Chapter 18: Abiotic Factors 429 Chapter 18: How have abiotic factors, Abiotic Factors including environmental stressors such as air pollution, influenced the overall health of the South’s forests, and what are future Jennifer A. Moore, John G. Bartlett, Johnny L. Boggs, effects likely to be? Michael J. Gavazzi, Linda S. Heath, and Steven G. McNulty USDA Forest Service, Southern Research Station ■ Continued increases in ozone specific and integrated broad-scale Key Findings concentrations will likely have stress responses at forest ecosystem, significant negative impacts on community, and species levels. ■ Sulfur deposition will continue pine forests in the South. to decrease and subsequently have ■ Forest area and growth rates Introduction less of a negative impact on forest could increase across the South ecosystem nutrient cycling, whereas with moderate increases in air future nitrogen deposition will be The sustainability of southern temperatures and carbon dioxide beneficial to most southern forests, forests could be threatened by the concentrations during the 21st century. which are nitrogen limited. interactions of biotic and abiotic Severe temperature increases could ■ High-elevation spruce-fir negatively affect forest productivity stressors (McLaughlin and Percy 1999). Environmental factors forests in the Southern Appalachian and area, especially if precipitation such as temperature, precipitation, Mountains are the only forests rates do not increase to compensate H atmospheric carbon dioxide (CO ) in which significant damage is for increased water demands. 2 and O concentrations, and acid linked to acid deposition. ■ Carbon storage in southern forest deposi3tion affect forest processes such E ■ The overall health of hardwoods, ecosystems, including public, private, as carbon, water, and nutrient fluxes. A oak-pine, and southern pine forests and industrial forests, could make These processes are the foundation has not been shown to be adversely a significant contribution to carbon of forest ecosystems, and abnormally L affected by acid deposition. sequestration. Future policies, large variability in their size, timing, T ■ Regionally, there is no evidence incentive programs, and forest or location may influence forest H that acid precipitation is causing management intensity will affect sustainability. Therefore, from an significant damage to stream chem- carbon sequestration rates. ecosystem perspective, changes in istry in the Southern United States. ■ Land use change, not climate forest processes may be indicators of Water quality in some streams change or atmospheric chemistry, long-term forest function and health. in the Southern Appalachian has been and probably will continue Sulfur and nitrogen deposition have Mountains is decreasing. to be the most important determinant been indicted as contributors to forest ■ Ozone-related annual growth of carbon storage, uptake, and release degradation, especially in the high- reductions for pine seedlings across in terrestrial ecosystems. elevation red spruce and Fraser fir the South are probably between ■ Existing climate change models forests that occupy the ridges of the 2 and 5 percent. Tree-water stress do not provide adequate information Appalachian Mountains (McLaughlin or forest drought is thought to to forecast changes in location, extent, and Kohut 1992). In an effort to protect seedlings from the negative frequency, or intensity of extreme manage and sustain spruce-fir and effects of ozone. Any protective weather events and their impacts hardwood forests in a way that does benefits provided by drought stress on forest ecosystems. Potential not compromise the ability of future for seedlings are likely offset by increases in air temperature and generations to meet their needs, the growth and productivity reductions. changes in precipitation patterns current and future impacts of sulfur ■ Southern pines typically do not may contribute to increased freq- and nitrogen deposition on overall show visible symptoms of ozone uency or intensity of some events. forest health in the Southern United States must be addressed. (O ) injury under ambient O ■ Detailed spatial and temporal 3 3 conditions, but growth of mature predictions of abiotic stressor Ground level (tropospheric) O is 3 southern yellow pines is being effects on forest sustainability are an air pollutant that affects U.S. forests reduced by current ambient ozone not possible without long-term (U.S. Environmental Protection Agency levels at annual rates that vary improvements in regional monitoring 1996). At current ambient levels, O 3 from 0 to 10 percent per year. and studies designed to understand can decrease tree growth, increase the 430 Southern Forest Resource Assessment probability of mortality, cause visible potential to capture CO and 2 foliar damage, and alter forest store carbon. Acid Deposition successional patterns (Flagler and Climate change also could generate Chappelka 1996, McLaughlin and forest stress, and extreme weather Downing 1995, Teskey 1996). For events can cause disturbances that Acid Deposition these reasons, current and projected shape forest systems by influencing Methodology: Current O impacts on southern forests are their composition, structure, and Conditions 3 addressed in this Assessment. functional processes. We discuss the Acid deposition occurs when Climate influences the establishment effects of these disturbances and their emissions of sulfur dioxide (SO ) and growth of forest trees, affecting the relationship to changing temperature 2 and oxides of nitrogen (NO) react extent and quality of forest ecosystems. and precipitation patterns. x with atmospheric water, oxygen, and The spatial and temporal distribution Biotic stressors such as insects and oxidants to form acidic compounds. of air temperature and precipitation pathogens have major negative impacts Mild solutions of nitric and sulfuric is the primary climatic factor shaping on forest ecosystems; in the United acids are formed and fall as acid forests. Human activities contribute States, they cause severe damage on precipitation. Sulfur and nitrogen significantly to current global climate an average of more than 50 million deposition was first described as change (Dale and others 2000), acres per year, costing $2 billion a a problem in Europe in the early predominantly due to