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Abstract The wheat wild relativeAegilops tauschiiwas previously used to transfer theLr42leaf rust resistance gene into bread wheat.Lr42confers resistance at both seedling and adult stages, and it is broadly effective against all leaf rust races tested to date.Lr42has been used extensively in the CIMMYT international wheat breeding program with resulting cultivars deployed in several countries. Here, using a bulked segregant RNA-Seq (BSR-Seq) mapping strategy, we identify three candidate genes forLr42. Overexpression of a nucleotide-binding site leucine-rich repeat (NLR) gene AET1Gv20040300 induces strong resistance to leaf rust in wheat and a mutation of the gene disrupted the resistance. TheLr42resistance allele is rare inAe. tauschiiand likely arose from ectopic recombination. Cloning ofLr42provides diagnostic markers and over 1000 CIMMYT wheat lines carryingLr42have been developed documenting its widespread use and impact in crop improvement.