For the Love of Bees and Beekeeping archives

April 2015

First Came the Nest

by Keith Delaplane


Before I go much further into the evolutionary history behind honey bees and beekeeping I need to pause and consider something so obvious it almost escapes notice. In fact, it did escape my notice until just recently when I realized my near-mistake. And I’m talking of course about the nest – or more accurately, the nest and the ability to nest, taking license to use the word as both noun and verb.

It seems to me fundamental that a society of any kind presumes a certain shared space. That presumption may not hold anymore for modern human societies, given our obsession with technology-assisted global connectedness, but even so we still tend to think of societies in terms of assemblies of individuals attached by cultural bonds to a place in geography – American society, British society, the Eastern Apicultural Society, Western Apicultural Society, and so on. The colonial insects seem to think so too – or at least the entomologists who study them, because they invariably fit the description of individuals sharing a place – a nest.

Ever since January in these pages I have been telling a story about the evolution of the honey bee and why it matters for beekeepers. And so this month we envision our ancient honey bee ancestor as a solitary reproductive female – no need to call her a queen – a female with the cognitive capacity for searching for and finding a suitable nest, recognizing landmarks, and navigating about the neighborhood and returning home. Now I am no neurobiologist but I’m sure what I have just described is a herculean feat for a brain of any size, and in fact the vast majority of insects can’t do it (my wife says I can’t either). Most insects are free living in their local habitats with at most only a brief and transitory association with a “place” of any kind, for most of them a sheltered spot to deposit their eggs.

But in the insect order Hymenoptera – the ants, wasps, and bees – nesting is well represented. It may not look like much – a hollow reed or a dead-end tunnel, but among the wasps it is common, and in the bees their descendants it is universal. It is in fact a shared ancient character and if not exclusive to social species (even solitary bees do it) it is nevertheless a necessary precondition to sociality. Nesting allows for parental care of young, defense, food storage, and shelter – all of which are stabilizing forces friendly to the evolution of colonial life. It’s hard to imagine sociality taking off without it. And under the most advanced insect societies nesting has evolved far beyond hollow reeds into elaborate triumphs of animal architecture, like the nest of the honey bee.

It is this nest, of course, that interests us most. But it didn’t just pop up out of nowhere. It is a product of historic precursors of much simpler designs and functions, but unfortunately we do not have a fossil trail back to those precursors. The nests of modern primitively social bees are our best clues, and from them we can infer that nests of our honey bee’s ancient ancestors were simple cavities or tunnels elaborated over time with increasingly complex individualized cells for food storage and developing young. Individualized brood cells seems to be an ancient character, suggesting that protection of the developing young against parasites, weather, or even trampling by nest mates was strongly selected for (Fig. 1). The partitioning of cells is accomplished by a variety of materials including mud, leaf material, or wood paste. Modern species exist in which the cells are separated by exquisitely thin walls of soil glued together by glandular secretions. It is glandular products like these that were the precursors to wax glands in the modern bumble bees (Fig. 2) and honey bees and the subsequent introduction of beeswax into the natural world.

We are on firmer ground when we get ...

March 2015


The Mysterious Case of the Self-Sacrificing Worker

by Keith Delaplane


Among the many “Wow I didn’t know that” moments that confront a new beekeeper is the lesson that a worker bee dies after she delivers a sting. It’s true. Her barbed stinger implants in the skin of her adversary like a harpoon, tears away as she leaves, and its attached poison sac keeps throbbing away under involuntary muscles pumping out every last dreg of venom (Fig. 1). Meanwhile, the mortally wounded worker renews her assault with more determination than ever. She flies recklessly at the intruder, burrowing into fur or hair with a buzzing designed to distract and deter. She flies straight at the eyes. It’s as if she knows she’s a gonner so she might as well give it all she’s got. If she survives the encounter and returns to the nest she has only a few more hours to live before her internal injuries catch up with her and she dies, only to be unceremoniously disposed of by an undertaker bee.

Now I must shamefully admit that more than once I have taken a measure of revengeful comfort in this knowledge, especially at the end of a hot day with my hands sore and swollen and my sweaty shirt “pinned to my back.” But if we can dissociate ourselves a moment we may appreciate the necessity and efficiency of such a repertoire of behaviors. From the perspective of a large predator the honey bee nest is a lottery prize of protein and carbohydrate, and this stationary target requires extraordinary protection. The detachable singer/poison sac essentially doubles the defensive output of every single worker, freeing its possessor to return to the attack with additional intimidations. Multiply this one terrifying performance by tens, scores, or even hundreds of assailants and the result can be effective – the successful deterrence of a predator. And lest we humans get too complacent thinking ourselves somehow detached from the rest of nature, it’s insightful to consider the co-occurrence of Apis mellifera with ancient humans and our immediate ancestors on the African continent. It’s not far-fetched to wonder if the bees’ defensive behaviors we find so effective today are in fact evolutionary adaptations to us, arguably the most dangerous predator they’ve encountered in their natural history.

But for humans of a more modern variety the detachable sting also has the advantage of being convenient for research. What better way to measure defensive behavior than count bee stings on a red leather patch? This is in fact the most common method used by scientists working on defensive behavior, especially in areas with Africanized bees,
 (Fig. 2). One season while I was a graduate student some of my lab coworkers were studying defensive behavior of Africanized bees in Venezuela. Things were going quite well until one day the beekeeper who had volunteered his apiary for their experiments angrily ordered them off his property. Come to find out, his bees were responding so well to repeated leather patch tests that my colleagues were killing his entire apiary!

The defense behavior of the honey bee is as firmly engrained in the minds of most people as the fact that they make honey. No greater impediment to recruiting new beekeepers is to be found; in fact, some old-timers celebrate the fact that bees sting, saying – not without reason – that if it weren’t for bee stings then everyone would be a beekeeper.

Whether this is a good thing or bad is fodder for another conversation, but for our present purposes the self-sacrificing worker bee is a bit of a mystery and a clue that bigger issues may be afoot. For not only is the worker bee self-sacrificing, she is reproductively sterile – which makes no sense at all from an evolutionary point of view. In human terms, what the worker bee is saying is, “Not only will I give up reproduction in order to help my mother reproduce, but I will die if necessary to do so.” This is altruism practiced to a pitch rarely seen anywhere. And at the beginning of things it was a serious affront to Charles Darwin’s theory of natural selection. How can one pass on any genes whatsoever, favorable or unfavorable, if one isn’t even capable of reproducing? After all, in Darwin’s economy the only coin of the realm is successful reproduction, that is – getting one’s genes passed on to the next generation.

