Beemaster's International Beekeeping Forum
BEEKEEPING LEARNING CENTER => GENERAL BEEKEEPING - MAIN POSTING FORUM. => Topic started by: The15thMember on June 06, 2020, 03:59:34 pm
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I have a small colony (1 medium box) that has gone laying worker on me. I also have a colony that is a little behind my others, but is queenright, and I'd like to combine the LW hive with this weaker colony to deal with the LWs and give the queenright colony a boost. I'm thinking about making myself a double screen board and putting the laying worker colony on top of the queenright one in the hopes that the appropriate brood pheromones from the bottom colony will correct the LW situation. Will just being on top of the queenright colony be enough to suppress the laying workers?
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Member I am thinking HoneyPump told us a while back, the best thing to do is shake them out and let the hives deal with the laying workers when they beg to enter into a good hive. If I am wrong I hope Mr HP will quickly correct me.
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... Will just being on top of the queenright colony be enough to suppress the laying workers?
15th - my understanding is "no, it's not enough". Phillip has the correct answer, to my understanding. Shaking them out causes them to beg into another hive, and sorts out the LW's naturally. At the same time, all other hives will benefit from an increased workforce.
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Ms. Member, I do not know if a laying worker can revert back to normal forager or nurse bee. The ovaries are enlarged and I do not know if the ovaries ever shrink is another way to put things. If you try to combine, you might lose the queen of the weak hive.
I would do as Ben Framed, Phil, suggest which; I have done the exact same thing in your situation: shake all the bees out of the laying worker hive.
Van
Cool and I posted at the same time.
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Thanks for the replies everyone. I have been reading through some older posts on the subject, and I'm aware that the shakeout is the most recommended method. What I was looking for was a way to give the resulting boost in population to only or mostly this weak queenright hive. Do you know of any way to do that? I'd be willing to take some moderate risk to the queen in this hive, since it's not building up well and she may be to blame.
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Ms. Member, your laying worker hive may have only 2-3 laying workers. Most likely there are 200-300 or in the thousands, we just don?t know. If the 2-3 was indeed fact, that might be workable, but 200-300+ would result in queen loss unless you shake out the bees.
If you want to risk it: go ahead and shake out the lw hive 30 ft away. After the bees return, then combine. The hope is the laying workers have never conducted an orientation flight and therefore do not know where the home is. Let?s hope the returning foragers have not gone LW.
Your main problem is a poor queen and a combine will not help that as the combine will result in mostly forager bees and not nurse bees. Nurse bees are needed just as bad, maybe more so than foragers. Anyway to obtain a queen?
If that bee place you told me about, Next Door or de la Cruz farms [sourwood honey] has queens :I can arrange for you to have a new queen from one of those stores. I have account at both. Check on queen availability and text back to this message if either has queens.
Van
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Your main problem is a poor queen and a combine will not help that as the combine will result in mostly forager bees and not nurse bees. Nurse bees are needed just as bad, maybe more so than foragers. Anyway to obtain a queen?
If that bee place you told me about, Next Door or de la Cruz farms [sourwood honey] has queens :I can arrange for you to have a new queen from one of those stores. I have account at both. Check on queen availability and text back to this message if either has queens.
Van
Thanks for the offer, Van. If I decide to requeen, I'd rather just have the bees make their own from a frame of eggs from a better queen. I don't have the cash to spare on a queen at the moment, and I like breeding from my own stock. I'm not sure that this queen is at fault, and I was planning on give her a little while longer. I had to treat this hive for varroa in the spring, and it seemed to set the colony back, so it's possible this colony took the treatment harder than my other hives.
Ms. Member, your laying worker hive may have only 2-3 laying workers. Most likely there are 200-300 or in the thousands, we just don?t know. If the 2-3 was indeed fact, that might be workable, but 200-300+ would result in queen loss unless you shake out the bees.
If you want to risk it: go ahead and shake out the lw hive 30 ft away. After the bees return, then combine. The hope is the laying workers have never conducted an orientation flight and therefore do not know where the home is. Let?s hope the returning foragers have not gone LW.
I'm inclined to give your suggestion a try; it seems like a good compromise approach, and if it doesn't work, I haven't really lost anything.
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Shake them out 5 to 10 feet in front of the LW hive. Most will go to it.
