Physiological and foliar injury responses of Prunus serotina,Fraxinus americana,and Acer rubrum seedlings to varying soil moisture and ozone

Sixteen black cherry (Prunus serotina, Ehrh.), 10 white ash (Fraxinus americana, L.) and 10 red maple (Acer rubrum, L.) 1-year old seedlings were planted per plot in 1997 on a former nursery bed within 12 open-top chambers and six open plots. Seedlings were exposed to three different ozone scenarios (ambient air: 100% O3; non-filtered air: 98% ambient O3; charcoal-filtered air: 50% ambient O3) within each of two different water regimes (nine plots irrigated, nine plots non-irrigated) during three growing seasons.During the 1998 and 1999 growing season, leaf gas exchange, plant water relations, and foliar injury were measured. Climatic data,ambient- and chamber-ozone-concentrations were monitored. We found that seedlings grown under irrigated conditions had similar (in 1998) but significantly higher gas exchange rates (in 1999) than seedlings grown within non-irrigated plots among similar ozone exposures. Cherry and ash had similar ozone uptake but cherry developed more ozone-induced injury (< 34% affected leaf area, LAA) than ash (<5% LAA), while maple rarely showed foliar injury, indicating the species differed in ozone sensitivity. Significantly more severe injury on seedlings grown under irrigated conditions than seedlings grown under non-irrigated conditions demonstrated that soil moisture altered seedling responses to ambient ozone exposures.     

Gravitational infusion of ethylenediurea (EDU) into trunks protected adult European ash trees (Fraxinus excelsior L.) from foliar ozone injury

Adult ash trees (Fraxinus excelsior L.), known to be sensitive or insensitive to ozone, determined by presence or absence of foliar symptoms in previous years, were treated with ethylenediurea (EDU) at 450 ppm by gravitational trunk infusion on six occasions at 21-day intervals in summer 2005 at Turin, Italy. At the end of the season, foliar ozone injury on EDU-treated trees was not complete, but was greatly and significantly reduced when compared to results from trees infused with water. Significant symptom reduction occurred at any crown level in the treated trees suggesting that EDU protected whole crowns. Gravitational infusion of EDU resulted in protection from ozone injury for ozone-sensitive ash trees. The amount of EDU needed to provide protection is assumed to be in the range 13-26 mg m(-2) leaf.     

Responses of hybrid poplar clones and red maple seedlings to ambient O(3) under differing light within a mixed hardwood forest

The responses of ramets of hybrid poplar (Populus spp.) (HP) clones NE388 and NE359, and seedlings of red maple (Acer rubrum, L.) to ambient ozone (O(3)) were studied during May-September of 2000 and 2001 under natural forest conditions and differing natural sunlight exposures (sun, partial shade and full shade). Ambient O(3) concentrations at the study site reached hourly peaks of 109 and 98 ppb in 2000 and 2001, respectively. Monthly 12-h average O(3) concentrations ranged from 32.3 to 52.9 ppb. Weekly 12-h average photosynthetically active radiation (PAR) within the sun, partial shade and full shade plots ranged from 200 to 750, 50 to 180, and 25 to 75 micromol m(-2) s(-1), respectively. Ambient O(3) exposure induced visible foliar symptoms on HP NE388 and NE359 in both growing seasons, with more severe injury observed on NE388 than on NE359. Slight foliar symptoms were observed on red maple seedlings during the 2001 growing season. Percentage of total leaf area affected (%LAA) was positively correlated with cumulative O(3) exposures. More severe foliar injury was observed on plants grown within the full shade and partial shade plots than those observed on plants grown within the sun plot. Lower light availability within the partial shade and full shade plots significantly decreased net photosynthetic rate (Pn) and stomatal conductance (g(wv)). The reductions in Pn were greater than reductions in g(wv), which resulted in greater O(3) uptake per unit Pn in plants grown within the partial shade and full shade plots. Greater O(3) uptake per unit Pn was consistently associated with more severe visible foliar injury in all species and/or clones regardless of differences in shade tolerance. These studies suggest that plant physiological responses to O(3) exposure are likely complicated due to multiple factors under natural forest conditions.     