The awkward problem of the sterile worker was not lost on Darwin himself, and in chapter 7 of his seminal book On the Origin of Species he confronts the problem, and I quote:

“I will . . . confine myself to one special difficulty, which at first appeared to me insuperable, and actually fatal to my whole theory. I allude to the neuters or sterile females in insect-communities: for these neuters often differ widely in instinct and in structure from both the males and fertile females, and yet, from being sterile, they cannot propagate their kind.”

Now it’s interesting at this point to pause and consider one of history’s great What Ifs? and that’s the fact that while Darwin was putting the fine touches on his great theory in the south of England, a contemporary of his, an Augustinian monk Gregor Mendel, was untangling the basic laws of hereditary genetics 1000 miles away in Brno in what is today the Czech Republic. The two never met, and it is unclear to what extent either was aware of the other’s work. As a result, Darwin went to his grave ignorant of the mechanisms of inheritance, the consequences of which he nevertheless understood so profoundly. More practically, this historical near-miss meant that the great neo-Darwinian synthesis between genetics and evolution had to wait to emerge until the 1930s and 1940s. But if Darwin had known even a rudimentary level of Mendelian genetics he might have propelled forward evolutionary biology by decades and solved his “insuperable problem” as well. As it stands, he still hit pretty close to the mark, and again I quote:

“This difficulty, though appearing insuperable, is lessened, or, as I believe, disappears, when it is remembered that selection may be applied to the family, as well as to the individual, and may thus gain the desired end.”

With this sentence Darwin anticipated the neo-Darwinian explanation for social insect altruism that would emerge over 100 years later in the form of two papers published by 26-year old graduate student W.D. Hamilton at University College, London in 1964. It was Hamilton, advantaged with Mendelian genetics, who mathematically showed that the self-sacrificing behavior of an individual can still be adaptive if it promotes the survival and reproduction of near-kin who possess the same genes, or to put it in his own words: “a gene may receive positive selection even though disadvantageous to its bearers if it causes them to confer sufficiently large advantages on relatives.” Hamilton called this “inclusive” fitness, in distinction to “individual” fitness, and in so doing opened up all the resources of Darwin’s theory to biologists working with social insects and their teeming populations of sterile workers. Overnight, Hamilton unloosed a new science onto the world – so-called Kin Selection – the idea that individuals in social units can be predicted to behave in a way that promotes transmittal of their genes, even if those genes are in the bodies of near-kin. A seemingly puzzling behavior like self-sacrifice might make sense if it promotes transmission of that worker’s genes – even if it’s copies of those genes possessed by the worker’s sister.

In the case of honey bees Hamilton’s math is quite simple – 8th-grade level – and I spell it out in the sidebar. Doing the exercise is worthwhile because it really paints the matter in black-and-white.

To begin, let’s remember that natural selection operates on variation in a population – “good” genes vs. less-good vs. bad. Variation in a breeding population is a necessary condition to ...

February 2015

The Road to Apis mellifera: Putting the “Honey” in Honey Bee

by Keith Delaplane


I’m convinced that beekeeping provides a place for nearly every kind of human personality and interest. There are as many reasons for getting into beekeeping as there are beekeepers, and whether the initial attraction in your case was pollination for your garden, a sideline income, a family tradition, a social outlet, a shared family project, or a taste for honey, I hope that at least part of the reason was a frank fascination with the colonial life of these social insects. The term “social insect” was coined by humans as a descriptor for ants, termites, and group-living bees and wasps because we saw in their teeming populations, appearance of order, and elaborate nests an appearance of our own societies and infrastructures. But as we will see in the next few months, the term “social insect” is falling under new scrutiny.

My purpose here is not to unpack the vocabulary of sociobiology but instead to begin laying out an evolutionary history of our bee, the western bee, Apis mellifera. I’m doing this with beekeeping in mind – so we can better understand why it does what it does and how it has solved life’s problems. I suggested last month that it’s time we beekeepers become students of this evolutionary history so we can mine it for clues to modern bee health management. This column and the ones to follow are written in context to opinions I’ve expressed in earlier months about the shortcomings and limits to the management paradigms that have prevailed in beekeeping up to this point.

At the beginning of a journey it’s helpful to know where we’re going; so let’s remind ourselves that the honey bee is an example of a so-called eusocial insect, as spelled out by the venerable (and still quite active) biologist E.O. Wilson1, and by definition must express three characteristics: (1) cooperative brood care, such that individuals in the same nest help tend the common brood, (2) reproductive division of labor, such that some individuals abandon reproduction in order to help others reproduce, and (3) overlapping generations, such that some or all offspring remain at the nest to help their mother rear another generation. It is possible to possess some but not all of these characters, thus giving rise to lower and higher expressions of sociality. Sweat bees, for example, often express characters (1) and (2), but not (3). Among the eusocial species it is possible to be eusocial only part of the year. The bumble bees and many wasps fit this pattern; over winter these species exist in a solitary state – represented by single mated queens in hibernation. These solitary queens emerge in spring and single-handedly found a nest, forage for food, and rear the first batch of workers. Only at that point do the workers take over foraging, the queen stay at home, and the colony achieve a truly eusocial state. We call these ...


January 2015


Honey Bee Origins and Why It Matters

by Keith Delaplane


It can be inconvenient when new information comes along and changes things. A few years ago this happened to me in a small way (ok, a very small way), and it forced me to revise the exams I give students and Master Beekeeper candidates. The original question and answer went like this:

(True or False) Honey bees are native to the North American continent. False

But now it looks like this –

(True or False) Honey bees are native to the North American continent. True

All this came about because of a new paper announcing a fossil honey bee, member of the genus Apis, recently found in a shale deposit in Nevada dating from 23-5 million years ago. It was named Apis nearctica by its discoverers1 (Fig. 1) and represents members of the genus that moved northeast from their native southern Asia, crossed the Bering land bridge during the early to mid-Miocene epoch, were cut off from retreat by rising seas, and settled down as residents of western North America for a few million years. Absence of additional fossils leaves us ignorant whether A. nearctica was the sole North American representative of Apis, but it seems unlikely that others didn’t make the trip or that the group didn’t spin off new species during its long tenure here. In any case, its history is a story of the power of contingent events, in this case weather. Just as climate-induced low seas allowed its arrival, climate-induced high seas cut off its retreat, and climate shifts from warm-wet to cool-dry spelled its demise. Bottom line – this native American honey bee went extinct and left no living descendants, leaving North America empty of honey bees until humans reintroduced another Apis in historic times, our beloved western honey bee, Apis mellifera.

