-wap/R&B vocal group but instead a quick follow-up to a recent post on drifting in honey bees. I discovered an interesting article in a 2011 issue of American Bee Journal in which Wyatt Mangum (Mangum, W. [2011] Varroa immigration and resistant mites ABJ 151:475) quantified mites introduced with bees from other colonies. The experiment was straightforward and quite clever … a number of colonies were prepared with very low mite numbers, overwintered and then miticides (unspecified, but from the remainder of the article I’m assuming Apistan) were applied continuously for the rest of the season. This would kill all the mites present. With a Varroa tray in place it was therefore possible to count newly introduced mites throughout the season. These must arrive with drifting workers, drones (not sure if drones ‘drift’ as such … perhaps there’s a better term for their itinerant wandering?), bees that have abandoned other colonies or potentially robbers. The newly infesting mites would of course be killed by the miticide after introduction and before reproduction. They could therefore easily be counted on the Varroa tray under the open mesh floor.

The results were striking … in one year between mid-May and early October an average of 1415 and 1001 mites were introduced to each of the seven ‘recipient’ colonies in two separate apiaries. Mite arrivals weren’t evenly spread, but peaked during a late summer dearth of nectar … perhaps, as suggested by the author, as other colonies started to run out of stores. The source colonies were not identified, but were not within the test apiaries. Whatever the cause, this represents a very significant influx of up to 7-10 mites per day. In Mangum’s experiment these mites could not replicate (due to the miticide that was always present). Had they been able to do so the impact on the recipient colony, in terms of numbers of mites transmitting viruses within the hive, would have been much greater.

The impact of drifting and mite reinfestation

The impact of drifting and mite reinfestation

Using BEEHAVE this impact can be modelled. In untreated colonies (solid lines), primed with 20 mites at the beginning of the year (and default conditions as previously described), the average mite level at the year end is ~430 (n=3) having reached a maximum of ~600. Using the same infestation period as reported by Mangum¹, with a mite infestation rate of 7/day (the lowest he observed), the average mite levels at the year end were ~2700 (n=3), with maximum levels reaching ~3800 in late summer (dotted lines). In this simulation the introduced mites can reproduce. Therefore, within just a few months, phoretic mites carried on workers and drones from other colonies, have the potential to raise mite levels in the recipient colony to dangerously high levels – significantly higher than the maximum recommended level of 1000/colony. This is potentially of fundamental importance in strategies to effectively control Varroa.  It should be noted that in a repeat of his study this large scale infestation was not observed. This suggests that this type of infestation – from outwith the apiary – may only be a problem in certain years or under specific conditions. One possibility that comes immediately to mind would be a collapsing feral colony or abandoned (or potentially not abandoned, but just completely ignored and untreated … or ‘abandoned‘ as some might say 😉 ) hive within foraging distance.

Ample opportunity ...

Ample opportunity …

Interestingly, a recent study has looked at the influence of a number of honey bee pathogens on drifting (or inter-colonial transmission as they rather long-windedly call it) behaviour. Of the viruses, Varroa and Nosema tested, only the presence of high mite levels influenced drifting … but not in the direction that might be expected. Distance between colonies in an apiary was the major factor that influenced drifting and ~17% of tested workers had drifted (with a third to half of these being apparently unrelated to other colonies in the test apiary). Surprisingly, colonies with high Varroa levels were more likely to acquire drifting workers, though the mechanism for this was unclear. The increased mixing through drifting would ensure that these colonies would likely end up with a greater diversity of viral and other pathogens though whether these colonies could, later in the season, act as a source rather than a sink for mites was not tested.


Drone …

Finally, returning to the subject of drifting bees and the ABJ … in the February 2016 issue there’s an interview with Tom Seeley (of Honeybee democracy fame … Sharashkin, L [2016], ABJ 156:157) in which he states that, when quantified, 34% of drones in his apiary colonies were from other hives. This article – on Surviving without Treatments: Lessons from Wild Bees – also discusses the importance of colony separation to coping with Varroa. The feral colonies Seeley studies are located at least half a mile apart in woodland. When recovered and relocated together in apiaries (‘beeyards’ as they’re called in the US) they rapidly succumb to mite-transmitted viral diseases, whereas those maintained some distance apart (30+ metres) survive. Seeley makes the point that pathogens evolving in closely-spaced colonies are likely to be more virulent, whereas those that are in distantly spaced colonies should be less virulent (or they’ll kill the host colony before being transmitted). Seeley is referring to the virulence of Varroa but I think his comments apply better to the viral payload carried by the mite. This is a relatively minor distinction but these observations further emphasise that drifting in honey bees is clearly a major factor in mite, and consequently disease, transmission … and therefore needs to be considered in control.

STOP PRESS – A recent Bee-L post highlighted a further study on the influence of re-infestation. Greatti et al., (1992) showed that ~2-14 mites/day/colony were acquired in their test apiary during June-August, and that this number rose to up to 75 mites/day/colony in September and October². This type of re-infestation can occur by drifting as already discussed, or by workers in the sentinel colonies robbing out mite-infested collapsing nearby hives or feral colonies.

¹In the Mangum study the mites did not infest the sentinel colonies at an even rate of 7+/day. Instead there was a marked peak in mid-season. I’ve not attempted to model this. Clearly if mites don’t arrive earlier in the season the overall levels would be lower (as they wouldn’t have the chance to reproduce). However, an influx of mites in mid/late-season might just arrive at the wrong time to damage the all-important winter bees … the topic of a future post.

²Greatti, M., Milani, N. and Nazzi, F., (1992). Reinfestation of an acaricide-treated apiary by Varroa jacobsoni Oud. Exp. Appl. Acarol., 16: 279-286