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I am working on a creature (tentatively calling them “beepipers”) that lives in hives (heavily based on honeybees). The drone (reproductive male) is born to mate with a queen, and will die shortly after doing so. He works very little and eats a lot, making him expensive to raise.

Here’s where the nuance starts. For a beepiper queen, one drone only provides enough seed for 50-60 fertilized eggs. As a result, a queen must continue to mate once a month or so throughout her life.

Beepiper drones mature reasonably quickly, but don’t leave the hive right away. This way, queens can formally trade their drones, giving all involved queens new genetic material. It is viable for a queen to mate with her own drones, but it’s obviously frowned upon, so the trade is the best way for nesting queens to maintain their hive.

A nesting queen wants a few foreign drones stocked up at a time, as she mates once a month. She guards this “hoard” jealously, since if she runs out of fertilized eggs, her workers replace her. The hoarded drones basically sit around and eat.

Shortly before winter, all of the hive’s drones and virgin queens will take their nuptial flights, allowing a new generation of queens to enter the picture. Queens are highly competitive; while nesting queens can tolerate each other well enough to trade, nuptial queens without hives are seen as a direct threat, particularly to their mothers.

The virgin queens are already allowed in and out of the hive, so they can easily get away, but the drones only otherwise leave the hive to be traded, so it is obvious when they are preparing for their nuptial flight.

Logically speaking, a smart nesting queen (who knows whenever anyone enters/leaves the hive) would kill any drones before they flee, as the drone could create a rival queen. However, if she succeeds in this, the species would end with her.

There are a few options I’m considering for addressing this problem.

  • The flighting drone emits a toxic chemical that prevents the queen from killing him. However, he must then mate with a younger queen, who would also be vulnerable to this chemical, so this could get complicated.
  • The workers actively defend the nuptial drones, keeping the queen from killing them or possibly sneaking them outside the hive. However, the workers are threatened from the drones leaving too, so this would have to be an instinctive measure.
  • The drones are protected by moral instinct/religion, and a queen killing them outside of mating is considered a cardinal sin. I’d prefer to avoid this, since I want these creatures to be largely without moral constraints.

Would any of these work well to address this issue? I’d prefer the drone’s escape method to be simple, since it isn’t a particularly central part of my story, but it would close the continuity of the drone’s life cycle.

TLDR: Smart bee queens realize letting drones on nuptial flights creates rival queens. Why would they let them go anyway?

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    $\begingroup$ /Logically speaking, a smart nesting queen... would kill any drones before they flee, as the drone could create a rival queen. / It is not clear to me why drones are any threat. I have to think that mostly drones create workers. If a rival queen occurs in some other nest that is their problem. If your queen creates a threatening and unwanted virgin queen that is who should be summarily killed but you have worked around that. Why are drones a threat? $\endgroup$
    – Willk
    Nov 19, 2022 at 17:44
  • $\begingroup$ @Willk I should have clarified- these hives actively compete for territory and frequently invade each other. The drone itself isn’t a threat, but it will mate with a new queen who may start a competing hive in the area (or supersede an older queen). Queens trading drones is about the extent of peaceful relations between hives- outside of that they are usually hostile. $\endgroup$
    – Mark Price
    Nov 19, 2022 at 18:12
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    $\begingroup$ I presume that while a drone dies after it has mated, the drones that haven't mated don't die? Or do all the drones that go on mating flights die whether they atually mate or not? The answer to this will make a very large difference to the species mating strategies. $\endgroup$
    – Monty Wild
    Nov 20, 2022 at 22:20
  • $\begingroup$ Honeybee queens need only mate once for their entire lives. They will mate with multiple drones, found a hive, then never leave the hive again or do anything other than produce eggs. Also, honeybees actively remove drones from the nest every fall. They die to the elements, whether having ever mated or not. This is not a problem because the queen can make new drones at any time. An unfertilized egg grows into a drone. In reality, drones are the cheapest resource. They're literally disposable. You can resolve this problem by changing the biology a little. $\endgroup$
    – user458
    Nov 22, 2022 at 19:54
  • $\begingroup$ @fredsbend but queens would prefer non-related drones to avoid inbreeding. Inbreeding is far less of a risk for them then say mammals, but they still will preferentially desire non-related mates if for genetic variety if nothing else. We can see this in herms capable of self impregnation and aphids which will mate with haploid sons if necessary in both cases we see a strong preference to mate with a foreign individual over inbreeding. This means that there is still real value to be had from trading drones with a foreign queen even if it's possible to produce and mate with drones yourself. $\endgroup$
    – dsollen
    Nov 25, 2022 at 15:57