the increasing year (Dale and others 2000). Biotic 19th century and has been studied concentration of greenhouse gases stressors are the focus of chapter 17. extensively in North America since such as CO . Since the beginning of 2 Each of the abiotic stressors— the 1970s (Blancher 1991). Sulfur the industrial revolution, CO levels 2 methods, data sources, results, and nitrogen deposition can impair have been steadily increased by fossil discussion, and conclusions—are tree growth in several ways. They can fuel burning and land use changes discussed separately. Current abiotic leach calcium and magnesium from (Sarmiento and Wofsy 1999; U.S. stressors have been described for soils where base cation stores are very Department of Energy, Office of Sci- different coarse-scale studies. Attempts low, and the ability of the ecosystems ence, Office of Fossil Energy 1999). at regional-scale characterizations to retain sulfur or nitrogen is minimal Even if changes in CO concentration 2 and future predictions are underway (McLaughlin and others 1998). Acid did not effect climate changes, they and are highlighted when feasible. deposition may also involve the release would affect plant growth. of toxic elements such as aluminum It is important to recognize the Independently developed climate from the soil, adversely affecting H integrated nature of these abiotic change scenarios are generated with biological processes and living organ- stressors and their cumulative effects transient general circulation models isms (Malmer 1976). Nutrient loss and E on forest ecosystems. This integration (GCMs) that simulate atmospheric soil degradation have been observed is referenced throughout the chapter. Adynamics under a gradual doubling in some hardwood forests (Swank It is imperative that readers consider in greenhouse gas concentrations from and Vose 1997). However, pine, hard- Labout 1895 to 2100. Emissions of CO cumulative integrated effects when wood, and mixed (oak-pine) forests 2 interpreting the results and conclu- Tto the atmosphere are predicted to experience slower losses of base cation sions from this chapter. increase from 7.4 gigatons per year in H 1997 to 26 gigatons per year by 2100 (U.S. Department of Energy, Office of Science, Office of Fossil Energy 1999). For this Assessment, these scenarios are used with ecological process models to investigate the potential effects of climate change on forest ecosystems. Forest carbon sequestration, the ability of forests to store and release carbon, is currently an important issue debated in the policy arena. Carbon stored in forests affects the amount of carbon contributing to the increasing atmospheric CO concentration. 2 Reductions in carbon emissions have been proposed as a mitigation strategy Pounds per acre for rising atmospheric CO , which 1.88–6.28 2 may be causing global warming. Rising 6.28–9.74 atmospheric CO levels could also 9.74–12.87 be mitigated by i2ncreasing carbon 12.87–15.31 sequestration through forestry and 15.31–21.80 other land management activities. Terrestrial ecosystems have enormous Figure 18.1—Current (1999) distribution of sulfate deposition in pounds per acre across the South (National Atmospheric Deposition Program 2000). Chapter 18: Abiotic Factors 431 They are produced primarily in industrialized States in the northern part of the South. Currently, forests in the South are exposed to a wide range of nitrogen deposition rates (National Atmospheric Deposition Program 2000) (fig. 18.2). The mean regional nitrogen deposition for 1999 was 10 pounds per acre, a 10-percent decrease in nitrogen deposition from 1994 (National Atmospheric Deposition Program 2000). The highest regional nitrogen values are generally located in the Pounds per acre northern part of the South (fig. 18.2). 2.06–5.94 5.94–7.97 Their sources are emissions from all 7.97–9.99 31 States east of the Mississippi River 9.99–12.40 (Nash and others 1992). 12.40–14.33 For this discussion, the South has Figure 18.2—Current (1999) distribution of nitrogen deposition in been divided into nine forest types pounds per acre across the South (National Atmospheric Deposition Program 2000). according to various factors that include geographic location, precipi- tation, minimum and maximum air nutrients and degradation because (National Atmospheric Deposition temperatures, and soil conditions (more of their ability to buffer sulfur and Program 2000) (fig. 18.1). The mean or less sensitive to acid precipitation). nitrogen deposition. These forests regional sulfate deposition for 1999 Five of these forest types are shown in generally have large calcium pools was 11 pounds per acre, which is a fig. 18.3. Sensitive soils have low base that increase their ability to buffer 13-percent decrease in sulfur deposi- cation stores, and the ecosystem has a acid deposition. tion from 1994 (National Atmospheric low ability to retain sulfur or nitrogen, There is a wide range of sulfate Deposition Program 2000). The highest or both. Less sensitive soils are ones deposition rates across the South regional sulfur values are in North with high concentrations of base H Carolina and Tennessee (fig. 18.1). cations, high buffering capacity to E sulfur and nitrogen deposition, and, normally, nitrogen deficiency. Within A the region, the high-elevation spruce-fir forests are most sensitive to sulfur and L nitrogen deposition. The least sensitive T ecoregions are those covered primarily Spruce–fir H by hardwood, pine, and oak-pine Longleaf–slash pine forests. The sensitivity of a given Loblolly–shortleaf pine region to acid precipitation depends Oak–pine on the ability of the rocks and soils Oak–hickory to neutralize or buffer the acid. Soils derived from granite, which are low in calcium, are highly sensitive. Soils VA derived from limestone, which are high in calcium, are much more capable of buffering the acid. Acid Deposition Methodology: TN Future Predictions Sulfur deposition is a primary contributor to acid deposition that indirectly affects forest decline by NC leaching base cations from the soil. Figure 18.3—Distribution Therefore, in 1990, Title IV of the of spruce-fir and other Clean Air Act set as its primary goal southern forest types the reduction of annual SO emissions in eastern Tennessee, 2 by 10 million tons below 1980 levels western North Carolina, (U.S. Environmental Protection Agency and southern Virginia (Eyre 1980). 