Size-mediated foliar response to ozone in black cherry trees

Local ozone concentration and visible foliar injury were measured over the 1994 growing season on open-grown black cherry (Prunus serotina Ehrh.) trees of varying size (age) within forest stands and adjacent openings at a site in north-central Pennsylvania. Relationships were determined between visible ozone injury and ozone exposure, as well as calculated between injury and ozone uptake expressed as the product of stomatal conductance and ozone concentration. In addition, simultaneous measurements of visible symptoms and leaf gas exchange were also conducted to determine the correlation between visible and physiological injury and ozone exposure. By September, the amount of leaf area affected by visible foliar ozone injury was greatest for seedlings (46%), followed by canopy trees (20%) and saplings (15%). A large amount of variability in foliar ozone symptom expression was observed among trees within a size class. Sum40 and Sum60 (ozone concentration > 40 and > 60 nl liter(-1)) cumulative exposure statistics were the most meaningful indices for interpretation of foliar injury response. Seedlings were apparently more sensitive to ozone injury than larger trees because their higher rates of stomatal conductance resulted in higher rates of ozone uptake. Seedlings also had higher rates of early leaf abscission than larger trees with an average of nearly 30% of the leaves on a shoot abscised by 1 September compared to approximately 5% for larger trees. However, per unit ozone uptake into the leaf, larger trees exhibited larger amounts of foliar injury. The amount of visible foliar injury was negatively correlated (r(2) = 0.82) with net photosynthetic rates, but was not related to stomatal conductance. Net photosynthesis and stomatal conductance thus became uncoupled at high levels of visible foliar injury.     

Foliar injury, leaf gas exchange and biomass responses of black cherry (Prunus serotina Ehrh.) half-sibling families to ozone exposure

Open pollinated families of black cherry seedlings were studied to determine genotypic differences in foliar ozone injury and leaf gas exchange in 1994 and growth response following three growing seasons. An O(3)-sensitive half-sibling family (R-12) and an O(3)-tolerant half-sibling family (MO-7) planted in natural soil were studied along with generic nursery stock (NS) seedlings. Ozone exposure treatments were provided through open top chambers and consisted of 50, 75, and 97% of ambient ozone, and open plots from May 9 to August 26, 1994. Ambient ozone concentrations reached an hourly peak of 88 ppb with 7-hour averages ranging from 39 to 46 ppb. Seedlings in the 50 and 75% of ambient chambers were never exposed to greater than 80 ppb O(3). Visible foliar ozone injury (stipple) was significantly higher for R-12 seedlings than MO-7 seedlings and increased with increasing ozone exposures. For the chamber treatments averaged over all families, there was no significant difference in stomatal conductance and net photosynthetic rates, but there was a significant decrease in root biomass, and a significant decrease in root/shoot ratio between the 50 and 97% of ambient chambers. Stomatal conductance and net photosynthetic rates were significantly different between families with R-12 seedlings generally greater than MO-7 seedlings. The R-12 seedlings had a 7.5 mmol m(-2) increase in ozone uptake compared to MO-7, and at the same cumulative O(3) exposure R-12 exhibited 40.9% stippled leaf area, whereas MO-7 had 9.2% stippled leaf area. Significant differences were observed in stem volume growth and total final biomass between the open-top chambers and open plots. Although R-12 had the most severe foliar ozone injury, this family had significantly greater stem volume growth and total final biomass than MO-7 and NS seedlings. Root:shoot ratio was not significantly different between MO-7 and R-12 seedlings.     

Ozone uptake (flux) as it relates to ozone-induced foliar symptoms of Prunus serotina and Populus maximowiziixtrichocarpa

Field studies were conducted during 2003 and 2004 from early June to the end of August, at 20 sites of lower or higher elevation within north-central Pennsylvania, using seedlings of black cherry (Prunus serotina, Ehrh.) and ramets of hybrid poplar (Populus maximowiziixtrichocarpa). A linear model was developed to estimate the influence of local environmental conditions on stomatal conductance. The most significant factors explaining stomatal variance were tree species, air temperature, leaf vapor pressure deficit, elevation, and time of day. Overall, environmental factors explained less than 35% of the variation in stomatal conductance. Ozone did not affect gas exchange rates in either poplar or cherry. Ozone-induced foliar injury was positively correlated with cumulative ozone exposures, expressed as SUM40. Overall, the amount of foliar injury was better correlated to a flux-based approach rather than to an exposure-based approach. More severe foliar injuries were observed on plants growing at higher elevations.     

