Varroa Resistant Bees

[sqkcrk]

Twenty years ago Steve Taber said that what we should do about Varoa mites is nothing, And in 30 years, or so, we will have bees that can exist with varroa mites.

Well we didn't do that did we?

What we have done is breeding stronger mites. When we use chemical treatments and managment techniques to "control" varroa mites, we seem to be selecting the mites that still survive those treatments.

What we need is a different tact. Maybe someone from outside our industry can see something that we, who are too close, can't see.

Since we are so successful at raising mites, maybe that is what we should be trying to find an economic value for.

I'm not really serious about that. But, perhaps, that's the kind of thinking that we need more of.

When I was in school, in Ohio, Dr. Tew told us about foraging bees who find a source, tell the rest of the foragers and exploit that prime source.

There are also renegade foraging bees who don't follow the pack. They are out foraging on obscure flowers and finding, what will often turn out to be, the next meal.

Thaty's what we need. Some renegade beekeepers to show us the way to our next meal.

Where are you? What are you doing? How does it work? Show us the way to our next meal.

[Michael Bush]

>Since we are so successful at raising mites, maybe that is what we should be trying to find an economic value for.

Sell them to the researchers. They are always needing more mites. smile.gif

[George Fergusson]

Tony Jadczak was telling me the other day that once upon a time he was overnight shipping capped drone combs to some place for research. I don't remember the details.

[Ian]

>>What we have done is breeding stronger mites

They arent stonger mites.

>>what we should do about Varoa mites is nothing,

Absoulutely wrong!! The work that has gone into breeding for the traits we are selecting for is mind numbing. Just look at what we have accomplished here, even just in the past 5 or so years. Not only have we been able to continously select for heavey honey producers, but we have also maintain all the rest of the traits beekeepers want, wintering, gentleness.
And now as we are selecting heavily for v and t mite tolerance, we have been able to successfully maintain all the other traits we want along side of the tolerance. Mother natures selection pressures arent so focused.

Prime example is what has been happening up here with our own queen breeders.
From their testing and breedings from hives in the area, they have developed an "area select" bee far supperiour in the ability to tolerate or suppress the mite. In fact, much supperiour that the russian stock that they are also testing. AND they have been able to bring all the other wanted traits with that line. Unlike the original russian lines!!

Here are a few number they were showing us,
Unselected hive showed 10-20% grooming behaviour
Russian hives showed a 50% grooming behaviour
While the area select line was continously showing 80% grooming behaviour

They dont like to make claims attatching to thier queens, but the numbers are speeking for themselves.

This is a long ways from what mothernature would have given us

[BjornBee]

The problem I see is just the amount of time it takes to get where I want to be. Every year, I have only so much time to do anything from a bees' perspective.

This will be the third winter without any treatments. I have over 400 hives going through winter without any treatments. Thats hard to do every year. I have seen some good results, but how many years will it take to get to where I want to be?

Although I am trying to have as good as a northern bred bee as possible, I also need to keep in mind that something 100% effective against the V-mite may be years away.

I am trying different over-wintering techniques, such as those employed by others having success. I am greatly interested in managing bee "systems" that take into account a certain amount of bee loss. I think splitting, queen rearing, and propogating your own queens goes a long way.

Everyone is focused on when the perfect queen is going to be bred. Or when the next "silver bullet" treatment is coming on the market. What if that never comes about? Then What? Can you achieve a much smaller loss due to management practices that could make the difference between 50-60% loss and bring it down to 15-25 % loss. Without treatments.

The bottom line is that many beekeepers do not do the proven things that give small advantages. How many of the mainstream beekeepers do mite counts? Hw many requeen with known stock every year? How many rotate/replace comb? How many pay attention to the small details that perhaps would improve the overall survivability and improve honey yields?

The industry is fragmented. The industry is going a hundred different directions. I can read articles and advertisements from 10 years ago that have breeders stating the "resistant' stock they claim to have. In many ways, we are no further along.

One thing I mention from time to time is the price and quality of bees. If you want to raise quality beef, or have a top notch milk herd, you start with good stock. This can be equated in many other farming commodities. Beekeeeprs are notorious of buying the queens that will make all the difference in thier hives by basing it purely on price. Many will buy a queen just a couple dollars less, without even asking about what they are getting. I see the same clubs order bees every year from the cheapest place they can obtain from.


We want the freedom to keep bees the way we see fit. We want to do as we please, all the while hoping someone comes along with all the answers.

Good luck looking.......

[sqkcrk]

Ian, how long before this type of bee is readily available and widely used by the majority of beekeepers? Do you think that the queens will cost $20.00 each by then? Everything else is going up.

So, in your opinion mites are not getting stronger or was that a poor choice of words? Should I have said that they are being selected for survival?

[Michael Bush]

>Twenty years ago Steve Taber said that what we should do about Varoa mites is nothing, And in 30 years, or so, we will have bees that can exist with varroa mites.

I've heard similar things from other "bee scientists". I think they are right, except that 30 years probably is longer than it would have taken.

>Well we didn't do that did we?

No. We didn't. At least not at first. smile.gif Now I have, except for letting them build their own comb.

Sounds like Bjorn is doing that now.

[Ian]

Mark, they already are avaliable from the queen breeders association that has been working in cooperation with any beekeeper here in the area who takes interest in this idea. I hope I stressed the bees are more "tolerant" and didnt imply they are resistant. There are so many factors and characteristic that influence mite tolerance that we are just brushing the surface. These bees tend to show an increased grooming behaviour over all other stock tested along side them. What does that mean? Well, I think it mean we are working in the right direction.

3$ for a cell, 20$ for an open mated queen.

These guys arnt alone you know. There are many breeders out there working towards this common goal, beekeepers alike.
We got to quit being so critical of the work done here and support it. Anyone who doesnt live up to their claim will be taged with it, just the same as it would happen with anyother service provided.

[sqkcrk]

Oh, I wasn't being critical, was I? I was just looking for some light at the end of the tunnel of dispare caused by Varroa destructor.

Is Medhat Nassar working in your area? I like what he has to say. He seems to have it together in regards to research and practical beekeeping. I think so anyway.

I wish I could get some of those queens down her in NY.

With all of the extra time I will have, not working for the state, I'm going to get my own queen rearing program going. I'd like to wean myself off of migratory beekeeping, too. If I can. Time will tell.

[Ian]

>>I'm going to get my own queen rearing program going

cool,

dont know Medhat Nassar,


>>what we should do about Varoa mites is nothing, And in 30 years
>>I've heard similar things from other "bee scientists". I think they are right,

To stand back and watch our stock die off is purly a waste of our beekeepers equity, searching for just one trait amungst the so many to be selecting for. We can find these traits, exploit them to propaget them through out our existing stock. Otherwise we would loose so much of the already developed traits in stock selected for other dieseases.
Having a line that is alone the way of resisting AFB, killed off becasue there they have no tolerance to varroa would be very discourageing. Not to mention all the beekeepers lost.

[Joel]

I greatly respect the success MBis having. I believe though Ian is right on target. Taber is making the assumption that nature will take it's course. As we have discovered with many species this Nature does not always bring about survival.

This brings us back full circle to the main coefficient in this problem some of us continually ignore. We are trying to attain natural selection in an un-natural environment that will constantly have pressure towards pest and disease to reduce the un-naturally large populations of bees we keep confined in any given area. In addition we make the assumption that survival will occur as a natural process. Many millions of species have gone extinct in the history of the world due to both natural and un-natural pressures of which both are very high for honey bees.

Natural selection is one tool in reaching a better level of control. Treatments will be another. In some isolated cases we may see success. However whenever an outside influence is introduced into a population (better mites, resisitant disease, genetically different stock) the pressure will be towards change, not necessisarily towards the positive.

[ February 12, 2006, 09:33 AM: Message edited by: Joel ]

[naturebee]

Mark Writes:
-Twenty years ago Steve Taber said that what we should do about Varoa mites is nothing,,,,

Mark, I would love to have a link to this reference or similar type references if they exist.

Mark Writes:
-That's what we need. Some renegade beekeepers to show us the way to our next meal.