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Make love, not war

The drones exist only to be sexy. The queen and drone pheromones stimulate each other in a feedback loop, where ultimately the queen has control - she can choose to dampen her own pheromones, which in turn leads the drones to tune down their own, and inhibit sexual characteristics. This allows the queen to get on with her queenly business and only go "in heat" once a month when her sperm store is low, while the rest of the time the drones remain in a relatively drab state that isn't too distracting.

Virgin queens, however, produce their own drone-stimulating pheromones - this is, in part, how they convince them to leave the safety of the hive to go on a dangerous nuptial flight. Drones surrounding a virgin queen have their sexual characteristics in overdrive - and the senior queen finds them irresistible. She can try to lower her pheromone production, but the drones are saturated with the virgin queens' pheromones and remain in ultra-sexy state. The senior queen might attempt to attack them, but it inevitably devolves into, ahem, distraction - they are just too damn hot. While she's preoccupied with one or a few of the sexy sexy drones, the others are able to fly off safely with the virgin queen.

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Creating Allies, Not Enemies

A new queen that has mated with one of your drones now has a connection to you, which makes her and her offspring from that drone less likely to want to challenge you. So you actually want to encourage your drones to go mate with new queens, because you're not creating rivals for yourself, you're creating rivals for all of the other nearby queens.

Flipping the question

If it's commonplace to swap out drones, but young un-mated queens are a threat, and especially to their mothers, then why are they allowed out of the hive? Seems like a queen that wanted to eliminate potential rivals would kill the young queens (or at least prevent them from mating), not let them roam free.

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    $\begingroup$ Making allies works well- I think I can use that for new queens forming alliances. Just to clarify- the nesting queen can and often does kill her un-mated daughters before the winter, but she also needs them for other tasks in the hive, so they must be allowed to come and go and thus can easily slip away without the queen noticing. This is a more central part of the story I’m still working on though. $\endgroup$
    – Mark Price
    Nov 19, 2022 at 21:51
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    $\begingroup$ Preventing all the young queens from mating and having offspring would effectively evolutionarily eliminate the genes responsible for such a behaviour. This behavioural pattern is hence bilogically impossible. $\endgroup$ Nov 21, 2022 at 17:20
  • $\begingroup$ Bees allow new queens to stay in the hive while the old queen leaves with half the bees in the hive to make a new one. Also, intelligent, eusocial creatures do not murder kin, usually. $\endgroup$ Nov 21, 2022 at 19:57
  • $\begingroup$ @mindwin A bit too strong of a statement. For instance in many ants it's possible for workers to lay eggs (males can be produced without mating). If an egg of anyone other then the queen is found the other workers will kill their nephew, even though they share as much as 37.5% of their genes with that egg, more genes then they share with drones produced by the queen (% genes shared with relatives is different in haploidy mating system) They do this because if workers could produce drones workers wouldn't work, the hive system would fall apart, and long run this would harm everyone. $\endgroup$
    – dsollen
    Nov 23, 2022 at 18:02
  • $\begingroup$ @dsollen be charitable. What strong statement did you see, when the first sentence is a truthful statement about Apis mellifera behavior, and the second is about intelligent creatures, which your ants are not. Also, it ends with "usually". Of course we admit there will be outliers. $\endgroup$ Nov 24, 2022 at 12:11
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Genetic Drives

One must presume that the behavior is genetically driven.

