> "Must" is a pretty strong word here Pete, especially since I know that you understand epigenetics.

I don't think anyone understands epigenetics, and the genetic underpinning of behavior is only partially understood. On the other hand, there is a mountain of evidence that particular behavioral traits are heritable (hygienic behavior, aggression, susceptibility to disease). 

And no evidence that I can think of where a specific behavior can be shown to be passed from one colony to another, or even from bees in the same colony via some sort of epigenetic mechanism. Unless you include learned behaviors, but that isn't really what is generally meant by epigenetic, which conventionally refers to cellular processes such as methylation. 

It may be discovered that learning also involves marking or transformation of dna as well, but to date we believe that learning is principally a neurological process, and not heritable in the conventional sense. Epigenetics tends to be used as a catch-all for anything in the gray area between inheritance and environmental influence. 

PLB

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