ratio. For each phenotype, meta-analyses of the results across the different cohorts were conducted in METAL with weights proportional to the square-root of the sample size for each study22. The effective sample size of each cohort was calculated based on the case-control ratio and the relatedness matrix. Ancestry-specific (African-ancestry and European-ancestry) and trans-ancestry meta-analyses were conducted. Heterogeneity was evaluated across all cohorts and between study designs.