This story is interesting in its own right, but it also reminds us that every species we know on Earth today is the outcome of a similar string of contingent events in the history of our planet, a history in which organisms and their genes are constantly being challenged and shaped by conditions or accidents of nature. Sometimes members of a population possess a “survivable” set of genes that lets their possessors reproduce and pass those “good” genes on to the next generation. In this manner, successful genes tend to accumulate in a population and over time literally shape the species, whether behaviorally or morphologically, into a finely-tuned member of its ecosystem. Natural selection operates in an extravegance of time which means that a species becomes adapted to most of the extremes its habitat can throw at it. This is what makes it different from non-natural selection, ie., human-guided breeding: in the window of our short lifetimes we cannot see all the ramifications of our selections which may work well for our narrow purposes, but fall short given a new turn in the climate or other natural event that to us appears anomolous, but is nevertheless within the range of normal for that environment in geologic time. Thus, outcomes of natural selection are comparatively stable and intimately connected to geography of origin whether that means a continental region if you’re a honey bee or this pool of water versus that one if you’re a snail.2 This is why organisms aren’t randomly dispersed around the world: zebras in America or elephants in Europe.

It is also true that individuals, as well as whole species, eventually miss the boat and fail in this high-stakes game. Individuals die; species go extinct. Death applies to all, and the vast majority of species that ever existed in Earth’s history are now extinct.

But the glory of natural selection is not death, but its participation in the effusive generativity of life. Just as individuals can give birth to new individuals, species can give birth to new species. This happens through a process called speciation. It begins when a subset of a species’s population is cut off from others.


December 2014


Survivor-stock Beekeeping

by Keith Delaplane


Last month I indulged in a bit of nostalgia, recalling an abandoned apiary just down the road from where I grew up. It was in a meadow of unmown grass, snuggled along a wobbly fence, and shaded under a big tree. Over the coarse of 18 years I watched its unpainted boxes molder slowly into the ground. In spite of the unpromising facade, this little scene was a happy place full of life, as every summer the air above those boxes shimmered with the comings and goings of the inmates, numerous, healthy, and thriving in complete absence of any human hand.

Fast-forward to today, and one can’t hang around beekeeping circles long before an idea like this comes up in conversation – the idea of minimal human footprint on honey bee management. The variety of its labels underscores the variety of perspectives of its practioners. Called natural beekeeping, organic beekeeping, let-alone beekeeping, or survivor-stock beekeeping, the movement is a reaction against the perception that things have gone too far when it comes to chemical mite control and Big Agriculture. Like so many mega-trends in beekeeping, this one can ultimately be laid at the feet of Varroa mites, given that no other disorder makes such persuasive arguments for chemical treatments inside bee hives. Overnight, the arrival of Varroa mites in the 1980s transitioned beekeeping from one of the most chemical-averse agricultural industries to one of its most chemical-dependent, and this state of affairs has been a preoccupation of these columns the last few months.
So this month I want to linger a little on the idea of natural, survivor-stock beekeeping and the murky mixture of science and ideology that surrounds it. And to begin, let’s note that honey bees occupy a peculiar niche straddling the world of agriculture and the world of natural ecology. In the centuries since its introduction by European settlers, the western honey bee has become naturalized throughout North America. Encountering similar latitudes, seasonality, and plant life here as it had in its native Europe, Apis mellifera was equally at home wild in our forests or managed in our hives. Unlike some other exotic newcomers, a strong case has never been made that A. mellifera is damaging to our native species. To the contrary, it is a credible pollinator for many plants cultured and wild. Overall, among exotic species it is safe to say that honey bees are beneficial at best and neutral at worst. This ambiguity about honey bees’s niche explains how North American beekeepers have become skillful at wearing the “wild animal” hat versus the “domesticated animal” hat, as warranted by political circumstances or municipal zoning regulations.

Now the phrase “survivor stock” borrows from a lexicon attached to Charles Darwin, the great 19th century English naturalist who along with Alfred Russel Wallace first articulated a coherent idea of natural selection and biological evolution. The great insight of these thinkers was that natural forces, things like weather, seasonality, available food resources, predators, parasites, competitors, and any number of interactions with other species affect the ability of any one species to survive and reproduce. Those individuals of a species that possess favorable gene combinations have a greater likelihood of surviving, reproducing, and passing those favorable genes along to a next generation. Thus, genes favorable to a local condition tend to accumulate in a local population. In this manner, over geologic time the environment literally “shapes” a species so that it is finely tuned to survive and thrive in that locality. Those individuals that are “fit” survive; those that aren’t, don’t. Hence the popular phrase “survival of the fittest” to sum up Darwin’s and Wallace’s great insight.

When it comes to survivor stock in modern beekeeping, proponents are usually thinking in terms of honey bees that can withstand Varroa mites without the assistance of any miticide applied by the beekeeper. The idea is that genes for Varroa resistance exist in the western honey bee and that chemical miticide applications, although winning the beekeeper short-term relief, are ultimately counterproductive because they delay the inevitable reckoning of the “unfit”’ bees with the natural reality of Varroa mites. Better that the unfit die off and remove their genes from the population. What this looks like on the ground is, proponents decide in advance that they will forego miticide applications and accept the catastrophic losses that follow, hoping that a fraction of the colonies survive, however small, and serve as breeding stock for a subsequent generation. It is these golden survivors that theoretically possess genes to withstand Varroa mites. If one has the courage to endure the carnage that ensues in the early stages, he or she will be rewarded with a bee line that can increasingly withstand Varroa depredation without the need for environmentally-damaging or ideologically-offensive miticides.

Let’s pause here to note that such a practitioner is clearly wearing the honey-bee-as-wild-animal hat. It’s hard to imagine a similar mind-set taking hold in dairy cattle, beef, or poultry, and my gut reaction is to think it’s a bit severe to single out honey bees for this kind of strong medicine. But as always in these kinds of reflections, a good starting point is to accumulate published data on the matter.


