Responses of oak seedlings to increased herbivory and drought: a possible trade-off?

Abstract

Anthropogenic disturbances are causing a co-occurring increase in biotic (ungulate herbivory) and abiotic (drought) stressors, threatening plant reproduction in oak-dominated ecosystems. However, we wonder whether herbivory could compensate for the adverse impact of drought by reducing evapotranspiration. Thus, we investigate the isolated and joint effects of herbivory and drought on oak seedlings of two contrasting Mediterranean species that differ in leaf habit and drought resistance. California oak seedlings from the evergreen, and more drought-resistant, Quercus agrifolia and the deciduous Q. lobata (n=387) were assigned to a fully crossed factorial design with herbivory and drought as stress factors. Seedlings were assigned in a greenhouse to 3-4 clipping levels simulating herbivory and 3-4 watering levels, depending on the species. We measured survival, growth, and leaf attributes (chlorophyll, secondary metabolites, leaf area and weight) once a month (May-Sep) and harvested above- and below-ground biomass at the end of the growing season. For both oak species, simulated herbivory enhanced seedling survival during severe drought or delayed its adverse effects, probably due to reduced transpiration resulting from herbivory-induced leaf area reduction and compensatory root growth. Seedlings from the deciduous, and less drought-resistant species, benefitted from herbivory at lower levels of water stress, suggesting different response across species. We also found complex interactions between herbivory and drought on their impact on leaf attributes. In contrast to chlorophyll content which was not affected by herbivory, anthocyanins increased with herbivory - although water stress reduced differences in anthocyanins due to herbivory. Herbivory seems to facilitate Mediterranean oak seedlings to withstand summer drought, potentially alleviating a key bottleneck in the oak recruitment process. Our study highlights the need to consider ontogenetic stages and species-specific traits in understanding complex relationships between herbivory and drought stressors for the persistence and restoration of multi-species oak savannas.