I’ve been doing assessments of ferals caught in remote locations next to domestics. For me, to run a varroa test out several years is not practical and I have found it not really necessary, as fit genetics will rapidly out perform right from the start. All my feral swarms are assessed in single deeps from time of capture, and bad performers culled throughout the season. Only the best performers (which usually averages about 50% of the swarms caught per season) are kept. These are then wintered into next seasons spring buildup when the initial assessment will be completed sometime in May.

Results for me so far are very encouraging, as I am seeing that the remote ferals, especially the remote woodland ferals are outperforming all other swarms on a consistent basis. These woodland ferals are assessing at a level of performance higher than that even of my average survivor colonies. This necessitated the decision to systematically destroy most genetic lines suspected of being from commercial bees and non acclimated stock, as these colonies were under performing in comparison to the woodland ferals. This season I will be only accepting swarms from areas that are remote woodland and farmland areas, and refusing swarms that are found too close to other beekeepers.

The exciting potential of remote feral bee colonies for Varroa coexistence
http://www.beesource.com/pov/wenner/varroaabstract.htm

[sqkcrk]

Pcolar, the link is in my head, my memory bank. I believe that I read it in ABJ. I have heard Steve Taber talk, but not for ages. I'd love to go to the queen rearing course in SC, but it costs $150.00 and is right in the middle of when i should be raising my own and making splits. So, sunday is usually my only day off.

[Joel]

Pcolar, how are you preventing the ferals from back breeding to yours or other commercial stock in the area? Am I understanding you to say you are concentrating not on catching swarms but catching woodland swarms? Are you seeing any particular traits (coloration, honey production, grooming etc.)consistant from these ferals. How long have you been working on this?

[Ian]

>>Only the best performers (which usually averages about 50% of the swarms caught per season) are kept.
>>remote woodland ferals are outperforming all


What exactly do you mean when you say they are outperforming all the rest?
Narrowing down to varroa tolerance is one thing, but varroa tolerance incorperated with all the other desired traits is what is trying to be achieved here.

We can find a varroa resistant bee, but if it doesnt produce the honey we are familliar with, we will go broke all the same,...

[ February 12, 2006, 07:18 PM: Message edited by: Ian ]

[Jonathan Hofer]

Hello Ian

I am from Manitoba, and am continiously looking for more varroa tolerant bees. Whom do I contact here?

Jonathan Hofer

[tecumseh]

joel sezs:
Natural selection is one tool in reaching a better level of control.

tecumseh replies:
seems to me natural selection and control are not terms that go hand in hand.

I do believe this question(s) is quite a bit like afb resistance and jared diamond's discussion on nut bearing (almond I think) trees. both are subject to two allels that come together in the individual's genetic code. for afb one allel that programs for cell cleaning and one for removal. in the diamond illustration two allels that determines that the nut is not bitter. as diamond might suggest (and mathmatical experience would rubber stamp) any more than the two allel requirement means that probablilites for a successful combination become incredible unlikely. once the perfect combination is obtained (isn't this the bases of the russian and buckfast bees???) an individual is unlikely to recongize this exceptional queen and the heavy odds are that when 'this perfect queen' is replaced her replacement will be no more perfect than the average queen.

[Ian]

Contact the extention office at the U of M,
they will direct you to the Queen Breeders Association here in Manitoba.

[Michael Bush]

>any more than the two allel requirement means that probablilites for a successful combination become incredible unlikely

Unless all the bees without that combiniation die. Which will never happen as long as we keep propping up the ones without those properties necessary for survival.

[TwT]

are there areas where the mites don't kill the hive's and coinside with the bee's or does my father have mite resistant bee's? Dumb question I know but my father (and I have said this before in other post) has 1 hive going on 12 years old and 4 other hive's going on 9 years old... I seen mites on some of the drones last year, he the type beekeeper that doesn't talk or know anyone that keeps bee's (except me now)and he's one of those that looks at the hives entrance 1or more times a week to see if they haven't left (so he says), he know's nothing about treating hive's until I told him last year and he said they will be ok with out those chemicals and he wasn't going to use them, if they die they die, (OLD HARD HEADED FART),, he has regular foundation and only takes some honey every now and then, he uses these hive's for his greenhouses and garden. he lives in Denham Spring Louisiania. just something I thought I might ask the resistant posters here......

[Ian]

>>OLD HARD HEADED FART

Ha ha!! sound like my father! smile.gif

[naturebee]

--Pcolar, how are you preventing the ferals from back breeding to yours or other commercial stock in the area?

Good questions! I went thru my colonies and requeened all queens that were suspected as being domestic. The plan this season is to transport mating nucs to remote woodland locations for mating, or mating in my ridge apiary as in the past which is a relatively isolated area. I believe that you do not necessarily need complete isolation to insure a majority of fit feral matings, all you need to do is tip the scale in favor of the ferals. Studies have shown that ‘fit’ colonies such as that found in the wild produce males which not only out-competed males from other colonies in mating, but also has sperm that was more successful in post-mating competition. The fit colonies produced drones with a higher reproductive success than those of the other, less successful colonies.

--Am I understanding you to say you are concentrating not on catching swarms but catching woodland swarms?

Baited traps in wooded areas is my strategy.

--Are you seeing any particular traits (coloration, honey production, grooming etc.)consistant from these ferals.

What some describe as a wild Itialian is what the queens tend to look like. A bit darker, and darker towards the tip of the abdomen. Workers look like dark Italians, a bit more propolis than Itialians do, and I have some ferals exhibiting a high degree of all grooming. They tend to build up earlier in the spring, and I am also pleased by the gentleness in these ferals. Brood patterns in these trapped woodland ferals are nicer than I have ever seen with excellent out mating! This indicates to me that there is a healthy viable population of ferals existing in these woodlands. I don’t use any treatments on these bees.

--How long have you been working on this?

I’ve been catching ferals for over 10 years. The last 5 seasons is when the best ferals started showing up. I’ve been noticing better ferals from remote farmlands and such, so I decided to place some traps last season in the woodlands and these swarms are outperformed all others, so I decided to start focusing my efforts on woodlands.

Opps, I forgot to add, I created a list a while back if you or anybody is interested. I have it unpublicized because I don’t want folks joining just to join, it's called 'Feral Bee Project'. You must have an interest in catching feral honeybees, acclimated bees to join. The list is designed to explore methods of trapping a accessing remote ferals. Send me a PM if interested.

[ February 13, 2006, 07:47 PM: Message edited by: Pcolar ]

[naturebee]

--What exactly do you mean when you say they are outperforming all the rest?

Hi Ian,

According to the dictionary:
Outdo, outmatch, surpass, exceed, To be or do something to a greater degree; "her performance surpasses that of any other student I know"; "She outdoes all other athletes"; "This exceeds all my expectations";
"This car ( or colony of bees) outperforms all others in its class"

--Narrowing down to varroa tolerance is one thing, but varroa tolerance incorperated with all the other desired traits is what is trying to be achieved here.

Good point! These woodland ferals seem to have most desired traits, they are good honey producers, very gentile thank goodness and good wintering and early build up being the most obvious here.

--We can find a varroa resistant bee, but if it doesnt produce the honey we are familliar with, we will go broke all the same,...

This is an excellent point you make! I’m not looking specifically at varroa tolerance here. What I want above all is good health OR fit genetics, and with fit genetics in ferals that fend for themselves comes varroa tolerance and other essential and necessary survival traits. When a fit and healthier bee, is achieved, then and only then will you will get productivity. So fitness come first for me, then I can breed and enhance traits that I believe might be lacking thru selection.

Assessments made in a very dry season last year with 4 swarms caught in towns and residential areas and 3 woodland ferals revealed in August the 4 swarms caught in towns on the verge of starvation and 3 woodland ferals outperforming with sufficient stores and superb brood patterns, so I need to look into this further. I’m not going to reveal just yet what traits I suspect are the cause for the remote ferals outperforming because first I want to try and develop some assessment techniques that I can use to determine just way they are foraging so well and also try and find some existing research that might back up what I am seeing. I will say that Bro Adam is only one of a few that I hear mentioning this and other traits that might be occasionally neglected in much of the bee breeding community.
Best Wishes,

[thekeeper]

I have found that the most mite resistante bees are the MN hygentics. We will never get ride of the pest. The best thing to do is keep them under control.If you wipe them out you will eventually get an infestation and have a treamendous loss in bees. I use the OA treat and founs that it works best. They dont become ressistant to it and it works on the female mite by suffacating her in the bee. forces her out before she lays her larva and forces her to drop.Best stuff I ever used. The fomic pad works the draw back there is it doesnt work under 60F and kills all eggs larva and steralizes the queen. If it goes over 70 the effects are catastrofic to the hive.

