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Precipitation pulses and carbon fluxes in semiarid and arid ecosystems.

In the arid and semiarid regions of North America, discrete precipitation pulses are important triggers for biological activity. The timing and magnitude of these pulses may differentially affect the activity of plants and microbes, combining to influence the C balance of desert ecosystems. Here, we evaluate how a "pulse" of water influences physiological activity in plants, soils and ecosystems, and how characteristics, such as precipitation pulse size and frequency are important controllers of biological and physical processes in arid land ecosystems. We show that pulse size regulates C balance by determining the temporal duration of activity for different components of the biota. Microbial respiration responds to very small events, but the relationship between pulse size and duration of activity likely saturates at moderate event sizes. Photosynthetic activity of vascular plants generally increases following relatively larger pulses or a series of small pulses. In this case, the duration of physiological activity is an increasing function of pulse size up to events that are infrequent in these hydroclimatological regions. This differential responsiveness of photosynthesis and respiration results in arid ecosystems acting as immediate C sources to the atmosphere following rainfall, with subsequent periods of C accumulation should pulse size be sufficient to initiate vascular plant activity. Using the average pulse size distributions in the North American deserts, a simple modeling exercise shows that net ecosystem exchange of CO2 is sensitive to changes in the event size distribution representative of wet and dry years. An important regulator of the pulse response is initial soil and canopy conditions and the physical structuring of bare soil and beneath canopy patches on the landscape. Initial condition influences responses to pulses of varying magnitude, while bare soil/beneath canopy patches interact to introduce nonlinearity in the relationship between pulse size and soil water response. Building on this conceptual framework and developing a greater understanding of the complexities of these eco-hydrologic systems may enhance our ability to describe the ecology of desert ecosystems and their sensitivity to global change.

Carbon↗

Respiration characteristics in temperate rainforest tree species differ along a long-term soil-development chronosequence.

We measured the response of dark respiration (R(d)) to temperature and foliage characteristics in the upper canopies of tree species in temperate rainforest communities in New Zealand along a soil chronosequence (six sites from 6 years to 120,000 years). The chronosequence provided a vegetation gradient characterised by significant changes in soil nutrition. This enabled us to examine the extent to which changes in dark respiration can be applied across forest biomes and the utility of scaling rules in whole-canopy carbon modelling. The response of respiration to temperature in the dominant tree species differed significantly between sites along the sequence. This involved changes in both R(d) at a reference temperature (R(10)) and the extent to which R(d) increased with temperature (described by E(o), a parameter related to the energy of activation, or the change in R(d) over a 10 degrees C range, Q(10)). Site averaged E(o) ranged from 44.4 kJ mol(-1) K(-1) at the 60-year-old site to 26.0 kJ mol(-1) K(-1) at the oldest, most nutrient poor, site. Relationships between respiratory and foliage characteristics indicated that both the temperature response of respiration (E(o) or Q(10)) and the instantaneous rate of respiration increased with both foliar nitrogen and phosphorus content. The ratio of photosynthetic capacity (Whitehead et al. in Oecologia 2005) to respiration (A(max)/R(d)) attained values in excess of 15 for species in the 6- to 120-year-old sites, but thereafter decreased significantly to around five at the 120,000-year-old site. This indicates that shoot carbon acquisition is regulated by nutrient limitations in the retrogressing ecosystems on the oldest sites. Our findings indicate that respiration and its temperature response will vary according to soil age and, therefore, to soil nutrient availability and the stage of forest development. Thus, variability in respiratory characteristics for canopies should be considered when using models to integrate respiration at large spatial scales.

Age Factors↗

Do carbon-based defences reduce foliar damage? Habitat-related effects on tree seedling performance in a temperate rainforest of Chiloé Island, Chile.

