Triticum aestivum chromosome "4A" is, like the B genome chromosomes, extensively heterochromatic while the remaining six A genome chromosomes are not. In the presence of the Ph gene it does not pair with any chromosome of einkorn wheats, T. monococcum and T. urartu, the source of the A genome. It is shown here that the same chromosome is also present in T. timopheevii which represents the other evolutionary lineage of wheats. The "4A" chromosomes of T. timopheevii and T. aestivum pair poorly with each other, like the B genome chromosomes of the two lineages, while the remaining A genome chromosomes, except for one arm, pair relatively well. Hence, in both lineages chromosome "4A" has the attributes of the B genome chromosomes, not of the A genome chromosomes. The C-banding pattern of chromosome "4A" of T. aestivum and T. timopheevii closely resembles the C-banding pattern of a chromosome of T. speltoides and less closely chromosome 4B1 of T. sharonense. On the basis of this and other evidence it is concluded that this chromosome was contributed by a species of the section Sitopsis and, consequently, belongs to the B genome. Additionally, there is evidence that the chromosome that was originally designated "4B" belongs to the A genome.