[Kieck]

"They dont become ressistant to it...." -Thekeeper, concerning OA treatments

This topic has been argued before on BeeSource. While we may not have documented resistant mites yet, that doesn't mean that mites could not become resistant to OA. Like any other selective pressure, mites could develop some form of resistance to OA. The greater the pressure, the more rapidly resistance spreads through the remaining population. In other words, if every beekeepers switches to using OA to control Varroa, resistance can be expected to "develop" and spread through Varroa than if only a few people use OA. That's the theory behind alternating treatments (such as Apistan and Checkmite+, as well as other treatments). The chances that mites that are resistant to Apistan will also be resistant to Checkmite+ are greatly diminished, and the chances that mites resistant to Apistan and Checkmite+ will also be resistant to (fill in the blank) are equally small. Rotating treatments will help.

[Aspera]

Pcolar, how do you know that you're not catching Bjorn's escaped Russians, or somebody's wayward SMR's. If you buy an SMR mated to russians drones, you will not see much varroa around that colony. The USDA selection programs work well, and probably faster than natural selection.

[Ian]

>>These woodland ferals seem to have most desired traits, they are good honey producers, very gentile thank goodness and good wintering and early build up being the most obvious here.


Looks like you have a good bee there Joe. I dont like reading between the lines when claims are being made. Breed those queens!!!

>>mites could develop some form of resistance to OA.

This is an issue of conterversy. First off I think we first have to agree with what actually it is about the Oxalic Acid that kills the mites. Then we can better understand how the mite will adapt to the leathal exposures of the OA treatments.
If it kills the mites by physically damaging the mite, how quick will that mite evolve to tolerate the exposure to survive OA treatments?

[Ian]

>>When a fit and healthier bee, is achieved, then and only then will you will get productivity

Your still talking about v mite tolerant bees here, being the reason of a healthier bee, right Joe?

[Kieck]

"This is an issue of conterversy. First off I think we first have to agree with what actually it is about the Oxalic Acid that kills the mites. Then we can better understand how the mite will adapt to the leathal exposures of the OA treatments." -Ian

To a point, right. Like you wrote about the source of the genetics, Ian, don't read between the lines.

If we understand HOW oxalic acid kills mites, we can hypothesize HOW different mechanisms might impart some resistance in the mites to OA. Now whether or not it will happen, but how it might happen.

"If it kills the mites by physically damaging the mite, how quick will that mite evolve to tolerate the exposure to survive OA treatments?" -Ian

This is the second part. How quickly resistance "develops" and spreads in mites to OA treatments depends only on the strength of the selective pressure, not on the mechanism. Unless we expect that a treatment will eradicate a pest (and, really, when has such a strategy ever worked in human history?), we have to respect the power of survival in pest organisms. Somehow, some way, mites will become resistant to the treatments, regardless of how the treatments work. That brings up the first question again (how might resistance work?), but figure that it will happen.

Otherwise, if we believe that mites couldn't become resistant to a "physical" assault, we're contradicting ourselves on this tread. If you believe that organisms can't develop resistance to physical influence, then honey bees couldn't develop resistance to Varroa mites. If honey bees can't develop resistance to Varroa mites, why are we wasting our time trying to breed Varroa resistance? (Even "small cell," by the way, can be counted as a mechanism of resistance in bees to Varroa.)

[George Fergusson]

>How quickly resistance "develops" and spreads in mites to OA treatments depends only on the strength of the selective pressure

For what it's worth, mites are particularly good at developing resistance because of the way they breed. Most mites inbreed- a mother mite produces a son and some number of daughters and they mate. Only when 2 or more foundress mites enter the same cell is there a possibility of a daughter mite mating with a male other than her own brother. Hence a foundress mite's genes are passed on directly to her offspring. If she happens to be resistant to a treatment, her children will be resistant to that same treatment. This is why resistance to certain chemicals happens so fast with varroa.

[Bill Ruble]

How long would it take a mite to become resistant to being squashed? How many mites would it take befor they became too hard to squash? And of all the squashed mites, non of them survived to breed, now could this happen? Read between the lines to get to where I am going with this.

Bill

[George Fergusson]

Well Bill, if you've got an IPM strategy worked out around squashing varroa mites (without flattening the bees) I'm all ears! Sounds labor intensive!

Now I'm sure that's not what you're talking about, but I'm notoriously bad at reading between the lines, I much prefer it when people state in no uncertain terms what they're driving at! I'm thinking you're suggesting that certain "physical" actions such as squashing, burning, dismemberment, freezing, drowning, electrocution, etc., are treatments with a mode of action for which no manner of resistance can be developed.

I agree with that to some extent, it's hard to imagine a mite developing resistance to getting squished but for what it's worth, none of the varroa treatments discussed thus far quite fall in the category of "a rock and a hive tool". So I can't rule out the possibility that varroa mites may some day, some how, develop resistance to some forms of treatment with so-called "physical" modes of action.

Part of the problem is the fact that nobody has figured out how Oxalic acid kills mites, though there is no shortage of speculation, some of it well-informed, some of it not so well informed. Until the mode of action of OA is known, talking about whether or not mites will develop resistance to it is fun, but not very helpful.

CSI: Where are you when we need you?

Squashing incidentally doesn't work with deer ticks, wood tics, or fleas. You've got to actually place them on a hard flat surface and press down on them with a knife blade or something similar to get them to pop open. This just isn't practical smile.gif

[Finman]

During decades varroa arrived from Siberia to Europe and to other places. But in what place varroa has dropped national honey yields?

In USA annual honey yield is almost same however some write that half of hives have died.

Yes - in my country Finland 30% beekeepers went away when varroa arrived. Still total yield is at same level.

What it means: Varroa attached in those hives which had minimum care. Now it is much more easier to nurse bees when those "national&rural" bees do not mix good bees's matings.

It is nonsence to speak that chemical control is bad thing. It is so easy to wise in that way but it has no reality. In every country someone try to rise "varroa tolerant" bees but how many has succeeded.

[naturebee]

--Pcolar, how do you know that you're not catching Bjorn's escaped Russians, or somebody's wayward SMR's.

Although SMR are nothing more than hygienic bees as explained by Marla Spivak, I don’t know that they are not escaped SMR’s. But I have more evidence to suggest that they aren't than I do that they are. But how do the "smr" breeders know that their queens arent gaining smr traits from wayward ferals? The smr trait must have existed in the honeybees to begin with in order for it to be developed thru selective breeding.

But it would seem contrary to an assumption that they are escaped SMR when you look at discussions by Wenner suggesting a potential of remote feral bee colonies for Varroa coexistence.
http://www.beesource.com/pov/wenner/varroaabstract.htm
And other studies suggesting fitness in feral colonies etc. Believe what you choose to.

I am in the business of bee removal, and I assess most swarms that I catch. In areas near to other beekeepers where one would expect to find varroa resistant bees and such. I find that these bees simply do not perform as well as the remote ferals. Of course, I will not argue that swarms from these beekeepers may have for some reason NOT settled in the prime lowlands where forage is good, and voids are abundant, but instead for some reason choose to fly 2 or more miles into remote woodlands to find a nest site.


--you will not see much varroa around that colony. The USDA selection programs work well, and probably faster than natural selection.

Well, consider that an estimated 90% of the feral population was wiped out by 1995 leaving only the varroa resistant ferals, and the USDA still hasn’t bred a varroa resistant bee yet by that date. Who was faster?

Remember that by the USDA importation Russians you mention above as from a USDA selection program. How can the USDA is claim credit for breeding a varroa resistant bee when the Russians that they have imported have had over 50 years of natural selection in Russia’s far east.

