performed in R (R Core Team, 2016). False Discovery Rate (FDR) analysis was performed using the “q-value” package (https://github.com/jdstorey/qvalue) using Bioconductor 3.2 (Huber et al., 2015). To define genomic bins for follow-up, we started with all markers with a q < 0.5. Markers were initially collapsed into bins if they were within 10 kb. Post-hoc inspection showed several adjacent bins <75 kb apart which were then collapsed into the reported bins.