smooth the estimation until the sum of read depth exceeded the average depth across all bins. The contributions were weighted by the distance to k as well as by the read depth. The smoothed error rate was computed as E = e × n × r, where e is the error rate, r is the reciprocal of the distance to read position being smoothed (for example, 1/2, 1/3) and n is the read depth. The smoothed error rate as a function of read direction, read position and substitution type is presented in Figure S8 in Additional file 1.