# An Option Towards Developing Treatment-Free Bees



## honeydrunkapiaries

Hey been working a great deal on my management plan for next year, incorporating a lot of Michael Palmers ideas. Another way of looking at it, is my clutch is broke and I have nothing better to do until the parts come in. This is something I wrote up on my blog today about an idea I had, not something I am currently practicing, but if you write something down its much easier to remember it down the road. You can see a better formatted copy at http://honeydrunkapiaries.blogspot.ca/ but anyway here it is.

* An Option Towards Developing Treatment-Free Bees*

Being a treatment-free beekeeper is a marathon, not a sprint. First you must learn how to keep bees, and keep them alive. Secondly, you must acquire the genetics that are resistant, or otherwise cope with the varroa mite. This second option can take a great deal of time. Many of the “successful” treatment-free beekeepers have been breeding their own stock for thirty years or better. It is possible to get their stock, but it is unlikely Dee Lusby’s or Michael Bush’s bees will survive a tough Canadian winter –if we even could import their stock, which we cannot. Finding survivor bees is actually the easy part of the equation. If they survive winter you don’t really have a choice but to breed from the bees that survive. Mites however, are not that easy.

Many people ascribe to the “Live, and let Die,” Bond philosophy of beekeeping. Let nature take its course of natural selection and what you are left with are your survivor stock. The largest problem with this philosophy is that you must ask yourself “Where do the mites go?” Weak colonies or dead outs will simply get robbed out, and the robbing bees will take them back to their parent colony. Perhaps that colony was a survivor stock until those robber bees brought back a disproportionate amount of mites, and will now also die because they have exceeded their mite threshold
_
“A key point to remember is that the relative infestation (percent, or mites per 100 bees) is more important than total mite population—a large colony can handle more mites than a small one. At much above a 2% infestation in spring, honey production drops off severely. At much above 5% in fall, colony winter survival suffers (although the fall “economic injury threshold” numbers by various authors range from 1% to 11%) (Currie & Gatien 2006). “_ (Randy Oliver, IPM 3 Fighting Varroa 3: Strategy – Understanding Varroa Population Dynamics)

This method has a great deal of risk; you could end up with no bees at all! Also, it does not really make for a good business model in the years that it takes to get to that point.

Another method is to treat your bees every spring and fall, do mite counts and breed from the bees with the lowest mite counts. This method gives the bees a bit of a crutch; are they surviving because of the treatment, or genetics? There is also not a 100% they will survive despite the treatments. Testing for hygienic behaviour can also be a key marker for your breeding selection. Hygienic behaviour also reduces other diseases like American Foulbrood, and Chalkbrood.

The purpose is then to develop a system creating a balance between two opposing ideologies: sustain a sound productive apiary whilst developing mite tolerant survivor stock; as well as keeping true to a more natural method of beekeeping.

My proposal is this. Do not treat your bees prophylactically; let them have normal and acceptable pressures on them at all times. However, if a hive is in danger of mite related collapse treat it as to not contaminate the hives around it. Keeping chemical contaminated comb, honey and stock out of your developing resources is important. 

How do we approach this?

There is a hive in danger of collapse, being robbed out and spreading mites. Treat it with an organic acid, mark the hive body and all the frames so that you know those hives have been exposed to this chemical. Once the mites have been ‘knocked down’ let it overwinter. If it successfully overwinters, split that hive up into nucleus colonies giving each a queen cell from the breeding survivor stock. A queen cell as opposed to a laying queen is important; it allows a short brood break to reduce the mite population, and give the new queen a chance to prove herself. Swarm cells in the spring should be rather easy to find.
The marked frames (contaminated,) are then pulled as time will allow and the wax is rendered. That wax is then put in a separate stock from your uncontaminated wax and used for purposes other than foundation.

What we will have accomplished is:

We have not endangered our existing hives by allowing them to rob out the infested hive
We have created nucleus colonies from a hive that would have otherwise died
We have perpetuated the genetics of our survivor stock
We have isolated any chemical contaminates from our treatment-free operation


How can we model this into our existing apiary management?

Firstly, you have to monitor for mites. Whether it is a sticky board, an alcohol shake, it doesn’t matter. You need a baseline to know which of your hives have a mite problem and which don’t. Which are candidates for breeder stock, and which hives are going to need help?

Secondly, hygienic testing; either begin testing for hygienic behavior or buy bees from breeders who are selecting for these traits and breed from them.

Fall I believe may be the best time for integration: the mite populations are at their peak, struggling hives will be apparent, and winter is coming (had to throw the Games of Thrones reference in there). You take your surplus honey off. At this point we are still treatment-free. We isolate which hives are over their mite threshold and apply a treatment. Those hives/frames are clearly marked, I am thinking green paint marker for plastic frames, or green thumb tacks (green kind of has a chemical connotation to it). Feed and maintain those hives as regular and let them overwinter.

Spring has come. Your non survivor stock has died through the great selector –winter. If your isolated hives have survived split them into as many nucleus colonies as you can, giving each brood, honey, and feed. Acquire queen cells by either: harvesting from survivor stock, or buying new genetics from a breeder. If you let them raise their own queen, you will simply be perpetuating unwanted genetics and you have just exponentially increased them by the amount of nucleus colonies you have made. By giving them a queen cell you create a brood break. This allows the mites developing inside the brood to hatch out, and gives the existing mites no larvae or eggs in which to create more mites. It will not get rid of the mites, but it will decrease their numbers. By the time the queen hatches, mates, and begins laying 16 days +/- will have passed where no eggs have been laid, and the majority of mites will be in the phoretic stage. Nucleus colonies generally do not suffer the effects of mites as severely as large colonies (more bees = more mites). The queen can be evaluated, and the nucleus colony can be put to various other uses. 

Summer is business as usual more or less. Monitor mites, and test for hygienic behavior. This is also the time you want to start removing that contaminated comb, melting down the wax, and using it for anything other than new foundation. Any honey harvested from these frames can be sold commercially, but cannot be sold as treatment free honey, or fed to your own bees. Basically treat any product of that frame as like chemical waste, isolate it, get rid of it; minimize the chances of it re-entering your treatment free operation. Remove the marker from the frame and put it back into circulation.

Next fall, rinse and repeat.

Incorporating this into your treatment-free operation will allow you to remain profitable, sustainable, and still retain your core philosophical beliefs without vicariously endangering it by allowing mites to spread from dead hives.


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## enjambres

How do you "test for hygenic behavior"? 

And what do you do with the queen in the treated hives? Pinch her?

And about the drones (from these hives)-how do you make sure their genetic contribution to the newly installed, or created, queens doesn't simply perpetuate whatever genetically-borne, pro-varroa characteristics that the original queen had? (The drones would be the orginal queen's sons, right?)

Somehow, I am skeptical of the notion of hygenic bee-breeding our way out of the varroa crisis.

And as an aside: I have three hives. All three were 2013 swarms that were then cut out of my barn walls in late June. (I apparently have a swarm-magnet barn!) Two of the three lost their queens in the cut-out or ensuing trouble I had keeping the bees in their new digs after the cut out. The other queen also didn't like her new hive, but survived, though she stubbornly remained living outside the hive for another ten days - no matter how many times I re-captured her and re-installed her bees and their newly-drawn combs back in the hive. (Eventually she tired of the game and let me "win".).

So I had an unusually long brood break on the two almost queen-less hives that only barely had time to convert previously laid eggs into queen cells.

Of the three colonies can you predict which one(s) the developed a varroa problem in the late Fall? 

It wasn't the one that didn't have a brood break. It was the other two. 

So I'm also somewhat skeptical about the powerl of brood breaks as varroa-vanquishers. I think it may just annoy them or reduce their numbers, but I don't think it's as effective a tool as we may (wish to) believe. Varroa are wily beasts.

I treated the two brood-broken hives last Fall with only moderate success (with Apiguard). The one that never had a brood break still has almost no varroa and I haven't treated that one at all. 

I have no idea where my swarms orginated from so their genetics are a mystery: they could be true ferals or simply first or second year elopers from other bee-yards. On one hand I could surmise that the un-treated, non brood-broken, almost no varroa, hive has some superior genetic make-up, but I'm doubtful of that. I think it most likely is some other (probably combination of) factor(s). But darned if I can tell what that might be. (I try to treat all my bee-children the same, like any good parent.) Perhaps it's the color of that hive: purple!

Enj.


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## honeydrunkapiaries

> How do you "test for hygenic behavior"?


Liquid nitrogen testing, here in Ontario they do have a program for that where people will come out and do it for you. It is however much cheaper to do it yourself.



> And what do you do with the queen in the treated hives? Pinch her?


"requeening" is a nice way of saying pinch her, yes.



> And about the drones (from these hives)-how do you make sure their genetic contribution to the newly installed, or created, queens doesn't simply perpetuate whatever genetically-borne, pro-varroa characteristics that the original queen had? (The drones would be the orginal queen's sons, right?)


Nucleus colonies typically wont make alot of drone comb, and youve already requeened her anyway. If you are doing this in the fall chances are you arent doing any breeding, as it is too late.


> Somehow, I am skeptical of the notion of hygenic bee-breeding our way out of the varroa crisis.


I dont think it is THEE answer either, but it is part of the equation. You may not believe it will cure varroa, but hygienic behavior will help prevent chalkbrood, as well as AFB http://www.apidologie.org/articles/apido/abs/2001/06/spivak/spivak.html

I think at the end of the day, between mites, the weather, etc sometimes it really is just a roll of the dice. I am just trying to present a model that I am thinking of adopting. The big thing that got me thinking was the transfer of one hive to another while getting robbed out, and the possibility of infecting (largely through neglect) the rest of your colonies. I think it is probably the colour of hive


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## Fusion_power

Or you could take a different approach and find bees similar to mine that have a bit of Apis Mellifera Mellifera genetics. They overwinter better than Carniolans and some of them have decent levels of mite tolerance.

The major item I see missing in your plan is management of drones and management of a mating area so the number of mite tolerant colonies can be dramatically increased in 2 or 3 years. Also, in my experience, finding a mite tolerant colony usually involves a lot of unwanted genetics that take years to reduce to a tolerable level. I found a queen whose colony was mite tolerant in 2004/2005. The unwanted genetics turned out to be a very high level of hive defensiveness. They could not be worked without a suit. And no, they were not Africanized, they were just typical AMM stock for the area. It took 7 years to get them toned down enough to work in short sleeves.


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## Specialkayme

For the sake of argument, I will say your answers here are likely to be biased. Your plan, which I don't think is foolish in the least, involves treatments (to at least some of your hives). This runs directly counter to the purpose of the treatment free section. Most, if not all, that post in this section have a problem with treatments (hence the name). I've had many discussions here before about using treatments as a method of getting to a treatment free status. Most of the time the conversations don't end well. Just throwing that out there.

But, being as it may, I think your system would need to be run by someone who is very capable of keeping EVERYTHING of the two apiaries (treated and non-treated) separate. I applaud you if that's your approach, as it would involve alot more note taking and work than I'm willing to put into it (honestly). 

But you would need to make sure that your treated and untreated hives don't mix, not only equipment wise, but location wise. If you have a treated hive collapse (for whatever reason) and a treatment free colony robs the dead out, taking the honey (that was stored in comb that was contaminated with treatments) it may skew your results. I don't think much, but I'm not one of the people on here that think so low of treatments. That would likely result in at least two apiaries, probably more. One that is pure treatment free, one that is treated (as needed) at a minimum, and likely at least a third for your "transition" nucs (the ones that were treated, but requeened with a TF queen, as the combs are contaminated with treatments, as are the bees, but you are working to get the treatments out). Just some thoughts.



honeydrunkapiaries said:


> _
> “A key point to remember is that the relative infestation (percent, or mites per 100 bees) is more important than total mite population—“_ (Randy Oliver, IPM 3 Fighting Varroa 3: Strategy – Understanding Varroa Population Dynamics)





honeydrunkapiaries said:


> Nucleus colonies generally do not suffer the effects of mites as severely as large colonies (more bees = more mites).


For the sake of editorial purposes, I understand what you are trying to say, but the two statements appear to contradict. I would agree that Nucleus colonies generally don't suffer AS MUCH of the effects of mites as large colonies. I have had nucs collapse from mites though. It happens, not when the number of mites gets too high, but when the % of infestation gets too high. This lack of collapse doesn't happen because nucs have less bees (and thereby less numbers). It usually happens because nucs are brood making factories, and generally speaking they increase larger than the mites do, keeping the % of infestation within manageable levels. When you transfer the nuc to a full size colony, eventually the brood rearing levels off. But the mites continue to reproduce. This is what causes the % infestation to rise, and why more colonies can collapse from mites than nucs. At least my observations.



honeydrunkapiaries said:


> Secondly, hygienic testing; either begin testing for hygienic behavior or buy bees from breeders who are selecting for these traits and breed from them.


Selecting for hygienic behavior is important, but I wouldn't say it's the magic bullet. If you have a conversation with Marla Spivak ("creator" of the Minnesota Hygienic strain) she'll tell you that VSH and hygienic behavior are not the same thing. They are similar, but if you select for hygienic behavior your are not, from a genetic perspective, selecting for a mite resistant stock. You are instead selecting for a "cleaner" bee. They will clean out noticeable diseases faster than a non-hygienic bee, if they can. 

Mite resistant stock is a complicated area, that involves a number of behaviors. Those that have mite resistant stock don't usually know why, and quite honestly don't care why. But it has been found that there are a number of characteristics that can lead to mite resistance, including VSH qualities, hygienic behavior, "mite chewing" behavior, forager grooming behavior, increased swarming tenancies, increased absconding tenancies, and shorter brood gestation periods are just a few. Most don't want the latter three (usually found in AHB colonies, although not necessarily). Selecting for one of the above traits may improve your selection process. But you may select for hygienic behavior to the determent of breeding against grooming behavior. Just a word of caution, as I don't think hygienic behavior, while it is important, is the golden bullet to the mite problem.



enjambres said:


> I have three hives. All three were 2013 swarms that were then cut out of my barn walls in late June.
> 
> . . .
> 
> So I'm also somewhat skeptical about the powerl of brood breaks as varroa-vanquishers.


Brood breaks don't "vanquish" the mites. It just helps keep their % infestation below acceptable levels. I wish I had a chart showing the increased populations of mites and bees over the course of the summer. I've seen it around, but can't locate it at the moment (last saw it at 2012 EAS). But generally speaking, mite levels increase as bee populations increase, although right behind them in the spring. As more bees are reared, more mites are able to be reared. Eventually, at some point in the summer the colony hits critical mass (so to speak). The population of bees starts to slow down, if not level off. Eventually the bees will raise enough brood to maintain their colony size, if not increase it slightly, rather than increase it exponentially. But the population of mites continues to increase as long as brood is reared. During this slow down, the mite % investation skyrockets, as you have more mites but fewer cells for the mites to hide in. This results in multiple mites to be found in every cell (if left untreated) which can cause a collapse. 

The brood break doesn't work if it is done too soon. It also doesn't work if it is done too late (as the mite levels have already increased too much). It only works when you can hit it on the upswing. As the levels of brood are increasing, and so are the mite levels, you stop brood rearing for a window of time. This causes the mites to become exposed, allowing the bees to (hopefully) harass them. The mites (in theory, at least some of them) will die or leave the colony. Then when the bees start ramping brood production back up, to levels equal to or greater than they were pre-brood break, the mites don't have populations large enough to pick right back up where they were before. But again, if done too early or too late, it doesn't work.

I would suggest giving it more of a chance than to two out of three hives in one summer (and probably not actually timed that way, just happened).



Fusion_power said:


> Or you could take a different approach and find bees similar to mine that have a bit of Apis Mellifera Mellifera genetics.


I've found alot of people that claim to have pure, or close to it, AMM genetics in the states. I've never found anyone that can substantiate this claim other than based on "looks" or conjecture. One person appeared to support his claim of having almost pure AMM genetics, based on years of selection and back crossing. It ended up this person was running a fraudulent enterprise (and was probably a borderline Ponzi scheme), wasn't breeding any of his own stocks, and was buying stock from halfway across the country, relabeling it, and reselling it as something else (if he wasn't pulling queens out of local hives that didn't belong to him of unknown genetic makeup).

So, I'm skeptical to anyone who claims to have AMM stock, or a large makeup of it, unless they have a reason to support it (other than they are "dark" and "mean"). Not lumping you in that category, just explaining my skepticism. 

So, how did you come to the conclusion that your local stock is AMM?


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## Barry

Specialkayme said:


> Your plan, which I don't think is foolish in the least, involves treatments (to at least some of your hives). This runs directly counter to the purpose of the treatment free section.


I've updated the unique forum rules to reflect what I've always considered acceptable treatment discussion here.

"Any post advocating the use of treatments, according to the forum definition of treatment will be considered off topic and shall be moved to another forum or deleted by a moderator, unless it is employed as part of a plan in becoming treatment free."


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## Specialkayme

Helpful Barry. Thanks.

Lets hope the comments are as helpful and supportive.


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## Fusion_power

> So, how did you come to the conclusion that your local stock is AMM?


