Tree mortality in the eastern and central United States: patterns and drivers

Substantial uncertainty surrounds how forest ecosystems will respond to the simultaneous impacts of multiple global change drivers. Long‐term forest dynamics are sensitive to changes in tree mortality rates; however, we lack an understanding of the relative importance of the factors that affect tree mortality across different spatial and temporal scales. We used the US Forest Service Forest Inventory and Analysis database to evaluate the drivers of tree mortality for eastern temperate forest at the individual‐level across spatial scales from tree to landscape to region. We investigated 13 covariates in four categories: climate, air pollutants, topography, and stand characteristics. Overall, we found that tree mortality was most sensitive to stand characteristics and air pollutants. Different functional groups also varied considerably in their sensitivity to environmental drivers. This research highlights the importance of considering the interactions among multiple global change agents in shaping forest ecosystems.     

Concentrations,deposition, and effects of nitrogenous pollutants in selected California ecosystems

Atmospheric deposition of nitrogen (N) in California ecosystems is ecologically significant and highly variable, ranging from about 1 to 45 kg/ha/year. The lowest ambient concentrations and deposition values are found in the eastern and northern parts of the Sierra Nevada Mountains and the highest in parts of the San Bernardino and San Gabriel Mountains that are most exposed to the Los Angeles air pollution plume. In the Sierra Nevada Mountains, N is deposited mostly in precipitation, although dry deposition may also provide substantial amounts of N. On the western slopes of the Sierra Nevada, the majority of airborne N is in reduced forms as ammonia (NH3) and particulate ammonium (NH4+) from agricultural activities in the California Central Valley. In southern California, most of the N air pollution is in oxidized forms as nitrogen oxides (NOx), nitric acid (HNO3), and particulate nitrate (NO3-) resulting from fossil fuel combustion and subsequent complex photochemical reactions. In southern California, dry deposition of gases and particles provides most (up to 95%) of the atmospheric N to forests and other ecosystems. In the mixed-conifer forest zone, elevated deposition of N may initially benefit growth of vegetation, but chronic effects may be expressed as deterioration of forest health and sustainability. HNO3 vapor alone has a potential for toxic effects causing damage of foliar surfaces of pines and oaks. In addition, dry deposition of predominantly HNO3 has lead to changes in vegetation composition and contamination of ground- and stream water where terrestrial N loading is high. Long-term, complex interactions between N deposition and other environmental stresses such as elevated ozone (O3), drought, insect infestations, fire suppression, or intensive land management practices may affect water quality and sustainability of California forests and other ecosystems.     

Impacts of acid deposition,ozone exposure and weather conditions on forest ecosystems in Europe: an overview

Background

In 1994, a “Pan-European Programme for Intensive and Continuous Monitoring of Forest Ecosystems” started to contribute to a better understanding of the impact of air pollution, climate change and natural stress factors on forest ecosystems. The programme today counts approximately 760 permanent observation plots including near 500 plots with data on both air quality and forest ecosystem impacts.

Scope

This paper first presents impacts of air pollution and climate on forests ecosystems as reported in the literature on the basis of laboratory and field research. Next, results from monitoring studies, both at a European wide scale and related national studies, are presented in terms of trends and geographic variations in nitrogen and sulphur deposition and ozone concentrations and the impacts of those changes in interaction with weather conditions on (i) water and element budgets and nutrient-acidity status, (ii) forest crown condition, (iii) forest growth and carbon sequestration and (iv) species diversity of the ground vegetation. The empirical, field based forest responses to the various drivers are evaluated in view of available knowledge.

Conclusions

Analyses of large scale monitoring data sets show significant effects of atmospheric deposition on nutrient-acidity status in terms of elevated nitrogen and sulphur or sulphate concentrations in forest foliage and soil solution and related soil acidification in terms of elevated aluminium and/or base cation leaching from the forest ecosystem. Relationships of air pollution with crown condition, however, appear to be weak and limited in time and space, while climatic factors appear to be more important drivers. Regarding forest growth, monitoring results indicate a clear fertilization effect of N deposition on European forests but the field evidence for impacts of ambient ozone exposure on tree growth is less clear.     