November 2014

What’s next for Varroa control?

by Keith Delaplane


In my column last month I spelled out what I think is a fundamental tension facing beekeeping today: (1) controlling Varroa is priority #1, and pesticides have proven the best way to do this; however (2) it is impossible to use pesticides without getting unintended consequences in the form of non-target die-off, unknown lethal or sublethal synergies, or degradation of environmental assets. This tension can be extrapolated upward to apply to all of agriculture in general as we face the challenge of feeding 9 billion human beings by mid-century. In last month’s column I also asserted that these tensions must be resolved as co-priorities with neither one marinalized. The first part, that pests must be managed, is non-controversial. But it’s the second, environmental side of the argument that typically has the uphill battle in these kinds of discussions. What’s at stake here is the erosion of those environmental assets I mentioned, sometimes called “ecosystem services,” those free gifts of nature upon which all human enterprise depends. When I talk to farmer groups I ask them, What dollar value can you place on rainfall? soil fertility? erosion control? Well, pollinators fall into that same camp as do the thousands of beneficial organisms that combat the damaging ones. Nature is full of unsung assets like these, the value of which is not only incalculably huge but downright obligatory – we couldn’t live without them. But even though my second priority is at least as important as the first, it’s also the easiest to be set aside, given that we humans tend to favor immediate results at the expense of the deferred.

I was recently reminded of this when I read a review article on the environmental risks of neonicotinoid insecticides  - currently making headlines as both the most widely used class of insecticides in the world as well as a special hazard to pollinators. Neonics are systemic – available in all the plant’s tissues, which gives them 24/7 action against herbivorous pests but also gives them their peculiar hazard to pollinators who are exposed to the toxin when they collect nectar and pollen of treated crops. Neonics are regarded as comparatively safe to mammals and other vertebrates. They are usually applied as seed-coatings which further minimizes their contact with non-target species and reduces applicator labor. Patent expirations have rendered them relatively cheap. But it’s their efficacy that ultimately carries the day as neonics are famously effective against a wide range of insect pests. So effective, in fact, that in certain cropping systems neonics have ushered in a sort of anti-IPM backlash where their economy, ease of use, and efficacy have pushed growers away from cumbersome IPM sampling schemes in favor of chemicals to a degree reminiscent of the 1950s and 60s . Years’ worth of gains in IPM-oriented grower education and behavior modification were dropped overnight, so to speak, with the arrival of a sufficiently cheap and effective insecticide. This experience underscores the fact that any sustainable pest control scheme has to be efficacious, affordable, and practical or it will never get off the ground.

Perhaps I’m overstating things here, but I want to stress the co-priority of these two dynamics in my attempt to remove partisanship from the discussion. There’s no need for a “pro chemical” side versus a “green side” when it comes to the future of Varroa control. The winning formulas will possess the efficacy of chemicals and the sustainability of nature in equal measure. The trick is how to get there, and I’m not sure the trajectory of current research and education is sufficiently on target.

For staters, I’m not interested in a search for better or safer acaricides insofar as they are just another variation of acute toxin unloosed into the environment. If CCD and bee decline have taught us anything it’s that broad-spectrum toxins involve negative synergies and unintended consequences.

Second, my confidence is shaken in a classical IPM approach to Varroa based on thresholds. As I have written earlier, in some cropping systems the tolerable pest threshold is essentially zero. Given our growing knowledge of the tight relationship between Varroa and dangerous viruses like DWV, I suspect that our published Varroa treatment thresholds, already rather low, are still too liberal and should be adjusted downward and earlier in the year. Moreover, at a commercial level it has proven difficult for beekeepers to practice mite sampling at any meaningful scale. Should it be done at the level of colony? how about apiary? Given the inconvenience of counting mites and the importance of controlling mites it usually boils down to treatment by calendar instead of treatment by sampling.

And last but not least, I am on record expressing my ...


October 2014

The Messy Picture of Honey Bee IPM

by Keith Delaplane


Last month I spelled out an IPM regimen for Varroa control that I think does a good job summarizing the state of the science on this important topic. I also made it clear that in my opinion the state of this science is not very good. I can say this in good conscience because I’m an active contributor to this literature and acutely aware the limits of my own work. The truth is, it’s all too common nowadays for beekeepers to practice more or less exactly what I spelled out – and still experience catastrophic losses. Moreover, I reiterate the point I made last month that the fundamental premise of IPM – delaying chemical treatments until the pest reaches a critical population threshold – may be unrealistic if the pest causes serious damage at even small densities. I raised the specter of this for Varroa given the fact that Varroa is almost always confounded with one or more bee viruses, the most dangerous being Deformed Wing (DWV).

In other columns I have also laid out the conundrum that chemical-based pest control is a double-edged sword: the right miticide at the right time can cause immediate and dramatic mite control, but the miticides themselves cause sublethal damage to bees, interact dangerously with agri-chemicals and other Varroa miticides, and contribute to the toxic stew of background environmental contamination that’s earning agriculture a bad rap in the popular press. What’s the way out of this mess?

Lacking an immediate and brilliant answer, I default to what I think is the best move – getting real data on all the pieces of the puzzle so the discussion can proceed as much as possible from a factual basis. Fortunately, I am able to contribute new information on the state of IPM thanks to funding from the EPA and the USDA Managed Pollinator CAP – a four-year inter-institutional consortium, many members of which contributed a series of columns in this magazine from 2009-2012. I’d like to describe a project where my lab conducted a survey and boots-on-the-ground hive inspections in an effort to associate specific beekeeper practices with subsequent bee colony health.

Twenty beekeepers in the Southeast participated in an on-line survey that was designed to associate bee management and IPM practices with actual colony survival, queen replacement rates, and colony parasite and pathogen loads. Four of these participants were commercial beekeepers and the rest were hobbyists or small sideliners. Adult bee samples were collected from participants’ hives during Spring 2011, Fall 2011, Spring 2012, and  Fall 2012. All samples were processed for Varroa mite levels, pesticide residues for 7 organic compounds in hive materials, and pathogen loads. Statistical associations among these variables was done with multivariate analysis, with some of the significant associations shown in the Figure. As with any correlation analysis, these associations do not necessarily show cause-and-effect relationships, but rather show simultaneous movements between two variables which can be either negative – such that one decreases while the other increases, or positive – such that the direction of movement is the same. In the figure I have used iconic graphs to illustrate these relationships – positive or negative. These lines are not the actual computed lines; they are simply showing the direction of movement for simplicity of discussion.

