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Lumpers will lumps. I'm no taxonimist ot geneticist, but I am always suspicious when such broad lumping is done on the sole basis of mtDNA analysis. The Southern Mapleleaf doesn't get all those extra pustules all over it's sulcus from it's mitocohondrial DNA, it gets them from it nuclear (nucular if you're Republican) DNA . But since analyzing that is way more expensive, most phylogenetic studies are done with mtDNA, hence the shortcomings and doubt. And when I read this sort of stuff, I wonder whatever happened to to notion of sub-species?, or species complexes? I should read more sciency stuff, but it's more fun to muck around looking for - highly variable phenotypes of - critters.
Alright I'll leave as a draft for now. You're correct that they don't have much analysis of anything other than DNA. They briefly mention shape and range, but not sculpture which seems odd to me. Subspecies and species complexes would require distinct clades, which doesn't seem to be the case according to this paper (Q. quadrula is paraphyletic). And yes, whole genome sequencing would be nice to have. :) Mitochondrial analysis does have the benefit of (generally) having a constant mutation rate in cytochrome b.
I found it, it's "A taxonomic study of Quadrula quadrula (Rafinesque)" by J.K. Neel, from 1941.
PDF available.
http://hdl.handle.net/2027.42/56887
It did place Q. fragosa in with Q. quadrula quadrula.
It also placed T. nobilis in with Q. quadrula apiculata (perhaps due to Conrad's own mixed collection in the original paper).
Thanks! Sorry for my late response. It does say that tubercles/pustules are generally distinct and geographically isolated in Q. apiculata, but that that is also highly variable and there are intergrades to Q. quadrula sensu stricto. Given the genetic evidence I think I'll go ahead with the merge here and elsewhere if there are no objections. Best to stay in line with MB anyway.
https://www.inaturalist.org/taxon_merges/92562
@dbarclay @redgarter Thoughts on this swap?