Observation of traits.
1. Flying and foraging at low temps and very late in the day, specifically @45 degrees on a bright sunny day.
2. Excessive stinging behavior, typical AMM, sting if you get within 20 ft of the colony.
3. Highly aggressive when the colony is manipulated, not like Africanized, just hotter than any other bees around.
4. Overwintered on less than 20 pounds of honey, foraged early, still built up faster than any other colony I had.
5. Very fast to swarm, typical of AMM, not typical of Italian or Carniolan.
6. Very good tolerance to mites which is NOT typical of AMM and tells me they were crossbred.

I am not claiming they were pure AMM, just that they had some in them. I caught them as a swarm near Rainsville Alabama in an area that I have caught AMM for years. I used them to produce queens that were mated to drones from queens I got from Purvis. In other words, I mated a mite tolerant queen to mite tolerant drones, then let them sort out over the next several years.


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## honeydrunkapiaries

> Or you could take a different approach and find bees similar to mine that have a bit of Apis Mellifera Mellifera genetics. They overwinter better than Carniolans and some of them have decent levels of mite tolerance.


As far as survival and overwintering go, I am not as worried about the race of bee. Italians overwinter just fine. My current bee was developed by Tibor Szabo over the past 40+ years from feral stock. My guess is that it is predominately Italian, but he was a scientist and wrote the book on colour and bee genetics... so who knows? I have ordered a few Olivarez queens for early splits this spring. I know a very good breed who says they dont have problems overwintering, good genetics, but he still prefers to develop local stock. If I could find a Apis Mm, I would love to try them out even just for curiosity sake -however, I dont believe it is to be readily found.



> The major item I see missing in your plan is management of drones and management of a mating area so the number of mite tolerant colonies can be dramatically increased in 2 or 3 years


.

Always the other half of the equation! Other then artificial insemination, or owning your own private island you cannot be 100%. The theory is that the hives that are doing well, are TF, are the big hives and will produce more drones. You have split the weak colonies up, by the time the queen emerges from her cell those weak genetics will be far and few. Over many years I think the genetics, and drones would get better. I dont think any part of this is an 'overnight' thing.



> I think your system would need to be run by someone who is very capable of keeping EVERYTHING of the two apiaries (treated and non-treated) separate. I applaud you if that's your approach, as it would involve alot more note taking and work than I'm willing to put into it (honestly).
> 
> But you would need to make sure that your treated and untreated hives don't mix, not only equipment wise, but location wise. If you have a treated hive collapse (for whatever reason) and a treatment free colony robs the dead out, taking the honey (that was stored in comb that was contaminated with treatments) it may skew your results. I don't think much, but I'm not one of the people on here that think so low of treatments. That would likely result in at least two apiaries, probably more. One that is pure treatment free, one that is treated (as needed) at a minimum, and likely at least a third for your "transition" nucs (the ones that were treated, but requeened with a TF queen, as the combs are contaminated with treatments, as are the bees, but you are working to get the treatments out). Just some thoughts.


I think you would definitely have to develop a system of marking, or keeping track for sure. Multiple locations would be helpful, if you were creating your splits in early spring it makes sense to move them to a new location anyway. However, I wouldn't put too much stock into robbing out treated colonies if they steal contaminated honey it will be either in the spring, or late fall and that honey is going to get used up. It will not be sold AND you probably havent applied treatments by then. Sure doesn't sound optimal but nothing about beekeeping is 100%, It would be no different then your TF bees robbing out another apiary.

Woodenware I could care less about contamination, the chemicals are not stored as they are in the wax. 

I do however thank everyone for the positive input and it is helping me develop and hone this idea. Where does this idea come from? Well I have a very small apiary in Ontario, I have however decided that I would rather keep bees then slug it out in work I loathe so I need to think about how to increase my numbers, stay profitable, and stay true to my philosophy. Currently all my hives are treatment-free. Last year was a bad year for bees up here. The fall flow was a dud and the goldenrod did not come through (which is a big flow here), and we had a warm autumn so the bees despite no pollen and nectar remained prolific. I left a ton of honey on the hives thinking that would do them for winter, by October it was all gone. We have also had an exceptionally cold winter. Many beekeepers around here I know are estimating a 70% die off. Because of the pesticides bee breeders are charging an absolute mint for nucleus colonies. I cannot afford to let hives die off, and buy more to make my increases. Michael Palmer's presentations gave me a great deal to think about, and next year I will set myself up to mimic his system on a smaller scale the year after.



> I've updated the unique forum rules to reflect what I've always considered acceptable treatment discussion here.
> 
> "Any post advocating the use of treatments, according to the forum definition of treatment will be considered off topic and shall be moved to another forum or deleted by a moderator, unless it is employed as part of a plan in becoming treatment free."


Thanks Barry!

I am sorry if my answers aren't quite as satisfactory, as I am still thinking them over myself.


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## Juhani Lunden

honeydrunkapiaries said:


> There is a hive in danger of collapse, being robbed out and spreading mites. Treat it with an organic acid, mark the hive body and all the frames so that you know those hives have been exposed to this chemical. Once the mites have been ‘knocked down’ let it overwinter. If it successfully overwinters, split that hive up into nucleus colonies giving each a queen cell from the breeding survivor stock. A queen cell as opposed to a laying queen is important; it allows a short brood break to reduce the mite population, and give the new queen a chance to prove herself. Swarm cells in the spring should be rather easy to find.
> 
> Firstly, you have to monitor for mites. Whether it is a sticky board, an alcohol shake, it doesn’t matter. You need a baseline to know which of your hives have a mite problem and which don’t. Which are candidates for breeder stock, and which hives are going to need help?
> 
> Secondly, hygienic testing; either begin testing for hygienic behavior or buy bees from breeders who are selecting for these traits and breed from them.


Good plan, but because I have done all this, few comments:
1. The collapses come so quickly, that in practice, you don´t have time to react. They collapse in less that two weeks. From a top hive to dead hive. In order to this plan to work, monitoring needs to be done at least once a month.
2. If treatments, or any other measure by the beekeeper, are done differently to so some hives, this reduces the sc. "effective size of the population"(breeding term), this makes breeding selection more uncertain
3. Matings must be controlled. If they are not, breeding is a gamble: one step forward and two steps back. No way you can make right decisions, when selecting breeder queens , if they don´t have uniform matings. To make queens from swarm cells is a mistake.
4. Making selection according to measured hygienic behavior, might slow down the process. All measurements have error and all assays are only the best ASSUMPTIONS, what might be important. In real life, bees don´t need good results in hygienic behavior, they just need to survive. There are no quarantines that breeding according to hygienic behavior results will get us any further, there is no breeder, who has done varroa resistant bees by having hygienic behaviour in a major role.


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## honeydrunkapiaries

Juhani Lunden said:


> Good plan, but because I have done all this, few comments:
> 1. The collapses come so quickly, that in practice, you don´t have time to react. They collapse in less that two weeks. From a top hive to dead hive. In order to this plan to work, monitoring needs to be done at least once a month.
> 2. If treatments, or any other measure by the beekeeper, are done differently to so some hives, this reduces the sc. "effective size of the population"(breeding term), this makes breeding selection more uncertain
> 3. Matings must be controlled. If they are not, breeding is a gamble: one step forward and two steps back. No way you can make right decisions, when selecting breeder queens , if they don´t have uniform matings. To make queens from swarm cells is a mistake.
> 4. Making selection according to measured hygienic behavior, might slow down the process. All measurements have error and all assays are only the best ASSUMPTIONS, what might be important. In real life, bees don´t need good results in hygienic behavior, they just need to survive. There are no quarantines that breeding according to hygienic behavior results will get us any further, there is no breeder, who has done varroa resistant bees by having hygienic behaviour in a major role.


I am currently reading your journal of your breeding program, quite interesting stuff and it is giving me a bit to think about. Thanks for writing all that stuff down!

Being out in the beeyard is not a problem for me. It is a beautiful location, and gives me some respite from the rest of my life -I am usually out there at LEAST twice a week.


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## Fusion_power

> there is no breeder, who has done varroa resistant bees by having hygienic behaviour in a major role.


This statement does not show that it can't be done, just that the two traits are probably not linked. In other words, if heavy selection is made for hygienic behavior, you may exclude most colonies that show high levels of mite tolerance. This emphasizes that VSH is not the same as hygienic trait though it is highly probable that at least 1 gene is common between them. An alternate route would be to simultaneously select for both high levels of varroa tolerance and high levels of hygienic behavior and develop two lines that can be intercrossed in the future to combine the traits.

I would also add that when Marla Spivak tested VSH bees, she found they were highly hygienic, almost off the scale. This indicates there is significant overlap between the two traits.


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## mike bispham

I think its a good plan. Its best points are:

A) Its an intelligent, detailed, well thought-through plan. 

B) Its based on raising resistance. Its a breeding plan.

C) Its well informed, cognizant of the bio-evolutionary realities, rationalised according to the known relations between mites and bees and up to date research about what works and why.

D) Its based on using locally adapted bees.

To the remarks about drone hives I'd add: try to be remote from large treating apiaries, and be aware of the likely benefit of expanding to a good number of apiaries in order to claim more matings. 

I don't see anything other than a plan of this kind succeeding - unless 'succeed' is defined in a way that doesn't including enabling the bees to manage their own mites. 

The one caution I might have has been raised already - breeding narrowly toward a single trait. I think working toward a diversity of mite management traits might be a better aim. By mostly taking out the weakest that's happening automatically, as long as you're not pushing too hard at the other end - the strongest. I'm interested in the mite-management mechanisms but wary of interfering with nature's choices. 

Somebody said the record keeping was too much work. On the contrary, the plan uses marks to make notes right on the hives themselve - excellent lazy data managment after my own heart. But I think detailed records, while a chore, are good. I use numbered hives, a notebook recording origins and current states, and make a to-do list on a clipboard as I go through the hives. The to-do list gets entered to a speadsheet, priority ratings attached, sorted to bring the priorities at each site to the top; printed to take. At each site I know what's to be done (and can take stuff I need), and how important it is. This seems unnecessary in the winter, but is essential in the working months. 

The bigger benefit should be when I get to learn which of my queens is best at making good new queens - by analysing the records. That is said to be essential in breeding.

Mike (UK)


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## honeydrunkapiaries

Thanks everyone for the supportive input! I will probably be writing some more longer pieces as I try and fit this into my management plan for next year. I will be adopting some of Palmers ideas, but realistically be a few years down the road before I can truly try and mimic them. Next year is going to be all about getting to that point -and hopefully making enough honey to pay for it.


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## Juhani Lunden

honeydrunkapiaries said:


> I will be adopting some of Palmers ideas.


Who is he? Has he made varroa resistant bees?


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## honeydrunkapiaries

Michael Palmer, did those presentations on the Sustainable Apiary, adopting his ideas for management.


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## rhaldridge

honeydrunkapiaries said:


> Michael Palmer, did those presentations on the Sustainable Apiary, adopting his ideas for management.


I think that while he is not treatment free, his ideas on what he calls the sustainable apiary are very well-suited to someone whose goal is TF beekeeping. He is almost certainly breeding for better resistance, by breeding from his strongest hives and using his middling hives for resources. Over time, that's bound to result in better bees or his climate in all respects.
s


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## Dominic

I'm not convinced about feral swarms being any more resistant to varroa than any average stock. When swarming, the bees ditch all of their brood, and thus capped varroa, in the old colony. When they finally settle up after a few days, they need to build all of their foundation from scratch. That's a pretty significant brood break. Rinse & repeat and you can easily have a lot of feral swarms prospering despite varroa, even if their genetics against them are so poor that they'd die in a single season if prevented to swarm. Same for productiveness: robbing is an easy way for a colony to increase food stores.

As swarming is easier than managing mites and as robbing is easier than harvesting, I tend to think that while feral swarms might APPEAR to be more resistant stock, natural selection favors the easy solutions that yield the most results, in this case swarming and robbing. Two traits that are good for the bees, not for the beekeeper. On the other hand, VSH and hygienic behavior results in brood loss, and thus slower population growth. Natural selection may even work against these traits.

This does not apply to treatment-free beekeeping, however, but it's my view on using feral swarms to start one's stock. The only advantage I see is that they are adapted to local climate, but you can get that from purchasing from local breeders...


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## Fusion_power

Dominic, if this were true, the parent colonies would periodically die out from mite buildup in the fall. I know of several feral colonies in this area that are at least 10 years old. They have been continuously occupied for that length of time.


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## heaflaw

Juhani Lunden said:


> To make queens from swarm cells is a mistake.
> .


Juhani,

Why do you say that?


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## honeydrunkapiaries

heaflaw said:


> Juhani,
> 
> Why do you say that?


If you follow his website on his breeding plan http://www.saunalahti.fi/lunden/varroakertomus.htm, and it is quite interesting I suggest you do. One of the first things he suggests is "There will be more instrumental insemination in the coming years. This is because the risks are getting bigger: one wrong choice, one drone-line that is not working as expected, would be a big hit back. "

What I think he is eluding to, is once youve sort of hit the nail on the head with what you want for genetics, if you have a wild card drone in there it will inject those genetics into possibly your whole apiary. To that effect I think he is right. His breeding plan is actually quite a well thought out operation, but very intensive.


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## Juhani Lunden

heaflaw said:


> Juhani,
> 
> Why do you say that?


Well at least in my country it says in every book of beekeeping, that if you use swarm cells, you increase swarming behavior in your stock. 

But: I have not done it, so there is nothing I really can base on own experience. Good point. You pointed out to me, that I believe in something I have read, but not done myself, which is stupid, in a way. But very often we bump into "facts" in beekeeping, which in fact are only old habits. For instance: In Scandinavia beekeepers are taught, in every book of beekeeping, that they should clean their hive bottoms in spring. Why? I haven´t done that for decades, and when I stopped doing it I saw no effect.

But as Honeydrunkapiaries pointed, if you are making a plan for breeding, make sure there is as little space as possible for chance. And if we still talk about swarmcells, it usually means, that besides you maybe are increasing swarming tendency, you use them just putting them in some neighboring colony and let them mate freely. There comes chance. Of course you can also mate swarm cells in a controlled manner, but I have not seen such a beekeeper.


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## Juhani Lunden

Dominic said:


> I'm not convinced about feral swarms being any more resistant to varroa than any average stock. When swarming, the bees ditch all of their brood, and thus capped varroa, in the old colony. When they finally settle up after a few days, they need to build all of their foundation from scratch. That's a pretty significant brood break. Rinse & repeat and you can easily have a lot of feral swarms prospering despite varroa, even if their genetics against them are so poor that they'd die in a single season if prevented to swarm. Same for productiveness: robbing is an easy way for a colony to increase food stores.
> 
> As swarming is easier than managing mites and as robbing is easier than harvesting, I tend to think that while feral swarms might APPEAR to be more resistant stock, natural selection favors the easy solutions that yield the most results, in this case swarming and robbing. Two traits that are good for the bees, not for the beekeeper. On the other hand, VSH and hygienic behavior results in brood loss, and thus slower population growth. Natural selection may even work against these traits.
> 
> This does not apply to treatment-free beekeeping, however, but it's my view on using feral swarms to start one's stock. The only advantage I see is that they are adapted to local climate, but you can get that from purchasing from local breeders...


I agree. 

To increase variation I suggest that a breeding plan like this should be started with your own bees. Catching swarms is taking a chance. You know nothing about them. At least they should be monitored for a year ( or preferably longer) to make sure varroa is actually building up slower in the swarm. 

Start with your own bees and mate them with some varroa resistant stock. By doing so, you eliminate chance and get at least 12 years head start.


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## Dominic

Fusion_power said:


> Dominic, if this were true, the parent colonies would periodically die out from mite buildup in the fall. I know of several feral colonies in this area that are at least 10 years old. They have been continuously occupied for that length of time.


Colonies known to have remained for long do have this in for them. If you know a colony has been there for quite a while, then you eliminate the chance that they survive by swarming alone.

However, I still think it's valid question to ask yourself why these bees survive, even in such cases. There are many potential mechanisms of survival, but not all are practical or desirable when one seeks to make a living with the strains that have developed them. "Why is that feral colony surviving?" It's obvious it is doing something that's good for them, but that doesn't mean that it's doing something that would be good for you, or that would work in a standard langstroth hive. Swarming doesn't just bring a brood break to the offcast colony either, even the mother colony needs the new queen to emerge, fly out and mate. Maybe their queens stop laying when food sources are scarce, like my carnies do, resulting in more brood breaks. Is a brood break something you want? Some want them, but I suspect most beekeepers don't. The environment is different too: a tree hunk isn't the same as big square wooden boxes. Maybe there's something about the dead trees the bees live in that is just outright hostile to mites.

In my book, feral swarms aren't bad by definition. They could have good genetics. But one just can't know: surviving in the wild is not the same game as surviving in a managed environment and not only making the colony live but the beekeeper as well. Bees adapted extraordinarily well to live in the wild are not guaranteed to do well in our hives. There's just too much unknown, so I'd personally only resort to using swarms for breeding if I didn't have access to stock that is both winter and pest hardy, which isn't my case.