[ March 12, 2006, 02:57 PM: Message edited by: Pcolar ]

[naturebee]

--Your still talking about v mite tolerant bees here, being the reason of a healthier bee, right Joe?

Hello Ian,

Varroa tolerance is an essential characteristic or trait as wintering ability etc. is. But as in the essential trait known as wintering ability, it is not the only reason for a healthier bee. Yes, varroa tolerance one essential trait of many that should be included the over all equation for a healthier bee. But in order to establish the foundations towards developing a healthier bee I believe it is essential to start with the over all fitness or fecundity of the colony, which according to Brother Adam “is an essential prerequisite for any exceptional performance“. Only after fecundity has been shown in a colony, can we go on assessing desired traits. In other words, you cannot get healthier bees from varroa tolerance alone. Over all fecundity as reflected in essential traits affecting fitness seems to be a determining factor as far as health of the colony goes.

Research shows that varroa tolerance is determined by fitness at the colony level. For instance, daughter queens from the most excellent queen determined to be varroa tolerant may show all the genetic markers for varroa tolerance. But, the colonies degree of varroa tolerance and subsequent survival will be affected if the daughter queen lacks in traits that influences fitness and fecundity, which can only be achieved thru mating with that of fit and non related genetics. Research shows that colony’s phenotype thru multiple mating is a direct reflection of the tasks performed by its workers and subsequent fitness of a colony, and has a significant effect hygienic behavior. Therefore, it might be better to say that colony fitness or fecundity is the reason for a healthier bee.

Research has found a significant relationship between many fecundity characteristics and most colony measures of fitness specifically during the growth phase of colonies. I am also seeing a more rapid growth rate and good brood viability and other traits affecting fitness during the colony initiation and development stage more often in the isolated feral types. This is why I am focusing the assessment of all ferals during the colony initiation and growth stage (about 18 weeks) and then on thru till spring buildup.

Joe Waggle
Feral Bee Project:
http://groups.yahoo.com/group/FeralBeeProject/

[ March 11, 2006, 01:43 PM: Message edited by: Pcolar ]

[Ian]

What would you say determines a "fit bee"?

[naturebee]

Hi Ian,

Sorry for such a long post. But this is my favorite topic.

Fitness is the capability of an individual of certain genotype to reproduce. Fitness is manifested through an organisms phenotype. A honeybee colony’s phenotype is a reflection of the tasks performed by its workers. So when we are evaluating colonies for brood viability, winter hardiness, disease resistance and other factors in the area of fecundity (traits that are known to enhance the health of a colony and it’s subsequent ability to successfully compete) we are evaluating the colonies fitness.

The fitnesses of different individuals with same genotype are not necessarily equal, but depend on the environment in which the individuals live, for example, bees from the south may not do as well in the northern climates and therefore are less fit for the environment in the north and this will be reflected in mating success. As differences in individual genotypes affect fitness and subsequent mating success, then the frequencies of the genotypes will change over generations; the genotypes with higher fitness become more common. This process is called natural selection, and fitness of the phenotypes of colony determines the direction natural selection will take.


I am using much of this information in developing swarm assessing techniques.
I’ve done much research in this area because I believe it very important for effectively assessing feral swarms and weeding out the duds. Here are some quotes that I’ve collected from researchers pertaining to fitness in honeybee colonies:

Fecundity:
There is a significant relationship between many fecundity characteristics and most colony measures of fitness during the growth phase of colonies (Tarpy & Page)

Disease Resistance & Winter Hardiness:
Increased genetic diversity has a direct influence on task diversity, disease resistance and other factors determining colony fitness (Oster and Wilson) and provides a buffer against fluctuations in the environment (Crozier and Page).

Brood Viability:
Multiple mating promotes colony fitness by lowering the probability that the queen will produce a high proportion of unviable diploid males within her brood (Tarpy & Page).

Colonies with a high level of polyandry will have a substantial fitness advantage because of differences in growth rate during colony development (Cole & Wiernasz).

Worker population effects a colonies fitness because a larger colony is able to collect more nectar and store more honey during the active foraging season, thereby increasing the food reserves that are necessary for it to survive the winter (Seeley).

There are significant correlations of brood viability to winter survival (Tarpy & Page).


A colony’s phenotype is a reflection of the tasks performed by its workers and a significant concave relationship exists between brood viability and worker population . Therefore, worker population is an important indicator of colony fitness and is arguably the best variable to distinguish non-linear effects of brood viability (Tarpy & Page).

Drones of a ‘fit’ colony not only had a higher probability to mate with a queen but
also the number of offspring sired in the mated queen was higher than that of its competitors. As the overall male fitness of a colony is the product of its mating
success (the number of drones mating successfully) and the average siring success of these drones the differences between the drone colonies are even enhanced. The fit colonies produced drones with a higher reproductive success, while weaker colonies are known to cease drone rearing altogether. (KRAUS, NEUMANN, SCHARPENBERG, VAN PRAAGH, MORITZ)

[MountainCamp]

I have refrained from getting into this thread as I did not want to repeat the How High Can Bees Fly thread of last year. - www.beesource.com/cgi-bin/ubbcgi/ultimatebb.cgi?ubb=get_topic;f=13;t=000171;p=9 (http://www.beesource.com/cgi-bin/ubbcgi/ultimatebb.cgi?ubb=get_topic;f=13;t=000171;p=9)

So I will apologize to all for jumping in.

However Joe, Pcolar, or NatureBee, I have the same problem with your assertions in the thread as I did with your comments in that one. You have changed your name, but not your ways.

You play very lose with facts; make assertions based on previous assertions given as facts.

We went back and forth on your claims and assertions, but you never answered the questions posed to you. You ducted, moved, mis-directed and provided the same double talk as here.

Last year you were an Organic Beekeeper who found the light in “regressed small cell” bees. Who’s jobs, if I remember correctly, finally came out as a musician and teacher?

Now you claim that “I am in the business of bee removal, and I assess most swarms that I catch.”

You talked of how you never got a honey crop, except from a hive or two. You said that you had to move hives to the farm land from your woodland yard so they could forage.

Now, a year later your “remote” woodland yard, just outside town in Pennsylvania has super feral bees that are out performing everything in sight.

Unfortunately only half of the story remains as you deleted most of your posts on that thread.

Well, I will be wearing my boots the next time I read this thread.

[ March 12, 2006, 07:49 PM: Message edited by: MountainCamp ]

[naturebee]

Thanks for your input mountain camp, You’re a sweetie.

Yes, I am on small cell and am having success in keeping bees without treatments. Most of the past years were spent regressing bees. And you either build comb and bees or make honey, it’s difficult to do both. Much of this time was also spent assessing my existing colonies and replacing with outperforming ferals that I catch and not concentrating on gaining surplus, but with only a few colonies.

I have been doing bee removals for over 10 years, I’m a musician and do other jobs and frankly that’s my personal business what I do!

I did move hives from the woodlands to the farm lands to forage and I am now getting good surplus in these areas. But here in the woodlands I have found an excellent spot to assess ferals as the poor genetics will fail and good will actually obtain some surplus even though forage is sparse here. This makes for an excellent location to assess swarms.

I routinely assess all my ferals and have for 10 years or so, but Only in the last year have I went out with trap placements in remote woodland areas and started assessing these ferals along side the ferals I catch form my routine bee calls in town and such. This is an evolving process, and the performance of the woodland ferals is fascinating to me and all I wish to do is share it with others and discuss the issue.

I am here to simply discuss the issue and hopefully gain some insight from others about ferals and ways to assess them and breeding techniques. The title of this thread was breeding varroa resistant bees which is why I got involved in seeing the woodland bees coping with varroa. You can go ahead with your personal grievance against me, but I really wish we would stick to discussing bees as the list is intended to do.

[ March 12, 2006, 09:24 PM: Message edited by: Pcolar ]

[naturebee]

Mountain Camp,

I am enjoying the talk with Ian which is coming along very well and polite I believe. Ian asked for a description of fitness and I did my best to provide information. I wasn’t trying to deceive anyone, just providing the info requested to the best of my ability. I enjoy this topic and am glad that Ian is interested also to engage in productive dialogue.

I'm not selling anything, it makes no difference is somone disagrees with me, I want to here differing opinions because of the potential to help me in what I am doing with the ferals.