Air pollution: forest health and passive sampling

32nd Annual Air Pollution Workshop
Auburn University in Auburn, AL, USA, April 2000
Air pollution has profound effects on agriculture, forests and natural ecosystems. The first Air Pollution Workshop was held over 30 years ago, and the most vital issues have always been highlighted within this forum. This year, forest health and passive sampling of air pollutants were two key areas of interest.     

森林火灾污染物质释放及其影响研究进展

森林火灾是大气中气体污染物和颗粒物的重要来源,可对全球气候系统、大气环境以及生态系统产生重要影响,对全球温室气体和含碳颗粒物释放具有重要的贡献,是推动全球气候变化的重要因素。森林火灾释放污染物已成为区域乃至全球范围内重要污染源之一,这些污染物质与辐射、能见度以及温室效应等问题直接相关。准确地描述森林火灾释放的气体和颗粒污染物释放机理、释放总量、时空分布特征、不同尺度的扩散过程模拟,以及对区域大气环境的影响,对于量化森林火灾释放污染物总量及区域影响具有重要意义。基于森林火灾污染物质释放方面的国内外文献,从火灾释放的污染物质对环境的影响、森林火灾释放污染物定量化和传输路径监测的研究方法、污染物质的扩散和运输模型以及跨区域影响等几个方面进行了综述。森林火灾释放的CO、PM₁₀和PM_2.5对环境和人的生命安全造成巨大威胁,而且森林火灾释放的污染物质能够随气流长距离传输,不仅对当地的空气造成污染,污染物也能够随着气团进行长距离传输,并在传输过程与当地气溶胶混合,形成跨区域污染。森林火灾释放污染物扩散、传输模拟通过不同模型相互耦合完成,包括可燃物载量估算模型、可燃物消耗和释放模型、污染物扩散传输模型,以及污染物预测和可视化模型等。总结了国内外森林火灾释放污染物质主要研究方法,并展望了今后研究重点:目前我国关于森林火灾释放物质相关的研究尚不足以支撑我国森林火灾温室气体释放、污染物释放等方面的研究,并且我国目前还没有发展出适合于我国的森林火灾污染物释放模型,以及污染物扩散、传输系统。森林火灾排放因子库大多数引用国外研究结果,在一定程度上增加不确定性,缺乏森林火灾对区域大气环境影响的定量化研究。因此,今后我国应加强对森林火灾污染物质释放与影响的研究,尤其是污染物质扩散和传输模型的预测和可视化研究以及排放因子的测量。     

On the importance of incorporating forest edge deposition for evaluating exceedance of critical pollutant loads

The concept of critical load (CL) was defined to express the tolerance of natural and semi‐natural habitats for anthropogenic air pollution. Correct evaluation of the exceedance of critical loads is fundamental for the long‐term protection of ecosystems by limiting emissions of potential acidifying and eutrophying pollutants. For forest ecosystems, the exceedance of critical loads is often calculated using deposition data measured in the forest interior. However, several studies report forest edges acting as ‘hotspots’ of acidifying and nitrogen deposition, showing up to fourfold increases in atmospheric deposition compared to the forest interior. This paper estimates the relevance of considering the higher deposition load in forest edges for calculating exceedance of critical loads for nitrogen and potential acidifying deposition. If measures to control and reduce atmospheric deposition are based on mean deposition fluxes within forest stands, deposition reductions will not be enough for preventing adverse effects. In fact, emission reductions should be adjusted to deposition values at the forest edge, since these zones are most threatened. We thus conclude that there is an urgent need to reconsider the calculation of exceedance of critical loads, taking into account edge enhancement of deposition. This is an issue of high relevance, particularly in highly fragmented regions, such as Flanders (Belgium).     

Impacts of air pollution and climate change on forest ecosystems--emerging research needs

Outcomes from the 22nd meeting for Specialists in Air Pollution Effects on Forest Ecosystems "Forests under Anthropogenic Pressure--Effects of Air Pollution, Climate Change and Urban Development", September 10-16, 2006, Riverside, CA, are summarized. Tropospheric or ground-level ozone (O3) is still the phytotoxic air pollutant of major interest. Challenging issues are how to make O3 standards or critical levels more biologically based and at the same time practical for wide use; quantification of plant detoxification processes in flux modeling; inclusion of multiple environmental stresses in critical load determinations; new concept development for nitrogen saturation; interactions between air pollution, climate, and forest pests; effects of forest fire on air quality; the capacity of forests to sequester carbon under changing climatic conditions and coexposure to elevated levels of air pollutants; enhanced linkage between molecular biology, biochemistry, physiology, and morphological traits.     