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Your main problem is a poor queen and a combine will not help that as the combine will result in mostly forager bees and not nurse bees. Nurse bees are needed just as bad, maybe more so than foragers. Anyway to obtain a queen?
If that bee place you told me about, Next Door or de la Cruz farms [sourwood honey] has queens :I can arrange for you to have a new queen from one of those stores. I have account at both. Check on queen availability and text back to this message if either has queens.
Van
Thanks for the offer, Van. If I decide to requeen, I'd rather just have the bees make their own from a frame of eggs from a better queen. I don't have the cash to spare on a queen at the moment, and I like breeding from my own stock. I'm not sure that this queen is at fault, and I was planning on give her a little while longer. I had to treat this hive for varroa in the spring, and it seemed to set the colony back, so it's possible this colony took the treatment harder than my other hives.
Ms. Member, your laying worker hive may have only 2-3 laying workers. Most likely there are 200-300 or in the thousands, we just don?t know. If the 2-3 was indeed fact, that might be workable, but 200-300+ would result in queen loss unless you shake out the bees.
If you want to risk it: go ahead and shake out the lw hive 30 ft away. After the bees return, then combine. The hope is the laying workers have never conducted an orientation flight and therefore do not know where the home is. Let?s hope the returning foragers have not gone LW.
I'm inclined to give your suggestion a try; it seems like a good compromise approach, and if it doesn't work, I haven't really lost anything.
Member, I will pay for the queen, will cost you nothing but travel to pick the queen up. That is why I ask if either store has queens. I have accounts at both bee stores texted above. I do not know if they have queens available? If the bee stores have queens, check Monday, I?ll buy a queen for you. You were nice to offer your time when I wanted real sourwood honey, I am just returning the favor I owe you.
Van
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Member, I will pay for the queen, will cost you nothing but travel to pick the queen up. That is why I ask if either store has queens. I have accounts at both bee stores texted above. I do not know if they have queens available? If the bee stores have queens, check Monday, I?ll buy a queen for you. You were nice to offer your time when I wanted real sourwood honey, I am just returning the favor I owe you.
Van
Again, thank you for the offer, but you do not owe me anything, and if you did, it wouldn't be something that would cost you anything, as my help to you did not cost me a dime. You are very generous and kind, Mr. Van, but it's really not necessary. I'm quite content with things as they stand. :smile:
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... What I was looking for was a way to give the resulting boost in population to only or mostly this weak queenright hive. Do you know of any way to do that? ...
Pull 2 frames of capped brood from another hive over to this hive and drop them into the center of the brood nest. They will hatch shortly and boost the population.
... and shake the others out ... :cool:
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Pull 2 frames of capped brood from another hive over to this hive and drop them into the center of the brood nest. They will hatch shortly and boost the population.
... and shake the others out ... :cool:
Yeah, I know that's an option. I was just looking to combine weak and weak into strong if possible, before I weakened the strong to strengthen the weak. :wink:
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How about combining in stages if you want an experiment. If you shake in a frame a day or so, would the queen right hive fight off the workers and accept the nurses? Or paper combine a medium with only two or three frames in it then, once combined, add more paper and a couple new frames?
Just throwing it out there. I have no idea.
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Usually by the time a colony has gone LW the bees are old and not of much value other than a very temporary foraging force boost. Think of them as that and that only. Not as a box of bees capable of boosting all functions of another weak colony. They can help, just not as much nor in the way you may be thinking.
What is the comparative size of the LW colony to the queenrite colony?
If the LW is 1/3 the size or less, then you could just newspaper combine. Spritz the frame bars and bees with sugar water first. Put the LW on top. The bees will sort the bees. A day or three later there will be a bunch of bees dead and curled on the ground out front. Those are the LWs. The combine does pose risk to the queen. However if you are mindful of the size ratio given that risk is reduced.
By far the best way to deal with LW is to shake out and completely remove their hive equipment so they have no place to go back to and must beg into the other hives. Again use the bees to sort out the bees. If there is a preference of which hive to be boosted by them, do the shake nearer that hive than the others, but do be mindful of comparative volume to spread them so the one hive is not overwhelmed by the misfits. (Ratio)
Hope that helps!
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We have had some success with shaking out the LW hive 50+ yards away putting the hive back with a frame of open brood and nurse bees.
The hive, brood and nurse bees are dusted with icing sugar. As the LW bees return they find a new smell in the hive and tend to accept the brood and nurse bees. Success about 75% of the time.