Explaining Things Through Evolution

Whenever a question of the form "Why does this mating strategy exist?" the answer must always be because strategies with small changes will produce fewer copies of the genes that produce them in the next generation.

When explaining such things, the usual thing is to explain it in terms of what the organism's genes know and want. This is as old as the book "The Selfish Gene" by Dawkins. It should be understood to be a metaphor, not actual anthropomorphism. I don't actually think that genes have knowledge and desire. They behave as though they do, since when they behave selfishly they get copied more.

Known Cases in Other Animals

Some quick examples.

If a male cat (house cat, leopard, tiger, lion, etc.) encounters a kitten under a certain age, he will kill it. He won't eat the body, just bite it until it is dead. This is because male cats are nomadic. A kitten under that age is unlikely to be his kitten. And if mother cat loses her kitten, she will be receptive to mating much sooner. So Mister Cat will have a receptive female, and remove rivals from the gene pool. As a genetic strategy, if he does not kill that kitten, the rival genes stay around, and Mother Cat stays unreceptive until the kitten is weaned. Months at least, by which time he likely has wandered away.

Males that are "good dads" to other male's kittens get erased from the gene pool.

From the standpoint of Mother Cat, it is ALSO a successful strategy. Not for her, but for the part of her potential genetic descendants that will be male. That is, her male offspring will be more successful if they do this also. So her genetics find it an attractive thing for a male to do this. She may be very upset that she has lost her kitten. She might even attack the male, scratching the heck out of him. Then she will go sulk for a day. Then she will come back and mate with him.

Because it is a genetically successful strategy, kitten-breath is a turn-on for female cats.

Consider chimps. A young male chimp must attract a mate. One thing he can do to be considered attractive is to show he is a good father. So young bachelor chimps will pester mothers with babies to be allowed to babysit. And they will do it in a way to be apparent to young females, to be seen caring for a baby. The problem is, a happy baby chimp will stay quiet and often go to sleep. A sleeping baby is not going to attract any attention.

So young bachelor chimp will pinch the baby to make it cry. Then he will make a big show of comforting the baby. The females will watch and judge his efforts. They will do this on their way to judging him a suitable mate.

The thing is, it works. It works even when he gets caught pinching the baby. To a female chimp, a male who creates a big show of comforting a baby, even when he mistreats it, is a male with a strategy to attract females. If his genes include this, then his male offspring will do the same. So her genes, which are interested in getting male genes that will be successful, will find this male attractive.

Because it is a successful genetic strategy, an abusive and showy male is attractive to a female chimp. (And consider, if you dare, the huge number of stories of how step-children get treated.)

The Beepiper Case

So finally, let's turn to our queen beepiper permitting drones to mate with her offspring. She has determined through the usual process that they are suitable to mate with her. Presumably they are fresh, not the fathers of her offspring. So they are suitable mates for her offspring.

(There might be some interesting drama here over keeping them fresh. Perhaps after about the time it takes for a newborn beepiper to reach adulthood, male drones become unattractive. That might be the life-expectancy of a male drone. Slam the door on them before they get a chance to mate with their own daughters.)

So her genes know that these mates are going to provide good genetic material for her offspring, and therefore will be good for making copies of her genes. Even if her offspring turn out to be rivals. Her genes will override any thoughts of her own well being. She will have drives to do things that are good for her genes regardless of how bad that turns out to be for her.

Genetically, her offspring have 50% overlap with her. So something that produces an increase of chance of reproduction for one of her offspring that has less than 50% chance of killing her, will be genetically attractive. If she can help multiple of her offspring at one time, it will approach the situation that her own death will be outweighed by that help.

So these mating flights, which set her offspring on their way, will be a genetically successful strategy, and so will be attractive to the queen.

As they will be attractive to every individual in the hive that is an offspring of the queen, even the non-breeding workers. Their genes have a genetic interest in the success of the mating flight, even if the rival queen has a chance to come back and burns down this hive. This is the case with bees in our world, where worker bees will assist "colonizing" queens that are about to leave the hive to form new colonies.

