participant; (2) one member of each monozygotic twin pair; (3) samples with questionable identity or low read depth (<98% of variant sites at depth ≥ 10×); and (4) samples with consent types inconsistent with analyses presented here. The ‘unrelated’ sample set consisting of 40,722 samples refers to a subset of the 53,831 samples of individuals who are unrelated with a threshold of third degree (less closely related than first cousins), identified using the PC-AiR method84. Exact numbers of samples used in each analysis are listed in Supplementary Table 18.