In the cross-ancestry meta-analysis of all available datasets, we identified 100 independent variants at 90 loci (Fig. 1f and Supplementary Table 1); 80 have not been previously reported in association with PAU. Of these, 53 variants were in protein-coding genes, of which 9 are missense variants: GCKR*rs1260326; ADH1B*rs75967634, rs1229984 and rs2066702; SCL39A8*rs13107325; OPRM1*rs1799971; SLC25A37*rs2942194; BDNF*rs6265 and BRAP*rs3782886. The cross-ancestry meta-analysis identified 24 more risk variants than the EUR meta-analysis, but 9 EUR variants fell below GWS (P values ranging from 5.26 × 10−6 to 9.84 × 10−8). In total, 110 unique variants were associated with PAU in either the within-ancestry or cross-ancestry analyses (Fig. 1b and Supplementary Table 1).