Genome sequences of five Sitopsis species of Aegilops and the origin of polyploid wheat B subgenome

Common wheat(Triticum aestivum,BBAADD)is a major staple food crop worldwide.The diploid progenitors of the A and D subgenomes have been unequivocally identified;that of B,however,remains ambiguous and controversial but is suspected to be related to species of Aegilops,section Sitopsis.Here,we report the assembly of chromosome-level genome sequences of all five Sitopsis species,namely Aegilops bicornis,Ae.longissima,Ae.searsii,Ae.sharonensis,and Ae.speltoides,as well as the partial assembly of the Amblyopyrum muticum(synonym Aegilops mutica)genome for phylogenetic analysis.Our results reveal that the donor of the common wheat B subgenome is a distinct,and most probably extinct,diploid species that diverged from an ancestral progenitor of the B lineage to which the still extant Ae.speltoides and Am.muticum belong.In addition,we identified interspecific genetic introgressions throughout the evolution of the Triticum/Aegilops species complex.The five Sitopsis species have various assembled genome sizes(4.11-5.89 Gb)with high proportions of repetitive sequences(85.99%-89.81%);nonetheless,they retain high collinearity with other genomes or subgenomes of species in the Triticum/Aegilops complex.Differences in genome size were primarily due to independent post-speciation amplification of transposons.We also identified a set of Sitopsis genes pertinent to important agronomic traits that can be harnessed for wheat breeding.These newly assembled genome resources provide a new roadmap for evolutionary and genetic studies of the Triticum/Aegilops complex,as well as for wheat improvement.