What I am saying is that honeybee colonies as a prerequisite for exceptional performance, they must score high in the areas of fecundity and other traits concerning fitness. There are out performers in every apiary that beekeepers should breed from. Outperforming is defined as a colony that performs above the rest, there is no assumption that this designates super bee status, but simply that it is performing in the top of the pac. That I am seeing this in some of the remote type ferals is no different than others that see outperforming within their own apiaries. That some breeders produce lines of bees that outperform others is no different. My point is as long as fecundity and other traits affecting fitness in any colony, they have the foundations for exceptional performance this according to Brother Adam and other supporting research.

[ March 13, 2006, 06:01 AM: Message edited by: Pcolar ]

[Kieck]

I know I'm a little behind the current direction of the thread here, but I wanted to address the issue of resistance to "physical" methods of control a little further.

"How long would it take a mite to become resistant to being squashed? How many mites would it take befor they became too hard to squash? And of all the squashed mites, non of them survived to breed, now could this happen? Read between the lines to get to where I am going with this." -Bill Ruble

Let's see, George already pointed out examples of ticks and fleas -- how about cockroaches and house flies? Both roaches and flies are "squishy" enough that they can be easily squashed, but how easy is it to do? Try it sometime. Do effective methods of escape (such as reflexes that allow roaches or flies to detect air movements and avoid "squashings") count as "resistance" to you?

I've talked to several evolutionary biologists (the type of people who study these sorts of things for livings) about the idea that organisms can only develop resistance to chemical methods of control, and not to physical methods of control, and not one of them that I talked to agreed with that opinion. Seems as though some beekeepers believe it, but the experts in the area of evolution don't.

So once again, even for those beekeepers out there who believe that resistance to physical attacks can never develop in Varroa mites, if organisms can not develop resistance to physical attacks, why should we even bother talking about breeding resistance in honey bees to Varroa?

Now about this fitness issue, I'm not sure that beekeepers are keeping bees with the highest "fitness," unless the selective pressures are entirely in the hands of the beekeepers.

"What I am saying is that honeybee colonies as a prerequisite for exceptional performance, they must score high in the areas of fecundity and other traits concerning fitness. There are out performers in every apiary that beekeepers should breed from." -Pcolar

Some of the most fecund bees are also the least productive in terms of building large colonies or storing large amounts of honey. AHB, for example, are particularly fecund -- they swarm frequently, and swarming is a consequence of fecundity -- yet few if any beekeepers find those traits particularly useful. Perhaps that's why we have so many problems with the bees we do keep -- we try to keep bees with relatively low natural fitness, then complain when these unfit bees become hosts to all sorts of problems.

[Hillside]

It's easy to fall into the trap of thinking there is one thing called fitness. Fitness always has to be considered in context. Fitness for cold survival, fitness for long distance flight, etc.

A given bee has many different levels of fitness. There is no super bee that is well fit for everything. Although, we all hope that such a bee exists and that it will show up in one of our hives.

[naturebee]

Thanks Kieck for the productive discussion!

Great insight! I see what you mean. are you a bee breeder? I do agree that that a colony can be fecund and not have the essential traits to be productive. When assessing ferals and swarms of unknown origin, for the moment I am going on the assumption that it I need to assess the colony for fecundity during the initiation and growth stage which (not sure which researcher said Tarpy of Page) is easily assessed during these 18 weeks of growth. Then as Brother Adam mentions after this prerequisite of fecundity has been identified in a colony it can be further assessed for traits of economic value.

The reason I post these things to stimulate conversation on the matter is that I am seeing wide range ofperformance of these ferals during the initiation and growth stage. It’s fascinating how some are complete duds and some swarms show brood viability and other traits that rival that from the best breeders in the country. We had a good drought last year in my area and the ferals with good brood viability seemed to be affected very little by the drought, and others were nearing starvation. I’m attempting to develop some kind of assessment chart that I can coordinate with the locations that the swarms originated and see if I can get some kind of idea what’s going on here, or if a pattern develops. I'm learning as I go, but I have to do something.

[naturebee]

Thanks Hillside, excellent input to the discussion!

I see your point that fitness is relative to the environment or context. Sorry for speaking it in such a general term. But it is interesting you mention fitness for long distance flight. I assessed about 10 ferals last season. I assess ferals in singles and also winter them the same way as part of my assessment process, no treatments and no feeding and end up eliminating on average about 50% along the way due to sub standard performance.

During the summer here in the ridge where I assess them, the nearest good forage after the tree bloom is probably 1.5 to 2 miles away and the lack of forage in the summer due to the area around the colonies being trees makes it that much tougher for a small colony during the dry summer months here in the ridge. The woodland ferals for some reason did much better collecting stores during the dry time. Do you think it is possible that the woodlands may have better traits for long distance foraging? I know that bees forage 5 miles or more, but most of the forage is done probably within a mile and a half, long distance foraging would certainly give a colony an advantage in the woodlands, what do you think here?

Great point that there is no super bee fit for everything! I agree! When I say a colony or swarm is outperforming, I am meaning it is among the top 10 maybe 20 percent in its class. I don’t mean to elude that they are a super bee, just doing well enought to grab a persons attention.

[Ian]

>>bees from the south may not do as well in the northern climates and therefore are less fit for the environment in the north and this will be reflected in mating success

Would this not be expressed along the wintering traits? I doubt the reason the southern bees winter poorly in northern area becasue they are less fit than the northern bee.

If you ment fit for the area, then I dont understand. You mean adapted for the area, right?

>>(quote)Worker population effects a colonies fitness because a larger colony is able to collect more nectar and store more honey during the active foraging season, thereby increasing the food reserves that are necessary for it to survive the winter
>>quote>>worker population is an important indicator of colony fitness and is arguably the best variable to distinguish non-linear effects of brood viability

There are so many different variable with worker population that I cant see how this fellow could corrilate larger colonies to fitter bees. And then corrilating it to winter better becasue it stores more food.

Perhaps I am missunderstanding the quote, and the logic, but tell me, are my five frame increase nucs less fit than my booming honey production hives?

Like I said, there are many varriable to wintering bees up north, strengh being ONE of them.

[naturebee]

----->
Would this not be expressed along the wintering traits? I doubt the reason the southern bees winter poorly in northern area becasue they are less fit than the northern bee.
----->

Thanks for having patience with me as I am still learning, but I will explain as best as I can please someone correct me if I am wrong.

According to the dictionary for biological terminology the components of fitness are traits that contribute to survival & reproduction. They also say that fitness it relative to environment, therefore bees that are fit for the north may do bad in the south because they are not fit for a southern environment and vice versa. Ether way, well bred north and south bees would both be fit providing they are in the environment that they have adapted the traits needed to survive. If they are in a climate that they have not adapted fitness traits too, they would be less fit compared to the bees that are adapted in the environment.

----->
There are so many different variable with worker population that I cant see how this fellow could corrilate larger colonies to fitter bees. And then corrilating it to winter better becasue it stores more food.
----->

That one was from Seeley. I guess he means that the larger colonies would store more food and have less chance of starveling to death. I guess Seeleys saying colonies that survive winter are fitter than the dead ones that might run out of stores. I can see a second advantage, and that would be more stores for spring build up and enable more drone rearing which would contribute to the reproduction component of fitness.

----->
Perhaps I am missunderstanding the quote, and the logic, but tell me, are my five frame increase nucs less fit than my booming honey production hives?
----->

Good question. The logic is strange but since a component of fitness is reproduction, the larger colony would have more mating success by being able to produce many more drones, and by the rules of fitness it would be more fit. Your right Ian, it is strange logic because a five frame nuc could contain excellent genes that might not be reflected in fitness of mating success until the colony is allowed to mature a bit.

----->
Like I said, there are many varriable to wintering bees up north, strengh being ONE of them.
----->

How you are able to winter bees up there I’ll never know as my winters are tough but must be mild compared to yours. Have maple starting to bloom here now.