The macroinvertebrates in a sewage-polluted aquifer

A quantitative evaluation of stream temperature alterations due to a commercial forest harvesting practice and a research treatment is presented. Summer maximum stream temperatures averaged 1 ° C higher in the commercial clearcut and 9 °C higher in the clearcut-herbicided watershed than in the forested control. The largest average monthly temperature increase on the commercial clearcut (2.2 °C) occurred during April; on the clearcut-herbicided basin it occurred during June (10.5 °C). Significant changes in stream temperature were observed on both watersheds as early as February and as late as November. Changes in minimum stream temperatures are presented in detail along with the impact on diel temperature fluctuations. Changes in the stream temperature regimes of the clearcut watersheds from the headwaters to the mouth of the watersheds are also given. Potential impacts of the stream temperature alterations on aquatic ecosystems are summarized in relation to stress limits for brook trout and other organisms. School of Forest Resources, Pennsylvania State University, University Park, PA 16802, U.S.A. USDA Forest Service, Northeastern Forest Experiment Station, University Park, PA 16802, U.S.A.     

亚热带典型流域C、N沉降季节变化特征及其耦合输出过程

大气湿沉降是流域生态系统水体中碳氮的重要来源,对生态系统的健康及稳定性有很大的影响。通过对江西千烟洲典型亚热带流域降雨过程的碳、氮湿沉降和径流过程的季节性动态特征进行监测分析,探讨流域沉降、径流输出的C、N耦合及平衡关系。结果表明:千烟洲香溪流域降雨径流中碳氮浓度明显低于雨水,流域大气降水中DOC浓度和TN浓度呈极显著正相关关系。香溪河流域常规水体C∶N均值为2.81,远低于根据Redfield比率得出的适宜浮游生物生长的C∶N(6.6左右),说明外源性N输入导致该流域水体环境处于N过量的状态,长期输出会提高下游鄱阳湖水系的营养化程度。降雨过程对流域碳输入输出平衡影响较小,对氮输入输出平衡的影响较大。流域湿沉降DOC年输入量为69.41 kg hm~(-2)a~(-1),TN湿沉降通量为77.23 kg hm~(-2)a~(-1),碳氮沉降水平受区域降雨量及空气污染情况控制。香溪流域生态系统截留的沉降TN占当地氮肥年均使用量的33.13%,大气降水对亚热带流域生态系统的大量营养物质输入不容忽视。     

A Pilot Application of Regional Scale Risk Assessment to the Forestry Management of the Upper Grande Ronde Watershed,Oregon

An issue in forestry management has been the integration of a variety of different information into a threat analysis or risk assessment. In this instance, regional scale risk assessment was applied to the Upper Grande Ronde watershed in eastern Oregon to examine the potential of risk assessment for use in the management of broad landscapes. The site was a focus of study for the U.S. Forest Service through the Interior Northwest Landscape Analysis System (INLAS) project. In the study, a range of stressors, habitats, and endpoints were identified from previous studies in the watershed, and endpoints were determined from meeting with the primary stakeholder, the U.S. Forest Service. These endpoints were focused around the historic range of variability (HRV) defined for the area. The relative risk model (RRM) incorporating a Monte Carlo analysis was used as the analysis tool. The risk model output showed the HRV fire regime was the endpoint most at risk. The results of this analysis were compared to the Wallowa-Whitman National Forest prioritization of watershed restoration analysis. The RRM demonstrated similar results but with a better accounting for uncertainty. From this trial the RRM has proven to be a potential management tool for forestry management.     

Impacts of tropospheric ozone and climate change on net primary productivity and net carbon exchange of China's forest ecosystems

