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Research Highlights: Excess N from atmospheric deposition has been shown to decrease plant biodiversity of impacted forests, especially in its effects on herbaceous layer communities. This work demonstrates that one of the mechanisms of such response is in N-mediated changes in the response of herb communities to soil resources and light availability. Background and Objectives: Numerous studies in a variety of forest types have shown that excess N can cause loss of biodiversity of herb layer communities, which are typically responsive to spatial patterns of soil resource and light availability. The objectives of this study were to examine (1) gradients of temporal change in herb composition over a quarter century, and (2) spatial patterns of herb cover and diversity and how they are influenced by soil resources and canopy structure. Materials and Methods: This study used two watersheds (WS) at the Fernow Experimental Forest, West Virginia, USA: WS4 as an untreated reference and WS3 as treatment, receiving 35 kg N/ha/yr via aerial application. Herb cover and composition was measured in seven permanent plots/WS from 1991 to 2014. In 2011, soil moisture and several metrics of soil N availability were measured in each plot, along with measurement of several canopy structural variables. Backwards stepwise regression was used to determine relationships between herb cover/diversity and soil/canopy measurements. Results: Herb diversity and composition varied only slightly over time on reference WS4, in contrast to substantial change on N-treated WS3. Herb layer diversity appeared to respond to neither soil nor canopy variables on either watershed. Herb cover varied spatially with soil resources on WS4, whereas cover varied spatially with canopy structure on WS3. Conclusions: Results support work in many forest types that excess N can decrease plant diversity in impacted stands. Much of this response is likely related to N-mediated changes in the response of the herb layer to soil N and light availability.

Excess Nitrogen in Temperate Forest Ecosystems Decreases Herbaceous Layer Diversity and Shifts Control from Soil to Canopy Structure