This write-up really points the finger at not solving occlusion culling or having good LOD discipline.
Give a person a dedicated optimization mandate and you can avoid most of this. One of the first things I do when I'm profiling is to sort assets by tris count and scan for excess. I wonder if they had somebody go through and strategically disable shadowcasting on things like those teeth? I am guessing that they made optimization "everybody's responsibility" but nobody had it as their only responsibility.