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Usually by the time a colony has gone LW the bees are old and not of much value other than a very temporary foraging force boost. Think of them as that and that only. Not as a box of bees capable of boosting all functions of another weak colony. They can help, just not as much nor in the way you may be thinking.
Oh, I get it now. That makes sense, since it's the older bees who are most probably not LW's. That definitely changes how I'm thinking about this situation.
What is the comparative size of the LW colony to the queenrite colony?
If the LW is 1/3 the size or less, then you could just newspaper combine. Spritz the frame bars and bees with sugar water first. Put the LW on top. The bees will sort the bees. A day or three later there will be a bunch of bees dead and curled on the ground out front. Those are the LWs. The combine does pose risk to the queen. However if you are mindful of the size ratio given that risk is reduced.
By far the best way to deal with LW is to shake out and completely remove their hive equipment so they have no place to go back to and must beg into the other hives. Again use the bees to sort out the bees. If there is a preference of which hive to be boosted by them, do the shake nearer that hive than the others, but do be mindful of comparative volume to spread them so the one hive is not overwhelmed by the misfits. (Ratio)
Hope that helps!
The LW colony is about 1/2 the size of the queenright colony, so probably pretty risky to just do a combine. Hmm. . . . Maybe I'll do this: Since there is still some capped worker brood in the LW colony, I could shake out the bees and give the struggling colony the capped brood as a booster.
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"""The LW colony is about 1/2 the size of the queenright colony, so probably pretty risky to just do a combine. Hmm. . . . Maybe I'll do this: Since there is still some capped worker brood in the LW colony, I could shake out the bees and give the struggling colony the capped brood as a booster. """
Hold on a minute, member. Something isn't jiving here. LW's don't develop when there is worker brood present. I think you need to re-evaluate this hive.
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"""The LW colony is about 1/2 the size of the queenright colony, so probably pretty risky to just do a combine. Hmm. . . . Maybe I'll do this: Since there is still some capped worker brood in the LW colony, I could shake out the bees and give the struggling colony the capped brood as a booster. """
Hold on a minute, member. Something isn't jiving here. LW's don't develop when there is worker brood present. I think you need to re-evaluate this hive.
Just to clarify, there is only capped worker brood, no open worker brood. There were a handful of larvae, but they were either VERY young, or very old. I was seeing capped drones here and there amongst spotty capped worker brood, so my assumption is the larvae are all drones. I did see some cells with only 1 egg in the bottom, but I also saw cells with many eggs, like 8-10 per cell.
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Member is seems like you have everything but a visible Queen.
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If you have some larvae and few eggs and capped ... An entirely possible (and highly probable) situation is the hive raised a queen and the LWs promptly killed her right shortly after she laid a few eggs. Happens often, nearly every time.
Only way to know for sure is to take the hive out a ways. Put and empty box on the ground with 3 empty frames in it. Put on a queen excluder. Put another empty box on the QE, no frames. Next shake them all into the open box. They will crawl through the QE. If there is a queen. You will find her. If there is no queen, finish up shaking everything out and take all the equipment away.
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Just to clarify, there is only capped worker brood, no open worker brood. There were a handful of larvae, but they were either VERY young, or very old. I was seeing capped drones here and there amongst spotty capped worker brood, so my assumption is the larvae are all drones. I did see some cells with only 1 egg in the bottom, but I also saw cells with many eggs, like 8-10 per cell.
Bee careful not to shake out a new queen. Young queens will often lay multiple eggs per cell (which the workers clean up), in the first few days. Shaking them out, kills her. ... ask me how I know ...
LW's lay on the walls of the cells mostly. New/young queens lay multiple eggs on the bottom of the cells. So - location of the eggs in the cell tells you what's really going on. (Lesson learned the hard way :grin:)
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If you have some larvae and few eggs and capped ... An entirely possible (and highly probable) situation is the hive raised a queen and the LWs promptly killed her right shortly after she laid a few eggs. Happens often, nearly every time.
Only way to know for sure is to take the hive out a ways. Put and empty box on the ground with 3 empty frames in it. Put on a queen excluder. Put another empty box on the QE, no frames. Next shake them all into the open box. They will crawl through the QE. If there is a queen. You will find her. If there is no queen, finish up shaking everything out and take all the equipment away.
