We evaluated the 14 WHR-associated loci for their relationships with related metabolic traits using GWA data provided by trait-specific consortia19–21 as well as our de novo genotyped follow-up studies. As expected, given the sample overlap between this GWA data with our WHR GWA data as well as known trait correlations (Supplementary Table 4), we observed directionally consistent enrichment of associations (P < 0.05) between the 14 WHR-associated alleles and increased triglycerides, LDL-cholesterol, fasting insulin, and HOMA-derived measures of insulin resistance (binomial P from 3.2 × 10−4 to 1.8 × 10−8; Table 3 and Supplementary Table 5a). For example, the WHR-increasing allele at GRB14 shows strong associations with increased triglycerides (P = 7.4 × 10−9), fasting insulin levels (P = 5.0 × 10−6) and insulin resistance (P = 1.9 × 10−6). Eleven of 14 WHR-associated loci showed directionally consistent associations with T2D, three of these (ADAMTS9, NISCH-STAB1, and ITPR2-SSPN) reaching nominal significance (P < 0.05) (Table 3 and Supplementary Table 5a,b). Because the association signals for correlated traits in this analysis are vulnerable to overestimation given the overlap in GWAS samples