Here are the results in bullets:

●    Varroa counts decreased as the likelihood increased that the beekeeper would use hygienic queens as an IPM Varroa control.
●    Varroa counts decreased as the number of simultaneous IPM practices increased.
●    Varroa counts decreased as the number of educational beekeeper meetings attended increased.
●    Colony winter mortality increased as the hive concentration (ppb) of coumaphos increased.
●    Varroa counts decreased as the hive concentration of coumaphos increased.
●    Queen supersedure increased as the likelihood increased that the beekeeper would use powdered sugar dusting as an IPM Varroa control.
●    The beekeeper’s likelihood of using thymol as a Varroa miticide decreased as the the hive concentration of coumaphos increased.
●    The beekeeper’s likelihood of using screen floors as an IPM Varroa control decreased as the the hive concentration of coumaphos increased.
●    The beekeeper’s likelihood of using survivor queens as an IPM Varroa control decreased as the the hive concentration of coumaphos increased.

These results support some of the premises and arguments in favor of IPM, namely the use of simultaneous IPM practices and use of genetically mite-resistant queens. I thought it was interesting that beekeepers who attend educational meetings are likely to have fewer mites in their hives. The results for coumaphos were typically paradoxical, with increasing chemical residues associated with both lower mite levels, as well as higher colony winter mortality. Unfortunately, powdered sugar dusting gains little traction from these data, showing a positive relationship with queen supersedure.

The chemical residue data give interesting insights into ...

September 2014

What Does Honey Bee IPM Look Like?

by Keith Delaplane


When it comes to honey bee health it’s almost become a platitude to extol the virtues of IPM – Integrated Pest Management – an approach to pest control that has dominated agricultural research, teaching, and extension since the 1960s. Over the years, IPM has made impressive advances into mainstream crop culture and animal husbandry. Its adherents can be found at all strata of agriculture from mega-farms to community farmers’ markets. The term began popping up in beekeeping magazines shortly after the arrival of Varroa mites in the 1980s, and today one can find IPM sections in beekeeping catalogs and copious literature about sustainable methods of controlling mites and keeping bees healthy. But it seems to me that there still remains no small confusion over what exactly IPM is. Or at least what it looks like in the case of beekeeping.

For starters, IPM is not the same thing as organic or natural beekeeping – philosophies of food production that, among other things, share an aversion to the input of toxic chemicals. It is not the same thing as let-alone beekeeping or the use of survivor stock. It does not mean an exclusive adherence to “soft” pesticides like formic acid or thymol. It’s almost, but not quite, synonymous with “sustainable” agriculture.

Now it’s true that IPM shares features of all of these, but what makes Integrated Pest Management distinctive is its, well, integration of multiple pest limiting tactics in an effort to keep pests at non-damaging levels. IPM is not explicitly anti-chemical but rather treats chemicals as a last resort after a string of prior measures. If the prior measures work, then there’s no need to use pesticides. If they don’t, then pesticides can always be called into service.

What all this means is that IPM leans heavily on the idea of treatment thresholds – research-derived pest levels that are known to represent the highest pest density that is tolerable without tipping over the crop – or bee colony in our case – into an irrecoverable decline. It’s at this point, and not before or after, that IPM says acute pesticides should be applied. But the emphasis – the whole point of it all – is on delaying that tipping point, ideally forever. It’s a subtle difference that, in my opinion, represents the most responsible way to include acute toxins in our modern food production system.

Now treatment thresholds are variable depending on time of year, location, and the past experience of scientists and beekeepers. Ideally, they are derived from experiments that set up a range of Varroa mite levels, monitor and treat at different times of the season, and retrospectively determine which mite levels and treatment episodes resulted in optimum colony strength and survival. In the table I have put together some published treatment thresholds for representative parts of North America. The most common values reported are number of mites counted from a 24-hour sticky sheet on the bottom board or else mites per 100 bees recovered from an alcohol-wash or sugar-shake method.

Once a beekeeper is armed with a locally-relevant treatment threshold, then the name of the game is to keep the treatment threshold from occurring. This is done by using those “prior measures” I talked about. Those prior measures in classical IPM, developed largely with crop plants in mind, have traditionally been lumped into three categories or triads: biological pest controls, cultural pest controls, and genetic host resistance.[7] Biological pest control refers to the use of beneficial predators, pathogens, or parasites to control a pest, and in row crops they play a huge role in pest insect IPM. But except for some positive evidence with pathogenic fungi[8] and a few anecdotes about pseudo-scorpions and predatory Hypoaspis mites, the literature on natural enemies of Varroa is pretty thin. With reflection this is not surprising, as the large nests of social insects constitute buffered safe habitats (sometimes called “homeostatic fortresses[9]”) in which the parasites themselves may not have evolved many natural enemies.

But when it comes to cultural controls and genetic host resistance, beekeeping IPM fits well with the classical model. Cultural controls refer to steps a grower can take to create growth and rearing conditions that discourage pest populations. In the case of Varroa we have a couple examples. One group showed that ...


August 2014


Mice, Old Combs, and the Reliability of Bee Science

by Keith Delaplane


I f one writes a monthly column on science-based recommendations it shouldn’t be a surprise if the idea of science-based recommendations comes up for discussion. This happened to me recently.

A reader emailed me with objections to my May column on screened bottom boards. How, this person asked, could I recommend screened bottom boards for Varroa control when no study has shown them to be statistically different [there was actually one that did] and all I could cite were “trends” for beneficial effects? “We must accept research based on significantly different levels to have any reasonable assurance that the effects of the study are true or accurate.” Could it be possible there were errors in the experiments, too small sample sizes, or other unaccounted factors? “Maybe,” he concludes, “the test should be repeated to verify the results.”

This is a lot to think about. These are indeed questions that should concern every practitioner or consumer of science. They strike at the heart of how we (especially in the developed world) make judgments about knowledge claims and use the time-tested scientific method.

