1) The species are probably not truely separated groups 2) The data (from the obsoleta/allegheniensis study alone) suggests *some* cladism within the entire group based on one gene 3) Studies based on molecular and morphological data are more reliable
But, does he successfully refute the subspecies? Based on his data (alone), I'd have to say it does. I'd like more proof though.
Also, what does convergence (as far as color/appearance) add to this? The suggested clades would imply that the groups converge to darker phenotypes as they go further north. Is this not a factor at all or is it just unlikely? Due to gene flow?