1997a). To achieve these reductions by 432 Southern Forest Resource Assessment 2010, the law invoked a restriction on Currently, high-elevation spruce-fir competitive inhibition of calcium power plants fired with fossil fuels. forests are the most susceptible to the uptake by aluminum. Dendroecolog- By 1995, nationwide emissions of SO effects of sulfur deposition (McLaughlin ical- and plot-based data have shown 2 were reduced by almost 40 percent and Percy 1999) because they lack the declines in radial growth of red below their required level. In addition, ability to buffer sulfur deposition and spruce (LeBlanc and others 1992) monitoring sites throughout the United are low in base cation pools. Future and canopy-crown deterioration States found statistically significant rates of sulfur deposition are expected during the mid-to-late 1980s in reductions in precipitation acidity and to decrease, which could lead to the Southern Appalachian Mountains sulfate concentrations (National Acid a reduction in the effects of sulfur (Peart and others 1992). Precipitation Assessment Program deposition on base cations in high- Whereas acid deposition has affected 1998). Attempts to reduce nitrogen elevation spruce-fir forests. Recent tree growth in spruce-fir forests of the deposition were initiated in 1996. evidence indicates that most Southern Southern Appalachians (McLaughlin Although Title IV initiated a reduction Appalachian soils supporting spruce- and others 1998), damage to these in annual nitrogen deposition, new fir ecosystems are poorly buffered, high ecosystems is not limited to acid concentrations are expected to have in aluminum, and nitrogen saturated deposition. Reams and Van Deusen potential impacts on forests across (Johnson and others 1991). Nitrogen (1993) reported that stand disturbances the South. Modeling future projections saturation occurs when ammonium and changes in stand dynamics have and impacts of nitrogen and sulfur (NH ) and nitrate (NO ) are present in resulted in radial growth declines in 4 3 deposition on forested ecosystems in quantities that exceed total combined spruce-fir forests. In addition, the the Southern Appalachian Mountains plant and microbial demand. Excess balsam woolly adelgid was introduced is an ongoing research objective of levels of nitrogen have been found to into North America at the beginning the Southern Appalachian Mountains affect soil and plant calcium:aluminum of the 20th century, and the exotic Initiative (SAMI). The North Carolina ratios (Johnson and others 1991), insect has been active in the Southern General Assembly is reviewing a bill cause aluminum toxicity (Shortle and Appalachians since the late 1950s that would reduce nitrogen oxides Smith 1988), and decrease calcium (McLaughlin and others 1998). The and sulfur oxides generated by coal- uptake and leaching of base cations damage to mature Fraser fir in the powered utility plants by more than (McLaughlin and others 1998) in Southern Appalachians by the woolly 70 percent (North Carolina General these sensitive forests. A lack of adelgid has been extensive over Assembly 2001). Governor Michael calcium changes the wood structure the past 15 years (Dull and others Easley supports this legislation and has of spruce and fir and may change 1988). Although heavy infestation begun to discuss regional air pollution the ability of branches to withstand H is unquestionable evidence that the reduction initiatives with lawmakers stress (McLaughlin and others 1998). adelgid plays a major role in killing around the South (North Carolina Furthermore, excess levels of nitrogen E these trees (see chapter 17 for more Department of Environment and decrease the rates of some critical details), it is also important to consider ANatural Resources 2001). functions of soil microorganisms, the influence of predisposing factors, including decay of forest floor material LAcid Deposition including abiotic stressors such as acid (Drohan and Sharpe 1997). These deposition, on the susceptibility of TData Sources effects on forest soils are most dramatic forests to pathogens (Manion 1981). H Primary data sources for sulfur in the sensitive soils under spruce-fir Hardwood forests in the South are forests. Conversely, in an oak-pine and nitrogen deposition were the considered less sensitive to nitrogen forest in the North Carolina Piedmont, National Acid Deposition Program deposition than spruce-fir forests Johnson and others (1995) predict (National Acid Deposition Program because they still have adequate stores that forest floor nutrient contents 2000) and cited literature. of base cation nutrients, and the soils will be virtually unaffected by a 50- still maintain considerable capacity to Acid Deposition Results percent reduction in sulfur deposition retain the deposited nitrogen (National over the next 20 years. Although sulfur is an essential Acid Precipitation Assessment Program Effects of acid deposition on tree nutrient for soil and plant metabolic 1998). In most hardwood forests, growth have been associated with processes, sulfur deposition can virtually all nitrogen deposition is nutrient limitations caused by increases contribute to degradation of soil either adsorbed in the soil or used by in soil aluminum concentrations. chemistry (Reuss and Johnson 1986). vegetation and microorganisms. Much Studies of historical tree-ring chemistry Long-term increases in soil acidity of this nitrogen may be removed later (Bondietti and McLaughlin 1992) have resulting from sulfur deposition by forest harvesting. These systems shown that calcium concentrations are believed to affect nutrient cycling therefore have not shown negative in stemwood increased as growth by leaching nutrients, such as calcium effects from increases in nitrogen increased during the late 1940s and and magnesium (Fenn and others deposition and may respond with 1950s. However, decreases in tree 1998). Research has also shown that increased growth. Research has shown growth were associated with increases sulfur deposition provides the stimulus that 22.8 pounds per acre per year in aluminum:calcium ratios in the to mobilize aluminum in soil solutions of nitrogen fertilizer increased basal wood, suggesting that the availability (Reuss and Johnson 1986). Dissolved area growth of trees by 67 percent of calcium