I think most readers of this magazine would agree that scientific explanations for beekeeping practices are to be preferred over other contenders like tradition or hearsay. But I hasten to add that just because an explanation is “scientific” doesn’t necessarily mean it’s right. The strength of a scientific explanation is greater or lesser depending on the quantity, quality and consistency of the evidence, and the best kind of evidence is numerous independent studies, each directly asking the question and each finding results consistent with the others. Repeatability is a good thing and increases our confidence that the conclusion is in fact true.

Sometimes the evidence is “scientific” only in the sense that the evidence was collected in a scientific context, secondary to some other purpose. Let me give an example: I’m pretty sure there’s a correlation between the solidness of a queen’s brood pattern and her overall brood quantity. In other words a queen that lays a solid pattern will make more brood than a queen that lays a spotty pattern, even if the spotty brood is spread all over the place. This hunch is based on years’ worth of collecting both types of data – but always in the context of other experiments and questions. The data are certainly good in the sense that I am confident of their accuracy, but they have never been used to directly challenge the hypothesis, so I should stay tentative on that count.

The literature is rich in “orphaned” data like this – hence the term “data mining” and the statistical technique of “meta analysis” which is nothing more than a question catching up with pre-existing data to save the investigator time, enlarge the statistical sample size, and broaden one’s inferences across time and space. But for meta analysis one must take more than the usual precautions against bias and advocacy. We are all humans, and I think it’s a fiction to think that any scientist can be truly unbiased. I admit every time I do an experiment, deep down inside me (or not so deep down!) there’s an answer I want to be right. One can imagine a ...


July 2014

More about Bees and Chemicals

by Keith Delaplane


It was a typical bee meeting that could’ve been Anywhere, USA. It was the annual state convention on a brilliant Saturday afternoon and I was guest speaker. I was wrapping up what I thought had been a successful lecture to a friendly crowd. The Q&A time was lively and prolonged – always a good sign – and the subject if not a perennial favorite, at least a perennial priority, Varroa Management. The next questioner was a sophisticated looking lady in one of the middle rows; she had hung on my every word with a concentration so intense I had vainly read it as supreme engagement bordering on admiration. I couldn’t have been more wrong.

Standing up, composing herself, and drawing a deep breath, she declared what my sketchy memory records as the following:

“How dare you – an employee of a reputable institution – stand up there and advocate chemicals that harm bees and the environment. How can you do that when bees are dying all over the world and it’s perfectly possible to control mites without any chemicals at all? Shame on you! You should be pointing the way to controlling mites naturally. But, of course, you can’t do that because the big Chemical Companies are funding your research.”

These are not happy moments for a speaker. In this case, I responded with some lame, over-worked, and over-conditionalized response that left me looking evasive. Any argument that puts “bees” and “chemicals” together in the same sentence already has the advantage, and my battle was uphill from the start. In the end I was rhetorically defeated, and I walked away frustrated, feeling that emotion had won over reason.

But sitting here in the safety of my office I can reflect and appreciate, if not perhaps her facts – for the record, big industry comprises less than 2% of my career funding – at least the intent behind her emotional words. It is true that the quantity and diversity of pesticide exposure endured by our bees is staggering. A survey of bee and hive samples across 23 states found residues of 121 different pesticides and their metabolites. The average number of residues per sample was 6 and the highest had 39.1 This solemn news is compounded when one considers the infinitude of lethal synergies possible in a witch’s brew like that. For example, we already know that fluvalinate – the active ingredient in Apistan – results in lethal synergies when it is comes in contact with coumaphos – another Varroa miticide – or chlorothalonil – a widely used agricultural fungicide. Yet another bad synergy happens when the Varroa active ingredient thymol comes in contact with chlorothalonil.2 What other devilish combinations exist out there yet to be uncovered?

Moreover, not all toxicity is the same. As I mentioned in my last column, there are acute toxicities which result in ....


 June 2014

Varroa Mites: Is the Cure Worse than the Disease?

by Keith Delaplane


It’s a moment for reflection when one learns that something one has always believed has been blown to bits by a piece of new information. This has been happening to me a lot lately, especially as new research keeps coming out shedding light on the mysterious web of causes surrounding bee decline. An example came out not long ago at the hands of Dennis vanEngelsdorp and his co-authors in a paper describing factors contributing to bee mortality in the U.S.1. It was shown that increasing levels of the miticide coumaphos in brood were associated with colonies less likely to express signs of Colony Collapse Disorder (CCD) and die. Coumaphos is registered for Varroa control in the U.S. under the product name Check-Mite+, but it and other synthetic miticides have been implicated by some as problems in their own right – vilified as just so many more man-made chemicals that interfere with the bee’s biology, encourage chemical resistant mites, and probably do the bees more harm than good. I’ve never been extreme in accepting this point of view, but I was certainly sympathetic to it and I had to look over vanEngelsdorp’s results twice to make sure I was reading them right.

Now the most obvious explanation for results like this is that commercial beekeepers are no slouches when it comes to feeding colonies, managing queens, and controlling mites. With livelihoods at risk and financial stakes high, these professionals are going to make sure that all the easy explanations for mite outbreaks are eliminated. In light of intense mite control, the stress of a move to California becomes comparatively tolerable. And if it is true that coumaphos does bad things to bees, it is also true that Varroa mites do worse, and I came away from this paper thinking it was less an endorsement for chemicals as an indictment against Varroa.

So, if things reduce to some kind of mathematical formula with Risk from Mites on one side of the equation and Risk from Miticides on the other, then it’s important that we do everything in our power to minimize the risk from miticides. And the way to begin that is to get accurate information.

There has been interest in low-grade or so-called sublethal effects of bee hive chemicals for a while now. Some of this comes from the fact that high-grade or acute toxicity is easy to diagnose – piles of dead bees. But what about low-grade toxicity that exacts a slow, mysterious, and cryptic cost on colony health? Until bee decline has been fully understood, it is important that we uncover as many of these “sleeper” toxicities as possible. There is, for instance, evidence that fluvalinate (Apistan) increases mortality of young drones, reduces their body weight, and decreases their sperm count2 and that coumaphos (Check-Mite+) reduces queen larva acceptance rate, body weight of virgin queens3, and longevity of adult queens.4 We know that in-hive acaricides affect gene expression5 and immune response6, rendering the bees susceptible to diseases and other pesticides.

We also know that fluvalinate and coumaphos can interact with each other to elevate the bee toxicity of fluvalinate to damaging levels.7 This bit of news was particularly startling to this bee expert who has spent decades advising beekeepers to “rotate your miticides to reduce chemical resistance to mites” – only to find out that the practice could lead to damage to the very bees we are trying to protect. What’s an advisor to do?