So the mating flights are allowed, even supported, by every individual in the hive who is genetically related to the soon-to-mate queens. They will find some means by which the flights can happen. Because, by so doing, they produce more copies of themselves in future generations.

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  • $\begingroup$ You seemed to miss the point that drones die after they have mated. No living drones are fathers. $\endgroup$
    – Monty Wild
    Nov 20, 2022 at 22:18
  • $\begingroup$ This now makes me wonder why humans are more different, and most of us tend to care for foster children on a similar level as genetic ones, at least there is a strong social pressure to do so. Is it merely that living in a larger community (village, city-state) gives more advantages than the pure genetic advantages provided by murdering all the children in the local community? Here, the competition between villages, and later city-states, was more crucial than competition between individuals, so that the city-state with more cooperative inhabitants got an edge over the others? $\endgroup$
    – vsz
    Nov 21, 2022 at 5:38
  • $\begingroup$ I agree in general, though I think this could be simplified to a TL/DR if she didn't let her children mate her genes die with her, so she is better off allowing both male and female children of hers out to mate to spread her genes. $\endgroup$
    – dsollen
    Nov 23, 2022 at 17:54
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Bee hives, the dumb ones at least, are democratically led. Oh sure, they can't have sex or make babies, but every other decision? The workers put it to a vote and majority rules. Where to get food? A few scouts go out, they come back and dance to indicate what they found, the rest of the hive vote by following the scout of their choice. Need a new place? A few scouts go out, they come back and dance, the rest of the hive votes by exploring, then comes back and dances with their choice until the whole hive agrees on one place to stay. Really need a new place to stay? The whole swarm goes out and votes en masse.

As the beepipers evolve, it would be easy for these dances to evolve into proper debates. And since the workers are the ones who actually go out and do stuff, they are the ones with all the information and therefore power. Why would they inform the queen, who just sits there and lays eggs all day? She doesn't have to do anything, it's us who make this hive go round! The workers trade the drones. The workers allow the nuptial flights. The queens do nothing and are as commodified as the drones.

So why would queens allow drones to go on nuptial flights? Answer: It doesn't matter because it's the workers who decide. The queens are just as useless and powerless as the drones.

As for why the hive as whole would allow it: Daughter hives are easier to ally with than fully foreign hives. For at least one generation, probably more, it would expand their effective territory.

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    $\begingroup$ Indeed, it's an urban myth that honeybee hives are 'ruled' by the queen. The workers feed the queen larva, clean her, and help her go out on a mating flight... then they feed her so much that she is too fat to fly again and is hive-bound, while leading her around the hive to the honeycomb cells that they have prepared to have eggs laid in them. If the workers collectively decide to rear a new queen (either for supercedure or to swarm) they will create a large queen cell and entice the queen to lay in it even though it's obvious that it's a queen cell, then... $\endgroup$
    – Stephen
    Nov 21, 2022 at 10:59
  • $\begingroup$ actively drive the queen (who seeks out the pheromone trail of other queens to attack and kill them) away from the larva until it's developed. If they want to swarm they will starve the queen until she's light enough to fly again. In short, while the queen might be the most important bee in the hive, she's far from the most powerful. $\endgroup$
    – Stephen
    Nov 21, 2022 at 10:59
  • $\begingroup$ @Stephen I'd argue that bees are a great example of instinct only species. There is no thought or procedure. Artificial introduction of pheromones and other chemicals produces these behaviors with very high rates of success. Neither the queen nor the worker contemplates any decision. They act on chemical cues. As such, one should view the hive as the organism, not the individual bees, and individual bees as tissues and organs within the organism. $\endgroup$
    – user458
    Nov 22, 2022 at 20:01
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    $\begingroup$ Very true; bees are a great example of complexity arising from the interplay of many individually-simple actions. When worldbuilding an individually-intelligent species from such a base it'd be essential to acknowledge that. A hollywood-style anthropomorphisation of an imperial court is not going to feel realistic. $\endgroup$
    – Stephen
    Nov 23, 2022 at 14:56
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It seems to me the workers would be the easiest solution. If the queen of their hive is replaced by a younger/ stronger rival that probably doesn't change very much for them, they will continue to be workers under the new queen.

