A fresh assessment of reproductive behaviour and anatomy, combined with new allozyme data, results in a phylogenetic hypothesis for the 23 polygyrid genera that differs substantially from previous attempts. Analyses based on this phylogeny suggest that (a) evolutionary transition from internal to external sperm exchange occurred only once in polygyrids, with unusual functional intermediates still extant; (b) polygyrid biogeographic history paralleled that of plethodontid salamanders, with hypothesized vicariance events at about 145, 120, 65 and 40 MA, and, dispersal events at about 55 and 40 MA; and (c) iterative shell evolution occurred despite a phylogenetic constraint on ontogenetic whorl-expansion rate.