My group weighed in on the question of sublethal effects on bees with a study comparing three legal bee hive chemicals – the miticides fluvalinate (Apistan) and coumaphos (Check-Mite+) and the popular wood preservative copper naphthenate8 used to protect woodenware against decay and termites. This paper is publicly available at the open access journal PLoS One.

Now, a personal note here about copper naphthenate. Back in the early 1990s I wrote and hosted a PBS television show and book on beginning beekeeping called A Year in the Life of an Apiary. Of all the footage filmed and ink printed on the details of building hives, installing bees, managing growth, controlling disease, harvesting honey, and marketing the crop – the one sequence that has garnered more comments than all the rest combined is a brief section where I belabor the point of treating woodenware with copper naphthenate. I can’t count how many times I’ve been asked “What was that green stuff you were painting hives with?” so it is with pardonable bias that I included this compound in our study as I consider the original toxicology work done on it rather sketchy and I had lingering curiosities whether the material was in fact safe for bees.

In order to eliminate beginning miticide residues, we set up experimental bee colonies with factory-new equipment and took the extra precaution of starting each comb with a one-inch strip of wax-less plastic foundation (beeswax foundation may harbor residues of bee hive chemicals). Each colony was ....

 May 2014

On Screen Hive Floors

by Keith Delaplane


If my columns the last couple months have had a theme it might be called: Great Ideas that Didn’t Work. So it’s high time I interject some optimism on the matter. Now, it may not be a lot of optimism, but when it comes to low-chemical Varroa mite control, the name of the game is numerous simultaneous pest control components, any one of which may be insufficient to keep mites at bay, but when used together might do the trick.

So for this month I’m happy to bring up the subject of screen hive floors. Only a decade ago most hive floors were simply solid, but nowadays it is common for suppliers to offer screened hive floors, and their use has widely increased. It is generally thought that screen hive floors allow improved ventilation, increase brood production, and give a measure of Varroa mite control.

Screened hive floors have been used in some parts of the world for years1, but it is safe to say they have never been mainstream until lately: the only mention they garner in Eva Crane’s authoritative 1990 Bees and Beekeeping2 is a 1981 reference to “netting” floor boards used in Norway. Aside from their use in ventilation, they have also been tested for their effects on overwintering3 and moisture content in honey4.

The arrival of Varroa mites in North America suggested new purposes for screen hive floors. Tibor Szabo noted their usefulness as a means to protect bees from sticky sheets placed on hive floors to catch and count dropping mites5. Noting that up to 50% of the mites dropping onto bottom boards are still alive, Pettis and Shimanuki6 reasoned that if a barrier could be placed between these mites and the colony that they would not be able to reinfest the bees. What they came up with was a screen insert that fit between a standard (solid) bottom board and the hive body (Fig. 1). The device was not properly a screen floor, per se, rather a screen suspended above the floor. It did not open to the ground and debris (and presumably mites) could accumulate beneath it. The authors monitored mite populations every month with standard sticky sheets and found a downward trend in mite numbers in colonies with screens in June and July, but screens did not prevent an exponential jump in mite numbers in fall. Curiously, they found that brood production was significantly higher in colonies with screen floors. Pettis and Shimanuki concluded that screen floors could not stand alone as a mite control measure, but could be a helpful in an integrated control approach.

My lab got in on the action with a project my then-student Jamie Ellis published in 20017. We joined the floor modification of Pettis and Shimanuki with two miticides to examine their combined effects on Varroa. Like the previous authors, we found a slight increase in brood production with screen floors and, similarly, a slight but non-significant drop in mite levels, in our case 15%. We had evidence that Apistan was not working in our test apiaries, but in those colonies treated with Apistan plus a bottom screen mite control was restored to an average of 44%. We concluded that the modified screen floor exerted a small but beneficial downward pressure on mite populations and helped compensate for chemical resistance in mites.

Up to this point the published research on screen floors in the U.S. had centered around the modified design of Pettis and Shimanuki. It was Harbo and Harris8 who foregrounded the idea of the simpler design known today in which the bottom is fully screened and open to the ground (Fig. 2). These authors compared mite population growth and brood production in colonies with solid bottom boards or an open screen and found a trend for fewer mites in colonies with screen floors. Interestingly, the open screen group also had a lower fraction of their mite population in brood cells – a proxy indicator of the fecundity of the mite population, given that mites can only reproduce in brood cells.

Another paper came along describing the usefulness of screen floors integrated with mite-resistant queens9. There was a consistent downward effect of screens on mite populations, albeit rarely statistically significant, but this time there was also evidence that screens interacted with mite-resistant queens in a way to elevate mite control above that found with only one component – an effect sometimes called a synergy or a positive interaction. But not all was rosy with screens. In one experiment this same paper detected a significant reduction in stored pollen and honey in colonies with screen hive floors.
This discussion by no means exhausts the literature on screen hive floors, but it is clear that consistent trends are detectable. Screens rarely provide a dramatic level of mite control, but they do predictably put a downward pressure on mites and tend to increase brood production. I put together a summary below of some of the published findings about screen hive floors (Table 1).

Now for me a curiosity remains – ...

April 2014

On Small-Cell Foundation

by Keith Delaplane

The worldwide spread of the Varroa mite has morphed beekeeping from one of the most chemical-averse agricultural industries to one of the most chemical-dependent. It is widely thought that the only practical control for mites is the use of toxic miticides inside the hive. It is equally agreed that this is a sorry state of affairs because it’s a fine line between killing one arthropod and sparing another when both live in the same space.

As a result, there has been an outpouring of creativity among scientists and beekeepers with the aim of controlling mites without pesticides. One of the most visible of these projects has been the use of small-cell foundation.

The idea behind small-cell foundation is the fact that mites can only reproduce in bee brood cells, and a few studies have shown that, if given a choice, Varroa mites prefer comparatively large brood cells. This preference seems to hold whether mites are given a choice of brood cells made by European bees (larger cell size) versus African bees (smaller cell size)1 or if mites are experimentally given a choice of three cell sizes: 4.8 mm, 5.2 mm, or 5.3 mm2. It is reasonable to assume that colony mite population growth would be correspondingly reduced in colonies with brood cells smaller than the mites’ natural range of choices.

