The effect of shading and drought on (a) maximum quantum yield of PSII (Fv:Fm) and (b) efficiency of heat dissipation (i.e. non-photochemical quenching, NPQ) of the four study species. The control is indicated by black bars, drought is indicated by white bars, shade is indicated by dark gray bars, and shade + drought is indicated by light gray bars. Means (±SE) of all species include 16 replicates per treatment, and individual species means (±SE) include 4 replicates per treatment. Means not connected by the same letter are significantly different (Tukey HSD, p<0.05).