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## beekuk

Dominic said:


> surviving in the wild is not the same game as surviving in a managed environment and not only making the colony live but the beekeeper as well. Bees adapted extraordinarily well to live in the wild are not guaranteed to do well in our hives. There's just too much unknown, so I'd personally only resort to using swarms for breeding if I didn't have access to stock that is both winter and pest hardy, which isn't my case.


 I agree, they are not being continually opened up and manipulated like most bees kept in hives, so retain the nest warmth and humidity, which is detrimental to mites, and maybe have other creatures within a wild nest which prey on varroa to some extent...and no doubt several other reasons.


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## mike bispham

Dominic said:


> They could have good genetics. But one just can't know: surviving in the wild is not the same game as surviving in a managed environment and not only making the colony live but the beekeeper as well.


The question is, if you want to be treatment free, can you do that without 'survivor' colones? 'Managed' bees tend to be treatment-addicted bees; 'survivors', by definition are not. If you can buy resistant queens and requeen your whole treatment-addicted apiary fine, and if you can steadily raise resistance in your own bees fine - lots seem to find this hard, but that doesn't mean it can't be done.  Bringing in resistant genes from 'survivor' colonies offers a valuable contribution to the effort to go (properly) treatment free for those of us unable to source bred resistant bees.

Mike (UK)


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## Solomon Parker

The "treat and breed from the best" has never produced treatment free bees to my knowledge. The ones I see succeeding are the ones who simply don't treat and let things fall where they may. There is so much fear in this method, so much worrying and angst, no peace, no freedom to let things happen as nature intends them. Also, so much work. 

My experience militates against ideas like "mites from deadouts end up in neighboring hives." If the neighboring hive can deal with it, then end of story. Additionally, if you have a decent winter, mites are going to die long before a hive can get robbed out. And if a hive cannot deal with the infestation, then it ought to die too, and along it goes down the line. However, I have not seen it happen that way. I have not seen chain crashes and I have had plenty of hives die and get robbed out or just get robbed out. These crashes have not happened to me.

Now I'm adding an additional stressor, I don't feed anymore. Didn't feed at all this winter.


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## mike bispham

Thanks for breakdown of culling Solomon,



Solomon Parker said:


> Now I'm adding an additional stressor, I don't feed anymore. Didn't feed at all this winter.


That involves a judgement about how much honey to leave... which amounts to the same thing - in raw calorific terms anyway. Take too much a good hive will die.

Also places restrictions on how late to make nucs. And no robbing screens...?

Seems on the surface too many factors to make a true test.

Mike (UK)


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## Solomon Parker

It does involve a judgment on how much honey to leave, that's good beekeeping. For instance, I do not do a fall harvest. Another beekeeper I know here does. He still makes less honey per hive than I do.

My mating nucs have a screened ventilation hole which kinda serves as a robber screen, a la a design I found from Bushkill Farms.

It's not really a test, it's a strategy, a way of life. It produces stronger bees which is the only way forward I see. No other method does, as far as I can tell, make ultimately strong bees. You've got to put them through the real-world conditions you want them to be able to survive.


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## honeydrunkapiaries

Solomon Parker said:


> Now I'm adding an additional stressor, I don't feed anymore. Didn't feed at all this winter.


I think that is completely dependent on the year. Typically up here we get a good heavy goldenrod flow for autumn and that might give them enough honey to keep through the winter. This year it did not come. I left two medium supers of honey on for autumn/winter but autumn ended up being unusually warm so by the end of October they ate through all of that. Now if I didn't feed I would have 100% losses. Right now, after -40C ice storms and the lot, I still have all my hives alive (topped up dry sugar yesterday). Not feeding is a choice to be made at the end of the year, not a criteria in my opinion.


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## Solomon Parker

Could you clarify what you mean by criteria?


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## honeydrunkapiaries

I dont think feeding should be a criteria for raising TF bees, at least not at the moment. Here anyway, honey is dependent on the climate and getting a fall flow. If they have enough sure, dont feed em. But if you have to feed them....


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## Solomon Parker

I see. Not feeding is probably the next step up, but I see it the same way I see treating. If it's there when they "need" it, then they're apt to "need" it without pressure to not need it. And the same with treating, when I don't do it, I expect the ones that "need" it to die, that's kinda what I'm going for. I'm looking for brutally durable bees. And there is the argument that feeding is treating for starvation. But I'm not interested in pushing that view too hard.


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## D Semple

Good post Dominic (post # 24)


Don


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## Juhani Lunden

Solomon Parker said:


> I see. Not feeding is probably the next step up, but I see it the same way I see treating. If it's there when they "need" it, then they're apt to "need" it without pressure to not need it. And the same with treating, when I don't do it, I expect the ones that "need" it to die, that's kinda what I'm going for. I'm looking for brutally durable bees. And there is the argument that feeding is treating for starvation. But I'm not interested in pushing that view too hard.


I agree, unnecessary feeding will "soften" the bees.
I think it is a good practice for a breeder to feed as little as possible. No little portions, only for winter, large storage amounts. Good durable bees need to get along by themselves. In our climate winter feeding is necessary, because bees are not natural animals here and wintering with honey is very uncertain (6 months winter, no flying, honey has too much minerals in it, it fills the gut ) 

It is a matter of money too, honey sells for 10€/ kg, sugar is about 1€/kg.


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## jonathan

Fusion_power said:


> Observation of traits.
> 1. Flying and foraging at low temps and very late in the day, specifically @45 degrees on a bright sunny day.
> 2. Excessive stinging behavior, typical AMM, sting if you get within 20 ft of the colony.
> 3. Highly aggressive when the colony is manipulated, not like Africanized, just hotter than any other bees around.
> 4. Overwintered on less than 20 pounds of honey, foraged early, still built up faster than any other colony I had.
> 5. Very fast to swarm, typical of AMM, not typical of Italian or Carniolan.
> 6. Very good tolerance to mites which is NOT typical of AMM and tells me they were crossbred.


I keep AMM (in Ireland not the US) 2, 3 and 5 are not characteristic traits of AMM.
They are traits of poor bees.
Any decent breeder will select against those traits.
My stock is not like that at all.

In Ireland we have started a mite tolerance programme based on AMM which involves selecting the colonies which deal better with mites. They are not naturally more resistant to mites than other subspecies per se.


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## Dominic

jonathan said:


> I keep AMM (in Ireland not the US) 2, 3 and 5 are not characteristic traits of AMM.
> They are traits of poor bees.
> Any decent breeder will select against those traits.
> My stock is not like that at all.
> 
> In Ireland we have started a mite tolerance programme based on AMM which involves selecting the colonies which deal better with mites. They are not naturally more resistant to mites than other subspecies per se.


From what I gather, I suspect that these traits, which resemble A.m.l/A.m.s (AHB) hybrids, are the results of A.m.l/A.m.m. hybrids rather than A.m.m. purebreeds. Makes one wonder if AMM has the problem genes, or if AML does.


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## mike bispham

Dominic said:


> From what I gather, I suspect that these traits, which resemble A.m.l/A.m.s (AHB) hybrids, are the results of A.m.l/A.m.m. hybrids rather than A.m.m. purebreeds. Makes one wonder if AMM has the problem genes, or if AML does.


I'd say neither. If the traits are in the local population its because a pressure in recent predecessors has caused an adaptation. If they are in individuals its may be because they come from a population like the above, or that they are a random occurrence, a throw of the dice. 

Trait 5, swarminess, may be an early response to the pressure supplied by mites. 

Any race, and mongrel populations, can be pressed toward different propensities. Nature does it, breeders can do it.

Aggression is (so I've been told) more common in early hybrids - just as you get hybrid vigour so you get hybrid aggression. But you can raise aggression in any population by, for example, introducing a predator that the bees can, if stroppy enough, see off. The genes coding for gentleness will fade in the population.

Mike (UK)


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## Dominic

mike bispham said:


> I'd say neither. If the traits are in the local population its because a pressure in recent predecessors has caused an adaptation. If they are in individuals its may be because they come from a population like the above, or that they are a random occurrence, a throw of the dice.
> 
> Trait 5, swarminess, may be an early response to the pressure supplied by mites.
> 
> Any race, and mongrel populations, can be pressed toward different propensities. Nature does it, breeders can do it.
> 
> Aggression is (so I've been told) more common in early hybrids - just as you get hybrid vigour so you get hybrid aggression. But you can raise aggression in any population by, for example, introducing a predator that the bees can, if stroppy enough, see off. The genes coding for gentleness will fade in the population.
> 
> Mike (UK)


Yet, A.m. carnica / A.m. caucasica hybrids are not known to be aggressive.

Are all hybrids being generalized as being aggressive just because most hybrids are AML mutts?

Sure, after a few generations of being feral, natural selection will favor certain traits over others regardless of race, but I'm talking of F1 hybrids that have yet to be affected by any selection pressure. I don't have the answer, really, I'm just adding food for thought. With the Old World stock imported by the WSU, it could be possible to do experiments with this, to see what results of each crosses, though sadly they didn't seem to think it worthwhile to import old world AMM germplasm.


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## jonathan

Dominic said:


> From what I gather, I suspect that these traits, which resemble A.m.l/A.m.s (AHB) hybrids, are the results of A.m.l/A.m.m. hybrids rather than A.m.m. purebreeds. Makes one wonder if AMM has the problem genes, or if AML does.


I don't think either subspecies has a problem but as you point out the hybrids are often aggressive. Ruttner found that the The Amm x Carnica hybrid was the most aggressive, and these subspecies are both dark, so many people wrongly assume that a dark bee is AMM when it is actually a dark hybrid between AMM and Carnica. Heterosis can produce a vigorous healthy bee but when the vigour is expressed as aggression this is not the bee which most beekeepers want to work with. Heterosis is one reason why simply breeding from the most vigourous colonies can be misleading as this is an F1 effect which is often lost in subsequent generations.


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## honeydrunkapiaries

I would really like to get my hands on some A.m.m. as far as I know they are extinct here. In _Beekeeping at Buckfast Abbey_ Brother Adam points out some of the potential problems with certain crosses.


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## jonathan

A.m.m. has been largely displaced from its natural range in Europe by the popular commercial bee types such as carnica, ligustica and Buckfast.
We still have some decent A.m.m. stock in Ireland and you can find groups of A.m.m breeders in Scotland and Wales as well.
There are a few breeding groups in Europe and Poland still has a decent amount of A.m.m. There is also a pure population in Tasmania which is descended from stock brought from England in the 18th Century.
I think A.m.m has great potential as it has rarely been part of an intensive selection process like the more commercial strains, and as such there is great room for improvement.
Where serious selection has taken place, the bees have proved to make gentle and productive colonies and are not at all like the bees described in the list above.
As I mentioned earlier, we are working in a coordinated way to see if we can develop varroa tolerant A.m.m stock in Ireland. Still early days.


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## Juhani Lunden

jonathan said:


> A.m.m. has been largely displaced from its natural range in Europe by the popular commercial bee types such as carnica, ligustica and Buckfast.
> We still have some decent A.m.m. stock in Ireland and you can find groups of A.m.m breeders in Scotland and Wales as well.
> There are a few breeding groups in Europe and Poland still has a decent amount of A.m.m. There is also a pure population in Tasmania which is descended from stock brought from England in the 18th Century.
> I think A.m.m has great potential as it has rarely been part of an intensive selection process like the more commercial strains, and as such there is great room for improvement.
> Where serious selection has taken place, the bees have proved to make gentle and productive colonies and are not at all like the bees described in the list above.
> As I mentioned earlier, we are working in a coordinated way to see if we can develop varroa tolerant A.m.m stock in Ireland. Still early days.


There is a club for A.m.m. beekeepers in Sweden. They have very nice bees. Buckfast breeder Ulf Gröhn once said, that he has, in his long career, not seen such enormous honey gathering ability which he experienced with the crosses of buckfast and the Läsö island A.m.m. It is probably something what Brother Adam experienced with Sahariensis crosses: Bees which have been kept in isolation can create unbelievable hybrid vigor when the inbreeding is released.


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## rhaldridge

honeydrunkapiaries said:


> I would really like to get my hands on some A.m.m. as far as I know they are extinct here. In _Beekeeping at Buckfast Abbey_ Brother Adam points out some of the potential problems with certain crosses.


You may find this interesting. Delaney found a number of feral amm colonies in her research.

http://www.youtube.com/watch?v=bDQNoQfW-9w


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## WLC

That was from her thesis research and represented feral colonies collected between 1980 and 1992.

28% of the ferals colonies were of an Amm haplotype/mitotype.

The post Varroa invasion %age is likely very different, but we don't have a clear picture just yet.

Delaney's grad student found Amm mitotypes in 3 states only in sampled umanaged colonies.

NC had 3%, GA had 8.9%, and DE had 6.1%.

So, you can find Amm mitotypes if you know where to look. But, we don't know much else about them.


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## JWChesnut

Delaney found 4 "M-lineage" mitotypes in southern breeders (Delaney 2009).
M3 0.00 
M4' 0.013
M7 0.019
M7' 0.026

M3, M7, M7' are exclusively Iberian mitotypes (Irati, 2007). They are not queen mitochondria associated with northern Europe.

The southern M-lineage may represent 1) M-lineage co-invading with A-lineage AHB (as hypothesized by Pinto), or relict Spanish colonial or pre-1920 Spanish imports. The vigorous importation in the early 20th century of many, many different lineages seems the most likely source of Egyptian, Syrian, Cyprus, Basque (M7') bees. It does not represent "German Black Bees" or any northern European colonial relict as the mito-type is wrong.

(The usual caution that mitotype does not represent nuclear inheritance or expression, only clonal descent of the queen).

Citation: 
Apidologie 38 (2007) 141–155 141
DOI: 10.1051/apido:2007007
Gene flow within the M evolutionary lineage of Apis
mellifera: role of the Pyrenees, isolation by distance
and post-glacial re-colonization routes in the western
Europe*
Irati Miguel, Mikel Iriondo, Lionel Garnery, Walter S. Sheppard,
Andone Estonba http://hal.archives-ouvertes.fr/docs/00/89/22/56/PDF/hal-00892256.pdf
First, 14 mitotypes detected in northern Iberia were
absent in French and Belgian populations (M3, M5, M7, M7’, M8’, M11’, M19, M20’, M27,
M27’, M29, M32, M36, M37).

In the same direction, M7’ was present only in the western and central
part of the Pyrenees, where M7 occurred in high frequency.​
Delaney cites this geographic origin in her thesis/paper -
Several of these M haplotypes,
M3, M7, and M7, are known from western Europe,
speciÞcally the Iberian Peninsula (Franck et al. 1998,
2001).​
Genetic Characterization of Commercial Honey Bee
(Hymenoptera: Apidae) Populations in the United States by Using
Mitochondrial and Microsatellite Markers
D. A. DELANEY, M. D. MEIXNER, N. M. SCHIFF, AND W. S. SHEPPARD
Ann. Entomol. Soc. Am. 102(4): 666Ð673 (2009)
http://www.researchgate.net/publica...oc_dl&origin=publication_detail&inViewer=true

Illysov (2011) notes that Italian and Sicilian bees are admixed with an unspecified M mitotype. Consequently the southern M-type may represent resortment from the orignal "Italian" importation.

PHYLOGENETIC RELATIONSHIPS OF DARK EUROPEAN
HONEYBEES APIS MELLIFERA MELLIFERA L.
FROM THE RUSSIAN URAL
AND WEST EUROPEAN POPULATIONS
Rustem A. Ilyasov1,
Vol. 55 No. 1 2011 Journal of Apicultural Science P. 67
http://amellifera.narod.ru/publicat..._relationships_of_dark_european_honeybees.pdf


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## mike bispham

jonathan said:


> A.m.m. has been largely displaced from its natural range in Europe by the popular commercial bee types such as carnica, ligustica and Buckfast.
> 
> We still have some decent A.m.m. stock in Ireland and you can find groups of A.m.m breeders in Scotland and Wales as well.


While higher concentrations of native blood may well exist at the land margins, I think there is deserved skepticism about much of the UK dark bee breeding effort. Selecting for particular wing vein patterns and dark colour will result in dark bees with particular wing vein patterns - but I'm not sure there is any good reason to think that anything else about them is authentic. I'm not saying that's always the case, but I would like to hear the rationale for believing that selecting for two characters supplies a pressure that also works on the rest. 

I'd always go with what is working in your area - the locally adapted ferals, or the nearest ones you can find as a first choice. 



jonathan said:


> There are a few breeding groups in Europe and Poland still has a decent amount of A.m.m.


Do you refer to wild Amm here Jonathan? If so are you talking about 'survivor' bees? Or only 'dark' bees, some of which may be largely native, and others that may be camouflaged mongrels?



jonathan said:


> There is also a pure population in Tasmania which is descended from stock brought from England in the 18th Century.


Is it adapted to varroa?



jonathan said:


> I think A.m.m has great potential as it has rarely been part of an intensive selection process like the more commercial strains, and as such there is great room for improvement.


In general terms _anything_ would be better than commercial strains. They're bred to perform with crutches - the last thing a tf beekeeper is looking for. I'd have thought that in general terms well mixed blood, subjected to local pressures for a period, has a better chance of thriving than any foreign pure strain. There's so much more diversity there for natural selection to work with. Someone will probably tell me I'm wrong about that.



jonathan said:


> As I mentioned earlier, we are working in a coordinated way to see if we can develop varroa tolerant A.m.m stock in Ireland. Still early days.