These observations ultimately led to a commercially-available product, a small-cell foundation that measures 4.9 mm per cell compared to conventional foundations ranging around 5.2 mm to 5.4 mm. This product is available in bee supply catalogs and many beekeepers use it as part of an overall Varroa control strategy.

The trouble is, the practice has not held up to experimental challenge. I was part of a team that tested small-cell foundation as a means for reducing colony mite populations3, and I want to give an overview of our experience here.

We set up three independent studies with colonies with one of two brood cell types: small-cell (4.9 mm cell width) or conventional-cell (5.3 mm). In one study, ending colony bee population was significantly higher in small-cell colonies than conventional-cell. However, the main interest for small-cell foundation is its effects on mites, and on this count small-cell didn’t do so well. In fact, small-cell colonies were significantly higher for mite population in brood, percentage of mite population in brood, and mites per 100 adult bees. We were forced to conclude that small-cell foundation does not slow Varroa population growth. This conclusion is reinforced by the fact that the experiment was replicated independently three times with start dates ranging between spring and fall and test periods ranging from 12-40 weeks.

The work of Martin and Kryger4 seemed to support small-cell foundation when they observed that mortality of male offspring mites was increased under conditions which constrict the space between the bee pupa and male. However, these same authors pointed out that, “reducing cell sizes as a mite control method will probably fail to be effective since the bees are likely to respond by rearing correspondingly smaller bees.” We found some evidence to support this. In one of our trials we compared average body weight of bees reared in the two cell types and found that average bee weight was smaller in small-cell colonies.
These results were pretty convincing to me, but other authors have chimed in too. Small-cell foundation was shown to be ineffective in reducing mite population growth in independent tests in Florida5 and New Zealand6. In fact, I am unaware of any publicly-accessible peer-reviewed papers that directly support it.

If our results were convincing to ...

March 2014

On Comb Replacement

by Keith Delaplane

One of the brightest spots on the American beekeeping landscape is the Bee Informed Partnership BIP is a USDA-funded consortium of university and USDA bee scientists focused on surveying the beekeeping industry to identify management factors that affect colony health and survival. The nice thing about BIP surveys is that the data are summarized and the trends publicized on the BIP website. One need only peruse the BIP homepage to get an appreciation of the power of data and its ability to strip away guesswork and make sense of complicated phenomena. There one finds not only annual colony loss reports, but clear and succinct summaries of noteworthy trends in the surveys that give clues to good management.

It is one of these succinct summaries that caught my attention the other day. The online BIP article, “Brood comb management and treatment of dead outs: National management survey 2011-2012” informs us that beekeepers who replaced 50% or more of the brood combs in their colonies experienced 30.7% colony overwinter loss; beekeepers who replaced 10% of brood combs lost 21%, and beekeepers who replaced none of their combs lost 22%. The 30.7% loss rate suffered by the 50% replacers was statistically significant.

I admit these results surprised me, as I count myself among the many voices down the years who have advocated that beekeepers should regularly replace their brood combs. My opinions on this matter draw mainly from the research of my first graduate student, subsequent lab manager, and now authority in her own right, Jennifer Berry. One of the chapters of Jennifer’s master’s thesis dealt with the effects of old brood combs on colony strength.1 In each of three years she set up apiaries of 21-24 four-frame Langstroth nucleus colonies and assigned each colony one of two treatments – establishment on first-year, newly-drawn beeswax brood combs or establishment on old black combs of unknown age (Fig. 1). She then tracked colony brood production, brood survivorship, emergent adult weight, and adult populations. We measured brood production by overlaying a plexiglass grid marked in square centimeters over the brood and visually adding it up (Fig. 2). We measured brood survivorship by overlaying a sheet of transparent acetate onto a comb and marking on it the location of 10-40 cells of live, uncapped larvae (Fig. 3). Three days later we returned the sheet of acetate to its corresponding frame and made note of surviving brood to determine percentage survivorship.

Colonies housed on new comb produced more brood and heavier ...

February 2014

The Rise and Fall of the Dust-ructor

by Keith Delaplane


   It was an idea whose time had come. Varroa mites were raging as the front-runner of beekeeping problems; the synthetic chemicals used to control them were coming under scrutiny as problems in their own right, and non-chemical alternative remedies were looking smarter and better all the time. And what could be a safer alternative than powdered sugar?

 The idea of treating bees (and mites) with finely ground dusts such as wheat flour or confectioner’s sugar had been around a while, the idea being that dust impedes temperature-sensing organs on the mite’s forelegs that it uses to locate bee hosts, or impairs the mite’s ability to keep its grip on the bee1, or induces a grooming response from bees that dislodges mites2. Moreover, once a mite is dislodged and falls onto a dusty hive floor, it may have trouble moving around and eventually die of starvation. A handful of early studies suggested a degree of efficacy in dislodging mites with dust, either for diagnostic purposes or outright control, but these references were for the most part hidden away in non-English literature or obscure conference proceedings. But by the early 2000s there were new studies giving the matter more exposure in mainstream journals. Mite dislodging rates between 77%3 to more than 90%4,2 were being reported, and American beekeepers and bee scientists were taking notice. But no one can pretend that it was a revolution taking place. For starters, there was no consensus on such details as mode of delivery, quantity of dust, timing and intervals of treatment, or even the basic question whether dusting worked.

A convincing field-scale study finally came out in 2009 from Florida – and the results were not promising5. Amanda Ellis and her co-workers dusted the top bars of brood combs every two weeks from April until the following February and found no difference in colony strength or mite populations between dusted colonies and non-dusted controls. When I read these results, I was ready to write off powdered sugar once and for all, but that was not to be.

My intrepid staffers Brett Nolan, Ohad Afik, and Jennifer Berry weren’t quite as pessimistic as I and reasoned that several questions remained unresolved. To begin, they argued that (1) the efficacy of dusting had not been adequately tested in the context of a brood-free period (bee colonies in Florida are rarely brood-free), and it was exactly a brood-free period when one could expect maximum control when the whole mite population was on adult bees and vulnerable to dislodgement. They also argued that (2) more than one delivery method should be tested – especially one that could work at a commercial scale, and finally, they thought that (3) more than one treatment interval should be tested. In short – they dreamed up a whole new experiment6, and who was I to resist such youthful initiative?