• Y.L., W.R.T., E.V.L., F.X.C., D.M.L. and T.B. are equally contributing second authors; and B.F., S.J.H., Z.H., G.M.H., M.F.J., B.K.L., M.M., E.W.M., M. Pippel, S.J.P. and N.B.S. are equally contributing third authors, listed alphabetically. This study was conceptualized and led by the five corresponding authors: E.C.T., M. Hiller, L.M.D., S.C.V. and D.A.R. who led the scientific direction, manuscript writing and editing. Together with the corresponding authors, 18 co-authors led the data analyses and interpretation detailed in the Supplementary Notes and contributed to writing and editing: A.E.M., Y.L., W.R.T., E.V.L., F.X.C., D.M.L., T.B., B.F., S.J.H., Z.H., G.M.H., M.F.J., B.K.L., M. Mai, E.W.M., M. Pippel, S.J.P. and N.B.S. All authors had the opportunity to review and edit the manuscript. Z.A.A., I.A.V.T., D.V.A., L.M.A.H., P.B., A.V.B., J.B., N.C., W.O.Y.C., L.M.D., D.K.N.D., A.D., D.E., M.D.E., I.G., A.M.G., S.M.G., E.G.G., L.H., M.R.d.W., D.M.J., S.A.J., M.K., J.E.K., M.E.L., M.L., B.K.L., B.D.L., L.L., V.J.M., N.N., H.N., E.E.N., Z.N., S.H.O., A.O., B.A.O., J.O., K.L.P., M.Pieri, S.J.P., P.P.-S., D.A.R., B.R.-H., I.R., S.J.R., C.S., S.N.S., N.B.S., A.S., E.L.S., P.H.S., R.M.T., N.S.U., J.M.V., S.C.V., L.-F.W., L.C.W., D.W., Y.W., L.R.Y., H.Z., J.L.E., S.J.E., J.F., M.A.G., T.W.H., M.C.M.G., M.N., D.R., F.S., P.S., W.O.M., E.C.T., M. Hiller, M. Mai and P.V. acquired samples and data. S.W., D.W.M., C.G., A.B.H., A.M.G., D.K.L., E.H., F.J.P.-L., J.I.C., J.E.G., K.A.N., L.C., M. Meyer, M.W.G., M.Z., N.J.G., N.Z., W.O.Y.C., W.S.P., Y.G., S.S., N.A., B.P.O., L.-F.W., S.C.V., E.C.T. and M. Hiller performed genome sequencing. T.B., E.V.L., M. Pippel, T.S., L.A.L.A., K.M., S.T., B.K., G.F., E.D.J., L.U., E.W.M., S.C.V., E.C.T., M.B., M. Hiller and B.P.O. conducted the genome assembly. A.E.M., E.V.L., D.A.R. and M. Hiller performed the assembly statistics. A.E.M., A.G.-I., Y.V.M., L.H., S.L., D.-G.K., E.V.L. and M. Hiller conducted genome annotation and pairwise genome alignments. D.A.R. and L.M.D. performed the transposable element annotation. B.F., Z.H., S.P., M.U., M. Hackenberg, M. Mai and S.C.V. conducted the microRNA analyses. A.E.M., E.V.L., F.X.C., D.A.R. and M. Hiller performed the multiple genome alignments. A.E.M., E.C.T., M. Hiller, A.G.-I., G.M.H., S.J.P., Y.L., N.M.F. and W.J.M. undertook the phylogenomics. M. Prylutskyi, A.E.M., M. Hiller, H.A.L. and D.M.L. performed the ancestral karyotype reconstruction. W.R.T., M.F.J., F.X.C., G.M.H., S.J.H., N.B.S., L.M.D. and E.C.T. contributed fossils, total evidence and node divergence times, and biogeography. A.B.H., T.B., P.D., G.F., A.G.-I., J.G., M.F.J., B.K., M.E.L., E.V.L., Y.L., M. Mai, Y.V.M., A.E.M., M. Pippel, T.S., N.B.S., W.R.T., M. Muffato, G.Q., L.U., E.C.T., M. Hiller, L.M.D., S.C.V. and D.A.R. curated the data and performed the archving.