Manuscript Number: 17-1153R1 Author(s): Erin L. Boespflug, Hiroko H. Dodge, Yasushi Higashiuesato, Yuriko Katsumata, David Lahna, Junko Nishihira, Yusuke Ohya, Nutta-on Promjunyakul, Lisa Silbert, Takashi Tokashiki