Am I right in thinking varroa came later to Ireland than to the UK Jonathan?

Mike (UK)


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## Oldtimer

Thanks for that JWC it's a subject I'm pretty interested in but I'm not a geneticist, your summary is very helpful for lay persons like me.

It's also one of the things I really enjoy about Beesource, there are experts available in just about any field.


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## jonathan

mike bispham said:


> While higher concentrations of native blood may well exist at the land margins, I think there is deserved skepticism about much of the UK dark bee breeding effort. Selecting for particular wing vein patterns and dark colour will result in dark bees with particular wing vein patterns


Wing patterns and body colouration is yesterday's news. AMM can be identified by microsatellite markers and SNP work.
The more I have read about wing venation, the less confidence I have in it. I think wing pattern and colour can only tell you if a bee is *not *AMM. ie a yellow bee could not possibly be pure AMM but a pure black bee could be Amm or a mixture of whatever. A wing with a high cubital index value is more likely to be Carnica than AMM but a low CI value does not 'prove' a bee is AMM.

Ireland is very different from Kent. There are parts in the west that have had little or no introgression from non native subspecies. Wish it was like that in my area but unfortunately is is mongrelised.



> I'd have thought that in general terms well mixed blood, subjected to local pressures for a period, has a better chance of thriving than any foreign pure strain. There's so much more diversity there for natural selection to work with. Someone will probably tell me I'm wrong about that.


There is plenty of natural variation within any pure subspecies according to people like Marla Spivak. 
The Jensen and Pederson paper on AMM populations in Europe pointed out that the variation within a given AMM population is much greater than that between populations. Lack of variation is not the problem. identifying the various genes which confer resistance is the problem.



> Am I right in thinking varroa came later to Ireland than to the UK Jonathan?


First noted in 1998 on the Sligo Leitrim border, 2002 in the north. I think England was 1992.
Came in to Ireland with a guy who brought over a few colonies illegally from England.


----------



## WLC

"Wing patterns and body colouration is yesterday's news. AMM can be identified by microsatellite markers and SNP work."

I think that Geometric Wing Venation analysis is far more accessible to beekeepers than a lab that can do satellite/SNP, etc., work.

I was impressed by Dr. De Jong's work in Katherine Darger's Thesis paper. The clusters looked good to me.


----------



## jonathan

Much more accessible but maybe not much use.
This paper by Robin Moritz, The limitations of biometric control on pure race breeding in Apis
mellifera, does a fairly good job of pointing out the fallacies of using wing venation for distinguishing between subspecies.


----------



## WLC

Have you had a chance to read the Darger Thesis paper? She's Delaney's student.

I think that you'll find her thesis on geometric wing venation, and molecular methods, to be an eye opener.

http://udspace.udel.edu/bitstream/handle/19716/12667/Katherine_Darger_thesis.pdf?sequence=1

I hope that you change your mind about geometric wing venation.


----------



## Fusion_power

when DNA sampling becomes cheaper and more accessible, wing venation will be a thing of the past. It is a matter of using DNA direct versus the phenotype of the wing structure. There is no comparison.


----------



## jonathan

WLC said:


> I hope that you change your mind about geometric wing venation.


I think you may have missed the point made in the Moritz paper.

Germany has been using Carnica as its commercial bee for many decades.
The wing samples of the German bees show perfect Carnica wing venation patterns.
This is not surprising as bees without the perfect pattern are considered to be hybrids and are eliminated from the breeding population.

But does this mean they are pure carnica?

In fact Moritz found that when he looked at other morphometric variables such as tongue length and suchlike, the population was actually intermediate between A.m.carnica and A.m.mellifera. Germany was formerly part of the natural range of AMM.
The bees had perfect wings but this appears to be a selection artifact built up over many generations through active beekeeper selection based mainly on wing pattern. The other morphometric variables which were largely ignored did not show carnica characteristics. Ruttner has listed more than 30 morphometric variables which can be measured but selection was based on the wing pattern alone.

The paper you cite above looks at introgression of scutellata genetics into mellifera and is not really relevant to the main point made in the Moritz paper.
However, if US beekeepers start selecting on wing venation alone it will not be long before a similar effect occurs, ie bees with perfect mellifera wing pattern which unfortunately has no real discrimination value for showing introgression of scutellata genetics.


----------



## JWChesnut

WLC said:


> Have you had a chance to read the Darger Thesis paper? She's Delaney's student.
> I think that you'll find her thesis on geometric wing venation, and molecular methods, to be an eye opener.
> http://udspace.udel.edu/bitstream/handle/19716/12667/Katherine_Darger_thesis.pdf?sequence=1
> I hope that you change your mind about geometric wing venation.


WLC, It is impossible to tell what point you are trying to make. 

The Drager paper illustrates the exact opposite of the Idée fixe you seem preoccupied with-- the hypothetical persistence of German Black Bee's in Appalachia.

The wing venation data actually demonstrates that eastern North American bees are fully homogenized and inseparable from managed and unmanaged population using the venation metric.



This chart are the first two principal axis of a PCA reduction of the morphometric data. The Brazilian researcher who prepared the data compared the Drager collected sample with "control" data from European collections from the point of origin of the various races. They made three distinct clouds --- ALL the Drager data made one cloud -- Florida, North Carolina feral, and managed Southern colonies. The other two clouds are Caucasian and Mellifera (overlapping and indistinguishable) and the same indistinguishable cloud group of Italian and Yugoslavian bees.

Jonathan has repeatedly made the key point-- if you select a breeding population based on a, say, Cubital index -- all you are doing is creating, de novo, a lineage of bees with that cubital index --- no more, no less. Genes are not linked -- the alleles are fully independent -- and breeding for an expression just selects for the expression not some race sui generis.


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## jonathan

I have wing morphometry data from about 100 colonies of mine I have sampled over the past few years, usually 40-50 wings per colony.
I was very enthusiastic about this as a diagnostic tool at the start, but in all honesty I think I may be wasting my time as it takes about 45 minutes per colony to scan the wings and get the data via the drawwing programme and I am not even sure what the data is telling me.


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## Oldtimer

WLC said:


> I hope that you change your mind about geometric wing venation.


Based on the evidence and papers done on the subject, it would appear not.


----------



## WLC

The Darger thesis was able to show that Honeybees identified by other wing venation methods as AHB were false positives by using both geometric wing venation and molecular methods.

She was also able to provide evidence that those AHB false positives did in fact cluster separately from known AHB stock using geometric wing venation.

So, we know that the De Jong methodology is useful in that regard.

There's a long standing debate with regards to the use of morphometrics vs DNA based methods for species identification.


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## Oldtimer

Isn't the debate over? Would it be logical to conclude these studies show it sometimes works, sometimes doesn't?

Like everyone's been saying?


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## WLC

Well, the Moritz paper is a 1992 study, whereas the Darger Thesis is very recent.

"Modern morphometric methods for
identifying Apis mellifera subspecies include four techniques: Fast Africanized Bee
Identification System (FABIS), Universal System for Detecting Africanization
Identification (USDA-ID), Automatic Bee Identification System (ABIS), and
Geometric Wing Venation."

In terms of morphometric tests, Darger has found support for geometric wing venation as a useful method for identifying Am subspecies.

So, I wouldn't call it a debate. It's a technical piece of research supporting GWV (De Jong) over other methods.

We don't know as yet how well it stands up when examining the Amm issue.

But, GWV did well when it came to AHB identification. In fact, it upends current AHB morphometric methods used in the U.S. and creates a controversy with regards to AHB claims being made in the South. 

It looks like they're dealing with something other than AHB.

It's the false positive problem.


----------



## Oldtimer

WLC said:


> We don't know as yet how well it stands up when examining the Amm issue.


I think Jonathan and JWC covered that.


----------



## Fusion_power

Do you remember the old tale about 3 blind men feeling of an elephant and then describing it? One felt the trunk and said it was long and flexible like a snake. One felt of a leg and said it was tall and sturdy like a tree. Another felt the belly and said it was like a wall that moves.

Wing venation by necessity cannot reflect more than a small percentage of the genome of a bee therefore will always fail because it reflects so little of the genome. This is why full genome dna analysis will displace all previous ID methods.


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## mike bispham

Fusion_power said:


> Do you remember the old tale about 3 blind men feeling of an elephant and then describing it? One felt the trunk and said it was long and flexible like a snake. One felt of a leg and said it was tall and sturdy like a tree. Another felt the belly and said it was like a wall that moves.
> 
> Wing venation by necessity cannot reflect more than a small percentage of the genome of a bee therefore will always fail because it reflects so little of the genome. This is why full genome dna analysis will displace all previous ID methods.


What I don't understand is: why does anyone care?

What matters to tf beekeepers is which bees do well, on their own *regardless of ancestry.*

What matters is: *are they amenable to tf regimes*. That is dependent on the presence of genes (gene clusters) coding for effective mite management. 

Those genes can come from *anywhere* - any race, any local population. There is no 'Pure Amm, Russian, mongrels etc, are more resistant'; there's only '_*these*_ Amm, Russian, mongrels etc. are able to manage varroa.'

The whole business is a fat red herring as far as tf is concerned. Encouraged, as far as I can see, by people who want to flog microscopes, AI equipment, courses on how to peer into microscopes.

Mike (UK)


----------



## WLC

Why would someone who used the BeeWeaver option in obtaining TF stock want to resort to Geometric Wing Venation morphometrics?

Or, Why would someone resorting to ferals, etc.,... ?

Piece of mind.

It's not a red herring.

Nor does it require anything too fancy. It needs a computer with a good scanner/printer, and the right App's.


----------



## Oldtimer

But it's not accurate.


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## WLC

The morphometrics used by the ARS to determine AHB isn't accurate according to Darger. GWV didn't return a false positive in the study. Plus, the GWV results matched the microsatellite data rather well.

Here in the U.S., ferals are commonly used for resistant/TF stock.

So, I think that many U.S. beekeepers would agree, if all they need to do is use their scanner, and learn to work an App or two, it would make it not only accessible, but possibly very useful. Especially if a GWV library could be built.

There's a lot of talent here on Beesource alone.


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## Oldtimer

Problem is, it is not reliable.

To put it clearer, you do the work, and at the end of the day, do not know if you can believe your results.

So, what's the point?


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## Barry

The fun is in the scanning and drawing and nice pictures of bee wings! We'll figure out a use for them another time!


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## Oldtimer

That probably sums it. 

With the large degree of hybridisation of multiple breeds you have in the US, that in itself would make id using wing pattern a virtual impossibility in itself, there will be so many different influences on the wing.

Over here it's more simple. Effectively we have 2 breeds, Italian & carniolan, AMM pretty much extinct now.

So no messing with looking at wing vein patterns. We look at the bee is it black, or yellow, and it's that simple LOL.


----------



## WLC

Not really.

"Morphometrically, the data shows that there is an American bee, an amalgam
that is distinctive from the originating, European subspecies (Figure 2). The GWV
morphometric study further distinguishes between Africanized bees from Arizona and
those from Africa and Brazil. Interestingly, the samples from
Florida/Alabama/Georgia which were originally diagnosed as Africanized by the
USDA-ID technique were found to have European ancestry using mtDNA markers,
European morphology based on GWV and grouped with managed and unmanaged
honey bee colonies in a population structure analysis using microsatellite markers."

Darger thinks otherwise.

She's Delaney's grad student, and I'm not going to argue with Delaney on this one.


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## Oldtimer

WLC said:


> Darger thinks otherwise.


Unsurprising.



WLC said:


> I'm not going to argue


What have you done with WLC, and who are you?


----------



## WLC

REVIEW ARTICLE
Standard methods for characterising subspecies and ecotypes of Apis mellifera
Marina D Meixner1*, Maria Alice Pinto2, Maria Bouga3, Per Kryger4, Evgeniya Ivanova5 and Stefan Fuchs6

https://bibliotecadigital.ipb.pt/bitstream/10198/8946/1/meixner_etal_2013.pdf

Yes, it can be challenging.


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## Barry

Yep, as Oldtimer said, "Problem is, it is not reliable."

He's a student of commonsense, and I'm not going to argue with commonsense on this one.


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## WLC

I think it's better to use the phrase, "Current state of the Art."

Here's an abstract from a paper in press that suggests that GWV can be both automated and done online:

http://www.sciencedirect.com/science/article/pii/S1574954113001222

"Several studies have shown that features extracted from patterns of bee wings are good discriminatory elements to differentiate among species, and some have devoted efforts to automate this process. However, the automated identification of bee species is a particularly hard problem, because (i) individuals of a given species may vary hugely in morphology, and (ii) closely related species may be extremely similar to one another. This paper proposes a reference process for bee classification based on wing images to provide a complete understanding of the problem from the experts’ point of view, and a foundation to software systems development and integration using Internet services. "

I can always fire up the thermal cycler, but I'd much rather just put a wing on a scanner, and get the 'ballpark' estimate.

One is cheap and easy, the other is time consuming and pricey.


----------



## jonathan

Barry said:


> The fun is in the scanning and drawing and nice pictures of bee wings! We'll figure out a use for them another time!


Yep. I have hundreds. If you ever need a scan of a bee wing, I'm the man.


----------



## jonathan

WLC said:


> REVIEW ARTICLE
> Standard methods for characterising subspecies and ecotypes of Apis mellifera
> Marina D Meixner1*, Maria Alice Pinto2, Maria Bouga3, Per Kryger4, Evgeniya Ivanova5 and Stefan Fuchs6
> 
> https://bibliotecadigital.ipb.pt/bitstream/10198/8946/1/meixner_etal_2013.pdf
> 
> Yes, it can be challenging.


That Meixner et al paper relegates wing morphometry to a mere footnote in comparison to DNA techniques.
It is a nice overview though.

It is worth pointing out that several morphometric variables used in conjunction provide a powerful diagnostic tool whereas wings on their own are unreliable.

This paper by Strange et al covers it well.

Most of these references including the Meixner et al are together on the Irish Native bee website for anyone interested in chasing them up.


----------



## WLC

"A high degree of consistency between wing morphometry and molecular information has been demonstrated by Miguel et al. (2010). Therefore, wing geometry is particularly suitable to track phylogenetic relationships between subspecies, where the full "classical" character set can be misleading."

I wouldn't use the term 'footnote' to describe the treatment of morphometrics in that paper.


----------



## Oldtimer

WLC said:


> "A high degree of consistency between wing morphometry and molecular information has been demonstrated by Miguel et al.


Which means it works, sometimes.


----------



## mike bispham

WLC said:


> Why would someone who used the BeeWeaver option in obtaining TF stock want to resort to Geometric Wing Venation morphometrics?
> 
> Or, Why would someone resorting to ferals, etc.,... ?
> 
> Piece of mind.


I don't follow WLC. How, in what way, does this supply peace of mind?

Mike (UK)


----------



## mike bispham

WLC said:


> "Morphometrically, the data shows ...."


Don't the morphometrics involved in this case involve many more (20 odd?) features - not just wing patterns?

Mike (UK)


----------



## mike bispham

WLC said:


> REVIEW ARTICLE
> Standard methods for characterising subspecies and ecotypes of Apis mellifera
> Marina D Meixner1*, Maria Alice Pinto2, Maria Bouga3, Per Kryger4, Evgeniya Ivanova5 and Stefan Fuchs6
> 
> https://bibliotecadigital.ipb.pt/bitstream/10198/8946/1/meixner_etal_2013.pdf
> 
> Yes, it can be challenging.


There you go. Page 6:

*3. Available methods and markers
3.1. Morphometry
There is no morphological "key" to honey bee subspecies, no simple logical tree based on a sequence of single discriminating characters. Instead, measurable morphometric characters show gradual changes and their ranges mostly overlap between subspecies. Thus, subspecies often differ only slightly in the mean values of several body characters, and therefore advanced statistical methods are required for discrimination of groups. The concept of numerical taxonomy was introduced into honey bee taxonomy by DuPraw (1964, 1965) and further elaborated by Ruttner et al. (1978).
* [1]


The table on page 7 shows 31 different markers, and with those "advanced statistical methods are required for discrimination of groups"

I would imagine that the more markers you have, the less challenging is the math required to reach a (more probable) result. Given, however, that beekeepers have been selecting for particular wing pattrerns for some time, I would guess that one is close to useless!

From the same report, the bit we should really take notice of: 

*"Honey bees show considerable geographical variation, resulting in adaptation to regionally varying factors of climate and vegetation, but also to prevailing pests and pathogens. However, this natural heritage is increasingly subject to diffusion by human beekeeping efforts at a worrisome speed. The demand for high economic performance of bee colonies, combined with desirable behavioural characteristics, has led to considerable changes caused by systematic bee breeding. Thus, the original geographic distribution pattern is being dissolved EU-wide by mass importations and an increasing practice of queen trade and colony movements. These activities endanger regional races and ecotypes by promoting hybridisation (De la Rúa et al., 2009; Meixner et al., 2010), and by adding various breeder lines with distinct properties to the picture. Yet another dimension is added by the deliberate replacement of native subspecies in some regions by non-native bees with more desirable characters and greater commercial interest (for instance, the replacement of A. m. mellifera in northern and central Europe by A. m. carnica or A. m. ligustica) (Bouga et al., 2011).