was reduced at the same aluminum interferes with the uptake (McNulty and Aber 1993). time aluminum concentrations of calcium and other root functions Impacts of nitrogen deposition on increased. McLaughlin and Kohut (Johnson and others 1991). forest health have not been detected (1992) have shown evidence for the Chapter 18: Abiotic Factors 433 in the pine and oak-pine forests of Table 18.1—Acid neutralizing capacity (ANC) categories for brook the South (National Acid Precipitation trout response Assessment Program 1998). However, nitrogen is a major contributor to the depletion of base cations in many meq/L Classification Biological response buffered soils supporting southern pine and oak-pine forests. Therefore, >50 Not acidic Reproducing brook trout populations over the course of decades, nitrogen expected where habitat is suitable deposition is likely to reduce pine 20-50 Transitional Extremely sensitive to acidification; forest productivity (National Acid brook trout response variable Precipitation Assessment Program 1998). Increases in growth are expected 0-20 Episodically acidic Sublethal and/or lethal effects on for some nitrogen deficient soils, brook trout likely whereas negative effects are expected to be limited to the most acidic soils. <0 Chronically acidic Lethal effects on brook trout likely In the future, nitrogen deposition will Source: Bulger and others 1998. continue to impact the structure and function of high-elevation spruce-fir forests. In addition, some hardwood, of brook trout can be altered by ANC drought, loss of foliage, insects, and pine, and oak-pine forests that are (table 18.1). Furthermore, the Southern diseases. Hardwood, pine, and mixed sensitive to nitrogen deposition could Appalachian Assessment has shown oak-pine forests are less sensitive than respond with reduced growth rates and that 70 percent of sampled streams spruce-fir for several reasons, including accelerated tree mortality over the long have suffered moderate to severe fish biological nitrogen demand, higher soil term. However, research has predicted community degradation, and about cation exchange capacity, and faster that in oak-pine forests in the North 50 percent of the stream miles in West nitrogen cycling. Carolina Piedmont, vegetation will Virginia and Virginia show habitat Since most hardwood, pine, and respond positively to a 200-percent disruption (Southern Appalachian mixed forests are nitrogen deficient, increase in nitrogen deposition over Man and the Biosphere 1996). they may experience increased growth the next 20 years. A 3- to 9-percent However, streams targeted by the rates in response to continued elevated increase in vegetation nutrient content NSS in the southeastern highlands, nitrogen deposition. Conversely, and a 10- to 30-percent increase (which includes the Ozarks/Ouachita, nitrogen deposition can significantly H in forest floor nutrient content are Piedmont, Southern Appalachians, degrade some of these forests over expected (Johnson and others 1995). Southern Blue Ridge, and ecological time (years to decades), especially E subregions in the States of Arkansas, Currently, the SAMI Class I in areas where nitrogen levels may A Georgia, North Carolina, and Wilderness Areas are much more be high and the soil has reached Tennessee) appear to be buffered sensitive to acid precipitation than any or is approaching saturation. L from sulfur deposition by a substantial other areas surveyed by the National Sulfate and nitrate concentrations T amount of sulfate adsorption in Stream Survey (NSS) in the Southern have increased in streams throughout watershed soils (Rochelle and Church H Appalachians (Herlihy and others the South, but not to levels that are 1987). As a result, sulfate concen- 1996). The wilderness areas of greatest considered regionally problematic. trations in these streams are low. concern are Otter Creek and Dolly Furthermore, sulfate and nitrate in Sods in West Virginia. There, the Acid Deposition Discussion some streams are low or near detec- percentage of acidic stream length and Conclusions tion limits (Swank and Vose 1997). is high, pH is low, and sulfate and inorganic aluminum concentrations Emissions of SO2 and NOx are Acid Deposition Needs are high. Additionally, stream nitrate decreasing. However, plant species for Additional Research concentrations, an indicator of acid structure and compositon, soil To address the indirect impacts of deposition effects, have been shown chemistry, and microbial activities nitrogen and sulfur deposition that to have a strong correlation with forest continue to change. Currently, the lead to soil and vegetation degradation age. The highest concentrations occur mortality and decline of Fraser fir in high-elevation spruce-fir and in old-growth forests, where biological and red spruce at high elevations in hardwood forests, continued intensive demand for nitrogen is lowest. The the Southern Appalachians are the only monitoring, modeling, and validating wilderness area of least concern is cases of significant ecosystem damage. of acid deposition and nutrient cycling the Sipsey in Alabama because sulfate Thus, less than 5 percent of the South is processes must occur across local concentrations are not increasing, currently being negatively impacted by and regional scales. Monitoring efforts and acid neutralizing capacity (ANC) elevated sulfur and nitrogen deposition should be supplemented with long- of streams in this area is high. (Fenn and others 1998). In addition, term regional experiments (greater atmospheric deposition reduces the ANC has been used to determine than 5 years) in which realistic acid number of microorganisms important stream quality because stream deposition effects on soil chemical to nutrient cycling and removes acidification affects fish and other properties and stream quality are important nutrients from the soil, aquatic species. Research in the South evaluated (McNulty and others 1996). making spruce-fir forests more has shown that the biological response susceptible to canopy deterioration, 434 Southern Forest Resource Assessment Ozone Methodology: Ozone Data Sources Ozone Future Predictions Ozone monitoring studies have identified different O exposure profiles Over the past century, industrial 3 Ozone Methodology: at high elevations (greater than 4,900 activity and automobile emissions