Relationship between environmental variables measured in the field and family-mean trait values: between distance from trees and rosette size under control conditions (A), dune position and time to flowering under control conditions (0 = top of dunes, 1 = bottom of dunes) (B), density of A. lyrata and time to flowering (C), and per cent coverage by herbs and grass and frost resistance after acclimation (D). Symbols depict family means (n = 40) and lines the linear regression relationship.