Carbon-based secondary compounds (CBSCs), such as phenols or tannins, have been considered as one of the most important and general chemical barriers of woody plants against a diverse array of herbivores. Herbivory has been described as a critical factor affecting the growth and survival of newly established tree seedlings or juveniles then, the presence of secondary metabolites as defences against herbivores should be a primary strategy to reduce foliar damage. We examined whether light-induced changes in leaf phenolic chemistry affected insect herbivory on seedlings of two rainforest tree species, Drimys winteri (Winteraceae) and Gevuina avellana (Proteaceae). Seedlings of both species were planted under closed canopy and in a canopy gap within a large remnant forest patch. Half of the seedlings in each habitat were disinfected with a wide-spectrum systemic insecticide and the other half were used as controls. Seedling growth, survival, and foliar damage (estimated by an herbivory index) due to insect herbivores were monitored over a period of 16 months (December 2001-April 2003). The total leaf content of phenols and condensed tannins were assessed in seedlings from both habitats. As expected, access to light induced a greater production of CBSCs in seedlings of both tree species, but these compounds did not seem to play a significant defensive role, as seedlings grown in gaps suffered greater leaf damage than those planted in forest interior. In addition, in both habitats, seedlings without insecticide treatment suffered a greater foliar damage than those with insecticide, especially 16 months after the beginning of the experiment. Canopy openness and herbivory had positive and negative effects, respectively, on seedling growth and survival in both tree species. In conclusion, despite the higher levels of defence in tree-fall gap, the higher densities of herbivore override this and lead to higher damage levels.

Animals↗

Responses of floodplain forest species to spatially condensed gradients: a test of the flood-shade tolerance tradeoff hypothesis.

Previous work in southeastern US floodplains led to the hypothesis that a tradeoff between flood and shade tolerance underlies species-specific responses to flooding and light, which drive forest regeneration. In systems where community turnover can occur with small-scale environmental changes, testing this hypothesis requires recognizing that turnover of species along the two gradients can be large relative to the spatial distances involved. We test the tradeoff hypothesis in an old-growth bottomland hardwood forest by (1) comparing shade and moisture profiles of woody juveniles versus random points and (2) using individual-based sampling of woody juveniles to model probability of occurrence in response to distance-to-water table and canopy openness gradients. We found that juveniles of all species combined occupied a similar range of distance-to-water table compared to measurements taken at random points, but average canopy openness above seedlings was significantly higher than at random points. On average, shade-tolerant species, with the exception of Acer rubrum, were found in shaded areas that were also drier, whereas less shade-tolerant taxa, plus A. rubrum, were found in wetter, more open areas, suggesting a tradeoff between flood and shade tolerance. Predictive models of species occurrence, which incorporate the availability of canopy and microtopographic conditions, indicated that three taxa (Fraxinus pennsylvanica, Quercus spp., and Ulmus americana) had patterns consistent with a flood-shade tolerance tradeoff. In contrast, Asimina triloba, Celtis laevigata, and Liquidambar styraciflua had positive responses when the joint stresses of flooding and shade were diminished. A. rubrum appeared to be the most tolerant to both stresses. Our work not only lends support to the flood-shade tradeoff hypothesis but also indicates that a more general model is needed that includes a "flood-shade release" component. We also suggest that responses to small-scale gradients should not be overlooked, particularly in systems where the gradients are ecologically steep.

Disasters↗

Limitations to seedling establishment in a mesic Hawaiian forest.

While invasive species may be visible indicators of plant community degradation, they may not constitute the only, or even the primary, limitation to stand regeneration. We used seed-augmentation and grass-removal experiments under different canopy conditions to assess the relative importance of dispersal limitation, resource availability, and competition on seedling establishment in the understory shrubs Sophora chrysophilla, Dodonea viscosa, and Pipturus albidus in a montane mesic forest in Hawaii. The study location was an Acacia koa-Metrosideros polymorpha forest at 1000-1500 m elevation on the leeward side of Hawaii Island; it is a closed-canopy forest historically subject to logging and grazing by cattle and sheep and currently dominated by the exotic grass, Ehrharta stipoides, in the herb layer. Seedling establishment after 1 and 2 years was strongly dispersal limited in Sophora and Dodonea, but not in Acacia, a non-augmented species in which abundant seedlings established, nor in Pipterus, in which only one seedling established in 2 years. Grass cover reduced seedling establishment in Acacia, Sophora, and Dodonea and, for the latter two species, seedling establishment was substantially greater in the warmer, more moist forest at the lowest elevation. Light, moisture, and resin-captured N and P were more strongly affected by elevation and canopy composition than by grass cover, but in most cases seedling establishment was not positively correlated with resource availability. Limitations to the establishment of woody seedlings in this forest-grassland mixture vary among species; however, both dispersal limitation and competition from a shade-tolerant grass are important deterrents to regeneration in these forests.

Ecosystem↗

Leaf anatomy and light acclimation in woody seedlings after gap formation in a cool-temperate deciduous forest.

The photosynthetic light acclimation of fully expanded leaves of tree seedlings in response to gap formation was studied with respect to anatomical and photosynthetic characteristics in a natural cool-temperate deciduous forest. Eight woody species of different functional groups were used; two species each from mid-successional canopy species (Kalopanax pictus and Magnolia obovata), from late-successional canopy species (Quercus crispula and Acer mono), from sub-canopy species (Acer japonicum and Fraxinus lanuginosa) and from vine species (Schizophragma hydrangeoides and Hydrangea petiolaris). The light-saturated rate of photosynthesis (Pmax) increased significantly after gap formation in six species other than vine species. Shade leaves of K. pictus, M. obovata and Q. crispula had vacant spaces along cell walls in mesophyll cells, where chloroplasts were absent. The vacant space was filled after the gap formation by increased chloroplast volume, which in turn increased Pmax. In two Acer species, an increase in the area of mesophyll cells facing the intercellular space enabled the leaves to increase Pmax after maturation. The two vine species did not significantly change their anatomical traits. Although the response and the mechanism of acclimation to light improvement varied from species to species, the increase in the area of chloroplast surface facing the intercellular space per unit leaf area accounted for most of the increase in Pmax, demonstrating the importance of leaf anatomy in increasing Pmax.

Acclimatization↗

Population and clonal level responses of a perennial grass following fire in the northern Chihuahuan Desert.

Relationships involving fire and perennial grasses are controversial in Chihuahuan Desert grasslands of southern New Mexico, USA. Research suggests that fire delays the resprouting of perennial grasses well after two growing seasons. However, such results are confounded by livestock grazing, soil erosion, and drought. Additionally, post-fire grass responses may depend on initial clone size. We evaluated the effects of fire, grazing, and clone size on Bouteloua eriopoda (black grama) in southern New Mexico grasslands. Four 2-ha plots were established in each of four sites. Fire and grazing were applied or not applied in 1999 such that four treatment combinations were assigned randomly to plots within each site. Within each plot, small (0-10 cm(2) basal area), medium (10-30 cm(2)), and large ( > 30 cm(2)) clones were initially mapped in five 0.91-m(2) quadrats where grass attributes and litter cover were evaluated before and at the end of two growing seasons following fire. Maximum fire temperature was also measured. At a population level, canopy and litter cover were each approximately 50% less in burned than unburned areas. However, compared to initial levels, canopy height had increased by 10% at the end of the study, regardless of fire. At a clonal level, basal cover reductions were attributed mostly to large clones that survived fire. Smaller clone densities had decreased by as much as 19% in burned compared to unburned areas, and fire reduced the basal cover of medium clones. Basal and canopy cover, recruitment, and clone basal area decreased with increased fire temperatures. Almost all responses were independent of grazing, and interactive effects of grazing and fire were not detected. Fire did not kill all perennial grass clones, regardless of size. However, rapid responses were likely influenced by above-average precipitation after fire. Future studies in desert grasslands should examine how perennial grass dynamics are affected by fire, precipitation patterns, and interactions with grazing.

Analysis of Variance↗

Highly predictable photosynthetic production in natural macroalgal communities from incoming and absorbed light.

Photosynthesis-irradiance relationships of macroalgal communities and thalli of dominant species in shallow coastal Danish waters were measured over a full year to test how well community production can be predicted from environmental (incident irradiance and temperature) and community variables (canopy absorptance, species number and thallus metabolism). Detached thalli of dominant species performed optimally at different times of the year, but showed no general seasonal changes in photosynthetic features. Production capacity of communities at high light varied only 1.8-fold over the year and was unrelated to incident irradiance, temperature and mean thallus photosynthesis, while community absorptance was a highly significant predictor. Actual rates of community photosynthesis were closely related to incident and absorbed irradiance alone. Community absorptance in turn was correlated to canopy height and species richness. The close relationship of community photosynthesis to irradiance is due to the fact that (1) large differences in thallus photosynthesis of individual species are averaged out in communities composed of several species, (2) seasonal replacement of species keeps communities metabolically active, and (3) maximum possible absorptance at 100% constrains the total photosynthesis of all species. Our results imply that the photosynthetic production of macroalgal communities is more predictable than their complex and dynamic nature suggest and that predictions are possible over wide spatial scales in coastal waters by measurements of vegetation cover, incoming irradiance and canopy absorptance.

