> colonies that do not swarm, despite their experiencing the dilution of queen pheromone due to their huge populations of workers.
This ties in with a topic I was about to raise. I have ten hives at my house and two of them had failing queens this summer. Neither made any attempt to raise a new queen. I do not believe that supersedure nor swarming are tied to quantity of pheromone.
The explanation that the workers perceive a failing queen via a reduction in pheromones doesn't add up, unless the actual value of the queen (laying capacity) and her ability to inhibit queen rearing via pheromones -- are decoupled. In other words, the bees regard her as the reigning queen and fail to replace her, causing the demise of the colony. The flip side is when perfectly good (to us) new queens are superseded by the bees.
Same for swarming. Large colonies in large hives often do not swarm, small colonies in small hives often do. The idea that cohesion is lost due to dilution of queen substances does not account for why and when bees swarm. The only thing that has ever made sense to me, based on what I have observed 40+ years is this statement:
> The success of social insect colonies lies in all members of the society acting in concert and in a well-organized manner. At the foundation of social insect self-organization are sophisticated means of communication, the chemical mode being at the center of it.
-- Conte, Y. L., & Hefetz, A. (2008). Primer pheromones in social hymenoptera. Annu. Rev. Entomol., 53, 523-542.
> A coercive control mechanism and an honest signal are likely to have the same outcome under all biologically relevant circumstances ...
the “queen control” and “queen signal” hypotheses cannot be distinguished empirically.
-- Ronai, I., Vergoz, V., & Oldroyd, B. P. (2016). The mechanistic, genetic, and evolutionary basis of worker sterility in the social Hymenoptera. In Advances in the Study of Behavior (Vol. 48, pp. 251-317). Academic Press.
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