Based on the fact that I was waiting for this hive to make a queen, I think your theory is very probable, HP. I know they made one because I found hatched queen cells, and I'm also pretty sure she laid at least a little bit because the population in the hive is the same and maybe even a little increased since last time I checked. I'll look for a queen the way you described tomorrow to make sure she's not in there somewhere. If I find her (unlikely, I imagine), how should I proceed? Also, why the 3 empty frames in the bottom box?
Bee careful not to shake out a new queen. Young queens will often lay multiple eggs per cell (which the workers clean up), in the first few days. Shaking them out, kills her. ... ask me how I know ...
LW's lay on the walls of the cells mostly. New/young queens lay multiple eggs on the bottom of the cells. So - location of the eggs in the cell tells you what's really going on. (Lesson learned the hard way :grin:)
I've seen new queens lay multiple eggs in a cell and this doesn't look like that to me. I'm not sure whether there were many eggs on the sides, as I forgot to specifically look for that, but many of the eggs looked kind of small and underdeveloped, and as I said, some cells were full of eggs, it wasn't just an extra one or two.
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"""An entirely possible (and highly probable) situation is the hive raised a queen and the LWs promptly killed her right shortly after she laid a few eggs. Happens often, nearly every time."""
HP, I don't disagree with you often, but I have to this time. If the hive raised a queen, where did the LWs come from. They don't develop in numbers as long as there is viable brood in the hive. If it were an introduced queen into a LW hive, I could see them killing her, but if the hive raised her, the LWs were not present.
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Iddee. The scenario I was thinking of is all brood is mature, capped and emerging, no larvae anywhere. LW develops. A frame of partial brood is later added. They raise a queen cell from the adder. The virgin is ignored and mates. When she starts laying she is killed. The missing link in the assumption only 15th can verify if a brood frame was given along the way.
15th. If you find the queen crawling on the excluder, temporarily cage her for safety. Then put a box back at the hive location with 1 or 2 empty frames, no bees. Release the queen. Now go back to the area you have the shaker stuff. Shake everyone out there. Take the bee-free frames back and put in the box with the queen. Finish and close up the hive. Alot of the bees will beat you back there. There will be quite a few bees flying. So suit up first so you do not get any in your hair. Wait 1 week then go back to see if things are looking normal.
The three frames below is to draw the bees down through the excluder, rather them wanting to climb out the shaker box.
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15th. If you find the queen crawling on the excluder, temporarily cage her for safety. Then put a box back at the hive location with 1 or 2 empty frames, no bees. Release the queen. Now go back to the area you have the shaker stuff. Shake everyone out there. Take the bee-free frames back and put in the box with the queen. Finish and close up the hive. Alot of the bees will beat you back there. There will be quite a few bees flying. So suit up first so you do not get any in your hair. Wait 1 week then go back to see if things are looking normal.
The three frames below is to draw the bees down through the excluder, rather them wanting to climb out the shaker box.
Thanks HP, will do.
Iddee. The scenario I was thinking of is all brood is mature, capped and emerging, no larvae anywhere. LW develops. A frame of partial brood is later added. They raise a queen cell from the adder. The virgin is ignored and mates. When she starts laying she is killed. The missing link in the assumption only 15th can verify if a brood frame was given along the way.
The LW hive is a swarm I caught on 5/2. Based on how they were acting (straggler bees moved from the swarm tree to the lid of my swarm bucket and then into the hive when I dumped them in) I assume they had a queen at the time. I put a frame of open brood in the box when I hived them to help them stick. On 5/10 I checked on them for the first time, and they had 2 queen cups and 1 elongated but not capped queen cell, all with larvae in them, on that frame I'd given them. I don't believe there was any other brood in the hive then. Yesterday I went in to check for a new queen, and found the LWs. Does that confirm or deny your suspicions?
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Confirms the scenario hypothesized. Thanks.
Let us know how it goes and plays out. Am hopeful you find a queen on the QE. Not to be at all surprised if you don't.
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Trying to follow this, are we saying that the evidence for LW is multiple eggs in cells? Is there drone brood yet?
As brood was added this would have finished hatching 5/26, not long for LW to get going!
I am going to back "a queen somewhere" getting her act together laying multiples.
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HP, with the new info, I now agree with you. I also agree with Oldbeavo. I think another week will show a nice laying pattern from the new queen.
Member, I would leave the hive as is for another week, then make a decision based on what it looks like then.
