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Phylogeography of the East Asian grassland plant, Viola orientalis (Violaceae), inferred from plastid and nuclear restriction site-associated DNA sequencing data


To elucidate the origin and migration history of the “Mansen elements,” a group of temperate grassland plants mainly distributed in northeastern Asia, phylogeographic analyses based on chloroplast DNA markers and double-digest restriction site-associated DNA sequencing (ddRAD-seq) data were performed on Viola orientalis, one of the representative species of the group. Phylogenetic analyses using ddRAD-seq data revealed that the populations of V. orientalis were clustered into five clades, among which the continental clades made of populations from Russia and Korea diverged more than 100,000 years earlier than the Japanese clades. The Japanese clade likely diverged during the last glacial period, followed by a further post-glacial divergence into the Kyushu and the Honshu subclades. Our study demonstrated that V. orientalis originated in the continental area of northeastern Asia and, during the last glacial period, has spread southward through the Korean Peninsula across the Japanese Islands. This finding supports the previously proposed evolutionary hypothesis regarding the origin and migration routes of the Mansen elements.

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We express our sincere thanks to M. Igari, S. Sei, K. Takayama, M. Watanabe, and Y. Yuasa (alphabetical order, title omitted) for sample collection of plant materials. We are grateful for the public institutions that permitted us permission to collect samples. We thank S. Kondo for assistance with ddRAD-seq experiments. We also extend our appreciation for the members of our laboratory in supporting our research. We would like to thank Enago ( for English language editing. Our cordial thanks are going to the referee who kindly gave us advices to revise our manuscript. This study was supported by JSPS KAKENHI Grant Number JP17H03721 and a grant from the Institute of Plant Science and Resources, Okayama University (3037, 3140, & R236).

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Correspondence to Noriyuki Fujii.

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Sata, H., Shimizu, M., Iwasaki, T. et al. Phylogeography of the East Asian grassland plant, Viola orientalis (Violaceae), inferred from plastid and nuclear restriction site-associated DNA sequencing data. J Plant Res 134, 1181–1198 (2021).

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  • DdRAD-seq
  • Genetic structure
  • Grassland
  • Japanese flora
  • Migration history
  • Population genetics