The response of dispersers to ruggedness. The relationship between the selection coefficients (beta values) for ruggedness. In a) dispersers are shown for the before, during and after periods of dispersal. Avoidance of ruggedness is stronger during and after dispersal compared to before dispersal. In b) the long step and short step groups, as derived from segmented regression (Figure 3), are shown for the dispersers during their dispersal period. Avoidance of ruggedness appears stronger for long steps than for short steps.