Latitudinal resource gradient shapes multivariate defense strategies in a long-lived shrub.

Plant defense against herbivores is multidimensional, and investment into different defense traits is intertwined due to genetic, physiological, and ecological costs. This relationship is expected to generate a trade-off between direct defense and tolerance that is underlain by resource availability, with increasing resources being associated with increased investment in tolerance and decreased investment in direct resistance. We tested these predictions across populations of the shrub Artemisia californica by growing plants sourced from a latitudinal aridity gradient within common gardens located at the southern (xeric) and northern (mesic) portions of its distribution. We measured plant growth rate, resistance via a damage survey, and tolerance to herbivory by experimentally simulating vertebrate herbivory. Plants from more northern (vs. southern) environments were less resistant (received higher percent damage by vertebrate herbivores) and tended to be more tolerant (marginally significant) with respect to change in biomass measured 12 months after simulated vertebrate herbivory. Also, putative growth and defense traits paralleled patterns of resistance and tolerance, such that leaves from northern populations contained lower concentrations of terpenes and increased N, specific leaf area, and % water. Last, plant growth rate did not demonstrate clear clinal patterns, as northern populations (vs. southern populations) grew more slowly in the southern (xeric) garden, but there was no clinal relationship detected in the northern (mesic) garden. Overall, our findings support the prediction of lower resistance and higher tolerance in plant populations adapted to more resource-rich, mesic environments, but this trade-off was not associated with concomitant trade-offs in growth rate. These findings ultimately suggest that plant adaptation to resource availability and herbivory can shape intraspecific variation in multivariate plant defenses.