Future species composition will affect forest water use after loss of eastern hemlock from southern Appalachian forests.

Infestation of eastern hemlock (Tsuga canadensis (L.) Carr.) with hemlock woolly adelgid (HWA, Adelges tsugae) has caused widespread mortality of this key canopy species throughout much of the southern Appalachian Mountains in the past decade. Because eastern hemlock is heavily concentrated in riparian habitats, maintains a dense canopy, and has an evergreen leaf habit, its loss is expected to have a major impact on forest processes, including transpiration (E(t)). Our goal was to estimate changes in stand-level E(t) since HWA infestation, and predict future effects of forest regeneration on forest E(t) in declining eastern hemlock stands where hemlock represented 50-60% of forest basal area. We used a combination of community surveys, sap flux measurements, and empirical models relating sap flux-scaled leaf-level transpiration (E(L)) to climate to estimate the change in E(t) after hemlock mortality and forecast how forest E(t) will change in the future in response to eastern hemlock loss. From 2004 to 2011, eastern hemlock mortality reduced annual forest E(t) by 22% and reduced winter E(t) by 74%. As hemlock mortality increased, growth of deciduous tree species--especially sweet birch (Betula lenta L.), red maple (Acer rubrum L.), yellow poplar (Liriodendron tulipifera L.), and the evergreen understory shrub rosebay rhododendron (Rhododendron maximum L.)--also increased, and these species will probably dominate post-hemlock riparian forests. All of these species have higher daytime E(L) rates than hemlock, and replacement of hemlock with species that have less conservative transpiration rates will result in rapid recovery of annual stand E(t). Further, we predict that annual stand E(t) will eventually surpass E(t) levels observed before hemlock was infested with HWA. This long-term increase in forest E(t) may eventually reduce stream discharge, especially during the growing season. However, the dominance of deciduous species in the canopy will result in a permanent reduction in winter E(t) and possible increase in winter stream discharge. The effects of hemlock die-off and replacement with deciduous species will have a significant impact on the hydrologic flux of forest transpiration, especially in winter. These results highlight the impact that invasive species can have on landscape-level ecosystem fluxes.

Can forest management be used to sustain water-based ecosystem services in the face of climate change?

Forested watersheds, an important provider of ecosystems services related to water supply, can have their structure, function, and resulting streamflow substantially altered by land use and land cover. Using a retrospective analysis and synthesis of long-term climate and streamfiow data (75 years) from six watersheds differing in management histories we explored whether streamflow responded differently to variation in annual temperature and extreme precipitation than unmanaged watersheds. We show significant increases in temperature and the frequency of extreme wet and dry years since the 1980s. Response models explained almost all streamflow variability (adjusted R2 > 0.99). In all cases, changing land use altered streamflow. Observed watershed responses differed significantly in wet and dry extreme years in all but a stand managed as a coppice forest. Converting deciduous stands to pine altered the streamflow response to extreme annual precipitation the most; the apparent frequency of observed extreme wet years decreased on average by sevenfold. This increased soil water storage may reduce flood risk in wet years, but create conditions that could exacerbate drought. Forest management can potentially mitigate extreme annual precipitation associated with climate change; however, offsetting effects suggest the need for spatially explicit analyses of risk and vulnerability.

Tsuga canadensis (L.) Carr. mortality will impact hydrologic processes in southern Appalachian forest ecosystems.

Eastern hemlock (Tsuga canadensis (L.) Carr.) is one of the principal riparian and cove canopy species in the southern Appalachian Mountains. Throughout its range, eastern hemlock is facing potential widespread mortality from the hemlock woolly adelgid (HWA). If HWA-induced eastern hemlock mortality alters hydrologic function, land managers will be challenged to develop management strategies that restore function or mitigate impacts. To estimate the impact that the loss of this forest species will have on the hydrologic budget, we quantified and modeled transpiration over a range of tree sizes and environmental conditions. We used heat dissipation probes, leaf-level gas-exchange measurements, allometric scaling, and time series modeling techniques to quantify whole-tree and leaf-level transpiration (E(L)) of eastern hemlock. We monitored trees ranging from 9.5 to 67.5 cm in diameter along a riparian corridor in western North Carolina, USA during 2004 and 2005. Maximum rates of daily tree water use varied by diameter and height, with large trees transpiring a maximum of 178-186 kg H2O x tree(-1) x d(-1). Values of E(L) could be predicted from current and lagged environmental variables. We forecasted eastern hemlock E(L) for inventoried stands and estimated a mean annual transpiration rate of 63.3 mm/yr for the hemlock component, with 50% being transpired in the winter and spring. In typical southern Appalachian stands, eastern hemlock mortality would thus reduce annual stand-level transpiration by approximately 10% and reduce winter and spring stand-level transpiration by approximately 30%. Eastern hemlock in the southern Appalachians has two distinct ecohydrological roles: an evergreen tree that maintains year-round transpiration rates and a riparian tree that has high transpiration rates in the spring. No other native evergreen in the southern Appalachians will likely fill the ecohydrological role of eastern hemlock if widespread mortality occurs. With the loss of this species, we predict persistent increases in discharge, decreases in the diurnal amplitude of streamflow, and increases in the width of the variable source area.

Soil DIC uptake and fixation in Pinus taeda seedlings and its C contribution to plant tissues and ectomycorrhizal fungi.

