Seasonal variation in effects of herbivory on foliar nitrogen of a threatened conifer.

Invasive herbivores can dramatically impact the nitrogen (N) economy of native hosts. In deciduous species, most N is stored in stem tissues, while in evergreen conifer species N is stored in needles, making them potentially more vulnerable to herbivory. In eastern forests of the USA, the long-lived, foundational conifer eastern hemlock (Tsuga canadensis) is under the threat of extirpation by the invasive hemlock woolly adelgid (HWA: Adelges tsugae). We assessed the impact of HWA infestation on the patterns of seasonal foliar N availability in hemlock planted in a deciduous forest understory. Over the course of a year, we sampled needles and twigs and measured N, carbon (C), C:N ratio, and total protein concentrations. Tissue sampling events were timed to coincide with key life-history transitions for HWA to determine the association between HWA development and feeding with these foliar nutrients. In uninfested trees, needle and twig N concentrations fluctuated across seasons, indicating the potential importance of N storage and remobilization for the N economy of eastern hemlock. Although N levels in HWA-infested trees also cycled annually, the degree to which N concentrations fluctuated seasonally in tissues was significantly affected by HWA feeding. These fluctuations exceeded N levels observed in control trees and coincided with HWA feeding. HWA feeding generally increased N concentrations but did not affect protein levels, suggesting that changes in N do not occur via adelgid-induced protein breakdown. Herbivore-induced mobilization of N to feeding sites and its rapid depletion may be a significant contributor to eastern hemlock mortality in US forests.