Concentrations of mercury in otters (Lutra lutra L.) in Scotland in relation to rainfall

The incidence and severity of visible foliar ozone injury on black cherry (Prunus serotina) seedlings and saplings and tall milkweed (Asclepias exaltata) plants in Great Smoky Mountains National Park (GRSM) were determined by surveys along selected trails conducted during late summer 1992. The incidence (% injured plants) of ozone injury on black cherry was 47% and the percent injured leaves/injured plant and average leaf area injured were 43 and 6%, respectively. Maximum severity (avg. leaf area of the most severely injured leaf) was 12%. Black cherry seedlings and saplings exhibiting ozone injury were taller than non-injured plants. When insect feeding was present, it occurred 96% of the time on plants with ozone injury. Significantly more injury (p=0.007) on black cherry (% injured leaves/injured black cherry) occurred in the NW section of GRSM compared with the other Park sections. Regression analyses showed no relationships in ozone injury with respect to aspect, slope or elevation. Tall milkweed was evaluated twice during August for ozone injury. The incidence (% injured plants) of ozone injury was 74 and 79% for the first and second survey, respectively. The percentage of injured leaves per plant from the first to second survey was 63 to 79%, respectively. Tall milkweeds showing ozone injury were taller than the non-injured plants. The percentage of insect-damaged plants was 50% among plants without ozone injury and 60% among ozone-injured plants. Non-injured tall milkweed had fewer flowers and/or pods than the injured plants. Mean leaf area injured increased over time, and mean maximum leaf area injured increased from 8 to 11% during the same period. Regression analyses showed no differences in ozone injury regarding aspect, slope or elevation. Our findings indicate that ozone injury is widespread throughout the Park on sensitive vegetation.     

Seasonal trends in reduced leaf gas exchange and ozone-induced foliar injury in three ozone sensitive woody plant species

Seasonal trends in leaf gas exchange and ozone-induced visible foliar injury were investigated for three ozone sensitive woody plant species. Seedlings of Populus nigra L., Viburnum lantana L., and Fraxinus excelsior L. were grown in charcoal-filtered chambers, non-filtered chambers and open plots. Injury assessments and leaf gas exchange measurements were conducted from June to October during 2002. All species developed typical ozone-induced foliar injury. For plants exposed to non-filtered air as compared to the charcoal-filtered air, mean net photosynthesis was reduced by 25%, 21%, and 18% and mean stomatal conductance was reduced by 25%, 16%, and 8% for P. nigra, V. lantana, and F. excelsior, respectively. The timing and severity of the reductions in leaf gas exchange were species specific and corresponded to the onset of visible foliar injury.     

Foliar response of black cherry (Prunus serotina) clones to ambient ozone exposure in central Pennsylvania

During late summer of 1996 and 1997 we examined ozone-induced foliar injury in a plantation of 111 black cherry trees (ramets) comprising 15 clones originating from wild ortets growing in the Allegheny National Forest, Pennsylvania, and the Monongahela National Forest, West Virginia. The experimental plantation was a clonal seed orchard in Centre County, Pennsylvania, started in 1971 using ortet buds grafted onto seedling rootstocks of mixed origin. Clones differed significantly in severity of foliar injury symptoms (F=31.83, p<0.001). One clone (R-12) had significantly more foliar injury with >50% leaf area affected than other clones during both years. In contrast, clone R-14, which is from the same area in northcentral Pennsylvania as R-12, exhibited significantly less injury (LAA<6%). Although ambient O(3) concentrations were similar in both years, foliar injury was significantly greater (15.7%) in 1996 than in 1997 (9.9%). This is probably explained by lower stomatal conductance in 1997 caused by drier and hotter weather patterns in June and July of that year. Despite very different weather patterns and overall levels of injury in 1996 and 1997, mean clonal injury was significantly correlated between both years of assessment (r=0.92, p<0.001). Within tree crowns, foliage in lower and inner crown positions was significantly more injured than foliage in upper and exterior crown positions. There was no evidence of geographically based population differences in sensitivity to foliar O(3) injury. On the contrary, results demonstrate that wild genotypes of proximal geographic origin may differ greatly in sensitivity.     

Influence of light fleck and low light on foliar injury and physiological responses of two hybrid poplar clones to ozone

Five-month old hybrid poplar clones NE388 and NE359 were exposed to square-wave 30, 55, and 80 ppb O(3) (8 h/day, 7 day/week) under constant high light (HL) and light fleck (LF) during 28 May-29 June 1999, and exposed to 30 and 55 ppb O(3) under HL, LF, and constant low light (LL) during 22 May-28 June 2000 within Continuously Stirred Tank Reactors (CSTR) in a greenhouse. Ramets of these two hybrid clones received similar total photosynthetically active radiation (PAR) within the LF and LL treatments. Visible foliar symptoms, leaf gas exchange, and growth were measured. More severe O(3) induced foliar symptoms were observed on ramets within the LF and LL treatments than within the HL treatment for both clones. The LF treatment resulted in significantly greater foliar injury than the LL treatment for NE388. The LF and LL treatments generally resulted in lower photosynthetic rates (Pn) for both clones, but did not affect stomatal conductance (g(wv)); therefore, the ratios of g(wv)/Pn and the O(3) uptake/Pn were greatest in plants grown under the LF treatment, followed by those grown under LL treatment; plants grown under HL had the lowest ratios of g(wv)/Pn and O(3) uptake/Pn. Greater ratios of g(wv)/Pn and O(3) uptake/Pn were consistently associated with more severe visible foliar symptoms. The negative impacts of the LF treatment on growth were greater than those of the LL treatment. Results indicate that not only the integral, but also the pattern of photo flux density, may affect carbon gain in plants. Increased foliar injury may be expected under light fleck conditions due to the limited repair capacity as a result of continuity of O(3) uptake while photosynthesis decreases under LL conditions.     

Protection of ash (Fraxinus excelsior) trees from ozone injury by ethylenediurea (EDU): roles of biochemical changes and decreased stomatal conductance in enhancement of growth

Treatments with ethylenediurea (EDU) protect plants from ozone foliar injury, but the processes underlying this protection are poorly understood. Adult ash trees (Fraxinus excelsior), with or without foliar ozone symptoms in previous years, were treated with EDU at 450ppm by gravitational trunk infusion in May-September 2005 (32.5ppmh AOT40). At 30-day intervals, shoot growth, gas exchange, chlorophyll a fluorescence, and water potential were determined. In September, several biochemical parameters were measured. The protective influence of EDU was supported by enhancement in the number of leaflets. EDU did not contribute its nitrogen to leaf tissue as a fertiliser, as determined from lack of difference in foliar N between treatments. Both biochemical (increase in ascorbate-peroxidase and ascorbic acid, and decrease in apoplastic hydrogen peroxide) and biophysical (decrease in stomatal conductance) processes regulated EDU action. As total ascorbic acid increased only in the asymptomatic trees, its role in alleviating O(3) effects on leaf growth and visible injury is controversial.     

Ambient ozone effects on gas exchange and total non-structural carbohydrate levels in cutleaf coneflower (Rudbeckia laciniata L.) growing in Great Smoky Mountains National Park

Ozone-sensitive and -tolerant individuals of cutleaf coneflower (Rudbeckia laciniata L.) were compared for their gas exchange characteristics and total non-structural carbohydrates at Purchase Knob, a high elevation site in Great Smoky Mountains National Park, USA. Photosynthesis and stomatal conductance decreased with increased foliar stipple. Sensitive plants had lower photosynthetic rates for all leaves, except the very youngest and oldest when compared to tolerant plants. Stomatal conductance decreased with increasing leaf age, but no ozone-sensitivity differences were found. Lower leaves had less starch than upper ones, while leaves on sensitive plants had less than those on tolerant plants. These results show that ambient levels of ozone in Great Smoky Mountains National Park can adversely affect gas exchange, water use efficiency and leaf starch content in sensitive coneflower plants. Persistence of sensitive genotypes in the Park may be due to physiological recovery in low ozone years.     

