Defoliated Trees Die Below A Critical Threshold Of Stored Carbon

1. Carbon starvation posits that defoliation- and drought-induced mortality results from drawing down stored non-structural carbohydrates (NSCs), but evidence is mixed, and few studies evaluate mortality directly. We tested the relationships among defoliation severity, NSC drawdown and tree mortality by measuring NSCs in mature oak trees defoliated by an invasive insect, Lymantria dispar, across a natural gradient of defoliation severity.2. We collected stem and root samples from mature oaks (Quercus rubra and Q. alba) in interior forests (n = 34) and forest edges (n = 47) in central Massachusetts, USA. Total NSC (TNC; sugar + starch) stores were analysed with respect to tree size, species and defoliation severity, which ranged between 5% and 100%.3. TNC stores declined significantly with increasingly severe defoliation. Forest edge trees had higher TNC stores that were less sensitive to defoliation than interior forest trees, although this may be a result of differing defoliation history. Furthermore, we observed a mortality threshold of 1.5% dry weight TNC.4. Our study draws a direct link between insect defoliation and TNC reserves and defines a TNC threshold below which mortality is highly likely. These findings advance understanding and improve model parametrization of tree response to insect outbreaks, an increasing threat with globalization and climate change. A free Plain Language Summary can be found within the Supporting Information of this article.