Current Conditions have increased the atmospheric con- feet) than at lower elevations (less than 1,600 feet) and near sea level (Aneja centrations of O precursors. As a Ground level O is created through 3 and others 1994). Levels of O in a complex series o3f atmospheric result, typical ambient O3 concen- mountains are lower than in lo3wlands trations have increased from 0.02- chemical reactions involving NO during the daytime. Near sea level, x 0.04 to 0.04-0.06 ppm—a trend that and volatile organic compounds O levels are very high during the is expected to continue into the 21st (VOC) in the presence of specific da3y , often exhibiting a distribution century (National Academy of Science climatic and weather conditions characteristic of the peak hours for 1992). Assuming a 1- to 2-percent (Chameides and Lodge 1992). Ozone automobile traffic. The concentrations annual increase in tropospheric O , exposure levels are influenced by 3 in the mountainous areas of the South as estimated by Fishman (1991), the factors such as temperature, time of have important implications for forest United States would achieve a 50- day, relative humidity, wind speed, health. The ambient O concentrations percent increase in ambient O in wind direction, and spatial proximity 3 are sufficiently high to 3induce injury 21 (base 1990) years and a doubling of anthropogenic and biogenic emission to sensitive native vegetation in the of O concentrations in 35 years. The sources (Schichtel and Husar 1999). 3 Blue Ridge Mountains (Skelly and National Academy of Science (1992) Ozone can reduce foliage, stem, and Hildebrand 1995). In addition, some estimated an increase of 40 percent root growth in trees by impacting leaf- areas in the region are downwind of by the year 2020. Thompson (1992) cell photosynthesis and gas exchange. significant NO and VOC emission used several computer models to x Allen and Gholz (1996) revealed predict that O concentrations will rise sources. For example, regionally high extensive spatial and temporal variation by 0.5 percent3 per year for the next 50 O levels found in the Blue Ridge 3 in O concentrations across the region. years, whereas Chameides and others Mountains and Shenandoah Valley of 3 For at least two reasons, accurate (1994) suggested that the frequency Virginia result from a combination of prediction of annual variability in of O events with concentrations high upwind emission sources located in the O levels for forested areas has not enou3gh to damage plants will triple industrial Midwest and specific weather 3 yet been achieved: (1) monitoring over the next 30 years. However, patterns (Wolff and others 1977). These sites in rural, forested areas are lacking; more recent ozone modeling efforts weather-related O episodes may be Hand (2) modeling O exposure is by SAMI predicted a 10- to 15-percent attributed to a com3bination of local- 3 very difficult because of weather reduction in maximum daily ozone and regional-scale factors: (1) higher Eand human-related conditions that levels between 1995 and 2010 for than normal ambient temperatures, (2) Acontribute to its annual variability the Southern Appalachians based wind speeds and directions associated (Allen and Gholz 1996). However, on current emissions controls with stationary high-pressure systems Lannual variation in O at select (Southern Appalachian Mountains that produce local air stagnation, and 3 Tmonitoring sites has been analyzed. Initiative 2001). (3) lower than normal relative humidity (Aneja and Li 1990). H Annual O variability for the United 3 States can be seen in figure 18.4, which shows 3-month maximum daily SUM06 O exposure levels for 3 1988 through 1992. A SUM06 value is the sum of all mean hourly daytime O concentrations that are at least 3 0.06 parts per million (ppm) over a continuous 3-month period (92 days) during the summer. The SUM06 exposure index represents the threshold ambient O level (0.06 ppm-hours) 3 below which many forms of vegetation can resist harmful cumulative O 3 effects. The SUM06 index may be particularly useful because negative effects of O exposure, especially 3 on tree photosynthetic capacity (Richardson and others 1992) and foliage production and retention (Kress and others 1992), may be Figure 18.4—Three-month maximum daily SUM06 ozone exposure cumulative and linear, extending grid for 1990 showing spatial variability in ozone concentrations. The over multiple growing seasons. exposure grid was calculated by U.S. Environmental Protection Agency using NHEERL-WED’s Geographic Information System model to spatially interpolate SUM06 values calculated from the AIRS monitoring network (Schichtel and others 1996). Chapter 18: Abiotic Factors 435 Ozone Results reductions for pine seedlings in the the location of a burn, and substantial South were between 2 and 5 percent. increases in O concentrations (greater To cause tree damage, O must 3 3 For mature loblolly pines, Dougherty than 50 percent above ambient) have be absorbed by the plant through and others (1992) used a process model been detected downwind of burned the stomatal openings found on the to estimate annual growth reductions areas and at the top of burn plumes. surface of leaves in a process known of about 3 percent under ambient as stomatal conductance. Stomates The important relationship between O concentrations. open during daylight hours to permit 3 soil moisture, stomatal conductance, the exchange of gaseous compounds Hogsett and others (1997) found that and tree sensitivity to O levels high- 3 (CO , O and water vapor) necessary black cherry has strong O sensitivity, lights the importance of climate in for p2hoto2,synthesis. At night, stomates and tulip poplar has mode3 rate O predicting future impacts of O on 3 3 close, preventing the transpiration sensitivity. Southern yellow pine species forest health. Under future climate of water. Because stomates are open showed little response to changes in scenarios, trees in areas of the South during the day, daytime O concen- SUM06 O concentrations, and sugar characterized by periods of persistent trations are most likely to 3d amage maple exh3ibited a threshold response drought and poor soil water storage trees. Rates of stomatal conductance in which annual biomass increased capacity will be more sensitive to vary by species and age, and these rates dramatically between 26 and 38 O pollution and will likely incur 3 directly determine both the quantity ppm-hour per year SUM06. substantial visible foliar damage of O uptake and the plant’s response Overall, it appears that the growth of (Maier-Maercker 1999) and growth to a g3iven concentration of O (Kelly mature southern yellow pines is being reductions (Southern Appalachian and others 1995). In general, 3ozone- reduced by current typical ambient O Mountains Initiative 2001). 