Best Wishes,
Joe

[db_land]

Hi Joe,
I also do a lot of removals (mostly in town but occasionally out in the 'boondocks'). Most of these "feral" colonies maintain small, very frugal clusters during the winter, but build rapidly in the spring. I've also observed that over the past 3 years my removal call rate has approximately doubled each year and that the colonies are smaller (rarely over 3 lbs). All of this is anecdotal, but seems to suggest small, frugal, fast build up, frequently swarming colonies are out-competing (at least in the central NC environment) larger colonies. AHB have these same attributes and clearly out-compete EHB in the survival game. Also, larger colony populations eat more and therefore are more likely to starve during hard times (like during a cold winter after a summer draught). If survival is the ultimate goal of "fitness", then anecdotal evidence suggest that small is better. Comments?
:cool:

[Kieck]

I don't want to turn all technical on you guys -- I've enjoyed this discussion, and hope to keep it going for a while. I know I've learned from it.

Having said that, I think "fitness," at least in an evolutionary context, needs some further explaining. The traditional concept of "fitness" in evolutionary biology doesn't include survival, except as survival long enough to reproduce.

Let me give an example, using humans for simplicity. We take two hypothetical humans: one with a congenital disease that kills her before age 8, and one woman with no chronic diseases who lives to be 95 and leaves behind three children. Which one has greater fitness? The second one, obviously.

Now we look at another two hypothetical humans: one woman who has two children and dies at age 25, and one woman who has two children and survives to age 80. Which one has greater fitness? They seem to be equal.

The problem is that fitness includes all future generations, and the ability to pass genes into future generations, and not just the ability to reproduce.

For example, again using hypothetical humans: 1) a woman has two children, but she dies at age 24; her two children each have two more children, and those four individuals have still more children; 2) a woman has nine children, and she lives to be 90; none of her children have any children. Which one has greater fitness? The first one, even though it isn't as immediately apparent.

Fitness includes direct fitness (producing offspring and future descendents) and indirect fitness (increasing the fitness of related individuals which are not direct descendents, such as caring for siblings or other kin). These two components get lumped into "inclusive fitness."

Adaptation and survival plays into fitness, because, obviously, individuals who aren't well enough adapted to reproduce or can't survive long enough to reproduce have lower fitness.

In the case of bees, fitness directly relates to casting swarms. The fitness of workers lies mostly in indirect fitness, unless those workers become laying workers. Taken as a whole unit (a colony), bees show greatest fitness by producing successful queens, which means swarming.

Again, hypothetically, 1) colony one survives eight years, superceding a single queen four times during the eight year period; the colony has, on average, 25,000 workers at any time during those eight years; 2) colony two casts two swarms during the first year, then dies; those two swarms build small colonies (less than 10,000 workers maximum), swarm four more times each, and die; those eight colonies also remain small, swarm between 2 and 6 times each, and die; and so on. Which hive has greater fitness? The second colony.

But, in a cold climate, if those two small colonies that started as swarms from "colony two" are too small to overwinter successfully and die as a result, then "colony one" would have greater fitness.

It's a complex issue, really, and not so easy to assess in real life as these hypothetical situations. Like db_land says, the evidence of smaller colonies becoming more commone suggests that smaller colonies more prone to swarming really are more fit in naturalized situations -- a greater proportion of small, "swarmy" colonies implies the ancestors of the successful bees were more fit than bees that produced large colonies which rarely swarmed. As beekeepers, we want those large colonies that will produce more honey for us and provide more workers for pollination. So we try to breed and manage bees to suit our purposes. But we don't necessarily select the most evolutionary "fit" bees.

[naturebee]

-- Most of these "feral" colonies maintain small, very frugal clusters during the winter, but build rapidly in the spring.

I’ve seen this small wintering clusters in some of the ferals, it was something of a curiosity to me, didn’t quite know what to make of it. I’m not sure if rapid attrition due to mites is the cause or what, but they do seem to winter just fine with smaller clusters.

--I've also observed that over the past 3 years my removal call rate has approximately doubled each year and that the colonies are smaller (rarely over 3 lbs).

I’m still seeing an average swarm just a bit larger than a football and some bigger. Woodland swarms seem smaller, but this could be a reflection of the size void the mother colony was in. Swarm calls averaged 30 per season till the mites, then went to zero for several years. I have also noticed the calls doubling each year. I think I got about 10 swarm calls and about 5 bee removal calls this year. I have good bosses at work that let me take my leave when ever I want for chasing swarms bee jobs.

--All of this is anecdotal, but seems to suggest small, frugal, fast build up, frequently swarming colonies are out-competing (at least in the central NC environment) larger colonies.

Yes, they seem to come out of winter brooding fast and early. I’m finally back to about 10% winter loss on average, I don’t use treatments so I think that’s pretty good for a bunch of mongrels..

--If survival is the ultimate goal of "fitness", then anecdotal evidence suggest that small is better. Comments?

I agree. Ferals are different and haven’t lost any productivity traits it seems, so the feral strategy seems to be working for them. These ferals you get DB, do the queens have wild Italian type markings? Mine are similar to the markings of the wild type shown on this link, except the black tip extends a bit further up the abdomen. These wild type Italians are surprisingly gentile and a pleasure to work, do you notice this in your ferals? I’m happy when I catch a feral with queens of similar markings because they seem to have similar traits that I like.
http://members.aol.com/queenb95/genetics.html

Kieck, might be able get to your letter tomorrow, the kids are kicking me off the computer. You’ve made some very interesting and though provoking comments and I want to think about is before I respond.

[Walt McBride]

Ian & Kieck, I believe I read that oxalic acid attacts the mouth part of the Veroa mite.
Walt

[Michael Bush]

>--All of this is anecdotal, but seems to suggest small, frugal, fast build up, frequently swarming colonies are out-competing (at least in the central NC environment) larger colonies.

I have ferals and I have no problems with swarming. But fast build up can certainly cause swarming if the beekeeper isn't doing their job.

>Ian & Kieck, I believe I read that oxalic acid attacts the mouth part of the Veroa mite.

This seems to be a nice theory. Heilyser Technology Ltd. Used to have a picture to this effect, but If you looked closely it was obviously two versions of the exact same photo that had been doctored.

As far as I know it's still a bit of a mystery how it works.

[Kieck]

I agree with Michael; I've been looking for real information about the mode of action of OA on Varroa, and the best I've come across seems to show that the pH of the OA is much more important than chelation effects once the oxalate gets into the mites.

"I have ferals and I have no problems with swarming. But fast build up can certainly cause swarming if the beekeeper isn't doing their job." -Michael Bush

Sure, but from an evolutionary standpoint, splits have equal fitness as swarms. If you split to reduce swarming, you, in essence, have done the equivalent in terms of making two colonies (and, more importantly, another queen) and perpetuating the genes.

And, of course, the adaptations that make some colonies more fit than others vary from one part of the country to another. For example, if small, "swarmy" colonies can't overwinter successfully in your neighborhood, I would expect to see larger, less prone-to-swarm, feral colonies.

Are your feral colonies more inclined to swarm (even if you manipulate them so they don't) than colonies from commercial stock?

[Michael Bush]

>Are your feral colonies more inclined to swarm (even if you manipulate them so they don't) than colonies from commercial stock?

Since I manage both to prevent swarming and since neither swarm unless I don't do my management, I'd say it's hard to say for sure. But neither swarm when I do my job. Both swarm if I don't.

[naturebee]

I see your point Keick! Yes, long term survival makes sense as a concept of fitness because a queen living longer would be able to produce more carbon copies of her self in the form of drones for several more years. As fitness is determined by the quantity of the copies of the genes of an individual in the next generation.


--In the case of bees, fitness directly relates to casting swarms. The fitness of workers lies mostly in indirect fitness, unless those workers become laying workers
--Taken as a whole unit (a colony), bees show greatest fitness by producing successful queens, which means swarming.

Fitness does relate to casting swarms. This has been the traditional view so far. Although the main selective force at the colonial level seems to be on the female side thru swarming, some fascinating research from Germany by NEUMANN and co. show that the often over looked male reproductive success appears to be a major driver of natural selection in honeybees. Usually drones 2000–5000 individuals per colony, per season outnumber the queens 2–3 per season. Perhaps, if a colony has to produce enough workers to facilitate reproductive swarming (also a major investment in queens), this might be at the expense of mating fitness on the male side.


