One of these SNPs, rs4846567, is in linkage disequilibrium (LD, r2 = 0.64, D’ = 0.84; HapMap CEU) with the previously reported WHR-associated variant (rs2605100) near the LYPLAL1 gene13. The remaining 13 loci were in or near genes not previously associated with WHR or other measures of adiposity: RSPO3, VEGFA, TBX15-WARS2, NFE2L3, GRB14, DNM3-PIGC, ITPR2-SSPN, LY86, HOXC13, ADAMTS9, ZNRF3-KREMEN1, NISCH-STAB1, and CPEB4 (Figure 2). These 14 loci explain 1.03% of the variance in WHR (after adjustment for BMI, age, and sex), with each locus contributing from 0.02% (ZNRF3-KREMEN1) to 0.14% (RSPO3) based on effect estimates in the follow-up stage.