Aim We investigated how ozone pollution and climate change/variability have interactively affected net primary productivity (NPP) and net carbon exchange (NCE) across China's forest ecosystem in the past half century. Location Continental China. Methods Using the dynamic land ecosystem model (DLEM) in conjunction with 10‐km‐resolution gridded historical data sets (tropospheric O₃ concentrations, climate variability/change, and other environmental factors such as land‐cover/land‐use change (LCLUC), increasing CO₂ and nitrogen deposition), we conducted nine simulation experiments to: (1) investigate the temporo‐spatial patterns of NPP and NCE in China's forest ecosystems from 1961–2005; and (2) quantify the effects of tropospheric O₃ pollution alone or in combination with climate variability and other environmental stresses on forests' NPP and NCE. Results China's forests acted as a carbon sink during 1961–2005 as a result of the combined effects of O₃, climate, CO₂, nitrogen deposition and LCLUC. However, simulated results indicated that elevated O₃ caused a 7.7% decrease in national carbon storage, with O₃‐induced reductions in NCE (Pg C year^?1) ranging from 0.4–43.1% among different forest types. Sensitivity experiments showed that climate change was the dominant factor in controlling changes in temporo‐spatial patterns of annual NPP. The combined negative effects of O₃ pollution and climate change on NPP and NCE could be largely offset by the positive fertilization effects of nitrogen deposition and CO₂. Main conclusions In the future, tropospheric O₃ should be taken into account in order to fully understand the variations of carbon sequestration capacity of forests and assess the vulnerability of forest ecosystems to climate change and air pollution. Reducing air pollution in China is likely to increase the resilience of forests to climate change. This paper offers the first estimate of how prevention of air pollution can help to increase forest productivity and carbon sequestration in China's forested ecosystems.     

High Foliar and Soil Nitrogen Concentrations in Central Appalachian Forests

Regional topography and climate variation yield differences in ecosystem attributes that make spatially scaled estimates of forest productivity challenging. Foliar nitrogen is a primary indicator of forest ecosystem productivity and is used in regional estimates of terrestrial productivity, but this characteristic has not been well described in the Central Appalachian region. Here we describe foliar and soil N variation among species and elevations at two spatial scales in the Central Appalachian region: (1) across the Elklick watershed in the Fernow Experimental Forest and (2) across the state of West Virginia. We found higher foliar N concentrations at both scales than those previously reported for other temperate forest regions. Canopy and soil nitrogen concentrations were also much greater in the Fernow than generally observed across West Virginia. Soil N concentrations in the Fernow were two times greater than those observed across West Virginia. Species-related differences were observed at both spatial scales, but were not always consistent. Canopy N ranges are generally consistent across elevations throughout the state of West Virginia, but should be scaled according to species-related elevation effects for studies that estimate productivity differences in response to harvest or changing species composition. The incongruence of foliar and soil N concentrations at the Fernow Experimental Forest are not explained by elevation or species composition, but are likely a consequence of greater historical N and H⁺ deposition relative to the surrounding West Virginia region.     

Nitrous oxide emissions in proportion to nitrification in moist temperate forests

Chronic elevated nitrogen deposition has increased nitrogen availability in many forest ecosystems globally, and this phenomenon has been suggested to increase soil nitrification. Although it is believed that increased nitrogen availability would also increase nitrous oxide (N₂O) emissions from forest ecosystems, its impact on N₂O flux is poorly known. In this study, 3-years monitoring of N₂O emissions was performed in a forested watershed receiving elevated nitrogen deposition and located in the suburbs of Tokyo, Japan. In addition, a comparative field survey was carried out in nine temperate forest sites with varying nitrogen availabilities. In the intensively studied forest site showing typical nitrogen saturation, the average annual N₂O emissions from the whole watershed were estimated to be 0.88 kg N ha⁻¹ year⁻¹, comparable to the highest observed levels for temperate forests except for some very high emission sites in Europe. Although no correlation was found for humid spots with WFPS > 60%, a clear positive correlation was noted between N₂O flux and net nitrification rate in situ for plots with water-filled pore space (WFPS) < 60%. The N₂O flux varied across nine forest sites almost in proportional to the stream water NO₃⁻ concentration in the watershed that ranged from 0.14 to 1.64 mg N/L. We conclude that N₂O emissions are related to nitrification in moist temperate forest, which may be associated with the magnitude of nitrogen saturation.

