> I suspect more due to the state of the colony than of the queen.

Yes, well this is sort of like the gene/environment interaction. Actually, genes can't be separated from environment; "genes" do nothing at all unless they are in an environment (the organism, which is also in an environment). 

Same with the queen bee, she is nothing without a colony of bees to carry out her reproductive program, which includes producing new colonies (swarms) and new daughter queens, at the favorable time. 

But this is referenced in Juliana's work. She refers to the colony as the "expanded phenotype" of the queen.

The present study provides evidence that, in honey bees,
queen reproductive potential affects several factors of her
colony’s phenotype. As such, we show that a honey bee
colony may be seen as the 'expanded phenotype' of its
queen. Thus, while the colony may primarily serve as the
unit of selection, the distinction between individual- and
colony-level selection (e.g., Tarpy et al., 2004) may actually
be more blurred.

Rangel, Keller, Tarpy
Insect. Soc. (2013) 60:65–73

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