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So I followed HP's instructions this morning and shook all the bees out through a queen excluder. They didn't all go through it as well as I expected them to, but with them all spread out and carefully inspecting each frame before I shook it, I'm quite sure there was no queen in there. However, I did find a capped queen cell. I sort of treated that queen cell as if it was a queen and followed those instructions, so I put all the bee-less frames in the hive back on its stand and shook out the bees into the woods a good ways away from the apiary. Also, sort of on instinct, I gave the LW hive a frame of eggs and young brood from another hive without shaking all the bees off it, in the hopes that the open brood will help and perhaps little boost of normal adult workers will too. I also dusted the LW hive with powdered sugar before closing it up to help with fighting between the added bees and returning foragers, and also because Oldbeavo recommended it. So now my question is this: when should I check on them again? If it was a normal situation, I'd wait 3 weeks for the queen to start laying, but I'm not sure I should do that since the LWs could get out of hand again in that time frame. What do you recommend?
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As you have open brood in the hive now, wait 3 weeks, as that will be when all the brood is hatched.
You will have eggs or polishing awaiting a queen to lay, or pollen and honey everywhere that brood should be. Decision should be easy then.
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Sorry to hear a queen did not materialize. Though not unexpected. By placing the cell frame, you have just begun take 3, round 3, of the endless cycle of false hopes of the LW condition. LW are wrought with efforts that end in despair and disappointment. They have killed their new queen they raised the previous brood frame given weeks ago and are now using one of her recently laid eggs to start another cell ... which will ultimately have the same fate. My advice --> Go squash that cell, and shake them all out now. Put the equipment away or redeploy it to other hives that are doing well. You will sleep better, and the other nearby colonies will thank you for the short lived boost in foragers.
If you wish to continue down the dark path, hoping to see light at the end, then you must now add a frame of open brood at minimum once per week for the next 6 to 8 weeks. That will be the time it will take for LW to be suppressed, the 2nd then the 3rd newly raised and mated queen to be killed, eventually one will be left alive, and the 3+ weeks for her own brood to start emerging.
IMHO.
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As you have open brood in the hive now, wait 3 weeks, as that will be when all the brood is hatched.
You will have eggs or polishing awaiting a queen to lay, or pollen and honey everywhere that brood should be. Decision should be easy then.
Sorry to hear a queen did not materialize. Though not unexpected. By placing the cell frame, you have just begun take 3, round 3, of the endless cycle of false hopes of the LW condition. LW are wrought with efforts that end in despair and disappointment. They have killed their new queen they raised the previous brood frame given weeks ago and are now using one of her recently laid eggs to start another cell ... which will ultimately have the same fate. My advice --> Go squash that cell, and shake them all out now. Put the equipment away or redeploy it to other hives that are doing well. You will sleep better, and the other nearby colonies will thank you for the short lived boost in foragers.
If you wish to continue down the dark path, hoping to see light at the end, then you must now add a frame of open brood at minimum once per week for the next 6 to 8 weeks. That will be the time it will take for LW to be suppressed, the 2nd then the 3rd newly raised and mated queen to be killed, eventually one will be left alive, and the 3+ weeks for her own brood to start emerging.
IMHO.
Ah, two opposing answers. Must be a beekeeping question. :wink: :cheesy:
Here's what I'm inclined to do. I agree that HP's description of the LW rabbit hole is risky and dismal, and I'm not really into sacrificing that much brood to prop up this hive when it might not even work. I'd rather give that brood to the weak queenright hive that I mentioned previously, if I'm going to give it to someone. I think I'll take Oldbeavo's advice and wait and see what happens. Maybe it'll work. If it doesn't take, then so be it, and I'll shake them out in 3 weeks and call it quits. I don't really have anything to lose by trying. Other than sleep, according to HP, but rest assured, I rather enjoy an experiment. This was kind of a bonus hive anyway; my goal was 6 hives this year, and this one was hive number 7, so no big loss.
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Hahaaa. Different views of what to dos from near global antipodes. :)
You have good input and advice throughout the thread in front of you however way you choose to proceed.
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I think it may have worked. I checked on the LW hive today and although I couldn't find a queen, I saw a lot of cells with single eggs in them. I saw 2 cells with 2 eggs, and 1 egg laid in a partially filled pollen cell, but nothing like it was with cells filled with eggs, and none of the eggs were on the sides of the cells. The hive has lots of small drones running around that developed from the worker eggs, so I easily could have missed a queen. I'll check again next week to see if I can find her and make sure the situation stays normal.
