9.N23 and SI1407-N23 (RLSE). MAN. acknowledges support from Office of Naval Research grant N00014.16.1.2914 and from the John Templeton Foundation. The Program fin Evolutionary Dynamics is supported in part by a gift from B Wu and E. Larson. Author contributions A.P.. IT.. K.C.. and M.A.N. performed the research and wrote and revi