More recent work by Nadja Steinmann also decouples long lifespan from pollen intake:

> Long-lived winter bees begin to be produced in August and September in temperate climate conditions until brood rearing ceases, usually at the end of October. To overwinter, they form the winter cluster enabling thermoregulation of colonies. From February on, they start to show a division of labor, similar to summer bees, to rear the new generation of summer worker bees.

Steinmann, Nadja, et al. "Overwintering is associated with reduced expression of immune genes and higher susceptibility to virus infection in honey bees." PloS one 10.6 (2015): e0129956.

[Comment: Our main honey and pollen flow takes place in August and September. Some hives even swarm at this time. On the other hand, in February there is no pollen coming in. Not until late March at the earliest is there significant pollen inflow. Some other signal must control the raising of long and short lived bees. ]

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