3 sensitive tree species under high levels at annual rates that vary from 0 Ozone Needs for O stress experience reduced leaf to 10 percent per year. Annual growth 3 Additional Research area, slower growth during drought reductions for pine seedlings in the conditions, and lower vertical South are probably between 2 and 5 Because expert predictions identify growth rates (Southern Appalachian percent (Teskey 1996). However, at O as a significant forest stressor well Mountains Initiative 2001). present there are no indications of in3to the 21st century (Heck and others It appears that O affects growth and community level changes (competi- 1998), scientists and policy experts vitality indirectly b3y predisposing trees tion dynamics, community structure, have jointly assessed critical research to injury from other biotic and abiotic and function, etc.) attributable to needs pertaining to effects on forested stressors (Chappelka and Freer-Smith O (McLaughlin and Percy 1999). systems. The Ecological Research 3 H 1995). For example, ponderosa pine Although O may be reducing annual Needs Workshop (U.S. Environmental 3 exhibits increased sensitivity to bark growth of trees in the South, other Protection Agency 1998) developed E beetle attack in the San Bernardino air-borne chemicals such as CO and one such set of research priorities. 2 Mountains following O damage (Cobb nitrogen and sulfur compounds may A summary of those priorities for A and others 1968). In th3e South, pines be simultaneously increasing growth forests and natural areas is provided typically do not show visible symptoms (Teskey 1996), thereby effectively here (Heck and others 1998): L odfi tOio3n isn (jBueryrr uanngd earn adm obthieenrst O193 9c6o)n- omna sokvienrga ltlh feo rneesgt ahteivaelt hef.fects of O3 1to. Cscoanlisnigd erreastuioltns ionf gfarcotwortsh rcehlaatmedbers TH except during extended periods of high Ozone Discussion to mature trees, stands, communities, O levels when injury is readily visible. and landscapes. 3 and Conclusions The amount and way that O affects 2. Measurement of selected endpoints trees depend on the age of the3 trees The growth impacts of ambient (growth, mortality, foliage injury, etc.) and the species. Given similar amounts O levels on southern pines appear in managed and natural ecosystems 3 of O exposure, immature hardwoods to be statistically significant at this such as loblolly pine plantations or gene3rally exhibit more growth loss than time (McLaughlin and Percy 1999, bottomland hardwood ecosystems softwoods (table 18.2) (McLaughlin Teskey 1996). Additional increases in across selected O gradients throughout and Percy 1999). Based on the limited tropospheric O will almost certainly the South, using r3esults to support 3 number of studies available, mature have negative impacts on the growth development of empirical and process- hardwood growth rates appear to be of pine species in the South (Southern based models designed to understand more sensitive to O exposure than Appalachian Mountains Initiative the mechanisms of plant response mature softwood gr3owth rates (table 2001, Teskey 1996). to O . 18.2). According to Dougherty and Another important consideration 3 3. Determination of utility of using others (1992), an average mature for future forest health is the frequency visible foliar injury and other biological loblolly pine tree growing in a plan- and intensity of forest fires. Forest indicators to interpret effects of O on tation experiences a 3-percent annual fires produce carbon monoxide (NO) 3 specific indices of ecosystem health. loss of gross primary production under and gaseous hydrocarbons that are x ambient O conditions in the South. the precursors of atmospheric O 4. Development of economic In a review3 of 19 studies measuring the (Bohm 1992). Therefore, forest f3ires techniques that measure changes influence of O exposure on growth of may contribute to O production in the value of managed and natural slash pine, sho3rtleaf pine, and loblolly in wilderness and ru3ral areas (Bohm ecosystems affected by O3. pine seedlings and saplings, Teskey 1992). Bohm (1992) observed that 5. Development of a reasonable O (1996) concluded that annual growth O has been found to accumulate near exposure index via defined relation3ship 3 436 Southern Forest Resource Assessment Table 18.2— Estimates of ambient O effects on growth of forest tree species occurring in the South 3 Growth Species reduction Conditions Source Percent Seedling/sapling studies Multiple species 0-10 Shoot growth Chappelka and Samuelson 1998 Southern pines 2- 5 Summary estimate of 19 field-chamber Teskey 1996 studies Loblolly pine 0- 3 Mean response to 50-200 ppm-hr Taylor 1994 1-10 Sensitive family response to 50-200 ppm-hr (synthesis-whole tree biomass) Hardwoods 13 Values derived from response Reich and others 1988 Conifers 3 surface at 20 ppm-hr Black cherry 10-24 Hogsett and others 1997 Yellow-poplar 5-13 Values derived from O exposure- 3 Sugar maple 0- 9 response functions and model- Red maple 0- 1 simulated tree and stand Loblolly pine 2- 5 responsea Eastern white pine 4- 8 Virginia pine 0- 1 Mature tree studies Loblolly pine 2- 9 Whole tree carbon model using branch Dougherty and others H chamber data (GA) 1992 E 3 Mean response 0-13 Mean annual weekly responses to O and McLaughlin and A 3 interactions of O and moisture stress, Downing 1996 3 L 5 years (TN) T 0- 5 Annual O effect—no water stress 3 0-30 Annual O effect—moderate water stress H 3 Hardwoods 3-16 Regional simulation with canopy-stand Ollinger and others 1997 model across moisture gradients. Highest reductions occurred in areas with highest O levels and on soils with high water holding 3 capacity where drought stress was absent. a Percent reduction in annual net primary production. Source: McLaughlin and Percy (1999), with additions provided. Chapter 18: Abiotic Factors 437 between O exposure