“…Third, current understanding of phylogenetic relationships in the Pungitius species complex resides heavily on mitochondrial DNA sequences (e.g., Bae & Suk, 2015;Takahashi & Goto, 2001;Teacher, Shikano, Karjalainen, & Merilä, 2011;Wang et al, 2015), and only a few studies-all based on either limited geographical or taxon sampling-have utilized nuclear genetic information (Aldenhoven, Miller, Corneli, & Shapiro, 2010;Bae & Suk, 2015;Haglund et al, 1992;Natri, Merilä, & Shikano, 2019;Shikano, Ramadevi, Shimada, & Merilä, 2010;Takahashi et al, 2016;Takata, Goto, & Yamazaki, 1987b;Wang, Shikano, Persat, & Merilä, 2017). Phylogenetic studies based on a large number of nuclear loci are motivated by the well-known limitations of phylogenetic and phylogeographical inferences based on mitochondrial information (e.g., Ballard & Whitlock, 2004;Hey & Machado, 2003;Toews & Brelsford, 2012), as well as by the evidence for weak post-zygotic isolation (Kobayashi, 1959;Takahashi, Nagai, & Goto, 2005;Tsuruta & Goto, 2006;Ziuganov & Gomeluk, 1985), and hybridization in this genus (Denys et al, 2018;Niwa, 1987;Takahashi et al, 2016;Takahashi & Takata, 2000;Wang et al, 2017;Natri et al, 2019).…”