Analysis of molecular variance (AMOVA)24, population pairwise genetic distances, and Mantel tests25 were performed with the package Arlequin 3.11.26 Admixture analyses were carried out with Admix2_0.27 Median-joining networks28 were calculated with Network 4.2 (Fluxus-Engineering). Such networks were highly reticulated, and we reduced reticulations by first weighting the loci according to the inverse of their variance in the dataset used29 and subsequently constructing a reduced-median network30 to form the input of the median-joining network. Male effective population sizes were calculated with BATWING31 with a demographic model that assumed a period of constant size followed by exponential growth; prior values were set for other parameters as described previously.20