The induction of spinescence in European holly leaves by browsing ungulates

Evidence is presented which suggests that the spinescence of leaves of European holly, Ilex aquifolium, deters feeding by ungulates and is induced by browsing. Spinescence decreased as leaf size increased; hence, spinescence may be achieved by reducing adult leaf size. Holly shrubs with very spiny leaves were browsed less often than less spiny shrubs. In the absence of browsing ungulates during a one year period, the spinescence of leaves of holly shrubs significantly decreased. Browsed shrubs exhibited reduced annual shoot growth, increased branching, and produced smaller leaves with high spinescence. The regrowth on browsed branches of holly trees was characterized by increased leaf spinescence relative to unbrowsed branches. Hence, the induced response was localized, thereby reducing the ability of browsing ungulates to exert selective pressures on holly trees.