---Again, hypothetically, ,,, those eight colonies also remain small, swarm between 2 and 6 times each, and die; and so on. Which hive has greater fitness? The second colony.

Good point. So many variables come in play concerning the mating aspects of the honeybee. Perhaps colonies that are investing energy on swarming are able to invest less in drone rearing, and have higher winter mortality. Also one needs to consider that weak or smaller colonies are known to produce small numbers of drones or abandon drone rearing altogether (Gary, 1992) decreasing mating fitness on the drone side.

In the case of the first colony you exampled that superceded it’s queen several times instead of swarming. Perhaps the mature status of this colony enabled it to achieve size, stability and strength needed to rear more drones and earlier drones than the swarming types causing an equalization of sorts with increased fitness on the drone side.

I agree with you that it is very complex. Hypos are a good ay to illustrate this, because there are so many variables to consider concerning honeybee fitness, that it might be pretty darn difficult to as you mentioned properly assess fitness to any type of accuracy. But I am focusing on ways to return competitive mating on the drone side, I’m thinking this may be achieved by moving mating nucs away from domestic beekeeping to the woodlands where a mating queen has a better chance of mating with fit drones rather than being mauled by hundreds of mediocre drones. Woodland mating I suspect might still benefit from the most fit domestic genetics, as the study by De Jong shows increased flight performance in fit honeybee drones enabling them to reach distant DCA‘s. This year I'm trying a few mating nucs in the woodlands and a few in my yards to assess them side by side to see what happens.
Best Wishes,

[Kieck]

Yeah, drones add a whole new layer of complexity to the fitness issue. Then we have to get into "game theory."

At first glance, raising drones seems to increase the fitness of honey bee queens more than raising new queens. Queens, being diploid, get half their genes from their mothers (ova) and half from their fathers (sperm). Drones, being haploid, get all their genes from their mothers (ova). So, mathematically, queens pass a greater proportion of their genes to future generations by producing drones than queens. That's not so obvious in the "F1" generation, but becomes more apparent in subsequent generations.

So, starting with that, it would seem that queens should produce more drones to increase their fitness. And that makes sense.

The "game theory" enters the picture at this point. What are the odds that a drone will successfully mate and pass those genes on to future generations? Obviously, those odds are pretty low. Think of all those drones that hang around in hives all summer, and maybe never even encounter a mate. Think of all the drones that pursue queens at DCAs and still fail to mate. The supply of queens, after all, is relatively limited. And, then think of the chances that sperm from a drone will be used to produce a queen in the next generation -- if all the sperm from a particular drone is used to produce workers, his fitness drops effectively to zero after that first generation(unless the workers lay unfertilized eggs, and those eggs develop into drones, and so forth; but those chances are pretty tiny).

So, using game theory, are the chances of passing genes to future generations higher by producing queens or drones? Do the higher proportions of genes passed to future generations by producing drones outweigh the smaller chances that drones will successfully pass those genes into future generations?

Of course, all of these speculations still haven't addressed the odds that new queens will successfully establish colonies. And that adds more variables to the issue.

Initially, I jumped in with some comments about fitness because I don't believe -- as beekeepers -- that we are really seeking lineages of bees with the greatest evolutionary fitness. As Joe points out, producing drones make increase the fitness of hives, too, but drones don't do much for beekeepers looking to produce large crops of honey or trying to raise large numbers of bees for pollination.

We can add our own selective pressures that influence fitness as well, though. Beekeepers who cull queens that aren't producing are limiting the fitness of those queens. Breeding only from the most productive stock increases the fitness of those bees. Treating for Varroa mites artificially increases the fitness of bees that might otherwise have such low fitness that they wouldn't leave any genes in future generations.

From a practical standpoint of managing colonies, though, "fitness" is a minor component in my decisions. Obviously, the bees have to be fit enough to perpetuate their genes, otherwise they're gone anyway. But I tend to base my breeding decisions on traits that, really, might lower the evolutionary fitness of bees. I select bees that are gentle (while aggressive bees might be more likely to pass genes into future generations), produce excess stores of honey (while the "most fit" bees should, theoretically, only produce exactly as much honey as they need to survive, and should devote more energy to producing reproductive bees), are less prone to swarming (clearly swarms spread more genes into future generations), etc.

[naturebee]

---The "game theory" enters the picture at this point. What are the odds that a drone will successfully mate and pass those genes on to future generations?

Pretty low for the unfit drones, when think that around 240 colonies on average are represented in a DCA and some estimates are around 12,000 drones visit a DCA daily the odds are low that they will mate successfully.

Hope you don’t mind the trivia, I’m not trying to be a smarty pants, I just enjoy trivia.
Consider the variability that exists with drones, and how fitness might affect them:

Up to 12 days to reach sexual maturity (Ruttner)
Drones experience selection pressure for fast adult maturation (Page)
Average flights 20 to 30 minutes (Witherell)
1-4 flights per day (Currie)
Fly about 3 miles from the hive, rarely do they fly more than 4 miles ( NEUMANN & Apis news letter)
Lifespan is variable 50 to 90 days (Page Fukuda & Ohtani)
Most drones (65%) initiated flight within three days (Pankiw & Page)
Assuming Varroa infestation affects fitness. A mating test flight of varroa partizised drones averaged less then 22 min. (DeJong)
Most of the drones parasitized by Varroa mites during pupal infestation will unlikely be able to reach a DCA (DeJong)
Drones have the ability of high acceleration (10 to 20 m/s2). Either by overtaking or leaving/entering the comet, drones seem to compete for more promising positions (Koeniger & Co)

The wide range of performance might suggest degrees of fitnees playing a part, fit drones would likely do the above more, faster, quicker, longer.

[naturebee]

--Initially, I jumped in with some comments about fitness because I don't believe -- as beekeepers -- From a practical standpoint of managing colonies, though, "fitness" is a minor component in my decisions. ….. But I tend to base my breeding decisions on traits that, really, might lower the evolutionary fitness of bees.

I agree. This is what I am attempting to do by breeding ferals. The ferals would have the fitness traits adapted to my climate, and once fitness is established in my genetics, more selection emphasis can be placed towards traits of economic value.

Breeding fitness is being highly emphasized now a days in the industry with all the talk about hygienic bees, SMR bees, allogrooming bees, autogrooming bees, disease resistance, foulbrood resistance ect. that are all traits that affect fitness. NONE of these traits were invented or developed by any scientist. All traits must have existed in the genetics for them to be brought forward. Many of these traits I am noticing occurring naturally in the feral population, so I am placing emphasis on breeding from the ferals to gain back fitness.

In Game Theory mentioned earlier, this is fascinating when you consider what a DCA might look like according to game theroy. Hypothetically now, consider the for example the big buck after the does, if threes dozens of smaller less fit bucks after the same doe, the big buck has less of a chance of mating even though he is much stronger and fitter by the over abundance of excess competition. Same example with long distance runners, they need to get to the front of the pac out of the crowd of les fit runners to have a chance to succeed. Now take a domestic queen visiting a DCA near commercial beekeeping, the DCA might be filled with thousands of drones that were given artificial fitness thru feeding, treatments, wintering in the south or what ever, many probably related due to common beekeeping practices to some degree. A fit drone in this DCA would have a tough time competing in this DCA, and the queens colony less chance of acquiring fitness thru mating.

If a queen were to go to a remote DCA, she might encounter a smaller DCA with a relatively low population of perhaps a thousand or so drones, but of higher overall fitness per drone due to the majority being feral, and possibly a wider range of genetic variability. In game theory, perhaps less is better here with the remote feral DCA’s because the available fit drones would have a higher chance of succeeding, and overall the queen would achieve a high degree of mating fitness. I’m still looking at this, but this is one theory I have that might explain the excellent brood viability and other fitness traits I am seeing in the remote ferals.

[Kieck]

I think we're still confusing some of the terms in this discussion, at least in the way that the people who came up with the terms defined them.

"Fitness" is determined by leaving offspring. It's measured by the number of offspring left in succeeding generations. Artificial fitness could be imposed on drones by deliberately selecting some for II; otherwise, as humans, we can't give "fitness" to drones. Fitness can't be measured by direct competition. For example, two white-tailed bucks fight, and one wins. Which one is more fit? We don't have a clue. Which one leaves more offspring? That's the real measure of fitness. Fit individuals don't and won't necessarily win head-to-head battles.

