General Recommendation: Accept with Minor Revisions

Overall Summary and Thoughts:

I recommend the article for publication with minor revisions, most pertaining to the introduction, which could use more detail and clarification for readers unfamiliar with Nabokov's research.  Although the abstact provides some of those important details, I feel that they should appear in the first pages as well. I have flagged phrases and words in the intro that I think need slight elaboration.  

It needs to be noted in the early paragraphs that Nabokov saw evidence of relationships and migrational patterns later confirmed by DNA analysis. The authors only mention DNA in the intro para, and given the title of the article, the argument (in the abstract) should be stated in the intro paragraphs. 

The first two+ pages are hard to follow because they seem to assume pre-existing knowledge in the reader. I’ve noted where clarifications might be helpful. But beyond this difficult entry, the argument is clear. 

So evidence for his dispersal theory existed prior to the DNA analysis, we just didn’t see it? And Nabokov “saw it” but did not remark on it explicitly enough for others to notice, until now.  

That evidence may have been available only through inference, but valid enough for a philologist or physical anthropologist, I’d say.

The authors mention, without elaboration, that "far from being an opponent of Natural Selection," Nabokov sought to find hard evidence for it. Readers may not be aware of the controversy and confusion about Nabokov's theory of the evolution and utility of wing patterns. Although he may have dismissed some arguments that various "mimicry" patterns might have affected reproductive fitness – “the lie to the seeming resemblance of wing patterns” – he absolutely recognized that genital shape and function likely would affect reproductive fitness. Obviously! He went for the evidence of Natural Selection for reproductive fitness that was the lowest hanging fruit; this is as uncontroversial as can be, even for Darwin's skeptics.  

I would like to know if Nabokov's conclusion that the "sheath around the aedeagus" was a primitive structure, likely lost in succeeding generations, was based on evidence of this structure existing in other primitive butterflies – or better, moths. If so, this would add to your argument that Nabokov wasn't merely a lucky guesser, but an empiricist. 

I am puzzled by the authors' referring to the morphology of genitalia as "arising from sexual selection" near the end of the article.   I think of "sexual selection" by the female as the selection of arbitrary characteristics, that she could detect (see?), that increase the male's fitness. Would not the shape of the genitalia tend to determine, pretty directly, reproductive fitness? That is, the morphology of the genitalia determines a good fit and consequent functionality. This is hardly an arbitrary association with reproductive fitness. The functionality of genitalia would be a clear case of natural selection not sexual selection, which requires interpretation on the part of the female.  

Overall, this is a very well-argued and fascinating paper.