Broadening the ecological context of ungulate-ecosystem interactions: the importance of space, seasonality, and nitrogen.

Spatial heterogeneity of soil resources, particularly nitrogen availability, affects herbaceous-layer cover and diversity in temperate forest ecosystems. Current hypotheses predict that ungulate herbivores influence nitrogen availability at the stand scale, but how ungulates affect nitrogen availability at finer spatial scales that are relevant to the herb layer is less understood. We tested the hypothesis that ungulate exclusion reduces the spatial complexity of nitrogen availability at neighborhood scales (1-26 m) apart from mean stand scale effects. This outcome was expected due to a lack of ungulate nitrogenous waste deposition within exclosures and seasonally variable ungulate habitat use. To test this hypothesis we examined spatial patterning of ammonium and nitrate availability, herb-layer cover and diversity, and under-canopy solar radiation using geostatistical models. Our study sites included six stands of eastern hemlock (Tsuga canadensis) forest: three where white-tailed deer (Odocoileus virginianus) were excluded and three that were accessible to deer. Where deer were present, patch sizes of ammonium availability, cover, and diversity were smaller compared to deer exclosures, whereas mean site-level effects were not significant. Within deer exclosures cover and solar radiation were more similar in patch size than were cover and nitrogen availability. Our results suggest that browsing ungulates affect spatial patterns of herb-layer cover and diversity through the excretion of nitrogenous wastes in small, discrete patches. Ungulate-excreted nitrogen deposition and herbivory were concentrated in the dormant season, allowing herb-layer plants a greater opportunity to benefit from nitrogen additions. Therefore, the impact of ungulates on nitrogen cycling in forest ecosystems varies with spatial scale and the seasonal timing of ungulate impacts. In this way, ungulates may function as a seasonally dependent link between fine-scale and landscape-level ecological processes.