Denmark↗

Measuring and modelling plant area index in beech stands.

For some beech ( Fagus sylvatica L.) stands with different stand densities the plant area index (PAI) was measured by means of a Licor LAI-2000 plant canopy analyser. The stands are located on the slopes of a valley in south-west Germany and had been treated by different types of silvicultural management (heavy shelterwood felling, light shelterwood felling, control plot). The analyser was used (a) to investigate the light conditions on plots of the same thinning regime, (b) to quantify the differences between the different treatments and (c) to obtain absolute values of PAI for interdisciplinary research. PAI was measured at three different phenological stages (leafless, leaf-unfolding and fully leafed season in 2000) and was found to be about 5.2 for the fully developed canopy on the control plots, 3.2 on the light fellings and about 2.0 for the heavy fellings. In the leafless period PAI was between 1.1 (control) and 0.4 (heavy felling). Measurements made in summer 2000 and summer 2002 were compared, and showed an increase of PAI, especially on the thinned plots. Measurements of photosynthetically active radiation (PAR) above and below the canopy in combination with measured PAI were used to apply Beer's Law of radiation extinction to calculate the extinction coefficient k for different sky conditions and for the different growing seasons on the control plots. The extinction coefficient k for the beech stands was found to be between 0.99 and 1.39 in the leafless period, 0.62 to 0.91 during leaf unfolding and between 0.68 and 0.83 in the fully leafed period. Using PAR measurements and the k values obtained, the annual cycle of PAI was modelled inverting Beer's Law.

Environmental Monitoring↗

The effect of urban ground cover on microclimate, growth and leaf gas exchange of oleander in Phoenix, Arizona.

We assessed how small patches of contrasting urban ground cover [mesiscape (turf), xeriscape (gravel), concrete, and asphalt] altered the microclimate and performance of adjacent oleander (Nerium oleander L.) plants in Phoenix, Arizona during fall/winter (September-February) and spring/summer (March-September). Ground-cover and oleander canopy surface temperatures, canopy air temperatures and pot soil temperatures tended to be lowest in the mesiscape and highest in the asphalt and concrete. Canopy air vapor pressure deficits were lowest in the mesiscape and highest in the asphalt plot. Rates of net photosynthesis of all oleander plants were highest in October and May, and declined through mid-summer (June-July), when rates tended to be highest in the cooler mesiscape, particularly when water was limiting. During fall/winter, oleanders in the mesiscape produced 20% less biomass, 13% less leaf area, and had 12% lower relative growth rates (R(G)) than those in the other ground covers. Lower nighttime temperatures in the mesiscape in December led to oleander frost damage. During spring/summer, oleanders in the mesiscape produced 11% more biomass, 16% more leaf area, and had 3% higher R(G) than those in the other cover types. The effects of urban ground cover on oleander performance were season-specific; while oleander growth was greatest in the mesiscape during spring/summer, it was lowest during fall/winter and these plants experienced frost damage. Because all oleander plants produced >10 times as much biomass during the spring/summer, on an annual basis oleanders in the mesiscape produced 5-11% more biomass than plants in the warmer ground covers.

Arizona↗

Arbuscular mycorrhizae of the palm Astrocaryum mexicanum in disturbed and undisturbed stands of a Mexican tropical forest.

Tropical forests are dynamic systems with extensive natural disturbance, gaps in the canopy being one of the most important types. Tree and branch fall are often the principal cause of natural disturbance. This research was done on adult individuals of a very abundant palm ( Astrocaryum mexicanum Liebm, Arecaceae), which is found in the understorey of the forest at Los Tuxtlas, Mexico. Percentages of colonization by arbuscular mycorrhizae were determined for individuals selected randomly from plots located both in gaps and under closed canopy. The highest percentages of total colonization, as well as those of hyphae and vesicles, were recorded for gaps. In forest with closed canopy, arbuscules had the highest percentages of colonization; on these sites the palm has been observed to grow less. The higher production of arbuscules may favour nutrient capture in this microenvironment, which is characterized by strong competition.

Arecaceae↗

Shoot morphology of Aucuba japonica incurred by anisophylly: ecological implications.