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Thanks for the update. I hope this works out and the new queen survives. Yes keep us updated, interesting.
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All right, I'm officially confused. I inspected this hive again today. There are 5 frames with sizeable patches of drone brood in all stages. I didn't see many eggs, but the ones I did see looked normal, single eggs in the bottoms of the cells. I went through the entire box twice and I didn't see a queen anywhere though, nor was there any worker brood to be found. Any idea what's going on in there now?
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It's time to shake all bees through an excluder and find the drone laying queen, tho I feel any more resources invested in this hive is a waste. I would just shake them all out onto the grass and put the equipment away, or add it to other hives.
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It's time to shake all bees through an excluder and find the drone laying queen, tho I feel any more resources invested in this hive is a waste. I would just shake them all out onto the grass and put the equipment away, or add it to other hives.
Ah, drone layer. That makes sense. I guess I just missed her because she is tiny. Yeah, I am thinking the same thing about shaking them out. Just no point putting more into them this late in the season. Thanks so much for all the help everyone! I have no idea what I would do without you all.
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I shook out the hive yesterday afternoon, and this morning there is a sizeable cluster of bees on the cinder blocks at the hive site. I assumed they'd disperse once it warmed up, but they seem quite content to just hang out here. I'm worried they are going to get rained on. Is this normal?
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I?m sure they just tried to go home, and there was no home. I wonder if you go dump the blocks off, if they would beg their way into another hive.
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Ignore them. In 2 days they?ll be gone. Having begged into the other hives.
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Ignore them. In 2 days they?ll be gone. Having begged into the other hives.
Thanks, HP. Will do.
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This has been very interesting and educational. I want to thank you contributing experts for your patience and non pressured persistence. The best kind of teaching IMHO. Member, this was a great thread. We are very fortunate to have these caliber of beekeepers here that have contributed with selfless helpful information. Your questions and clear answers of their questioners was a big help, in helping me understand more about laying worker hives. Sincere thanks to all.
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This has been very interesting and educational. I want to thank you contributing experts for your patience and non pressured persistence. The best kind of teaching IMHO. Member, this was a great thread. We are very fortunate to have these caliber of beekeepers here that have contributed with selfless helpful information. Your questions and clear answers of their questioners was a big help, in helping me understand more about laying worker hives. Sincere thanks to all.
Thanks, Phillip, and I whole-heartedly agree. I wouldn't be anywhere near as knowledgeable and successful in my beekeeping if it weren't for all the help I receive and the things I learn on this forum, both from my own questions and others'. It's so beneficial for us all to be able to learn from each other.
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The cluster of bees was still there this morning, so I decided to do as .30WCF suggested and try shaking them off the cinder block. When I did, I found they had built about a 3 in. by 3 in. comb in the center of the cluster! :shocked: The shaken bees are now reclustered on the other cinder block that was making up the hive stand. What should I do now?
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Is there a queen in there?
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Is there a queen in there?
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Well I'm assuming there was a drone laying queen in the hive, but before I shook out the hive I checked and double checked, and I didn't see a queen. Maybe it's an intercaste female or something. You'd have to think that something is holding them together like this.
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That?s right. I had forgotten that you shook the whole hive out
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FYI, clumping of bees does not mean a queen is present.
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FYI, clumping of bees does not mean a queen is present.
But does comb building?
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Normally, yes. However LW believe they have a queen; so same-same.
There is a misconception that a LW does not fly and are left behind at the shake out spot. I assure you they fly just as well as any other bee. This is why the method requires complete removal of the hive equipment when doing it so they have no place to go, no shelter, but into other nearby hives.
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3rd time was the charm. I shook them off the cinder block again today and removed both cinder blocks from the site, and now they are no where to be found. Thanks again, everyone.
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>Most likely there are 200-300 or in the thousands, we just don?t know.
"More than half of the bees in laying worker colonies have developed ovaries (Sakagami 1954)..."-- Reproduction by worker honey bees (Apis mellifer L.) R.E. Page Jr and E.H. Erickson Jr. - Behavioral Ecology and Sociobiology August 1988, Volume 23, Issue 2, pp 117-126
http://link.springer.com/article/10.1007/BF00299895