Chelcy R. Ford

Loss of foundation species: consequences for the structure and dynamics of forested ecosystems

In many forested ecosystems, the architecture and functional ecology of certain tree species define forest structure and their species-specific traits control ecosystem dynamics. Such foundation tree species are declining throughout the world due to introductions and outbreaks of pests and pathogens, selective removal of individual taxa, and over-harvesting. Through a series of case studies, we show that the loss of foundation tree species changes the local environment on which a variety of other species depend; how this disrupts fundamental ecosystem processes, including rates of decomposition, nutrient fluxes, carbon sequestration, and energy flow; and dramatically alters the dynamics of associated aquatic ecosystems. Forests in which dynamics are controlled by one or a few foundation species appear to be dominated by a small number of strong interactions and may be highly susceptible to alternating between stable states following even small perturbations. The ongoing decline of many foundation species provides a set of important, albeit unfortunate, opportunities to develop the research tools, models, and metrics needed to identify foundation species, anticipate the cascade of immediate, short- and long-term changes in ecosystem structure and function that will follow from their loss, and provide options for remedial conservation and management.

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.

Topographic and ecologic controls on root reinforcement

Shallow landslides are a significant hazard in steep, soil-mantled landscapes. During intense rainfall events, the distribution of shallow landslides is controlled by variations in landscape gradient, the frictional and cohesive properties of soil and roots, and the subsurface hydrologic response. While gradients can be estimated from digital elevation models, information on soil and root properties remains sparse. We investigated whether geomorphically controlled variations in ecology affect the spatial distribution of root cohesion by measuring the distribution and tensile strength of roots from soil pits dug downslope of 15 native trees in the southern Appalachian Mountains, North Carolina, United States. Root tensile strengths from different hardwood tree species were similar and consistently higher than the only native shrub species measured (Rhododendron maximum). Roots were stronger in trees found on noses (areas of divergent topography) relative to those in hollows (unchanneled, convergent topography) coincident with the variability in cellulose content. This cellulose variability is likely related to topographic differences in soil water potential. For all species, roots were concentrated close to the soil surface, with roots in hollows being more evenly distributed in the soil column than those on noses. Trees located on noses had higher mean root cohesion than those in hollows because of a higher root tensile force. R. maximum had the shallowest, weakest roots suggesting that recent expansion of this species due to fire suppression has likely lowered the root cohesion of some hollows. Quantification of this feedback between physiologic controls on root growth and slope hydrology has allowed us to create a curvature-based model of root cohesion that is a significant improvement on current models that assume a spatially averaged value.

Ecosystem Processes and Human Influences Regulate Streamflow Response to Climate Change at Long-Term Ecological Research Sites

Analyses of long-term records at 35 headwater basins in the United States and Canada indicate that climate change effects on streamflow are not as clear as might be expected, perhaps because of ecosystem processes and human influences. Evapotranspiration was higher than was predicted by temperature in water-surplus ecosystems and lower than was predicted in water-deficit ecosystems. Streamflow was correlated with climate variability indices (e.g., the El Niño—Southern Oscillation, the Pacific Decadal Oscillation, the North Atlantic Oscillation), especially in seasons when vegetation influences are limited. Air temperature increased significantly at 17 of the 19 sites with 20- to 60-year records, but streamflow trends were directly related to climate trends (through changes in ice and snow) at only 7 sites. Past and present human and natural disturbance, vegetation succession, and human water use can mimic, exacerbate, counteract, or mask the effects of climate change on streamflow, even in reference basins. Long-term ecological research sites are ideal places to disentangle these processes.

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.

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.

Water table depth affects productivity, water use, and the response to nitrogen addition in a savanna system

We investigated annual aboveground net primary productivity (ANPP) and transpiration (E) of the dominant plant life forms, longleaf pine (Pinus palustris Mill.) trees and wiregrass (Aristida stricta Michx.), in a fire-maintained savanna. Experimental plots spanned a natural hydrologic gradient (xeric and mesic site types) mediated by soil moisture (q) and water table depth (WTD), and received additions of either 0 or 100 kg Nha -1 � year -1 . Low rates of ANPP (1.3- 2.2 Mgha -1 ) and annual E (108-380 mm) were observed in these communities. WTD and N addition explained 95% of the variation in community ANPP, whereas site type and WTD explained 83% of variation in community E. Between tree and grass life forms, longleaf pine ANPP was more coupled to WTD than wiregrass. For any given leaf area sup- ported, ANPP of longleaf pine increased linearly with increasing water use and decreasing WTD. The longleaf pine ANPP response to N addition was greater in sites with high water use compared with those with low water use, indicat- ing that this savanna system is colimited by nutrient and water availability and that water table depth plays a role in reg- ulating savanna productivity. Resume´ : Nous avons etudiela productiviteprimaire nette de la partie aerienne (PPNA) et la transpiration (E) annuelles des especes vegetales dominantes (pin des marais (Pinus palustris Mill.) et aristide des pinedes (Aristida stricta Michx.)) dans une savane maintenue par le feu. Les parcelles experimentales couvraient un gradient hydrique naturel (types de sta- tion xerique et mesique), engendrepar la teneur en eau du sol (q) et la profondeur de la nappe phreatique (PNP), et ont recu des amendements de 0 ou 100 kg Nha -1 � an -1 . Nous avons observede faibles taux annuels de PPNA (1,3 a` 2,2

Early Successional Forest Habitats and Water Resources

Tree harvests that create early successional habitats have direct and indirect impacts on water resources in forests of the Central Hardwood Region. Streamflow increases substantially immediately after timber harvest, but increases decline as leaf area recovers and biomass aggrades. Post-harvest increases in stormflow of 10–20%, generally do not contribute to downstream flooding. Sediment from roads and skid trails can compromise water quality after cutting. With implementation of Best Management Practices (BMPs), timber harvests are unlikely to have detrimental impacts on water resources, but forest conversion from hardwood to pines, or poorly designed road networks may have long lasting impacts. Changing climate suggests the need for close monitoring of BMP effectiveness and the development of new BMPs applicable to more extreme climatic conditions.