     

Tree diversity mitigates defoliation after a drought‐induced tipping point

Understanding the processes that underlie drought‐related tree vitality loss is essential for anticipating future forest dynamics, and for developing management plans aiming at increasing the resilience of forests to climate change. Forest vitality has been continuously monitored in Europe since the acid rain alert in the 1980s, and the intensive monitoring plots of ICP Forests offer the opportunity to investigate the effects of air pollution and climate change on forest condition. By making use of over 100 long‐term monitoring plots, where crown defoliation has been assessed extensively since 1990, we discovered a progressive shift from a negative to a positive effect of species richness on forest health. The observed tipping point in the balance of net interactions, from competition to facilitation, has never been reported from real ecosystems outside experimental conditions; and the strong temporal consistency of our observations with increasing drought stress emphasizes its climate change relevance. Furthermore, we show that higher species diversity has reduced the severity of defoliation in the long term. Our results confirm the greater resilience of diverse forests to future climate change‐induced stress. More generally, they add to an accumulating body of evidence on the large potential of tree species mixtures to face manifold disturbances in a changing world.     

中国主要城市森林对大气PM2.5的削减量及其占大气PM2.5总量比例

空气中的细颗粒物(PM_2.5)是我国城市空气污染的主要污染物之一,严重威胁着城市居民的健康,限制城市发展的可持续性。PM_2.5去除的自然途径有两种,分别是干沉降和湿沉降,其中干沉降占据主导作用,且干沉降的过程和效率与城市森林紧密关联。目前针对城市森林对干沉降作用的研究主要是在小尺度中从不同树种、不同群落结构、不同景观类型等角度来估算并比较其滞尘量,较少关注其占空气污染总量的比率,从而可能影响对城市森林滞尘服务能力的判断。因此,利用城市森林效益(Urban Forest Effect, UFORE)模型中的大气污染干沉降模块的核心算法,以2015年为例,估算了我国主要城市辖区的城市森林一年内对大气中的PM_2.5削减量以及其占空气中PM_2.5污染总量的比重。结果显示：(1)2015年全国主要城市单位绿地面积日均滞尘量较高地区主要集中在华北地区、华东地区、以及东北地区。其中北京30.47mg/m²,苏州24.63mg/m²,沈阳28.55mg/m²     

Effects of nitrogen deposition on the greenhouse gas fluxes from forest soils

Nitrogen (N) deposition can alter the rates of microbial N- and C- turnover, and thus can affect the fluxes of greenhouse gases (GHG, e.g., CO₂, CH₄, and N₂O) from forest soils. The effects of N deposition on the GHG fluxes from forest soils were reviewed in this paper. N deposition to forest soils have shown variable effects on the soil GHG fluxes from forest, including increases, decreases or unchanged rates depending on forest type, N status of the soil, and the rate and type of atmospheric N deposition. In forest ecosystems where biological processes are limited by N supply, N additions either stimulate soil respiration or have no significant effect, whereas in “N saturated” forest ecosystems, N additions decrease CO₂ emission, reduce CH₄ oxidation and elevate N₂O flux from the soil. The mechanisms and research methods about the effects of N deposition on GHG fluxes from forest soils were also reviewed in this paper. Finally, the present and future research needs about the effects of N deposition on the GHG fluxes from forest soils were discussed.     

Molecular analysis of fungal communities and laccase genes in decomposing litter reveals differences among forest types but no impact of nitrogen deposition

The fungal community of the forest floor was examined as the cause of previously reported increases in soil organic matter due to experimental N deposition in ecosystems producing predominantly high-lignin litter, and the opposite response in ecosystems producing low-lignin litter. The mechanism proposed to explain this phenomenon was that white-rot basidiomycetes are more important in the degradation of high-lignin litter than of low-lignin litter, and that their activity is suppressed by N deposition. We found that forest floor mass in the low-lignin sugar-maple dominated system decreased in October due to experimental N deposition, whereas forest floor mass of high-lignin oak-dominated ecosystems was unaffected by N deposition. Increased relative abundance of basidiomycetes in high-lignin forest floor was confirmed by denaturing gradient gel electrophoresis (DGGE) and sequencing. Abundance of basidiomycete laccase genes, encoding an enzyme used by white-rot basidiomycetes in the degradation of lignin, was 5-10 times greater in high-lignin forest floor than in low-lignin forest floor. While the differences between the fungal communities in different ecosystems were consistent with the proposed mechanism, no significant effects of N deposition were detected on DGGE profiles, laccase gene abundance, laccase length heterogeneity profiles, or phenol oxidase activity. Our observations indicate that the previously detected accumulation of soil organic matter in the high-lignin system may be driven by effects of N deposition on organisms in the mineral soil, rather than on organisms residing in the forest floor. However, studies of in situ gene expression and temporal and spatial variability within forest floor communities will be necessary to further relate the ecosystem dynamics of organic carbon to microbial communities and atmospheric N deposition.     

