What they saying is that they haven't a clue how colony 'fitness' - thus which drones will get to mate - is determined:
We found clear evidence for an extensive diversity in
male mating success at the colony level. Some colonies
had significantly more matings (about an order of
magnitude) than other drone producing colonies in the
same year. This is remarkable, because we had taken
great care to standardize conditions for the drone
producing colonies. All drone mothers were sister
queens, all host colonies were of equal size and all had
similar numbers of drones in the colony (two covered
frames).
In our experiment the ‘fit’ colonies seem to have
produced males which not only out-competed males
from other colonies in mating, but also were more
successful in post-mating competition. The fit colonies
produced drones with a higher reproductive success than
those of the other, less successful colonies. We do not
know which effect caused the increased success of drones
from specific colonies.
Although the actual mechanism for this post-mating
selection in our study must remain open, this is to our
knowledge the first evidence for differences in the male
reproductive success for honeybees in a natural mating
situation.
LJ
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