The samples were assembled and genotyped as part of the larger POPRES project currently consisting of ~6,000 individuals from worldwide populations7. The subsample of European individuals used here is derived from two independent collections: the London Life Sciences Population (LOLIPOP) study26, which consists mainly of European individuals sampled in London, and (2) the CoLaus study27, which represents a broad set of European individuals sampled from Lausanne, Switzerland. The combined sample contains individuals with origins from across Europe (Supplementary Table 2), although origins from eastern Europe are generally less well represented (for example, Finland, Latvia, Ukraine, Slovakia and Slovenia) and some countries are not sampled at all (for example, Belarus, Estonia, Lithuania and Moldova).