离子树脂法测定森林穿透雨氮素湿沉降通量——以千烟洲人工针叶林为例

穿透雨是大气氮素输入森林生态系统的重要途径之一,穿透雨中氮素含量的定量评估在森林生态系统氮素循环研究中的作用不可忽视。穿透雨中氮沉降通量的空间异质性很强,传统降水收集法工作量大,且容易带来测定误差。分析了国产离子树脂测定大气氮素湿沉降的可行性,并以千烟洲人工针叶林为例探讨离子树脂法测定森林穿透雨的适用性。结果表明,离子交换树脂法和传统降水收集法测定值之间的相关性显著,离子交换树脂法可以很好的反映大气氮沉降通量和季节变化特征,并且在采样周期较长时也能准确测定氮沉降组分,是适用于野外站点林内穿透雨氮沉降通量的观测方法。千烟洲人工针叶林穿透雨的氮沉降通量为9.19 kgN?hm-2?a-1,夏季的5—7月份和冬季的1—2月份出现氮沉降通量高峰。夏季穿透雨氮沉降以铵态氮为主,而冬季以硝态氮为主。千烟洲人工针叶林的氮沉降通量与附近地区针叶林穿透雨氮沉降通量近似,低于临近区域阔叶林穿透雨的氮沉降通量水平,但已可能接近森林生态系统氮输出出现强烈反应的氮沉降临界值。     

森林生态系统碳循环对全球氮沉降的响应

森林土壤和植被储存着全球陆地生态系统大约46%的碳,在全球碳平衡中起着非常重要的作用。过去几十年来,森林生态系统的碳循环和碳吸存受到了全球氮沉降的深刻影响,因为氮沉降改变了陆地生态系统的生产力和生物量积累。以欧洲和北美温带森林区域开展的研究为基础,综述了氮沉降对植物光合作用、土壤呼吸、土壤DOM及林木生长的影响特征和机理,探讨了森林生态系统碳动态对氮沉降响应的不确定性因素。热带森林C、N循环与大部分温带森林不同,人为输入的氮对热带生态系统过程的影响也可能不同,因此指出了在热带地区开展碳氮循环耦合研究的必要性和紧迫性。     

Patterns of understory diversity in mixed coniferous forests of southern California impacted by air pollution

The forests of the San Bernardino Mountains have been subject to ozone and nitrogen (N) deposition for some 60 years. Much work has been done to assess the impacts of these pollutants on trees, but little is known about how the diverse understory flora has fared. Understory vegetation has declined in diversity in response to elevated N in the eastern U.S. and Europe. Six sites along an ozone and N deposition gradient that had been part of a long-term study on response of plants to air pollution beginning in 1973 were resampled in 2003. Historic ozone data and leaf injury scores confirmed the gradient. Present-day ozone levels were almost half of these, and recent atmospheric N pollution concentrations confirmed the continued air pollution gradient. Both total and extractable soil N were higher in sites on the western end of the gradient closer to the urban source of pollution, pH was lower, and soil carbon (C) and litter were higher. The gradient also had decreasing precipitation and increasing elevation from west to east. However, the dominant tree species were the same across the gradient. Tree basal area increased during the 30-year interval in five of the sites. The two westernmost sites had 30-45% cover divided equally between native and exotic understory herbaceous species, while the other sites had only 3-13% cover dominated by native species. The high production is likely related to higher precipitation at the western sites as well as elevated N. The species richness was in the range of 24 to 30 in four of the sites, but one site of intermediate N deposition had 42 species, while the easternmost, least polluted site had 57 species. These were primarily native species, as no site had more than one to three exotic species. In three of six sites, 20-40% of species were lost between 1973 and 2003, including the two westernmost sites. Two sites with intermediate pollution had little change in total species number over 30 years, and the easternmost site had more species in 2003. The easternmost site is also the driest and has the most sunlight filtering to the forest floor, possibly accounting for the higher species richness. The confounding effects of the precipitation gradient and possibly local disturbances do not show a simple correlation of air pollution with patterns of native and invasive species cover and richness. Nevertheless, the decline of native species and dominance by exotic species in the two westernmost polluted sites is cause for concern that air pollution is affecting the understory vegetation adversely.