Using game theory to evalute drone choices in DCAs could be very complicated, too. Game theory involves how organisms make "decisions." In the case of drones in a DCA, the decision would really come down to "stay" or "leave." If other drones are mating with virgin queens but the drone in question can't keep up at that DCA, he would be expected to leave (hopefully, by moving to a different DCA he might be more likely to mate). If no queens are showing up at the area, the drone should leave. Otherwise, he should stay.

"If a queen were to go to a remote DCA, she might encounter a smaller DCA with a relatively low population of perhaps a thousand or so drones, but of higher overall fitness per drone due to the majority being feral, and possibly a wider range of genetic variability." -Pcolar

Um, see, not entirely. At the time, the fitness of the drones would all be exactly equal. None of them would have mated, none would have left offspring, so the actual fitness at the time of any of these drones would be zero. The "higher overall fitness per drone," in this case, would actually come from the smaller number of drones at the DCA, rather than the origins of the drones. These drones might be less adapted to the climate than some drones in a larger DCA, but since fewer drones would be competing to mate at the smaller DCA, those drones would end up with a higher overall fitness per drone.

I think what you're really looking at in trying to assess these colonies that originated from feral stock is the adaptation to your situation, rather than fitness. "Fitness" gets thrown around as a term a lot, but assessing fitness can be next to impossible. Obviously, if the feral colonies were surviving without human help for a period of time, the most poorly adapted lineages would have been weeded out, and the survivors should be better suited to the local environment. They may actually have much less genetic variability than commercial stock, but the genes that remain are specifically the ones that can withstand whatever environmental conditions the area throws at them.

[naturebee]

---Quote--
--"Fitness" is determined by leaving offspring. It's measured by the number of offspring left in succeeding generations. Artificial fitness could be imposed on drones by deliberately selecting some for II; otherwise, as humans, we can't give "fitness" to drones.
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Most thought provoking point you make!

Please correct me if I am wrong, but isn’t fitness by definition given to drones by beekeepers in many ways? Things that directly affect the number of drones produced in a colony and subsequent fitness in mating success are feeding a colony, adding drone foundation, by splitting colonies beekeepers are giving fitness to a particular line, using treatments. Now, weather these things are good for the genetics is a different mater. But by definition, if it affects a colonies capacity for reproduction, it is a currency of fitness. Even though feeding and such may be as a result of a surrogate to fitness, these things will affect the capacity for reproduction, and therefore giving fitness, albeit through surrogate means.

In this quote by Seeley for example:

“Worker population effects a colonies fitness because a larger colony is able to collect more nectar and store more honey during the active foraging season, thereby increasing the food reserves that are necessary for it to survive the winter” (Seeley).

In light of Seeley’s quote, and obvious assumption that a colonies ability to survive directly affects it’s ability to reproduce (dead things can‘t reproduce).
Would it be correct in the assumption that a beekeeper can have an influence on the fitness of a honeybee colony simply by doing or not doing things that directly affect the colonies survival and or level of fitness and subsequent ability to reproduce? Ie, keeping bees in nucs rather than 3 deeps, breeding, splitting etc?

Other replies:

I agree that a buck winning a battle does not in itself indicate fitness because fitness is determined by the number of offspring.. BUT, a buck winning may be a currency of fitness because by winning, he get’s to hang out with the does.

--quote---
--- The "higher overall fitness per drone," in this case, would actually come from the smaller number of drones at the DCA, rather than the origins of the drones. These drones might be less adapted to the climate than some drones in a larger DCA, but since fewer drones would be competing to mate at the smaller DCA, those drones would end up with a higher overall fitness per drone.
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I see your point, and a good very one! The point I am trying to make is since fitness is directly determined by the “capacity for survival and reproduction variable in space & time; short-term and long-term”,,,

Wouldn’t one expect remote woodland DCA’s (away from domestic imported bees) to contain more phenotypes that possess characteristics of environmental adaptation (a currency of fitness) that are essential for survival in the environment? Am I correct in assuming, a queen mating at a remote woodland DCA would have a higher propensity to mate with phenotypes that are adapted to the climate? And considering polyandry in queen honeybees, by her mating there, wouldn’t this contribute to the fitness of her colony and capacity for survival and ability to reproduce as reflected in the phenotypes in tasks preformed at the colony level relative to other non adapted phenotypes?

Please correct me if I am mistaken, I understand that adaptation is a currency of fitness because it affects the colonies ability to survive and reproduce over the long term. I understand that genetic variability as far as fitness goes is relative to the environment and therefore for this reason you cannot include genetic variability in itself as a currency of fitness. Genetic variability that one may acquire thru successful matings in areas near beekeepers with bees and drones from the south, would not necessarily reflect in the adaptive characteristics at the colony level essential for long term survival and reproduction. So even though matings with these southern adapted drones might be successful and contribute to fitness for the short term, it does not meet the definition of fitness which is “survival and reproduction variable in space & time; short-term and long-term”.

Because fitness is ONLY relative to environmental conditions, mating success MUST have the end result of being beneficial to the survival and reproduction of the colony over the short and long term for it to be considered a currency of fitness. Therefore, wouldn’t successful matings of northern queens to southern drones (even though genetically diverse) not contribute to fitness because they will not be fit for the environment and will not meet the long term requirement?

[Kieck]

I think we're looking at two different viewpoints of fitness related to honey bees. I wrote earlier that humans can't really give drones additional fitness. The problem lies in what scale we look at for "fitness."

If we look at fitness from the perspective of the individual, I don't think beekeepers can really increase the fitness of drones. How could a beekeeper make a drone more likely to mate and successfully leave offspring, really, other than through instrument insemination?

However, I believe the fitness of a colony could be artificially improved by a beekeeper through production of more drones. In this case, viewing the colony as a single entity, additional drones could impart greater fitness to the colony as a whole.

"Would it be correct in the assumption that a beekeeper can have an influence on the fitness of a honeybee colony simply by doing or not doing things that directly affect the colonies survival and or level of fitness and subsequent ability to reproduce?" -Pcolar

I think it does influence the fitness of colonies -- at the colony level again, of course.

"Wouldn’t one expect remote woodland DCA’s (away from domestic imported bees) to contain more phenotypes that possess characteristics of environmental adaptation (a currency of fitness) that are essential for survival in the environment?" -Pcolar

Perhaps. Logically, it seems right. However, one of the points that gets missed in all this is the idea that the environment where your bees are kept is significantly different that the environment where any other bees might be kept, and the bees in either environment are ridgid enough in their adaptations that they gain some level of advantage. Many beekeepers seem convinced that their environments are extreme enough to quickly "weed out" any bees not adapted to the area once swarms revert to living on their own. I'm not sure that most environments are really extreme enough in the U.S. to see such selective force in such a short period of time.

Think about cats, another introduced animal that regularly reverts to feral life. Logically, long-haired cats should be better adapted to winter survival in the northern parts of the U.S. than short-haired cats. If that would hold true, those of us in the north should see mostly long-haired cats wandering around. Yet I don't see that -- the cats I see seem pretty evenly mixed. So, either some other selective pressure is working against long-haired cats, or the selective pressures that would seem to favor long-haired cats in the north aren't great enough to really reduce the numbers of short-haired cats in the same areas.

Same thing goes with honey bees. Can bees from southern stock successfully overwinter in [your] area? Are they about as likely to survive as stock from feral sources? If the differences in survival aren't significant statistically, the selective pressures aren't really working on those abilities.

Overall, I'm not sure what to expect from the ideas that honey bees can adapt rapidly. My gut instinct tells me that, like most insects, honey bees can evolve rapidly to adapt to new conditions. At the same time, the strongest selective pressures (the ones that kill the greatest percentages of individuals in populations) push evolution the most rapidly. From everything I've read and seen, Varroa should be a tremendously strong selective pressure. So, could feral bees really have adapted that significantly to their environments while Varroa resistance among bees lags so badly? Or, is Varroa really not such a strong selective pressure?