concentration, smoke can impair road visibility and in coastal and floodplain areas. On 3 uptake dose, and selected endpoints breathing in sensitive individuals. average, floods cause almost $4 (growth, mortality, foliar injury). billion dollars in damage each year Wildfire can substantially influence (National Oceanic and Atmospheric 6. Study of the interactions between forest structure and function. Ecological Administration 2000). Upland forest O and other abiotic or biotic stressors. effects of forest fires include mortality 3 ecosystems that experience flooding of individual trees, shifts in successional respond with reduced photosynthetic Climate Change and direction, induced seed germination, rates; over extended periods, changes acceleration of nutrient cycling, death Extreme Weather- in tree species composition are possible, of seeds stored in the soil, changes Related Events in surface soil organic layers and as some species are more flood tolerant that others (Burke and others 1999, underground plant root and Iles 1993). Most trees can withstand reproductive tissues, volatilization of 1 to 4 months of flooding duration Extreme Weather-Related soil nutrients, and increased landscape without significant injury (Bratkovich Event Methodology: heterogeneity (Whelan 1995). As a and others 1993). In extreme situations, result of these effects, the capacity of Current Conditions higher mortality rates may occur (Iles forests to provide wildlife habitat, 1993). Anaerobic soil conditions in Climate effects on forest conditions timber, and recreation may be flooded areas cause physiological are most strongly expressed by diminished (Flannigan and stress and influence nutrient availability extreme events such as fire, hurricanes, others 2000). (Burke and others 1999). Secondary tornadoes, floods, drought, and ice Hurricanes—Hurricanes disturb effects of flooding include elevated storms (Dale and others 2000). Each forests along the coastlines of the soil erosion and sedimentation rates type of event affects forests differently; South. Ocean temperatures and (Iles 1993). At the regional scale, some cause large-scale tree mortality, regional weather influence the path, there is high variability in the spatial whereas others, such as ice storms, size, frequency, and intensity of location and amount of disturbance impact community structure and hurricanes (Emanuel 1987). An average associated with floods. organization without causing of two hurricanes strike land every massive mortality. 3 years in the United States (Hebert Drought—Droughts occur in most forest ecosystems in the South. Occur- Wildfire—The frequency, seasonality, and others 1997). Some scientists have rence is irregular in forests east of the size, intensity, and type of wildfires hypothesized that hurricane impacts Mississippi River, occasional across depend on weather phenomena and on forests, including mortality, may most of the South, and more common H forest structure and composition. be related to soil characteristics in late summer on the Coastal Plain Fire initiation and spread also depend (Duever and McCollum 1993). (Hanson and Weltzin 2000). Conse- E on fuel availability, the presence of Tornadoes—Tornadoes are one quences of long-term drought or ignition agents, and topography. of the most important agents of abiotic A flooding are generally proportional Across the southern Coastal Plain, disturbance in eastern deciduous to the area affected; during the past L forest shrub and brush species can forests. Nearly 1,000 tornadoes occur few decades, an increasing portion T create highly flammable fuel conditions each year in the conterminous United of the United States has experienced in just 5 years under the right climatic States (Peterson 2000). In the South, either severe drought or flooding H conditions if fuel loads are not tornadoes are very common in (Karl and others 1995c). Drought managed. Therefore, fuel management Oklahoma and Texas and frequent effects are influenced by soil texture is necessary. Each year, across all land in Alabama, Florida, Louisiana, and and depth, exposure, species compo- ownership classes, 5.4 million acres Mississippi. Tornadoes can cause severe sition, life stage, and the frequency, are managed with prescribed fire. mortality, reduce tree density, alter duration, and severity of drought. Seventy-five percent of the prescribed stand-size structure, and modify local The immediate response of forests burning occurs in the States of environmental conditions via soil to drought is to reduce water use Alabama, Florida, and Georgia. All erosion or nutrient loss (Dale and and growth. Small plants, including 34 national forests in the region have others 2000). The resulting disturbance seedlings and saplings, are usually the prescribed fire programs, and, since may bring about the release of advance first to succumb to moderate drought 1944, approximately 21 million acres regeneration, seed germination, or conditions. Deep rooting and stored have been treated to minimize wildfire accelerated seedling growth (Peterson carbohydrates and nutrients make risk (Forest Health Protection Program and Pickett 1995). These effects can large trees susceptible only to severe 2000). Fire management would be change gap dynamics, successional droughts (Dale and others 2001). more prevalent were it not for smoke patterns, and other ecosystem level Ice Storms—Ice storms occur problems associated with controlled processes such as water use. The rela- throughout the South. They are burns. Criteria included in the U.S. tionship between wind strength and produced when rain falls through Environmental Protection Agency’s severity of disturbance varies by tree subfreezing air masses, freezing when National Ambient Air Quality species and forest type. Shallow-rooted contact is made with objects on the Standards for Particulate Matter (U.S. species and thinned stands tend to be ground. Ice accumulation varies with Environmental Protection Agency more vulnerable, but multiple factors topography, elevation, and area of 1997b) limit the amount and extent influence tree response to windstorms. exposure. Ice storms may sever twigs of prescribed fire programs because Floods—Floods occur throughout and bend or break stems, causing the South but are most concentrated 438 Southern Forest Resource Assessment moderate crown loss. Damage