The downside of these economically driven processes is an increasing trend towards uniformity of honey bee populations across Europe, leading to a loss of both genetic diversity and specific adaptations to local conditions (reviewed in De la Rua et al., 2009; Meixner et al., 2010).

Honey bees are particularly sensitive to inbreeding (Seeley and Tarpy, 2007 and references therein). Therefore, the loss of genetic diversity is of grave concern. It has been shown that colonies with reduced genetic diversity are less capable of controlling hive temperature (Jones et al., 2004) and more prone to develop diseases when challenged by parasites (e.g. Tarpy, 2003). This reduction in genetic diversity may also affect the capacity of honey bee populations to adapt to new threats, such as newly introduced parasites like varroa.

Thus, there is a widely recognised need to encourage regional breeding efforts to preserve local adaptation, and to maintain local strains in isolated conservation apiaries."*

Back to where we started: what matters is that we help the local variants - of whatever ancestry - to adapt to varroa, and thus thrive and preserve their diversity. To say that another way, we stop treating and/or manipulating against varroa. 

Mike (UK) 
[1] 
Standard methods for characterising subspecies and ecotypes of Apis mellifera
Marina D Meixner1*, Maria Alice Pinto2, Maria Bouga3, Per Kryger4, Evgeniya Ivanova5 and Stefan Fuchs6


----------



## mike bispham

WLC said:


> Here's an abstract from a paper in press that suggests that GWV can be both automated and done online:
> 
> http://www.sciencedirect.com/science/article/pii/S1574954113001222
> 
> "Several studies have shown that features extracted from patterns of bee wings are good discriminatory elements to differentiate among species, and some have devoted efforts to automate this process. However, the automated identification of bee species is a particularly hard problem, because (i) individuals of a given species may vary hugely in morphology, and (ii) closely related species may be extremely similar to one another. This paper proposes a reference process for bee classification based on wing images to provide a complete understanding of the problem from the experts’ point of view, and a foundation to software systems development and integration using Internet services. "


I don't think any of that says 'wing analysis alone tells you something about an individual's subspecies identification' - and if it did then it would be in direct conflict with Meixner et all (as just cited)

It seems to me to be talking about automating a process - that may then be of use to to people who know what they're doing, and know not to read (far) too much into a single morphometric.

Has anyone actually read the paper?

Mike (UK)


----------



## mike bispham

Interesting paper, thanks. 

It seems to me that a study of a particular population has to be done, to identify those characters that can be used as 'markers' for a subspecies or local ecotype. (This study concerned a local ecotype - late brood bees adapted - by beekeepers to some extent, and perhaps a lot) to a strong heather source.)

"Discussion
While no single diagnostic morphological or molecular
character was found to distinguish the Landes ecotype from
the broader A. m. mellifera population, the combined
morphological and molecular analyses provided a powerful
suite of characters for identification of the Landes population.
Morphological analysis seemed to be more informative
than molecular data for the characterization of the
Landes population (both ecotypic and non-ecotypic colonies)
from other populations of A. m. mellifera yet appears
to have little utility in differentiating ecotypic and nonecotypic
Landes colonies. Molecular data were quite useful
to distinguish A. m. mellifera from imported subspecies in
the study area but again provided little information to
distinguish among ecotypic and non-ecotypic Landes colonies."


"The similarity of morphometric results from Landes
colonies in the present study and the results obtained
by Cornuet et al. (1982) provide further evidence of the
continued presence of the Landes ecotype (Strange et al.
2007). *By focusing morphological analysis on the characters
identified as informative by discriminant analysis*
(wing venation and external abdominal characters) the
process may be greatly simplified. The statistically significant
differences in two morphological characters between
the ecotypic and non-ecotypic colonies (width of tomentum
light stripe and wing vein angle g18) must be viewed
cautiously with regard to their utility for selecting ecotypic
colonies. Moritz (1991) demonstrated that selection based
upon a few well-differentiated morphological characters
did not arrest significant hybridization between two sympatric
subspecies when all morphological characters were
analyzed. That is, the selection program produced hybrids
that went undetected when assays were based on only a few
morphological characters typical of the desired parental
population. Thus, it is critical to use a broad array of
informative characters in the selection of potential breeding
stock."

A can't see any indication here that selection based on a few markers would be likely to achieve anything other than to raise those markers in the population in any population unless that had been shown to be so by this sort of local close study.

I remain of the view that its a game that sells microscopes.

Mike (UK)


----------



## mike bispham

WLC said:


> "A high degree of consistency between wing morphometry and molecular information has been demonstrated by Miguel et al. (2010). Therefore, wing geometry is particularly suitable to track phylogenetic relationships between subspecies, where the full "classical" character set can be misleading."
> 
> I wouldn't use the term 'footnote' to describe the treatment of morphometrics in that paper.


OK, on the surface... However, its not clear exactly what is being said in that extract. The first sentence is not at issue - of course there is. What that 'therefore' is doing there however I don't know - what is written simply doesn't follow, and conflicts with what we've just seen. Can we read the paper?

Mike (UK)


----------



## mike bispham

Oldtimer said:


> Which means it works, sometimes.


Works at what? *All* physiological features correlate with dna coding (microsatellite data) - whether or not in any particular case that has been shown (i.e. the molecular features that correspond have been located.) That is where the 'design' originates - the dna is the 'blueprint', the instruction for making that pattern.

What has been shown is that the position of the particlar dna molecules responsible (for wing vein pattern) in this case have been found. *What that means* is: 'we' don't have to photograph and analyse the wings themselves, we can just refer to dna analysis to find out if that feature is present.

If it is... that doesn't tell you anything about the other 99.99% of the bee - _*unless*_.. other studies of *that particular local population* have indicated that there is a *high level of corrolation* between that marker and another feature, or group of features (like those typical of a particular subspecies) _*within that local ecotype*_ (and/or more generally).

For general application you need more markers, and then you need to be able to calculate the likelihood that all those markers would appear at once if the individual was not in fact in possession of a high proportion of the dna of a particular racial type - i.e. 'pure'. That's a complex undertaking. (see the first extract supplied in my post #81)

What the math ('stats') will do is give you a really quite complicated idea of how likely it is that the individual is mostly (and how much 'mostly') of a particular subspecies.

The question is, what does knowing a wing pattern/dna feature/s tell you about ancestry. This alone: either mum, or dad, or both, had that feature. (Someone will hopefully tell me why its more complicated/simple than that, and I'll be grateful for the detail)

Does that feature entail the presence of any other features - that might collectively demonstrate a level of racial purity. Nope. 

The best reason I can come up with for the conflict between what this paper says and what the others we've seen say is that the authors of this paper are software developers who lack proper advice on the detail of what-means-what here.

Mike (UK)


----------



## Oldtimer

It's rare indeed Mike, but I agree with your post. And pretty much because as previously stated the alleles that influence wing vein pattern do not have to be linked to race, as time goes along the method will become less and less applicable.

The conflict between which papers?


----------



## WLC

Fellas, I bought resistant stock from a breeder who open mates Italian/Buckfast queens in Texas where the 'hybrid swarm' is found.

So, my own particular need would be to see, as quickly and cheaply as possible, where a sample of workers from those open mated queens 'cluster' in a plot so that I could compare them to other Honeybee types.

GWV would also give me a chance to see if I've lost the original queen and hybrid workers.

Both GWV and molecular methods require statistical analysis. However, I think that using DNA markers has it's own set of caveats above and beyond what can be found using GWV.

In my opinion, it's six of one, and half a dozen of the other.


----------



## Oldtimer

WLC said:


> So, my own particular need would be to see, as quickly and cheaply as possible, where a sample of workers from those open mated queens 'cluster' in a plot so that I could compare them to other Honeybee types.


Why?


----------



## WLC

I think that using GWV cluster analysis to monitor degrees of hybridization, and how that changes over time in relation to resistance, is useful information for a TF beekeeper.

I paid for open mated hybrids. I want to be able to monitor that along with its relationship to resistance.

Think of it as a way to see how your queens are doing.


----------



## Oldtimer

I just find it surprising that of late you have been presenting yourself as if you are a genetics wiz kid, quite able to examine the DNA of your bees, but instead you opt for a debatable method based on wing design which may or may not be related to the breed.


----------



## jonathan

mike bispham said:


> I remain of the view that its a game that sells microscopes.
> Mike (UK)


No need for the cynicism about the plot to gain control of world microscope sales but it is definitely wise to be cautious.
Does Monsanto do microscopes!
Wing morphometry has some use, especially in populations where beekeepers have never used it as a selection criteria.
It is useful for telling you what something is *not* but not so clear at proving an individual bee is of a specific subspecies.
Picking up on some of your other points, I would argue strongly that it is important to preserve pure race subspecies where they are still free from hybridisation. When they are hybridised they cannot be unhybridised at a later date so the subspecies is lost for ever. 
In my case I believe strongly that A.m.m. is under threat due to the proliferation of traditional commercial bee types and as such it needs to be protected. Protection is not a static concept and there is no reason why A.m.m. cannot be a candidate for varroa tolerance - in the natural genetic variation within the subspecies.
Mellifera is not native to the US so this argument does not apply but it would be a shame to lose the native bee from North West Europe due to uncontrolled hybridisation with other subspecies.
In the South of England there is a complete mish-mash of different bee types so you might as well work away with your collection of swarms and cutouts.


----------



## mike bispham

jonathan said:


> No need for the cynicism about the plot to gain control of world microscope sales but it is definitely wise to be cautious.
> Does Monsanto do microscopes!


No need for cynicism about my cynicism! Seriously, a few folks who find a nice little line in sales patter, selectively presenting matters in a light that offers something of value, can and do shift billions of quids worth of gear. Never drop your guard near salesmen! Talk of high tech gear, and technical terms is sufficient to bamboozle most non-scientists into thinking they're getting something of value. As the shampoo ad says " here comes the science". Haha.



jonathan said:


> Wing morphometry has some use, especially in populations where beekeepers have never used it as a selection criteria.
> It is useful for telling you what something is *not* but not so clear at proving an individual bee is of a specific subspecies.


But Jonathan, what does it tell you except:

a) this bee has genes that code for this vein pattern.

b) (therefore) at least one of its parents had genes that code for this wing pattern.

By looking a lots from the same hive you can deduce that probably the queen had two alleles for that pattern _or_ that she mated with several drones with that allele, or even both. But without a number of other morphological traits to help reach a conclusion about likely make-up and purity of race/ecotype that offers no help at all. 

It doesn't tell you that a bee is not of a particular subspecies. You do realise that the longer this goes on, the more bees of all backgrounds will tend to have Amm wing patterns? The more people around you are doing it, the more likely it is that you are getting a false result?



jonathan said:


> Picking up on some of your other points, I would argue strongly that it is important to preserve pure race subspecies where they are still free from hybridisation. When they are hybridised they cannot be unhybridised at a later date so the subspecies is lost for ever.


Well, yes. But I suspect that the proper way of doing that is, first, by excluding - at least minimising - foreign genetic input; second by allowing natural selection to run its course. As soon as you start to interfere - especially if doing so badly because unequipped with the proper information to make selections, you become part of the problem - one of the factors that make the job harder. 

I can't see that making selections based on just two or three features is going in the right directionway at all - its just making more bees with the features that will fool more amateurs further down the line. Unless.. it has been shown that _*in that place*_ those features do tend to represent a high level of local native purity. 



jonathan said:


> In my case I believe strongly that A.m.m. is under threat due to the proliferation of traditional commercial bee types and as such it needs to be protected. Protection is not a static concept and there is no reason why A.m.m. cannot be a candidate for varroa tolerance - in the natural genetic variation within the subspecies.


I agree - although I might argue for stable thriving local ecotypes whatever their origins. Put it this way: if we disappeared tomorrow, the bees that would be around in fifty years would probably be as good as any that might result from exterminating all but high-purity native black bees, then disappearing. 



jonathan said:


> Mellifera is not native to the US so this argument does not apply but it would be a shame to lose the native bee from North West Europe due to uncontrolled hybridisation with other subspecies.


Again, I agree. I just don't think that having amateurs fiddling around with microscopes is assisting that aim, and it could well be making it harder. Amateurs who keep unadapted strains alive which then poison the genetically diverse natives around, on the ground that they exhibit native morphological features and so must be preserved at all costs, are worst of all. (Present company, if that applies, excluded of course!)



jonathan said:


> In the South of England there is a complete mish-mash of different bee types so you might as well work away with your collection of swarms and cutouts.


Sure. Not only that but a constant input of bees from heaven knows where. 

Mike (UK)


----------



## jonathan

You don't actually use a microscope for scanning bee wings.
All you need is a decent flatbed scanner and the software is free to download at drawwing.org.
I use the Epson scanner I use for general scanning of documents. Anything at 2400 dpi or over is sufficient.

Wing morphometry easily distinguishes between pure race subspecies.
The average cubital index of Carnica is about 4.0 whereas with A.m.m it is around 1.8.
A.m.m. has a negative Discoidal shift whereas Carnica is positive.
The problem is that it is of dubious use with regard to hybrids.
If you cross a pure A.m.m. queen with pure carnica drones does it produce a CI half way between the two? Who knows.
If you cross a mongrel queen with mongrel drones what do the wings tell you? Probably nothing


----------



## WLC

By using PCA, De Jong was able to study those kinds of issues.


----------



## mike bispham

jonathan said:


> Wing morphometry easily distinguishes between pure race subspecies.
> The average cubital index of Carnica is about 4.0 whereas with A.m.m it is around 1.8.


Jonathan,

What that says is... one subspecies is associated with one pattern, one with another.

If you have a hybrid - of the least sort - what it tells you is: the dna that coded this wing originally came from a particular subspecies.

It doesn't tell you how long ago the allele was passed on from a pure individual - it might be hundreds or thousands of generations ago.

That bee might be 99.99% Carnica, 0.001% Amm! It might be 99.99% Amm, 0.001% Carnica. And it might be anything inbetween, taking in several subspecies. 

Again:

3. Available methods and markers
3.1. Morphometry
*There is no morphological "key" to honey bee subspecies, no simple logical tree based on a sequence of single discriminating characters*. Instead, measurable morphometric characters show gradual changes and their ranges mostly overlap between subspecies. Thus, subspecies often differ only slightly in the mean values of several body characters, and therefore advanced statistical methods are required for discrimination of groups. The concept of numerical taxonomy was introduced into honey bee taxonomy by DuPraw (1964, 1965) and further elaborated by Ruttner et al. (1978).
Standard methods for characterising subspecies and ecotypes of Apis mellifera
Marina D Meixner1*, Maria Alice Pinto2, Maria Bouga3, Per Kryger4, Evgeniya Ivanova5 and Stefan Fuchs6
https://bibliotecadigital.ipb.pt/bit..._etal_2013.pdf



jonathan said:


> The problem is that it is of dubious use with regard to hybrids.
> If you cross a pure A.m.m. queen with pure carnica drones does it produce a CI half way between the two? Who knows.


It will draw genes/alleles from, yes, about half each. At random. 



jonathan said:


> If you cross a mongrel queen with mongrel drones what do the wings tell you? Probably nothing


That somewhere in its ancestry the bee has a particular subspecies.

Bear with me while I try to get a grip on this with an analogy.

Imagine we have three lego sets, all building the same house, but with red, blue, or yellow bricks and other parts.

We mix them all up, and a colour-blind person separates the parts back into three sets, now of mixed colours.

We build a house, and throw a cover over it. Someone comes along and says, if you take a single brick out of that house, I'll tell your whether the house belongs to the red, the blue, or the yellow set.

Eh? we say? That isn't, can't be, a legitimate question. You can't answer a question that isn't legitimate. 

It isn't legitimate because that house is made from a mixture of bricks from 3 sets. It doesn't belong to any.

So.. what you're saying is, you've found a village with 90% blue houses, all covered up, and you can, by making a little hole in the same corner, and looking at one particular corner-brick, tell if one house is more or less than than 90% blue. 

Bear in mind, each house is made from hundreds of thousands of bricks. 

There is, I agree, a relation: finding a yellow or red brick indicates the likihood of a lower blue component - which is to say more hybridisation. Is that the basis of the reasoning? 

(That doesn't btw equate, for a moment, with your claim "Wing morphometry easily distinguishes between pure race subspecies")

And: answer me this: how many deliberate selections made by beekeepers on this basis will it take to negate that diagnosis due to artificially enhanced levels? Have we already passed that threshold? If not, when are we likely to pass it?

At the moment I'm going to stick with the expert line against the amateur would-be pure-subspecies breeder: 

"*There is no morphological "key" to honey bee subspecies, no simple logical tree based on a sequence of single discriminating characters*."

Mike (UK)


----------



## mike bispham

WLC said:


> By using PCA, De Jong was able to study those kinds of issues.


Using technical acronyms tends to baffle rather than explain.

Mike (UK)


----------



## jonathan

Mike.
Post 95 makes no sense from start to finish!
Genetics is not a subject to be made up as you go along.
Stick to what you know.
We are largely in agreement about the limitations of wing morphometry.

