Effects of Atmospheric Nitrogen Deposition on the Herbaceous Layer of a Central Appalachian Hardwood Forest

Additions of nitrogen (N) have been shown to alter species diversity of plant communities, with most experimental studies having been carried out in communities dominated by herbaceous species. We examined seasonal and inter-annual patterns of change in the herbaceous layer of two watersheds of a central Appalachian hardwood forest that differed in experimental treatment. This study was carried out at the Fernow Experimental Forest, West Virginia, using two adjacent watersheds: WS4 (mature, second-growth hardwood stand, untreated reference), and WS3. Seven circular 0.04-ha sample plots were established in eachwatershed to represent its full range of elevation and slope aspect. The herbaceous layer was sampled by identifying and visually estimating cover (%) of all vascular plants. Sampling was carried out in mid-July of 1991 and repeated at approximately the same time in 1992. In 1994, these same plots were sampled each month fromMay to October. Seasonal patterns of herb layer dynamics were assessed for the complete 1994 data set, whereasinter-annual variability was based on plot data from 1991, 1992, and the July sample of 1994. There were nosignificant differences between watersheds for any sample year for any of the other herb layer characteristics measured, including herb layer cover, species richness, evenness, and diversity. Cover on WS4 decreased significantly from 1991 to 1992, followed by no change to 1994. By contrast, herb layer cover did not varysignificantly across years on WS3. Cover of the herbaceous layer of both watersheds increased from early in the growing season to the middle of the growing season, decreasing thereafter, with no significant differencesbetween WS3 and WS4 for any of the monthly cover means in 1994. Similar seasonal patterns found for herblayer cover—and lack of significant differences between watersheds—were also evident for species diversityand richness. By contrast, there was little seasonal change in herb layer species evenness, which was nearlyidentical between watersheds for all months except October. Seasonal patterns for individual species/speciesgroups were closely similar between watersheds, especially for Viola rotundifolia and Viola spp. Species richnessand species diversity were linearly related to herb layer cover for both WS3 and WS4, suggesting that spatialand temporal increases in cover were more related to recruitment of herb layer species than to growth of existingspecies. Results of this study indicate that there have been negligible responses of the herb layer to 6 yr of additions to WS3.

Effects of Harvesting on Herbaceous Layer Diversity of a Central Appalachian Hardwood Forest

Clearcutting is a common harvesting practice in many eastern hardwood forests. Among the vegetation strata of these forests, the herbaceous layer is potentially the most sensitive in its response to harvest-mediated disturbances and has the highest species diversity. Thus, it is important to understand the response of herbaceous layer diversity to forest harvesting. Previous work on clearcut and mature stands at the Fernow Experimental Forest (FEF), West Virginia, has shown that, although, harvesting did not alter appreciably herbaceous layer cover, it influenced the relationship of cover to biotic and abiotic factors, such as tree density and soil nutrients, respectively. The purpose of this study was to examine the response of species diversity of the herbaceous layer to harvesting at FEF. Fifteen circular, 0.04 ha sample plots were established in each of four watersheds (60 plots in total) representing two stand age categories: two watersheds with 20 years even-age stands following clearcutting and two watersheds with mature second growth stands. All woody stems ≥2.5 cm diameter at breast height were identified, tallied, and measured for diameter. The herbaceous layer was sampled by identifying all vascular plants ≤1 m in height and estimating cover for each species in each of 10 (1 m2) circular sub-plots per sample plot (600 sub-plots total). Species diversity for each plot was calculated from herbaceous layer data using the ln-based Shannon Index (H′) equation. Ten stand and soil variables also were measured on each plot. Mean herbaceous layer cover for clearcut versus mature stands was 27.2±14.3% versus 20.2±8.1% (P>0.05), respectively and mean H′ was 1.67±0.42 versus 1.55±0.48 (P>0.05), respectively. Herbaceous layer diversity was negatively correlated with cation exchange capacity and extractable Ca and Mg in the mineral soil in clearcut stands. In contrast, herbaceous layer diversity was positively correlated with soil organic matter and clay content. Although, 20 years of recovery after clearcutting did not have significant effects on the species diversity of the herbaceous layer when examining stand age means alone, harvesting did appear to influence the spatial relationships between herbaceous layer diversity and biotic factors (e.g. tree density) and abiotic factors (e.g. soil nutrients).

The Ecological Significance of the Herbaceous Layer in Temperate Forest Ecosystems

Despite a growing awareness that the herbaceous layer serves a special role in maintaining the structure and function of forests, this stratum remainsan underappreciated aspect of forest ecosystems. In this article I review and synthesize information concerning the herb layer’s structure,composition, and dynamics to emphasize its role as an integral component of forest ecosystems. Because species diversity is highest in the herb layeramong all forest strata, forest biodiversity is largely a function of the herb-layer community. Competitive interactions within the herb layer candetermine the initial success of plants occupying higher strata, including the regeneration of dominant overstory tree species. Furthermore, the herblayer and the overstory can become linked through parallel responses to similar environmental gradients. These relationships between strata varyboth spatially and temporally. Because the herb layer responds sensitively to disturbance across broad spatial and temporal scales, its dynamics canprovide important information regarding the site characteristics of forests, including patterns of past land-use practices. Thus, the herb layer has asignificance that belies its diminutive stature.