to forest and environmental scenarios. The For the continental United States, the stands can range from light and patchy several biogeography models used HadCM2Sul scenario includes a to the breaking of all mature stems, for this Assessment included the relatively modest 2.8∞ average increase depending on stand composition, past Mapped Atmosphere Plant Soil System in air temperature, a 20-percent average disturbances, and the amount of ice (MAPSS), BIOME3, and MC1 (Bachelet increase in precipitation, and effects accumulation (Irland 2000). Effects and Neilson 2000, Bachelet and others of doubled CO and altered sulfate 2 of ice storms on forest stands include 2001). Input datasets include latitude, aerosol concentrations (based on IPCC stem damage, loss of growth until leaf mean monthly temperature, wind projections of future greenhouse gases) area is restored, and possible shifts in speed, solar radiation, and soil by 2100 (Bachelet and others 2001). tree species composition toward trees properties such as texture and depth. The mean temperature increase for the more resistant to ice damage. All of these models project vegetation South is about 1.0∞ by 2030 and 2.3∞ responses to changes in CO but by 2100; this degree of warming is Recently thinned stands may have 2 through different mechanisms. smaller than that of any other region increased vulnerability to ice storm (National Assessment Synthesis Team damage because tree crowns have Climate Change and 2001). This scenario predicts that the spread into openings, but branch Extreme Weather-Related South will remain the wettest region strength has not yet increased. Potentially, there are several secondary Event Data Sources for the next century; mean annual precipitation increase will be about consequences of ice damage. Sus- To date, it is generally believed 3 percent by 2030 and 20 percent ceptibility to insects and diseases that hotter and more variable air by 2100. Other regions in the may be increased, and fuel loads temperatures will occur across the Eastern United States are predicted may accrue, heightening wildfire United States in the future (National to experience similar increases in risk in some areas (Irland 2000). Assessment Synthesis Team 2001). precipitation (National Assessment Climate Change However, the timing and distribution Synthesis Team 2001). of precipitation or other weather Methodology: Future Predictions of forest area, distribution, phenomena are much less certain Predictions and biodiversity used four equilibrium (Dale and others 2000). The transient (UKMO, GISS, GFDL-R30, OSU) climate change scenarios used for this The effects of climate change on and three transient (HadCM2Sul, Assessment do not adequately represent southern forest productivity and HadCM2GHG, CGCM1) climate extreme events because of their coarse hydrology across a range of climate scenarios as input for the MAPSS spatial and temporal resolution Hand site conditions were assessed biogeography and MC1 dynamic (monthly time step, approximately with the well-validated, physiologically global vegetation models. The range 1,000 square miles) (National Ebased forest process model PnET-II Assessment Synthesis Team 2001). in temperature increase is 2.8 to 6.6∞ (McNulty and others 2000). PnET-II for all scenarios, with changes in A Extreme events may last only minutes used four monthly climate variables precipitation varying greatly between or days, and their extents may range L(amir itneimmpuemra tauirr et,e mprpeecripatiutartei,o nm, aaxnimdum from local to small regional scales. tthraen sscieennta r(iaoss w, aitnhd HchadanCgMes2 Sinu lC) Oor2 T When the effects of extreme events solar radiation), forest-type-specific instantaneously doubling in the case are averaged over large periods of time Hvegetation parameters, and site-specific of the equilibrium scenarios. MC1 used and space, much information is lost. soil water holding capacity to predict only HadCM2Sul and CGCM1, whereas Therefore, very little quantitative data forest growth and drainage across MAPSS used all equilibrium scenarios on extreme weather events are available the South at a 0.5- by 0.5-degree and averaged the last 30 years of the to predict future forest impacts. Instead, (approximately 30- by 30-mile) spatial transient scenarios so they could be we will discuss the potential impact resolution. Atmospheric CO increases treated as equilibria. The BIOME3 2 of projected general trends in extreme were incorporated into PnET-II by model used only the transient climate weather events on forest structure entering the relationship between water scenarios (Bachelet and Neilson 2000). and function. use efficiency (WUE) and CO level. PnET-II results for pine and ha2rdwood Two climate datasets developed by Climate Change and forest types have been validated for the Vegetation/Ecosystem Modeling and Extreme Weather-Related the South (McNulty and others 2000). Analysis Project (VEMAP) were used Event Results with the PnET-II model to assess future Impacts of climate change on forest climate impacts on southern forest Wildfire—Because climate change area, distribution, and biodiversity were growth. The Historical Climate Series may alter the frequency, intensity, studied with biogeography models. includes monthly and daily climate distribution, or extent of wildfires, This type of model uses resource and data with interannual variability for the species regeneration patterns may be ecophysiological constraints such as conterminous United States from 1895 disturbed with species or communities available soil water and minimum to 1993 (National Assessment Synthesis at the edges of their natural range winter temperatures to simulate climate Team 2001). The Hadley Centre experiencing potentially severe effects. change impacts on forest ecosystems HadCM2Sul transient climate change at regional, continental, and global Model results from the fire scenario was used to represent climate scales (Bachelet and Neilson 2000). distribution module of MC1 predict variables from 1994 to 2100; other The biogeography models used here great variation in future fire-weather climate scenarios exist but were not predict the dominance of different patterns for the northern portion of available at the time of this analysis. plant species under different climatic North America (Bachelet and Neilson
Description: