I had to recalculate the "Bone Roll" of Piero's "Wing Feathers" to "Global Positive Y":
Since earlier tests weren't successful, I first turned off all "Bone Constraints".
After that I made Blender recalculate the "Bone Roll":
1) Result in "Edit Mode":
2) Result in "Pose Mode":
Did I do it right or did I overcomplicate it?