Plants can acquire carbon from sources other than atmospheric carbon dioxide (CO(2)), including soil-dissolved inorganic carbon (DIC). Although the net flux of CO(2) is out of the root, soil DIC can be taken up by the root, transported within the plant, and fixed either photosynthetically or anaplerotically by plant tissues. We tested the ability of Pinus taeda L. seedlings exposed to (13)C-labeled soil DIC and two NH(4)(+) availability regimes to take up and fix soil DIC. We also measured the concentration and distribution of the fixed soil DIC within the plant and mycorrhizal tissues, and quantified the contribution of soil DIC to whole-plant carbon (C) gain. Seedlings exposed to labeled DIC were significantly enriched in (13)C compared with seedlings exposed to unlabeled DIC (6.7 versus -31.7 per thousand). Fixed soil DIC was almost evenly distributed between above- and belowground biomass (55 and 45%, respectively), but was unevenly distributed among tissues. Aboveground, stem tissue contained 65% of the fixed soil DIC but represented only 27% of the aboveground biomass, suggesting either corticular photosynthesis or preferential stem allocation. Belowground, soil DIC had the greatest effect (measured as (13)C enrichment) on the C pool of rapidly growing nonmycorrhizal roots. Soil DIC contributed approximately 0.8% to whole-plant C gain, and approximately 1.6% to belowground C gain. We observed a slight but nonsignificant increase in both relative C gain and the contribution of soil DIC to C gain in NH(4)(+)-fertilized seedlings. Increased NH(4)(+) availability significantly altered the distribution of fixed soil DIC among tissue types and increased the amount of fixed soil DIC in ectomycorrhizal roots by 130% compared with unfertilized seedlings. Increased NH(4)(+) availability did not increase fixation of soil DIC in nonmycorrhizal roots, suggesting that NH(4)(+) assimilation may be concentrated in ectomycorrhizal fungal tissues, reflecting greater anaplerotic demands. Soil DIC is likely to contribute only a small amount of C to forest trees, but it may be important in C fixation processes of specific tissues, such as newly formed stems and fine roots, and ectomycorrhizal roots assimilating NH(4)(+).

Diurnal and seasonal variability in the radial distribution of sap flow: predicting total stem flow in Pinus taeda trees.

We monitored the radial distribution of sap flux density (v; g H2O m(-2) s(-1)) in the sapwood of six plantation-grown Pinus taeda L. trees during wet and dry soil periods. Mean basal diameter of the 32-year-old trees was 33.3 cm. For all trees, the radial distribution of sap flow in the base of the stem (i.e., radial profile) was Gaussian in shape. Sap flow occurred maximally in the outer 4 cm of sapwood, comprising 50-60% of total stem flow (F), and decreased toward the center, with the innermost 4 cm of sapwood (11-15 cm) comprising less than 10% of F. The percent of flow occurring in the outer 4 cm of sapwood was stable with time (average CV < 10%); however, the percentage of flow occurring in the remaining sapwood was more variable over time (average CV > 40%). Diurnally, the radial profile changed predictably with time and with total stem flow. Seasonally, the radial profile became less steep as the soil water content (theta) declined from 0.38 to 0.21. Throughout the season, daytime sap flow also decreased as theta decreased; however, nighttime sap flow (an estimate of stored water use) remained relatively constant. As a result, the percentage of stored water use increased as theta declined. Time series analysis of 15-min values of F, theta, photosynthetically active radiation (PAR) and vapor pressure deficit (D) showed that F lagged behind D by 0-15 min and behind PAR by 15-30 min. Diurnally, the relationship between F and D was much stronger than the relationship between F and PAR, whereas no relationship was found between F and theta. An autoregressive moving average (ARIMA) model estimated that 97% of the variability in F could be predicted by D alone. Although total sap flow in all trees responded similarly to D, we show that the radial distribution of sap flow comprising total flow could change temporally, both on daily and seasonal scales.

Assessing variation in the radial profile of sap flux density in Pinus species and its effect on daily water use.

We monitored sap flux density (v) diurnally in nine mature southeastern pine (Pinus spp.) trees with a thermal dissipation probe that spanned the sapwood radius. We found the expected pattern of high v near the cambium and decreasing v with depth toward the center of the tree; however, the pattern was not constant within a day or between trees. Radial profiles of trees were steeper earlier in the day and became less steep later in the day. As a result, time-dependent changes in the shape of the radial profile of v were sometimes correlated with daily changes in evaporative demand. As the radial profile became less steep, the inner xylem contributed relatively more to total tree sap flow than it did earlier in the day. We present a 3-parameter Gaussian function that can be used to describe the radial distribution of v in trees. Parameters in the function represent depth in the xylem from the cambium, maximum v, depth in the xylem where maximum v occurs, and the rate of radial change in v with radial depth (beta). Values of beta varied significantly between trees and with time, and were sometimes correlated with air vapor pressure deficit (D). We hypothesize that this occurred during periods of high transpiration when the water potential gradient became great enough to move water in the inner sapwood despite its probable high hydraulic resistance. We examined discrepancies among estimates of daily water use based on single-point, two-point and multi-point (i.e., every 20 mm in the sapwood) measurements. When radial distribution of v was not considered, a single-point measurement resulted in errors as large as 154% in the estimate of daily water use relative to the estimate obtained from a multi-point measurement. Measuring v at two close sample points (10 and 30 mm) did not improve the estimate; however, estimates derived from v measured at two distant sample points (10 and 70 mm) significantly improved the estimate of daily water use, although errors were as great as 32% in individual trees. The variability in v with depth in the xylem, over time, and between trees indicates that measurements of the radial distribution of v are necessary to accurately estimate water flow in trees with large sapwood areas.