Physiology, morphology, and ozone uptake of leaves of black cherry seedlings, saplings, and canopy trees

Patterns of ozone uptake were related to physiological, morphological, and phenological characteristics of different-sized black cherry trees (Prunus serotina Ehrh.) at a site in central Pennsylvania. Calculated ozone uptake differed among open-grown seedlings, forest gap saplings, and canopy trees and between leaves in the upper and lower crown of saplings and canopy trees. On an instantaneous basis, seedling leaves had the greatest ozone uptake rates of all tree size classes due to greater stomatal conductance and higher concentrations of ozone in their local environment. A pattern of higher stomatal conductance of seedlings was consistent with higher incident photosynthetically-active radiation, stomatal density, and predawn xylem water potentials for seedlings relative to larger trees. However, seedlings displayed an indeterminate pattern of shoot growth, with the majority of their leaves produced after shoot growth had ceased for canopy and sapling trees. Full leaf expansion occurred by mid-June for sapling and canopy trees. Because many of their leaves were exposed to ozone for only part of the growing season, seedlings had a lower relative exposure over the course of the growing season, and subsequently lower cumulative uptake, of ozone than canopy trees and a level of uptake similar to upper canopy leaves of saplings. Visible injury symptoms were not always correlated with patterns in ozone uptake. Visible symptoms were more apparent on seedling leaves in concurrence with their high instantaneous uptake rates. However, visible injury was more prevalent on leaves in the lower versus upper crown of canopy trees and saplings, even though lower crown leaves had less ozone uptake. Lower crown leaves may be more sensitive to ozone per unit uptake than upper crown leaves because of their morphology. In addition, the lower net carbon uptake of lower crown leaves may limit repair and anti-oxidant defense processes.     

Methane emissions from wastewater management

Gas exchange and ozone-induced foliar injury were intensively measured during a 6-day period in mid-August 1998 on leaves of Acer pseudoplatanus, Betula pendula, Corylus avellana, Fagus sylvatica, Fraxinus excelsior, Morus nigra, Prunus avium, Prunus serotina, Rhamnus cathartica, and Viburnum lantana at a forest nursery site in Canton Ticino, Switzerland. Plants were grown in four open plots (AA), four open-top chambers receiving carbon-filtered (CF) air, and four receiving non-filtered (NF) air. Significant variation in gas exchange (F > 12.7, P < 0.001) was detected among species with average net photosynthesis and average stomatal conductance differing by a factor of two. Species also varied significantly in foliar injury for those leaves for which we measured gas exchange (F = 39.6, P < 0.001). Fraxinus excelsior, M. nigra, P. avium, P. serotina, R. cathartica, and V. lantana showed more injury than A. pseudoplatanus, B. pendula, C. avellana, and Fagus sylvatica. Plants grown in CF chambers had significantly higher net photosynthesis (A) and stomatal conductance to water vapor (gwv), and lower foliar injury than plants grown in NF chambers and AA plots; interactions between species and ozone treatments were significant for all variables (F > or = 2.2, P < 0.05) except gwv (F = 0.7, P > 0.1). Although A and gwv decreased and foliar injury increased with leaf age, the magnitude of these changes was lower for plants grown in CF chambers than for plants grown in NF chambers and AA plots. Neither ozone uptake threshold (r = 0.26, P > 0.20) nor whole-plant injury (r = -0.15, P > 0.41) was significantly correlated with stomatal conductance across these species. It appears that the relationships between stomatal conductance and foliar injury are species-specific and interactions between physiology and environments and leaf biochemical processes must be considered in determining species sensitivity to ambient ozone exposures.     

Seasonal development of ozone-induced foliar injury on tall milkweed (Asclepias exaltata) in Great Smoky Mountains National Park

The goals of this study were to document the development of ozone-induced foliar injury, on a leaf-by-leaf basis, and to develop ozone exposure relationships for leaf cohorts and individual tall milkweeds (Asclepias exaltata L.) in Great Smoky Mountains National Park. Plants were classified as either ozone-sensitive or insensitive based on the amount of foliar injury. Sensitive plants developed injury earlier in the season and to a greater extent than insensitive plants. Older leaf cohorts were more likely to belong to high injury classes by the end of each of the two growing seasons. In addition, leaf loss was more likely for older cohorts (2000) and lower leaf positions (2001) than younger cohorts and upper leaves, respectively. Most leaves abscised without prior ozone-like stippling or chlorosis. Failure to take this into account can result in underestimation of the effects of ozone on these plants.     

