The only study-wide significant (P ≤ 2 × 10−9) binary trait association among the non-European populations was between PTVs in HBB and thalassaemia in individuals of South Asian ancestry (P = 2.7 × 10−46; OR = 176.4 (95% CI: 84.1–369.7)) (Supplementary Table 15). We next applied the Cochran–Mantel–Haenszel test to combine the results of the binary trait collapsing analysis across all four studied ancestral groups, including the European population (Methods). This pan-ancestry PheWAS identified 26 unique study-wide significant gene–phenotype associations that were not significant in the European analyses (Fig. 3a, Extended Data Fig. 6a, Supplementary Table 16). Conversely, 20 gene–phenotype associations that were significant in the European analyses did not reach the study-wide significance threshold in the pan-ancestry analysis.