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Thank you for visiting nature. You are using a browser version with limited support for CSS. To obtain the best experience, we recommend you use a more up to date browser or turn off compatibility mode in Internet Explorer. In the meantime, to ensure continued support, we are displaying the site without styles and JavaScript. These two types are found in localized groups, separated geographically by a boundary area predicted to lie somewhere around Lake Biwa in central Japan.

To determine this precise boundary, the heterogametic sex of 18 populations around Lake Biwa was examined by genotyping sex-linked genes. Phylogenetic relationships between the populations were also analyzed using mitochondrial 12S rRNA gene. As the main ZW group inhabiting northwestern Japan is known to have a different male heterogametic origin, this finding shows that change of heterogametic sex from male to female may have occurred twice, and independently, during the frog speciation.

In vertebrates, the two types are shared by the taxonomic groups and, particularly in lower vertebrates, they are shared even by closely related species Bull, ; Ezaz et al. It has been shown in amphibians that the heterogametic sex was originally female, and subsequently changed more than once to male at certain branching points in amphibian phylogenetic evolution Hillis and Green, In addition, there is evidence that the heterogametic sex returned to its previous female state in the frog Rana rugosa when its distribution range expanded across Japan Miura et al.

Therefore, it is plausible that heterogametic sex is not permanently fixed, but can change during evolution and speciation. To understand the evolutionary mechanisms of sex determination, it is important to determine the evolutionary reasons for and mechanisms of heterogametic sex change. Japanese Rana populations are divided into four genetic groups that inhabit four different geographic regions Figure 1. These groups differ in both sex determination and sex chromosome differentiation.