Face bones lose offset in animations

So I have my character’s face rigged with an armature, like I normally do (It lets me do a wider range of expressions than using blend shapes or shape keys or whatever they’re called.)

So after exporting, when the face bones aren’t animated, everything looks fine:

35256-offset.png

But when an animation has any of these bones moved, even something as simple as rotating the eyes, suddenly every face bone loses its offset, resulting in this:

35257-no+offset.png

I have checked and the bone positions are set in every single animation.

If it helps, whenever importing the model, it tells me the “Bone transform is different than original” error, so I figure that might be related to this problem.