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

Abstract

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 lifehistory 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 HWAinfested 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.