In fact if their queen is starting to get old, getting a new one might be in their best interest to ensure the long term survival of the hive. Even if she isn't old but just unpopular the workers could always hope for a better new queen (and possibly be disappointed).

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    $\begingroup$ One thing I just realized while reading this answer, which otherwise works well- the drones released from a hive won’t lead to a supersedure in that hive, since the drones would have to mate with their sister to make that happen (only a daughter can supersede her mother). Releasing drones would only result in new enemy neighbors, not a replacement for a queen. Possibly a nitpick, but the workers wouldn’t want an enemy hive springing up nearby any more than the queen does. At any rate, this answer definitely got me thinking about logistics! $\endgroup$
    – Mark Price
    Nov 19, 2022 at 18:19
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Evolution is funky

Survival of the fittest doesn't mean the strongest, or biggest, or most populous colony lives, it means that the species/mutations that can reproduce the most don’t go extinct.

So it might be beneficial for Queen Z to not allow nuptial flights, and maybe queen Z does stop them all. But it will be much harder for queen Z’s genes to spread, meaning whatever genes made queen Z do that won’t live on. However, Queen X did the opposite, and allowed nuptial flights. Now queen X, though maybe killed by rival queens, has spread her genes.

This might just seem like wild speculation, and that nothing like it has ever been observed. But it has! Kentrophoros is a genus of ciliate which has a symbiotic relationship with bacteria that live on it. It eats some of the bacteria, and in return the bacteria get access to the ideal living conditions. Getting eaten might be bad for the individual, but they can reproduce and spread their genes from the kentrophoros, and so they are able to survive.

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  • $\begingroup$ "Getting eaten might be bad for the individual, but they can reproduce and spread their genes from the kentrophoros, and so they are able to survive." Tell that to the farm chicken. $\endgroup$
    – user458
    Nov 22, 2022 at 20:03
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She Sends Her Own

The simplest solution is that she doesn't have to let her valuable traded drones out - she simply allows her own sons to go out. This will likely have a powerful instinctual urge behind it, as it's quite literally how the queens will propagate their own genes to future generations. The instinctual urge to send her sons out to make new hives would have to be just as strong as any animal's urge to have their own children.

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    $\begingroup$ And if these bee like creatures have some human traits, they might hope for some familial loyalty. Giving a daughter to a rival king's son was used all the time to sort of seal alliances. Depends on if the OP wants this Game of Thrones type stuff in the story. $\endgroup$
    – user458
    Nov 22, 2022 at 20:05
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It is not beneficial for the drone to mate starting from. A smart drone will never do, regardless of the pleasure. If that much happens, the rest may happen as well.

Behavior is not always driven strictly by the personal benefit, even if individual is otherwise sentient. Drones are children for a queen, understandable she does not want to kill them. You can make them more attractive than just lazy eaters, say drones are great artists that tell stories and decorate internals of the hive.

P.S. Real bee drones do leave the hive, flying longer distances away than worker bees. They return if they fail to mate, or may enter another hive where, unlike worker bees, they would be welcome.

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I strongly back up boba fit's statement, in the long run killing your offspring means your genes die with you. Even if a queen is spreading her genes partially through trading drones she still wants every way to spread her genes. The threat of increased competition is not enough to counteract the advantage of more genes spread.

I do have a few points to add as to why this is the case though.

Better a relative then a stranger for a neighbor

A queen is going to be fighting for resources with their neighbors, no matter who they are. However, if their neighbor already shares genes with them their neighbor has an increased incentive to play 'nice' with you. Both sides can help spread their own genes by aiding the related queen, by contrast someone that doesn't share genes from you benefits entirely from your death.

