There is no evidence for heterogeneity of effect between populations at HLA-DQB1*04:02 or HLA-B*49:01 (heterogeneity p = 0.8836 and p = 0.7049, Fig 4B), and the data best supports a model in which both HLA-DQB1*04:02 and HLA-B*49:01 modify risk of both paucibacillary and multibacillary leprosy (log10 Bayes factors = 2.22 and 0.43 respectively, Fig 4C and 4D). Here, HLA-DQB1 is linked to leprosy.