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The population mean iWUE value increased by 53 to 58% over the study period, much more than the 20 to 30% increase that has been measured in forests [J. Peñuelas, J. G. Canadell, R. Ogaya, Glob. Ecol. Biogeogr. 20, 597-608 (2011)]. Changes were associated with both increased CO2 concentration and increased water stress. Individuals whose lifetimes spanned the entire study period exhibited increases in iWUE that were very similar to the population mean, suggesting that there was significant plasticity within individuals rather than s