If your going to be competing for resources anyways it's better to do it with someone that has some genetic imperative to cooperate with you then one that has none. Better to have your daughters out compete and drive out an unrelated queen then have to deal with the less cooperative queen, even if that means you also have to compete with your daughter.

There will be drones regardless, so no point in handicapping yourself

There are already other queens with drones. Those queens are going to let their drones out to mate. If you don't let your drones out to mate the new virgin queens will mate with the other drones. The queens will still reproduce and start hives, the only difference is that those new haves have none of your genetics! Since new colonies are going to be started regardless of your choices might as well ensure more of your genes are spread when that happens.

You might claim this argument is begging the question how this habit started, why did the queens start releasing drones when they first started to evolve this system? Well you could say that the ancestors of the beepippers (that for some reason I want to call beedrills...) were doing it eons ago and it simple persisted as they evolved and changed into a new species for billions of years the exact same logic, that other's will release drones so I might as well, was sufficient motive to release your own drones and it traces back to an ancient ancestor that had no incentive to not release drones.

The virgin queens didn't really need your drones anyways

If your really basing this off of eusocial insects then they presumably share a single trait that pretty much all eusocial insects have and which was likely the driving factor in development of eusocial insects in the first place, namely they use a haplodiploidy sex determination system.

With this system females can produce males without mating, the males get exactly half the genes from their mother and no genes from their non-existent father. A female is produced from mating, getting a random selection of half the females genes an all the male's genes, since he only has one copy of each gene. This means that a workers may share as much as 75% of their genetics (50% of their mothers and 100% of their fathers) making sisters more closely related to each other then their mother or their own offspring. This oddity of sisterhood is likely why these insects became eusocial, there was a much stronger incentive to aid your sisters then to add your children since your more closely related to them, thus the reason the insects started working together (that's massively oversimplifying it but good enough for now).

The point of all this is that the virgin queens presumably can produce sons without needing to mate. If already present queens really refused to release drones the virgin queens would just produce their own drones and have a mating flight with them. Perhaps in a modern beepippers it would take too long for males to reach maturity, but in their evolutionary past males reached it very rapidly and this would have been a viable option driving the original decision for existing queens to release their drones. In fact it's kind of doubtful modern beepippers would have drones that take so long to reach maturity and cost so much resources to raise given how rapidly drones can/are produced in most eusocial insects of today.

Your daughters aren't a threat because they can't stand to be near you

Practically every species out there has some form of inbreeding avoidance techniques, since inbreeding is genially bad. Now admittedly inbreeding is actually far less of a problem for haplodiploidy insects (since males don't have a second copy of each gene bad recessive genes tend to be quickly weeded out of the gene pool when they cause the male to be non-viable). Still there are reasons, including simple preference for genetic diversity, that inbreeding is still preferentially avoided.

You said local queens trade drones. If a local queen was your daughter then their drones would be your grandchild and inbreeding would happen regularly when you traded drones. To avoid this queens would likely do what most animals in nature do to avoid inbreeding and disperse. Your daughters would not stay local to you, they would intentionally fly far far away from you to avoid inbreeding. Thus the colonies they start would be competing with other queens, not you. The natural result of inbreeding avoidance is that you don't have to worry about competition with your daughters.

For the record this wouldn't be true for drones. Usually only one sex bothers to disperse in nature and given the drone trading strategies and that drones are solely the children of their queen without a father it makes more sense for females to be the ones that disperse to avoid inbreeding.

If you really want daughters to stay local to the queen you may have a bit of an out here by arguing the harm of inbreeding is legitimately so much lower for haplodiploidy insects relative to most animals that the advantage of staying local to cooperate with genetically related queens outweigh it, but that still implies cooperation, not competition, between hives if daughters stay local.

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You already answered your question. The reason why she wouldn't kill them is because she knows that if she does, her species will die out. The drones are of great value to her; both in keeping her species alive and in market trade. She cannot let them go or let other queens take them from her. You could say they're precious.

Generated with minimal supervision by AI.

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