This plot shows wing samples from one of my A.m.m. colonies compared with wings from a New Zealand carnica queen colony in Scotland which I scanned. The CI difference alone makes it very easy to separate the two.
Like I said, I have data from about 100 colonies I have scanned and I am much less confident about the utility of this technique than I used to be - when you are dealing with hybrids.









Incidentally, the queen from the A.m.m colony was superseded in her 4th year and the colony never tried to swarm - for those who believe A.m.m. is naturally swarmy!


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## JWChesnut

I haven't gone back to look at the fundamental papers establishing the wing morphometric discipline. I played around with wing scans (and posted these to a previous thread). I see two big issues with its implementation. 

First is auto-correlation. The wing measurements are cross linked -- a shift in one, by necessity affects the intersections of its neighbors. This means rather than 7, or 18 INDEPENDENT measures you have a set of data that reflects the same underlying condition. A movement in one vertex affects its neighbors -- this amplifies apparent differences and distinctions. The statistical technique of Principal Component Analysis obscures the issue of auto-correlation because the x-y coordinates of a PCA are *not* reducible to any single "real-world" parameter. The axes are synthetic.

Second issue affects the "discoidal shift" -- which is the amateur-friendly measure of the position of a single vertex.

Discoidal shift is the measure of the angle of a line (d-e) relative to the perpendicular of the line (a-d-b). The constructed vertex d is erected from the wing vein intersection "c". (or a point of tangency on a circular arc "acb").

In my interpretation, "c" is a vague point (as the wing veins are broad at this juncture). The extremely short segment (c-d) defines the perpendicular, but is very short relative to the very long d-e ray. Inherent error in the Discoidal shift is enormous -- it is at least 3 degrees of freedom away from its definitional vertex's. Extending the baseline from the tangent point "c" which is very close to the chord line a-b magnifies its error in location. "a" and "b" are points of maximum rather than vertexes, and also have error associated with them. Their exact location can shift the tangent perpendicular. 

I work with principal component dataset all the time, and am in awe of their power *and* their ability to lead astray. Auto-correlation can be your friend (as it defines "neighborhoods" and linked parameters) or your worst enemy.


----------



## jonathan

mike bispham said:


> I don't think any of that says 'wing analysis alone tells you something about an individual's subspecies identification' - and if it did then it would be in direct conflict with Meixner et all (as just cited)
> 
> It seems to me to be talking about automating a process - that may then be of use to to people who know what they're doing, and know not to read (far) too much into a single morphometric.
> 
> Has anyone actually read the paper?
> 
> Mike (UK)


I have. And I cited it in an article I wrote last October for the December edition of the IBRA Bee World magazine!
It's worth joining IBRA to get access to the magazine and the Journal of Apicultural research.


----------



## jonathan

JWC.
You need a fairly large sample size to get any meaningful data about wing venation, and it may not even be that meaningful!
If a queen mates with say 15 drones and you want to sample a colony you need to take a reasonable sized sample to reflect this.
A large sample should even out some of the measurement error as well.
Drawwing places the points at the wing vein junctions automatically but you do need to correct some of the placements manually.
I agree with the caveats you are making about measurement.
Getting off topic for a treatment free thread.


----------



## JWChesnut

Jonathan,
Thanks for your real-world caveats and contributions. Yup, we've "bird-walked" way off-topic (I was trying to understand what point WLC was trying to make by deconstructing his gnostic comments). Good luck with the Amm breeding project.


----------



## WLC

GWV is a useful tool in developing TF bees.

Dr. De Jong, in Brazil, is the worldwide expert in its application.

Finally, the rest of us are waiting for a GWV app. for Honeybees to come out of Brazil. 

I'm a pragmatist.


----------



## mike bispham

jonathan said:


> Mike.
> Post 95 makes no sense from start to finish!


Jonathan,

I think it does. It would be useful if you could tell me what doesn't make sense. Then we could do some learning.



jonathan said:


> We are largely in agreement about the limitations of wing morphometry.
> 
> This plot shows wing samples from one of my A.m.m. colonies compared with wings from a New Zealand carnica queen colony in Scotland which I scanned. The CI difference alone makes it very easy to separate the two.
> Like I said, I have data from about 100 colonies I have scanned and I am much less confident about the utility of this technique than I used to be - when you are dealing with hybrids.


That seems to me to say: I spent a lot of time finding out something I could have found out by glancing sideways at the bees! 

I can see the utility when, like Ruttner, there are only two sub-species in play and you have the capacity to create and hold an island of foreign bees within the larger pond. (In his case he was attempting to wipe out the home bees using a massive scale centrally planned multi-beekeeper operation)

Anyway, its been an interesting conversation and I've learned a lot - thanks.

Mike (UK)


----------



## mike bispham

JWChesnut said:


> I haven't gone back to look at the fundamental papers establishing the wing morphometric discipline. I played around with wing scans (and posted these to a previous thread). I see two big issues with its implementation.


Angels on pinheads...?

Mike?


----------



## mike bispham

WLC said:


> GWV is a useful tool in developing TF bees.
> 
> Dr. De Jong, in Brazil, is the worldwide expert in its application.


This would tie us back to the topic ... can you direct us to relevant papers or reports WLC?

Mike (UK)


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## WLC

Most of the papers on the methodology have Francoy as the lead author.


----------



## WLC

In my opinion, the real value of using GWV in TF beekeeping is that it can give you information with regards to the degree of hybridization of your resistant stock.

You can also start to answer questions relevant to queen mating.

For example...

Are my queens mating with both domestic and feral/AHB stock?

Do queens mated earlier in the season show different hybridization patterns than later mated queens?

Do I have more than one laying queen in my colony?

So, the GWV/PCA 'app' does have it's limits, but it does have a range of questions that it can do well in exploring.


----------



## Barry

WLC said:


> In my opinion, the real value of using GWV in TF beekeeping is


Oh yeah, right up there with Thelytoky!


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## mike bispham

WLC said:


> Most of the papers on the methodology have Francoy as the lead author.


I found this one, thanks.

Variation morphogeometrics of Africanized honey bees (Apis mellifera) in Brazil
Lorena A. NunesI; Edilson D. de AraújoII; Luis C. MarchiniI; Augusta C. de C. C. MoretiI
(I imagine your author appears in the references)
http://www.scielo.br/scielo.php?script=sci_arttext&pid=S0073-47212012000300011

Having only scanned it I have a better idea of what is possible. Its not simple though is it? 

I really can't yet see a use for tf beekeeping. Unless you believe a particular race is predisposed toward mite management (I don't think there are any grounds for such a belief) and on the basis that locally adapted bees are best, then you're going to be working with hybrids, and there's little point in trying to do anything else. Unless you live in an area where a native race remains. But then you're doing something else, not doing something that aids tf. 

With that said, WLC's reasons (#107) are interesting, and I'd like to hear more. Maybe we could make them the starting point of a new thread - morphometry in the service of tf selection goals or somesuch.

I'd also like to hear what it is Jonathan finds objectionable in my housebrick analogy. I thought it worked rather well. We need to note that every brick is a certain shape/fits in a certain place to complete the picture. 

Mike (UK)


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## WLC

Barry:

Don't underestimate the Lusby/thelytoky issue with regards to TF beekeeping.

TF beekeepers do need some way to examine hybridization issues or at least know where their bees 'fit in' with regards to other stock.


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## Barry

mike bispham said:


> We need to note that every brick is a certain shape/fits in a certain place to complete the picture.


But the design of a brick is uniformity. Any brick can fit the same place.


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## WLC

Mike:

They're using a different statistical methodology in generating PCA plots than jonathan is using.

So, it's a different issue.


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## Barry

WLC said:


> TF beekeepers do need some way to examine hybridization issues or at least know where their bees 'fit in' with regards to other stock.


We do?

Humor me. Do you get on Beesource and reply to posts right there in your classroom in front of students, or do you step out of the room to the teachers lounge to reply? Curious how this happens when you're a teacher around students all day. Is Beesource part of the curriculum? :banana:


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## mike bispham

Barry said:


> But the design of a brick is uniformity. Any brick can fit the same place.


That's what I mean. We have to use the analogy maybe of a stone building, where each stone is different/and has a special position - a 3D jigsaw puzzle. 

For our three subspecies, each of tens of thousands of special stones comes in 3 colours. (We're still simplifying, but this'll do to get on with.)

When we peek through our hole in the covers of our newly built houses, we're looking at the same shaped stone each time. It may be any of three different colours.

The question is what can the knowledge thus gained tell us about the rest of the house/bee?

Say its the right colour - blue. Does it tell us that the rest of the stones are blue? No.

Does it tell us how many might be blue? No.

Say its the wrong colour. Does it tell us how many of the other stones are the wrong colour? No. 

It does tell us that its not a 100% pure blue house (and, if say the stone is yellow, that its also not a 100% red house)

It could be that *all the other stones are the right colour, and its just that one that's wrong*! (Very unlikely I know, but its still true.)

It *is* very likely that all the other stones are evenly distributed (in our case) between the 3 colours. And in that case the one with a blue stone (that we can see) is no more representative of its class than the rest.

Until you have an idea of the sorts of proportion of components held in any house, you can't evaluate significance of the presence of one particular stone. (We do have that knowledge here - they are evenly distributed from 3 sources)

You can get a picture of the proportions by making lots of holes (always in identical places) and combining the data statistically. *Then, and only then*, can you begin to evaluate matters - and *then and only then* can you assign a (statistical) significance to one particular stone. 

So you have to do an extensive survey, then evaluate what any particular feature might signify, before you start attaching significances - in order to make choices.

And you have to realise that as soon as you start making choices on the basis of a particular feature, you start undermining the significance you can attach to later instances of that feature (from your population).

Its pretty tricky stuff. I'm guessing, but I don't suppose the salesmen tell you that while they're trying to earn a living. 

Mike (UK)


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## D Semple

Anyway I can ban myself from this thread.

After some great earlier posts the last 2 pages have been a car crash

Don


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## Rader Sidetrack

Don, one option for "unsubscribe" from a thread is on your "Settings" page. Try clicking this link - it should take you to the correct spot:
http://www.beesource.com/forums/usercp.php

But, even _car crashes_ can be an interesting study in physics - and personalities! :lookout:



:gh:


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## mike bispham

WLC said:


> Mike:
> 
> They're using a different statistical methodology in generating PCA plots than jonathan is using.
> 
> So, it's a different issue.


Its a complex issue all round as far as I can see. Those who don't see that are almost certainly wasting their time and messing things up for those who do see it. 

Mike (UK)


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## mike bispham

D Semple said:


> Anyway I can ban myself from this thread.
> 
> After some great earlier posts the last 2 pages have been a car crash
> 
> Don


Ban yourself from this thread you can. Rollercoasters most threads are.

(UK) Mike : )


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## jonathan

WLC said:


> In my opinion, the real value of using GWV in TF beekeeping is that it can give you information with regards to the degree of hybridization of your resistant stock.


If you start with a pure race queen which has mated with drones from the same subspecies that is probably true. If you start with a hybrid queen which has mated with a selection of say 15 random drones how are you going to ascertain the genetics of each of her 15 virgin queen daughters which are half sisters then take into account the different drones and different numbers of drones each will mate with. Good luck with that one. Maybe with access to a lab and a budget of several thousand dollars per colony!
The scattergram of the wing plots from a hybrid queen is likely to be all over the place rather than a nice tight cluster and then you are looking at virgin queen daughters from bees with these varied wing patterns which mate with a wide selection of random drones. You would have more accuracy reading tea leaves.

The problem here is bee genetics. If a queen mated with a single drone, the wing patterns would be more straightforward to interpret but that's not the case.

Take this example. Each dot represents a scan of a worker bee wing from a colony. Any of these could potentially have become a queen.








You have wings which range from a Discoidal shift of -6 and a low cubital index of 1.5 to those which have a Discoidal shift of +1 and a Cubital index of well over 2.0. Which one of these could become a daughter queen?
In terms of appearance, a colony like this can look very uniform, ie all dark with no yellow banding.
That is not even an extreme example as I don't have that much hybridisation in my bees. I have seen scattergrams where the wing plots appear to be randomly distributed over the entire plot area. As I said, good luck with the tea leaves.


----------



## Juhani Lunden

jonathan said:


> As I said, good luck with the tea leaves.


 Well said, LOL.


----------



## WLC

Jonathan:

I don't think that De Jong is using Discoidal shift and Cubital index in the PCA. They seem to be using just 8 points for the GWV data.

I also wouldn't say that molecular data is that much 'reliable' when it comes to population studies.

I'm not relying on tea leaves. I'm citing peer reviewed studies.


----------



## mike bispham

WLC said:


> Are my queens mating with both domestic and feral/AHB stock?


How consistent are your readings from these two populations respectively? 

What I'm trying to establish is, how well do you know that what you are seeing is significant, based on real world information? And what grounds can you supply for that evaluation?



WLC said:


> Do queens mated earlier in the season show different hybridization patterns than later mated queens?
> 
> Do I have more than one laying queen in my colony?


Good questions, but can you properly justify the claim that you are able to answer them well?

Mike (UK)


----------



## mike bispham

WLC said:


> I'm not relying on tea leaves. I'm citing peer reviewed studies.


WLC,

I've offered what I think is a half decent criticism of one of your studies (that it makes a key statement that is inconsistent with the others, and with a reasoned position based on biological fundamentals) 

From my point of view, you ought to address that criticism if you expect me to accept your references.

A good question to ask of that (suspect) paper: who were the (peer) reviewers? More software designers?

I've been reviewing these issue in my copy of Ruttners book, and he is explicit: 40 different morphometric features are available for the two populations that interest him, and he wants a good number to be used in any interrogation of type. This criticism, against the usefulness of wing measurements alone, even for just two populations, needs answering with more than just one reference written by software designers, that is countered pretty much exhaustively by the more relevant fields.

Mike (UK)


----------



## jonathan

WLC said:


> Jonathan:
> 
> I don't think that De Jong is using Discoidal shift and Cubital index in the PCA. They seem to be using just 8 points for the GWV data.


It takes 8 points to get the data for Discoidal shift and Cubital index.


----------



## sqkcrk

But can't u just ACD me ASAP w/ a DVR STP?


----------



## mike bispham

jonathan said:


> It takes 8 points to get the data for Discoidal shift and Cubital index.


Before we talk about the number of angels that can dance on the head of a pin, can we establish the existence of angels in the first place?

Mike (UK)


----------



## jonathan

mike bispham said:


> Before we talk about the number of angels that can dance on the head of a pin, can we establish the existence of angels in the first place?
> 
> Mike (UK)


I don't think looking at wing venation is of any use at all in looking for treatment free bees. I am just providing background information as I do not think there are too many posting on the thread who have any practical experience of taking samples and using the software.
It can separate pure race subspecies - but as oldtimer said in a previous post, you can often do that on abdomen colour alone.
I get the odd colony which looks like pure race A.m.m., but when you check a wing sample there are clear signs of hybridisation. In that sense it is useful to avoid rearing queens from the colony if you want to keep pure race stock.

Much more useful for treatment free beekeeping would be the collating of accurate mite count data based on sugar shake or alcohol wash of a fixed number of bees.
The pitfall of many would bee treatment free beekeepers is just throwing bees into a box, not treating, then watching and waiting for a miracle.


----------



## Oldtimer

Yet another thread becomes *W*ay*L*aid *C*ompletely.


----------



## WLC

From Fusionpower:

"The major item I see missing in your plan is management of drones and management of a mating area so the number of mite tolerant colonies can be dramatically increased in 2 or 3 years. Also, in my experience, finding a mite tolerant colony usually involves a lot of unwanted genetics that take years to reduce to a tolerable level. I found a queen whose colony was mite tolerant in 2004/2005. The unwanted genetics turned out to be a very high level of hive defensiveness. They could not be worked without a suit. And no, they were not Africanized, they were just typical AMM stock for the area. It took 7 years to get them toned down enough to work in short sleeves. "

I think that geometric wing venation has an application to Fusionpower's experience and charting progress towards a TF apiary.

I might be in a similar situation with BeeWeavers.

Having used molecular methods in identifying species and hybrids, I would much prefer to use a simpler, faster, and cheaper approaches like De Jong's method.

Most of us have scanners, but most of us don't have a molecular biology lab or accounts with biotech companies established (although I do).


----------



## beekuk

WLC said:


> Most of us have scanners, but most of us don't have a molecular biology lab or accounts with biotech companies established (although I do).


 Have you got any good photographs of inside your lab, sounds like an interesting place.


----------



## WLC

Here's a paper that illustrates how Francoy and De Jong were able to follow hybridization between Africanized and Italian Honeybees:

"Rapid morphological changes in populations of hybrids between Africanized and European honey bees."

http://www.funpecrp.com.br/gmr/year2012/vol11-3/pdf/gmr2371.pdf

It look likes they were using MorphoJ software to do the statistical analysis.

Here's a peak at some analytical methods used in a different paper (Geometric morphometrics reveals morphological differentiation within four African stingless bee species)...