Anisophylly, having leaves different in size and/or shape, was quantified in adult Aucuba japonica and simulations were carried out to evaluate the effects of anisophylly on the extent of self-shading at the single-shoot level as well as at the whole-canopy level. Clear anisophylly was observed in the individual after switching from the single-stemmed juvenile stage to the multi-stemmed adult stage. In such plants, leaf area in the canopy abruptly increased. The effective display of adult foliage involved a variety of morphological changes in addition to anisophylly, most prominently reduction in leaf size compared to juveniles. The simulation results indicate that diversity of leaf size and shape is an effective means of minimizing self-shading as well as allowing the efficient exploitation of a larger canopy volume in adult plants. Anisophylly also increased the biomass use efficiency of individual plants at maturity. Taken together, having diverse leaf forms is superior to having a single leaf form for maximizing area acquisition and for efficiently filling the acquired area. We therefore conclude that the anisophylly expressed in A. japonica is adaptive.

Biomass↗

Microenvironmental preferences of oribatid mite species on the floor of a tropical rainforest.

The vertical distribution, microenvironmental preference, canopy cover, as well as monthly fluctuations of recently described species of three genera of macropyline oribatid mites, Mesoplophora, Bicyrthermannia and Nothrus and three genera of brachypyline mites, Scheloribates, Muliercula and Galumnella were investigated over a 9-month period in shaded and unshaded areas of a secondary regrowth forest floor in Ile Ife, Nigeria. Sampling was done monthly from August 2002 to April 2003 covering the two seasons of the annual cycle. There was an overwhelming predominance of mite populations in the 5 cm topsoil compared with the fermentation and litter layers. The preference of M. ifeana and B. nigeriana extracted from the topsoil for the shaded plot was attributed to highly conducive moisture conditions provided by the canopy. It is suggested that M. ifeana, B. nigeriana, N. lasebikani, S. mochlosimilaris, M. inexpectata and G. sonpona, that exhibited a single peak density in the wet season in this study, had similar ecological needs. The populations of M. ifeana and S. mochlosimilaris were relatively stable in the fermentation layer. The study concluded that canopy cover affected the sensitivity of soil dwelling mites on the floor of a secondary regrowth forest with regard to vertical gradients of light, microclimate and foliage quality in a complex way.

Animals↗

Interception and retention of Chernobyl-derived 134Cs, 137Cs and 106Ru in a spruce stand.

The time dependence of the specific activity of Chernobyl-derived 134Cs, 137Cs and 106Ru was determined in vegetation and soil samples from an old spruce stand within a period of 600 days after the beginning of the radioactive fallout. The results show that 70% of the total activity of radiocesium and 60% of radioruthenium deposited in the spruce stand were retained initially in the canopy. They were removed from the needles and twigs as a result of weathering (rain, wind, litter fall) and transferred to the forest floor, but only rather slowly (half-lives in the canopy: radiocesium, 90 days for the period 0-130 days, 230 days for the period 130-600 days; radioruthenium, 95 days for the period 0-200 days). The transfer of radiocesium and ruthenium to the forest floor by litter-fall was small when compared with that of weathering by rain or wind (radiocesium 7%, radioruthenium 8%, with respect to the total activity deposited in the canopy). The total deposition of radiocesium and ruthenium in the spruce stand was higher by 20 and 24%, respectively, than that observed in nearby grassland. The deposition velocity of radiocesium in the spruce stand was estimated at 5.5 mm s-1, higher by a factor of 10 than the figure for grassland. Similar values were found for radioruthenium.

Accidents↗

Flow rates of ions in waters percolating through a model ecosystem with forest trees.

From 1983-88 the long-term effects of low level exposure with O(3), SO(2) and simulated acid rain on mineral cycling in model ecosystems with spruce, fir and beech seedlings were investigated. Systems consisting of open-top chambers built above lysimeters were protected against the intrusion of ambient rain and dust. As part of the investigations on mineral cycling the fluxes of elements with water input and output of the canopy and soil compartments are presented. During the 5 year duration of the experiment, pronounced effects on canopy deposition and cation leaching were observed. Most noticeable were throughfall enrichment with sulfate through dry deposition of SO(2) as influenced by duration of needlewetting and factors promoting SO(2) oxidation. Depending on sulfur deposition, leaching of calcium, magnesium, manganese, zinc and ammonium from canopies was elevated, in total leading to enhanced soil input of acid. After 15 months, the water percolating the soils in the lysimeters of these treatments was acidified, with elevated flowrates of sulfate, manganese, calcium and magnesium. The results on canopy/soil leaching are compared to those from old conifer stands in the field.