Response to ambient ozone of two white clover (Trifolium repens L.cv. "Regal") clones, one resistant and one sensitive, grown in a Mediterranean environment

Two clones of white clover (Trifolium repens L.) differing in ozone tolerance were grown in southern Italy during 1997 and 1998 to study the effects of ambient ozone exposure on yield, leaf morphology and water use. Ambient ozone levels were high in both years with values exceeding the threshold for leaf injury reported in the literature. In both years ozone injury was observed on the sensitive clone (NC-S) but not on the resistant one (NC-R), and leaf and stolon dry matter production was significantly lower in NC-S than in NC-R. However, it cannot be excluded that other factors, such as high temperature, interacted with the effect of ozone on biomass production. The clones differed in morphological characteristics. Lower total leaf area in NC-S plants was due to a smaller number of leaves per plant, but the average area per leaf was higher in NC-S. Specific leaf weight and net assimilation rate were higher in the more productive clone (NC-R). Cumulative plant water use was higher in NC-R in each growth period because of the larger leaf area; by contrast, water use per unit leaf area was higher in NC-S, indicating higher leaf conductance to water vapour. The results suggest that ozone significantly reduces the yield of sensitive white clover plants under well-watered conditions, and that the difference in ozone tolerance between clover clones is related to differences in leaf morphology and water use.     

Environmental control of stomatal conductance in forest trees of the Great Smoky Mountains National Park

To determine if stomatal conductance (g(s)) of forest trees could be predicted from measures of leaf microclimate, diurnal variability in in situ g(s) was measured in black cherry (Prunus serotina), red maple (Acer rubrum), and northern red oak (Quercus rubra). Relative to overstory trees, understory saplings exhibited little diurnal variability in g(s) and ozone uptake. Depending on species and site, up to 30% of diurnal and seasonal variation in g(s )of overstory trees was explained by photosynthetically active radiation and vapor pressure deficit. Daily maximum g(s) was significantly related to soil moisture in overstory northern red oak and black cherry (R(2) ranged from 33 to 65%). Although g(s) was not fully predicted using instantaneous measures of leaf microclimate, ozone uptake of large forest trees was reduced by low soil moisture.     

Effects of air quality on foliar injury, growth, yield, and quality of muskmelon

An evaluation of the effects of ambient ozone (O3) on muskmelon was conducted with the use of open-top chambers (OTCs). 'Superstar' muskmelons grown in charcoal-filtered (CF) chambers compared to those grown in nonfiltered (NF) chambers showed significant differences in the severity of visible foliar O3 injury. Furthermore, plants grown in NF conditions had significantly less (21.3%) marketable fruit weight and fewer (20.9%) marketable fruit number than those from CF chambers. No differences were found in early biomass production, leaf area, or number of nodes after 3 weeks of exposure to treatment conditions. Ambient O3 did not affect soluble solids content of mature fresh fruit nor foliage fresh weight at final harvest. Results indicate that ambient concentrations of O3 in southwestern Indiana caused significant foliar injury and yield loss to muskmelons.     

The effect of ambient ozone pollution and acidic rain on the growth and chlorophyll content of green and white ash

Two- and three-year old green ash (Fraxinus americana L.) and white ash (Fraxinus pennsylvanica Marsh.) seedlings were exposed to combinations of ambient ozone and acidic ambient rainfall in New Brunswick, New Jersey. During the 3-year study the potted seedlings did not develop typical foliar ozone toxicity symptoms, despite the occurrence of as many as 78 h in exceedance of the National Ambient Air Quality Standard of 0.12 ppm. Although the pH of the rainfall was as low as 3.6 and averaged 4.1, no symptoms were observed resulting from the ambient precipitation. The rate of shoot growth in terms of height and diameter was generally not affected by either of the pollutants during the growing season. Although the chlorophyll content of white ash foliage was low following frequent rainfall in the early summer of 1984, there was no statistically significant evidence that acid raid or ambient ozone decreased chlorophyll in ash seedlings during the 3-year study.     

Observations of ozone-induced foliar injury on black cherry (Prunus serotina, var. capuli) within the Desierto de Los Leones National Park, Mexico City

A survey for ozone-induced foliar injury of black cherry was conducted in mid-June 1995 within the Desierto de Los Leones National Park located southwest of Mexico City. Evaluations of the upper and lower tree crowns of 18 trees revealed evidence of significant upper surface stipple, leaf reddening and premature senescence on 72% of the trees. A general survey of an additional 169 trees disclosed that 41% exhibited similar symptoms. A gradient of increasing symptoms with increasing elevation was also evident. For the most part, asymptomatic trees were observed to be situated within well-shaded coves at the lower elevations with very few symptomatic trees present in these areas.