"Statistical Analyses
The Cartesian coordinates of the landmarks were Procrustes aligned to evaluate existing shape variations among the different populations using MorphoJ software version 1.03 (45). Within MorphoJ software, further statistical
computations including principal component analyses (PCA), canonical variate analyses (CVA), discriminant
function analyses (DFA), Procrustes ANOVA and Regression analyses were conducted to discriminate populations
within each species against the different ecological zones. In addition, relative warp analyses was conducted to
summarize the variation among the specimens (with respect to their partial warp scores) using the tpsRelw software version 1.49 (46). The Mahalanobis square distances between the centroids of CVA were then used to construct a
neighbor joining dendogram with MEGA5 software version 5.05 (47)."

So, yes, you can examine hybridization with a different application in the same software package (MorphoJ).


----------



## Oldtimer

jonathan said:


> I get the odd colony which looks like pure race A.m.m., but when you check a wing sample there are clear signs of hybridisation. In that sense it is useful to avoid rearing queens from the colony if you want to keep pure race stock.


Actually that's a good point Jonathan, maybe I should even have a look at this myself.

We used to have a large AMM population here, and while no recognisable AMM bees are left their genetics live on in our Italian population, which pretty much all are hybridised even if only to a small extent. Despite my best efforts I will occasionally get a nice yellow looking bee, but they are nasty aggressive little sods. I always wonder if this is from a bit of AMM in them, wing morphology might be a way I could check this. First, I'll see if my scanner is up to scanning a bees wing, if it can, I'll post up here and ask to be pointed in the right direction to get the info to analyse.
To my knowledge wing morphology has never been used in my country for breeder selection so that would remove the element of it having been selected for as a trait.


----------



## julysun

pass


----------



## WLC

mike bispham said:


> How consistent are your readings from these two populations respectively?
> What I'm trying to establish is, how well do you know that what you are seeing is significant, based on real world information? And what grounds can you supply for that evaluation? Good questions, but can you properly justify the claim that you are able to answer them well? Mike (UK)


I bought 2 open mated queens from Texas. The first came with the package in April. The second was a replacement queen in June. I could ask the multiple queen question because I see different morphologies in the workers.

My point being, you should have questions that you could reasonably expect that wing morphometrics can help you to answer.

For example, in the paper, "Rapid morphological changes in populations of hybrids between Africanized and European honey bees", They make claims of seeing evidence of dominance and epistatic influences.


----------



## mike bispham

jonathan said:


> I don't think looking at wing venation is of any use at all in looking for treatment free bees.


Jonathan,

I'm interested to hear WLCs side of this. 



jonathan said:


> I get the odd colony which looks like pure race A.m.m., but when you check a wing sample there are clear signs of hybridisation. In that sense it is useful to avoid rearing queens from the colony if you want to keep pure race stock.


So you also have colonies that check out as pure Amm using a series of morphometric tests?



jonathan said:


> Much more useful for treatment free beekeeping would be the collating of accurate mite count data based on sugar shake or alcohol wash of a fixed number of bees.


I would have thought that - if you go along with that - it would apply just as much to your pure race/near pure race bees? Morphometrics doesn't help in any way with resistance traits does it? (Although I assume you will be working with the assumption that if you can get a pure race stable it will be easier to fix traits - including resistance behaviours - than in a mongrel population...)



jonathan said:


> The pitfall of many would bee treatment free beekeepers is just throwing bees into a box, not treating, then watching and waiting for a miracle.


I agree - though we have different recipies for fixing the problem

Mike (UK)


----------



## mike bispham

WLC said:


> Here's a paper that illustrates how Francoy and De Jong were able to follow hybridization between Africanized and Italian Honeybees:
> 
> "Rapid morphological changes in populations of hybrids between Africanized and European honey bees."


WLC,

Follow changes yes. Undo them (put humpty dumpty back together) no.



WLC said:


> ... further statistical computations .... were conducted to discriminate populations within each species against the different ecological zones.


Established ecotypes, yes. These may be 'hybrids', yes. But 'established' hybrids. This means you can find features that are common to all individuals. The absence of that feature then signals otherness - further hybridisation.

You have a base to measure against, just as you do with pure species.

None of this is available in the case of mongrels or higher levels of recent, unstable hybridisation between pure races or ecotypes. 

I would imagine that deep - forest/wilderness 'survivor' bees might show a good level of stability - and the ability to track that might be a vary useful thing.



WLC said:


> So, yes, you can examine hybridization with a different application in the same software package (MorphoJ).


I think you need to refine your terminology for statements like this. There is 'hybridization' and then there is 'hybridization'. A stable local ecotype may emerge from a hybrid population, and be amenable to morphological analysis. 

A deeply and recently hybridised ('mongrel') population won't be amenable - it'll be all over the place. But it may have a shared set of feature that are gold dust - the mite tolerance behaviours. And it may well be capable (if left alone) of becoming a thriving, stable local ecotype of the first sort.

Does all that fit with your understanding?

Mike (UK)


----------



## mike bispham

WLC said:


> I bought 2 open mated queens from Texas. The first came with the package in April. The second was a replacement queen in June. I could ask the multiple queen question because I see different morphologies in the workers.


Surely the queen could supply positve and negative results - on the assumption that she carries a gene set for the geometry you're looking for and another that doesn't code for that geometry. Second, it could be the sperm she carries that is giving the wrong results for you.

If you knew the queen was pure race (and thus both her gene sets would supply the right shapes) you could then blame the matings. 



WLC said:


> My point being, you should have questions that you could reasonably expect that wing morphometrics can help you to answer.


I'm not yet convinced that you are getting the answers you think you are. You're going to have to step me through the rationale at a basic level.

Mike (UK)


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## WLC

Mike:

While I think that breeders conserving 'pure' stocks of Honeybees is important, I believe that the vast majority of TF bees are hybrids of one kind or another. We already know that queens that are well mated with a diversity of drone stock can produce healthier colonies.

Since my TF bees are store bought (open mated in Texas), and many TF beekeepers use ferals, I think that GWV morphometrics provides important information with regards to the stock.

For example, I would want to know if loss of hybridization leads to a loss of resistance in my own bees. I can't open mate with the hybrid swarm here in NYC.

I don't have to wait for a miracle with BeeWeavers. I'm trusting Daniel Weaver's skill as a TF queen breeder/producer.

There's one key issue here in the U.S. that you don't have in the UK. We have to have a way to guard against AHB.

While we do use morphometrics to test for AHB, since the Darger Thesis, it looks like we're using a method that gives false positives, and we need to use GWV instead.

With regards to ferals in Florida which are being misidentified as AHB, it becomes a Honeybee conservation issue.

They may be destroying a valuable, resistant stock.


----------



## mike bispham

WLC said:


> Mike:
> 
> While I think that breeders conserving 'pure' stocks of Honeybees is important, I believe that the vast majority of TF bees are hybrids of one kind or another.


WLC,

Could you tell me whether you agree my distinction between stable local ecotypes that are 'hybrids', and 'mongrels'? If so, how would you place the dividing line between the two?



WLC said:


> We already know that queens that are well mated with a diversity of drone stock can produce healthier colonies.


Yes, although I'm not sure that entails hybridisation - well mated within subspecies works just as well as far as I know.



WLC said:


> Since my TF bees are store bought (open mated in Texas), and many TF beekeepers use ferals, I think that GWV morphometrics provides important information with regards to the stock.


I think it can only tell that there may be a fraction of the thing you fear - AHB genetics - that part responsible for WV pattern. I think you'd expect a lot of that in open mated queens from within the Texas hybrid 'survivor population', and there is no rationale for supposing that aggressiveness might accompany it.



WLC said:


> For example, I would want to know if loss of hybridization leads to a loss of resistance in my own bees. I can't open mate with the hybrid swarm here in NYC.


You would measure 'loss of hybridisation' by rising uniformity among workers? That may due to lower mating rates, or the presence of a stable survivor hybrid ecotype in NYC. 



WLC said:


> I don't have to wait for a miracle with BeeWeavers.


What would qualify as a miracle?



WLC said:


> There's one key issue here in the U.S. that you don't have in the UK. We have to have a way to guard against AHB.


You have to guard against undue aggressivness. So breed away when you see it. 



WLC said:


> While we do use morphometrics to test for AHB, since the Darger Thesis, it looks like we're using a method that gives false positives, and we need to use GWV instead.


Eh? Geometric Wing Veination is a morphometric. ...? 



WLC said:


> With regards to ferals in Florida which are being misidentified as AHB, it becomes a Honeybee conservation issue.


The problem here is that 'Africanised' is mumbo-jumbo. A scare-term. There's no such thing as an 'AHB.' There are hybrids with more or less African genetics, and that seems to roughly corrolate with aggressive behaviours. Fair do's. But finding a bit of African influence in a wing isn't a good reason to terminate the line. 



WLC said:


> They may be destroying a valuable, resistant stock.


Quite. So might you.

Mike (UK)


----------



## mike bispham

Oldtimer said:


> To my knowledge wing morphology has never been used in my country for breeder selection so that would remove the element of it having been selected for as a trait.


You've played in the slips haven't you.

Mike (UK)


----------



## Oldtimer

You speak in riddles but if you mean what I think you mean the answer is occasionally, more outfeild but never much enjoyed, roll on winter and something more physical.


----------



## WLC

mike bispham said:


> WLC,Could you tell me whether you agree my distinction between stable local ecotypes that are 'hybrids', and 'mongrels'? If so, how would you place the dividing line between the two? Mike (UK)


Managed Honeybees aren't considered a stable ecotype. That would apply to local ferals. While there have been reports of reproductively isolated feral populations, as a general rule, ferals and managed bees are in disequilibrium. There will be introgression of alleles in either direction.


> Yes, although I'm not sure that entails hybridisation - well mated within subspecies works just as well as far as I know.


I'm referring to hybridization regarding resistant vs non resistant stocks.


> I think it can only tell that there may be a fraction of the thing you fear - AHB genetics - that part responsible for WV pattern. I think you'd expect a lot of that in open mated queens from within the Texas hybrid 'survivor population', and there is no rationale for supposing that aggressiveness might accompany it.


It's not so much the aggressiveness. It's the other characteristics that make them challenging to work. As I've said elsewhere, I think that AHB drones mate earlier in the season than domestic ones do. I would expect to see that in a scatter plot of early vs late queens.


> You would measure 'loss of hybridisation' by rising uniformity among workers? That may due to lower mating rates, or the presence of a stable survivor hybrid ecotype in NYC.


I'd measure that by comparing the before and after analyses. What stable ecotype in NYC? They're managed colonies that get replaced frequently.


> What would qualify as a miracle?


Buying any old package bees, and having them actually survive.


> You have to guard against undue aggressivness. So breed away when you see it.


How about I just order a new queen from BeeWeaver?


> Eh? Geometric Wing Veination is a morphometric. ...?


They use a different method for determining AHB. But, it not the same as De Jong's method.


> The problem here is that 'Africanised' is mumbo-jumbo. A scare-term. There's no such thing as an 'AHB.' There are hybrids with more or less African genetics, and that seems to roughly corrolate with aggressive behaviours. Fair do's. But finding a bit of African influence in a wing isn't a good reason to terminate the line.


Africanized is, unfortunately, very real. Some of their traits do seem to be dominant. There are some folks who do hive feral swarms. If they fit the AHB morphotype, you can't keep them. Some southern beekeepers were told that there are AHB around, but Darger has shown that they're some else, and that the test used returned false positives.


> Quite. So might you.


My bees are store bought. They're chattel. Not a resource.


----------



## mike bispham

Oldtimer said:


> You speak in riddles but if you mean what I think you mean the answer is occasionally, more outfeild but never much enjoyed, roll on winter and something more physical.


I meant well caught! 

Mike


----------



## Oldtimer

Oh I get it but possibly lost on the non English non NZ audience!


----------



## mike bispham

WLC said:


> Managed Honeybees aren't considered a stable ecotype.


WLC,

Surely I think that would depend on who is doing the managing, where they got their bees, and how hard they work at keeping them pure. And who is doing the considering!



WLC said:


> That would apply to local ferals.





WLC said:


> While there have been reports of reproductively isolated feral populations, as a general rule, ferals and managed bees are in disequilibrium.


Are you talking about linkage disequilibrium? If so what does that mean for us?



WLC said:


> There will be introgression of alleles in either direction.


As far as I've seen (which isn't far) 'introgressions' refers to the insertion of alleles by repeated backcrossing into a pure strain. The commonly held notion of 'fixing' genes. So I'm struggling to see what you mean here. 

How does 'either' come ito it? When we're talking about mongrels we're talking about multiple genetic sources.

I'm not sure introgression is a usable idea outside of pure strains. In the feral and loosely managed populations there will be a spectrum of mash-up, with mongrels at one end.



WLC said:


> I'm referring to hybridization regarding resistant vs non resistant stocks.


Now I really can't see what you are getting at...?



WLC said:


> What stable ecotype in NYC? They're managed colonies that get replaced frequently.


I've heard of a developing feral population, and wondered if it might have managed to find the sort of reproductive barrier we've been hearing about. 



WLC said:


> My bees are store bought. They're chattel. Not a resource.


That's a big difference between us. Mine are in part the foundations of my future bees. I hope.

Are your pure strain? Is that what enables you to read them (morphometrically)?

Mike (UK)


----------



## WLC

mike bispham said:


> WLC said:
> 
> 
> 
> Managed Honeybees aren't considered a stable ecotype.
> 
> 
> 
> WLC,
> 
> Surely I think that would depend on who is doing the managing, where they got their bees, and how hard they work at keeping them pure. And who is doing the considering!
> 
> If they're in a hive body, I would say no. If you're considering ferals found in the desert, forests, etc., it would apply. For example, Paul McCarthy has hived 'stable ecotypes' (ferals) from mountains in New Mexico.
> 
> 
> 
> Are you talking about linkage disequilibrium? If so what does that mean for us?
> 
> No, I'm referring to gene flow. For example, at one time, AHB genes were believed to be flowing into managed EHB stocks, one-way. Now we know that EHB genes have been flowing the other way as well including feral AHB colonies.
> 
> I'm not sure introgression is a usable idea outside of pure strains. In the feral and loosely managed populations there will be a spectrum of mash-up, with mongrels at one end.
> 
> It's an important issue, especially when dealing with AHB or resistance. It has to come from somewhere into managed colonies. So, if you open mate queens, you're hoping for an introgression of favorable alleles into your stock.
> 
> Now I really can't see what you are getting at .
> 
> You can get resistant colonies by hybridizing your non resistant stock with resistant ferals when open mating.
> 
> I've heard of a developing feral population, and wondered if it might have managed to find the sort of reproductive barrier we've been hearing about.
> 
> The Arnot forest 'survivors' are one example.
> 
> 
> 
> That's a big difference between us. Mine are in part the foundations of my future bees. I hope.
> 
> No, the big difference between us is that I can order bees from BeeWeaver. I think that since they're open mating in their location (Texas), it's a key to getting resistant colonies.
> 
> Mike (UK)
> 
> 
> 
> Sorry about replying within your quotes, but it's a bother to do otherwise.
> 
> As for hybrids vs mongrels: I would call the BeeWeaver queens hybrid Italian/Buckfast. However, once they're open mated in Texas, you could consider the resulting workers to be 'mongrels'.
> 
> Click to expand...
Click to expand...


----------



## mike bispham

WLC said:


> mike bispham said:
> 
> 
> 
> 
> 
> WLC said:
> 
> 
> 
> Managed Honeybees aren't considered a stable ecotype.
> 
> [MB] Surely I think that would depend on who is doing the managing, where they got their bees, and how hard they work at keeping them pure. And who is doing the considering!
> 
> If they're in a hive body, I would say no. If you're considering ferals found in the desert, forests, etc., it would apply. For example, Paul McCarthy has hived 'stable ecotypes' (ferals) from mountains in New Mexico.
> 
> 
> 
> Eh!!!! How does putting bees in a hive body make them stable ecotypes!
> 
> 
> 
> WLC said:
> 
> 
> 
> [MB]Are you talking about linkage disequilibrium? If so what does that mean for us?
> 
> No, I'm referring to gene flow. For example, at one time, AHB genes were believed to be flowing into managed EHB stocks, one-way. Now we know that EHB genes have been flowing the other way as well including feral AHB colonies.
> 
> Click to expand...
> 
> Why not say that instead of using technical terms wrongly?
> 
> Yes of course they've gone both ways. They've been mixing up, natural selection has been sorting out the best adapted combinations, and these are moving toward stable ecotypes where beekeepers can't screw up the process.
> 
> 
> 
> WLC said:
> 
> 
> 
> [MB]I'm not sure introgression is a usable idea outside of pure strains. In the [unstable] feral and loosely managed populations there will be a spectrum of mash-up, with mongrels at one end.
> 
> It's an important issue, especially when dealing with AHB or resistance. It has to come from somewhere into managed colonies. So, if you open mate queens, you're hoping for an introgression of favorable alleles into your stock.
> 
> Click to expand...
> 
> Again you are using this technical term in an inadequate way. Go back and see what I just said - 'introgression' refers to the 'fixing' of an outside gene set in a pure population as a result of repeated backcrossing (in one of the two parent races). As far as I can see - I may be wrong. You don't have that context. So you can't use that technical term.
> 
> Genes coding for resistance behaviours can be found in all populations. They are at a low level in the population unless circumstances have been such as to promote them - natural selection or beekeeper selection. Getting initial stocks with higher levels is great. Keeping them there by constant selective propagation is great. None of that amounts to 'introgression'.
> 
> 'Introgression' of resistance genes, in the sense that they entirely replace alternative allelles would btw be a terribly thing. Your bees would be far too hygeinic.
> 
> 
> 
> WLC said:
> 
> 
> 
> [MB]Now I really can't see what you are getting at .
> 
> You can get resistant colonies by hybridizing your non resistant stock with resistant ferals when open mating.
> 
> Click to expand...
> 
> Yes, ok - though simply 'mating' is probably a better description. And you can lose it again just as fast through mating with treated bees....
> 
> Mike (UK)
> 
> Click to expand...
Click to expand...


----------



## WLC

No Mike, Paul found stable ecotypes on the mountains of NM, then he put them in a hive. 

No Mike, the fear was that AHB genes would swamp the genetic diversity of EHB colonies. Now we know that the flow has changed at the AHB front. In fact, in the South, it appears that AHB gene flow has been slowed down considerably by local feral/managed colonies to such an extent, that the flow may be going the other way.

Yes Mike, I can use the term introgression of alleles. Especially in the U.S. since AHB is an issue.


----------



## jonathan

mike bispham said:


> Again you are using this technical term in an inadequate way. Go back and see what I just said - 'introgression' refers to the 'fixing' of an outside gene set in a pure population as a result of repeated backcrossing (in one of the two parent races). As far as I can see - I may be wrong.


nothing to do with backcrossing and fixing. It is a straightforward term to describe when genes from one population get into the gene pool of another population. for example when Ligustica genes are found in an A.m.m. population.

Varying degrees of Apis mellifera ligustica introgression in protected populations of the black honeybee, Apis mellifera mellifera, in northwest Europe


----------



## Oldtimer

mike bispham said:


> Genes coding for resistance behaviours can be found in all populations.
> Mike (UK)


A citation to support that exists?


----------



## mike bispham

Oldtimer said:


> A citation to support that exists?


"We are now returning to our original goal of having queen producers and interested beekeepers select for this trait from among their own, tried-and-true stocks of bees. It is very important for beekeepers to have many stocks of bees to maintain a healthy level of genetic diversity [...] Fortunately, the hygienic trait is found in all races and stocks of bees."

The Hygiene Queen, Marla Spivak and Gary S. Reuter

http://www.apiservices.com/articles/us/hygiene_queen.htm

That does only specifically cover the 'hygeine' traits as Marla Spivak understood them at the time of writing. Its been a fast moving field and there might be cautions to add.

Modifying my statement: there may be further behaviours, or capacities that are helpful to internal mite management that are less common. But at least some are found in all populations, according to this paper.

Mike (UK)


----------



## mike bispham

jonathan said:


> nothing to do with backcrossing and fixing. It is a straightforward term to describe when genes from one population get into the gene pool of another population. for example when Ligustica genes are found in an A.m.m. population.
> 
> Varying degrees of Apis mellifera ligustica introgression in protected populations of the black honeybee, Apis mellifera mellifera, in northwest Europe


Jonathan,

I'm, learning as I go along, but I think my grumble is still there. The paper you reference talks about introgression of genes into pure race populations - its a study of the different degrees of introgression from apiary bees into a number of Amm ecotypes. The identified problem is that genes from apiary bees have entirely replaced the proper Amm genes - to varying degrees.

So the stable, pure race population has been compromised by introgression. The original genes are lost - if the same introgression occured in all Amm populations the original genes - a part of Amm would be extinct.

That is the *significance *of introgression. 

What WLC describes is simple mixing, the creation of hybrids. And as you both point out, its gone both ways - well actually every which way. And no gene-sets have been lost, no part of any population has been made extinct. In theory at least natural selection still has all the resources it ever had to locate the best possible strains from the admixtures.

Only a pure race or ecotype can be introgressed in a strict sense. What WLC describes is just hybridisation, mixture. I think if you read the abstract of the paper you instance very carefully you'll see the difference.

Its often difficult to speak clearly about aspects of highly complex systems. This is an example - see below - I'm not the only one thinking this term is problematic:

Mike (UK)

Introgression and Introduction
Online Biology Dictionary
http://www.macroevolution.net/introgression.html#.UuduLivFIgY

"Introgression, a word often used in discussions of hybrid variation, has had various definitions. As originally defined by Anderson and Hubricht (1938: 396), it was the "infiltration of germ plasm from one species into another through repeated backcrossing of the hybrids to the parental species." This definition, however, is problematic because it involves the word species, which is not clearly defined. More recently, there has been an attempt to limit use of the term to situations involving permanent infiltration of genes from one population into another (Heiser 1973; Stebbins 1959). Thus, Rieseberg and Wendel (1993: 7) define it as the “permanent incorporation of alien alleles into a new, reproductively integrated population system,” But here, too, a problem arises with the definition of the term "integrated population system." There is also the problem of saying what constitutes "infiltration." In the case of many crosses, the genetic influence of natural hybridization is very local and affects only individuals in the immediate vicinity of the contact zone. In the case of others, the effect may be detectable hundreds of kilometers away. How should one draw a line between these two cases? What is "infiltration" and what is not?

In fact, the main difficulty with "introgression" is that it is thought of as an introduction of genes from one entity (usually thought of as a "species") into another. Actually, natural hybridization between two populations treated as species generally has a genetic effect only on individuals in the vicinity of the contact zone. No genes are introduced into other individuals elsewhere. Over the remaining geographic ranges of the two interbreeding populations, all individuals remain pure and genetically unaffected. The same distinction exists with the breeder's term introduction, the transfer of a trait via hybridization from a donor stock to a target stock. Here, the trait is introduced into only a subset of the target stock. 

Indeed, the whole idea of introducing genes into a "genetic background" is probably misleading. In many cases, what is called "introduction into a target stock" is really crossing individuals of the target stock with individuals of some other type to produce hybrid individuals with genetic complements of mixed parental origin. Esuperanzi (2005: 302) suggests that birds produced by repeated backcrossing to a single parent should be regarded as belonging to that parental type only when they contain less than 1/16 blood from the other parent. But this is clearly arbitrary. Why not 1/8 or 1/32?

Even when the intention is to transfer a single gene to the target stock other genes often are transferred at the same time. For example, Carver and Taliaferro (1992: 131) note that even when the primary purpose is to transfer disease resistance to a crop, other traits, such as yield, may also be increased. If backcrossing occurs to one parental type or the other, the relative contributions of the two parents to the genetic complements of the resulting hybrids will vary in accordance with the amount of backcrossing that occurs. But no matter how many backcrosses to the target stock occur, so long as the traits of the offspring are discernibly mixed, they are hybrids, distinct from the target stock type.

For the foregoing reasons, the terms introduction and introgression, though widely used in the literature on hybridization, are avoided on this website."


----------



## beekuk

More genetic diversity needed.........

Increasing Genetic Diversity of

Honey Bees--A Necessity, Says

Bee-Breeder-Geneticist

Susan Cobey




Dept. of Entomology
University of California at Davis
DAVIS--Increasing the overall genetic diversity of honey bees will lead to healthier and hardier bees that can better fight off parasites, pathogens and pests, says bee breeder-geneticist Susan Cobey of the University of California, Davis and Washington State University.

Just as stock improvement has served the poultry, dairy and swine industries well, the beekeeping industry needs access “to stocks of origin or standardized evaluation and stock improvement programs,” Cobey said.

Cobey is the lead author of the chapter “Status of Breeding Practices and Genetic Diversity in Domestic U.S. Honey Bees” of the newly published book, Honey Bee Colony Health: Challenges and Sustainable Solution.

“The many problems that currently face the U.S. honey bee population have underscored the need for sufficient genetic diversity at the colony, breeding, and population levels,” wrote Cobey and colleagues Walter “Steve” Sheppard, professor and chair of the WSU Department of Entomology and David Tarpy of North Carolina State University, formerly a graduate student at UC Davis.

“Genetic diversity has been reduced by three distinct bottleneck events, namely the limited historical importation of a small subset sampling of a few honey bee subspecies, the selection pressure of parasites and pathogens (particularly parasitic mites) and the consolidated commercial queen-production practices that use a small number of queen mothers in the breeding population,” Cobey pointed out.

The honey bee, Apis mellifera, originated in the Old World where it diverged into more than two dozen recognized subspecies, they related. However, only nine of the more than two dozen Old World subspecies ever made it to the United States and only two of these are recognizable today.

European colonists brought one subspecies, Apis mellifera mellifera or “The Dark Bee” of Northern Europe, to America in 1622, establishing it in the Jamestown colony. The bee was the only honey bee present in the United States for the next 239 years (1622 until 1861).

The Italian or golden honey bee, Apis mellifera ligustica, was introduced into the United States in 1859 and is now the most common honey bee in the United States. “Currently available U.S. honey bees are primarily derived from two European subspecies, A. m. carnica and A. m. ligustica,” the bee scientists said.

The U.S. ban on the importation of bees in 1922 to ward off a tracheal mite (Acarapis woodi) further aggravated the genetic bottleneck. Today the No. 1 enemy of the beekeeping industry is the parasitic Varroa mite (Varroa destructor), which has played a major role in the crippling decline of the U.S. honey bee population.

Found in virtually all bee colonies in the United States, it feeds on bee blood, can transmit diseases, and generally weakens the bee immune system.

What’s being done? “In the U.S. the recognized need to increase genetic diversity and strengthen selection programs of commercial breeding stocks has resulted in collaborative efforts among universities, government researchers, and the queen industry,” according to Cobey, Sheppard and Tarpy. “The current challenges facing the beekeeping industry and new technologies being developed are pushing beekeeping into a new era.”

To increase genetic diversity in the U.S. honey bee gene pool, Cobey and Sheppard are importing honey bee germplasm or semen of several subspecies of European honey bees and inseminating virgin queens of domestic breeding stock. They are also working on diagnostic programs to assist beekeepers to assess colony health and to evaluate commercial breeding stocks.

Cobey, who teaches queen-bee rearing classes and queen bee instrumental insemination at UC Davis and WSU, joined UC Davis in May 2007. Her research focuses on identifying, selecting and enhancing honey bee stocks that show increasing levels of resistance to pests and diseases. Cobey developed the New World Carniolan stock, a dark, winter hardy race of honey bees, in the early 1980s by back-crossing stocks collected from throughout the United States and Canada to create a more pure strain.

Sheppard , who leads the Apis Molecular Systematics Laboratory at WSU, studies population genetics and evolution of honey bees, insect introductions and mechanisms of genetic differentiation. His work was featured in a recent edition of the Washington State University Magazine.

Tarpy, now an associate professor and Extension apiculturist, at North Carolina State University, received his doctorate in entomology from UC Davis in 2000. He studied with Robert Page, emeritus professor of entomology at UC Davis who later became the vice provost and dean of the College of Liberal Arts and Sciences and Foundation Professor of Life Sciences, Arizona State University.
__________________


----------



## Fusion_power

> Modifying my statement: there may be further behaviours, or capacities that are helpful to internal mite management that are less common. But at least some are found in all populations, according to this paper.


I would agree that hygienic behavior is found in all populations of bees. It is a general trait useful in prevention of brood disease. There should be a strong caveat that the trait is present in some populations at a higher frequency than in others. Italians for example have a relatively high level of hygienic traits where A.M.M. bees tend to have very little.

I would strongly disagree that this means that Varroa Sensitive Hygiene is found in all populations. The available evidence so far is that bees expressing very high levels of VSH also test as very highly hygienic, but that hygienic bees do not necessarily express VSH. In other words, you are stating that because A = B, of necessity B = C. You have not first proven that A = C. Either provide some evidence your statement is valid or else back off of it until you can prove it.

After 13 years of exposure to varroa and the death of most feral and many managed colonies in the U.S., I found one single colony that expressed moderately high varroa tolerance. The best I can estimate, 99.9% of our feral bees died up to that point. The only bees that remain from the original population are highly selected for traits that allow them to live with varroa. Unfortunately, this means that most of these bees swarm with an intensity never seen by me before. Serial swarming during summer allows multiple brood breaks which tend to reduce varroa pressure. Any varroa tolerance that relies on brood breaks to keep bees alive is fragile at best and will lead to compromised traits such as honey production.

There are three known mechanisms of varroa tolerance that do not rely on brood breaks. They are mite mauling behavior, varroa sensitive hygiene, and reduced days to worker maturity. VSH traits were present in Primorski bees to a high level as a result of 100 years of mite pressure. Both VSH and mite mauling are now present in feral colonies in the U.S. as a result of unrelentingly high selection pressure from varroa mites plus moderate levels of VSH genetics escaping into ferals.

Look for bees that survive and thrive in the presence of mites without expressing high levels of swarming. Don't hesitate to do what I did by purchasing some queens from a breeder who developed mite tolerant bees. The combination of traits from his queens and from my single queen gave me a very useful and highly mite tolerant line of honeybees.


----------



## mike bispham

Fusion_power said:


> I would agree that hygienic behavior is found in all populations of bees. It is a general trait useful in prevention of brood disease. There should be a strong caveat that the trait is present in some populations at a higher frequency than in others. Italians for example have a relatively high level of hygienic traits where A.M.M. bees tend to have very little.


I agree. Of course those populations exposed for a longer period have better developed and better balanced defence mechanisms - and are, all else being equal, better candidates for starting stock. Russians are supposed to be still better equipped. In all cases mite management ability will likely vary - according to historic exposure and (more recently, and likely dramatically) the use of treatments in progenitor stocks. 



Fusion_power said:


> I would strongly disagree that this means that Varroa Sensitive Hygiene is found in all populations. The available evidence so far is that bees expressing very high levels of VSH also test as very highly hygienic, but that hygienic bees do not necessarily express VSH. In other words, you are stating that because A = B, of necessity B = C. You have not first proven that A = C.


On second reading I think you're right: 

"Hygienic bees detect, uncap, and remove diseased brood from the combs before the disease becomes infectious. Hygienic behavior also is one defense against Varroa mites (Peng et al., 1987), and although it is not the main mechanism of resistance to the mites (Harbo and Hoopingarner, 1997), it appears to limit their reproduction and population growth to some degree. Our studies have shown that it is possible to select for hygienic behavior without compromising honey production or gentleness (Spivak, 1996; Spivak and Reuter, in press). The trait can be found in approximately 10 percent of the managed colonies found in the United States, in any race or stock of bees." http://www.apiservices.com/articles/us/hygiene_queen.htm

It seems likely that the specific intentions of the terms 'hygienic' and 'VSH' hadn't been separated out at the time this was written (the document is undated, but the latest reference is to a 1998 paper). The authors specify as follows:

"Although the common usage of the word hygienic denotes cleanliness, hygienic behavior is a specific response by the bees to diseased and parasitized brood. A colony that keeps its hive clean does not imply that it will be resistant to diseases. "

The authors go on to talk about the frozen brood assay for 'hygienic behaviour'. This test I agree doesn't specifically select for SMR/VSH.

Do you know of any studies that look at the specific issue of VSH across races and populations? 

Its worth noting that only 1 or 2 patrilines need be SMR/VSH equipped in order for the colony to be resistant - and too many more would be damaging. It follows that not all i.e. Russions in a functional natural population are so equipped - just a sufficient proportion. The same must be true of US 'survivor' ferals. 



Fusion_power said:


> There are three known mechanisms of varroa tolerance that do not rely on brood breaks. They are mite mauling behavior, varroa sensitive hygiene, and reduced days to worker maturity. VSH traits were present in Primorski bees to a high level as a result of 100 years of mite pressure.


There is also general resistance to the micro-oganisms that varroa creates vulnerability to. Grooming (auto and allo) without mauling may be separable behaviours. There may be others we don't know of. I don't necessarily agree that brood breaks are always a bad thing - breaks that are timed well with local flows might well be an advantage. The term 'brood break' might turn out to be a blunt instrument for what might be very nuanced and subtle responses to local environments, and we may want to speak more carefully about them. Of course those people who want mathematically programmed bees won't want anything to do with that kind of thing.



Fusion_power said:


> Both VSH and mite mauling are now present in feral colonies in the U.S. as a result of unrelentingly high selection pressure from varroa mites plus moderate levels of VSH genetics escaping into ferals.


It may be only the former reason. Genes coding for all kinds of mite-managment behaviours may well be present in all but the most narrowly diverse populations. I agree levels will be higher where mite exposure is longer (in ferals only).



Fusion_power said:


> Look for bees that survive and thrive in the presence of mites without expressing high levels of swarming. Don't hesitate to do what I did by purchasing some queens from a breeder who developed mite tolerant bees. The combination of traits from his queens and from my single queen gave me a very useful and highly mite tolerant line of honeybees.


That makes sense to me, but I'm increasingly thinking that possession of the right sort of mite in initial stocks would be a strong advantage too. A population of low fecundity mites supplies protection against a population explosion of flown-in mites from treated colonies. 

Mike (UK)


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## D Semple

Good post DarJones (# 153). Appreciate you sharing your experiences and knowledge. If time allows I will be assaying for mite mauling of my surviving hives. 

Mike, good to see less preaching and more thorough analysis in your posts of late. Good job.


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