Journal Article↗

Chemical climatology of high elevation spruce-fir forests in the southern Appalachian mountains.

The physical and chemical climatology of high elevation (> 1500 m) spruce-fir forests in the southern Appalachian mountains was studied by establishing a weather and atmospheric chemical observatory at Mt Mitchell State Park in North Carolina (35 degrees 44' 05" N, 82 degrees 17' 15"W). Data collected during the summer and autumn (May-October) of 1986, 1987, and 1988 are reported. All measurements were made on or near a 16.5 m walk-up tower extending 10 m above the forest canopy on Mt Gibbes (2006 m msl), which is located approximately 2 km SW of Mt Mitchell. The tower was equipped with standard meteorological instrumentation, a passive cloud water collector, and gas pollutant sensors for O3, SO2, NOx. The tower and nearby forest canopy were immersed in clouds 25 to 40% of the time. Non-precipitating clouds were very acidic (pH 2.5-4.5). Precipitating clouds were less acidic (pH 3.5-5.5). The dominant wind directions were WNW and ESE. Clouds from the most common wind direction (WNW) were more acidic (mean pH 3.5) than those from the next most common wind direction (ESE, mean pH 5.5). Cloud water acidity was related to the concentration of SO4(2-), and NO3- ions. Mean concentration of H+, NH4+, SO4(2-), and NO3- ions in the cloud water varied from 330-340, 150-200, 190-200 and 120-140 micromol litre(-1) respectively. The average and range of O3 were 50 (25-100) ppbv (109) in 1986, 51 (26-102) ppbv in 1987, and 66 (30-140) during the 1988 field seasons, respectively. The daily maximum, 1-h average, and 24-h average concentrations were all greatest during June through mid-August, suggesting a correlation with the seasonal temperature and solar intensity. Throughfall collectors near the tower were used to obtain a useful estimate of deposition to the forest canopy. Between 50-60% of the total deposition of SO4(2-) was due to cloud impact.

Journal Article↗

Micro-spatial variation of soil metal pollution and plant recruitment near a copper smelter in Central Chile.

Soil chemical changes produced by metal smelters have mainly been studied on a large scale. In terms of plant survival, determination of small scale variability may be more important because less toxic microhabitats may represent safe sites for successful recruitment and thus for plant survival. Three dominant microhabitats (open spaces and areas below the canopy of Sphaeralcea obtusiloba and Baccharis linearis shrubs) were defined in a heavily polluted area near a copper smelter and characterised in terms of microclimate, general soil chemistry, total and extractable metal concentrations in the soil profile (A0 horizon, 0-5 and 15-20 cm depth), and seedling densities. Results indicated a strong variability in microclimate and soil chemistry not only in the soil profile but also among microhabitats. Air/soil temperatures, radiation and wind speed were much lower under the canopy of shrubs, particularly during the plant growth season. Soil acidification was detected on top layers (0-5 cm depth) of all microhabitats while higher concentrations of N, Cu and Cd were detected on litter and top soil layers below shrubs when compared to open spaces; however, high organic matter content below shrubs decreased bioavailability of metals. Plant recruitment was concentrated under shrub canopies; this may be explained as a result of the nursery effect exerted by shrubs in terms of providing a more favourable microclimate, along with better soil conditions in terms of macronutrients and metal bioavailability.

Biological Availability↗

Epiphytic lichen growth abnormalities and element concentrations as early indicators of forest decline.

Thallus morphology and element concentrations (S, Al, Fe, Sr, Mn, Ni, Zn, Cu, Pb, As, F, and Cl) were compared in samples of the fruticose lichen genus Usnea at two heights of the Abies sibirica canopy in the East Sayan Mountains (Krasnoyarsk District, Russia) sampled from three stations at 15, 25 and 35 km from Krasnoyark. Usnea species with an abnormal morphology dominated on branches in the upper canopy, 15-22 m above ground level, and normal thalli on lower tree branches, 2-5 m above ground. Abnormal thalli at the tree-top level contained higher Al, Fe, Zn, F, Sr and Pb concentrations compared with normal thalli growing below, confirming a dust impact. No such clear trend was observed between sampling stations. Crown canopy architecture, surface microtopography and the balance between the processes of deposition and the movement and loss of particles play a major role in particle interception and in pollutant delivery to Usnea.

Air Pollutants↗