Do trade-offs govern plant species' responses to different global change treatments?

Plants are subject to trade-offs among growth strategies such that adaptations for optimal growth in one condition can preclude optimal growth in another. Thus, we predicted that a plant species that responds positively to one global change treatment would be less likely than average to respond positively to another treatment, particularly for pairs of treatments that favor distinct traits. We examined plant species' abundances in 39 global change experiments manipulating two or more of the following: CO2 , nitrogen, phosphorus, water, temperature, or disturbance. Overall, the directional response of a species to one treatment was 13% more likely than expected to oppose its response to a another single-factor treatment. This tendency was detectable across the global data set, but held little predictive power for individual treatment combinations or within individual experiments. Although trade-offs in the ability to respond to different global change treatments exert discernible global effects, other forces obscure their influence in local communities.

Quantifying the impact of ozone on crops in Sub-Saharan Africa demonstrates regional and local hotspots of production loss.

Tropospheric ozone can have a detrimental effect on vegetation, including reducing the quantity of crop yield. This study uses modelled ozone flux values (POD3IAM; phytotoxic ozone dose above 3 nmol m-2 s-1, parameterised for integrated assessment modelling) for 2015, together with species-specific flux-effect relationships, spatial data on production and growing season dates to quantify the impact of ozone on the production of common wheat (Triticum aestivum) and common beans (Phaseolus vulgaris) across Sub-Saharan Africa (SSA). A case study for South Africa was also done using detailed data per province. Results suggest that ozone pollution could decrease wheat yield by between 2 and 13%, with a total annual loss of 453,000 t across SSA. The impact on bean production depended on the season; however, estimated yield losses were up to 21% in some areas of SSA, with an annual loss of ~300,000 t for each of the two main growing seasons. Production losses tended to be greater in countries with the highest production, for example, Ethiopia (wheat) and Tanzania (beans). This study provides an indication of the location of areas at high risk of crop losses due to ozone. Results emphasise that efforts to reduce ozone precursors could contribute to reducing the yield gap in SSA. More stringent air pollution abatement policies are required to reduce crop losses to ozone in the future.

Reduced photosynthetic thermal acclimation capacity under elevated ozone in poplar (Populus tremula) saplings.

The sensitivity of photosynthesis to temperature has been identified as a key uncertainty for projecting the magnitude of the terrestrial carbon cycle response to future climate change. Although thermal acclimation of photosynthesis under rising temperature has been reported in many tree species, whether tropospheric ozone (O3 ) affects the acclimation capacity remains unknown. In this study, temperature responses of photosynthesis (light-saturated rate of photosynthesis (Asat ), maximum rates of RuBP carboxylation (Vcmax ), and electron transport (Jmax ) and dark respiration (Rdark ) of Populus tremula exposed to ambient O3 (AO3 , maximum of 30 ppb) or elevated O3 (EO3 , maximum of 110 ppb) and ambient or elevated temperature (ambient +5°C) were investigated in solardomes. We found that the optimum temperature of Asat (ToptA ) significantly increased in response to warming. However, the thermal acclimation capacity was reduced by O3 exposure, as indicated by decreased ToptA , and temperature optima of Vcmax (ToptV ) and Jmax (ToptJ ) under EO3 . Changes in both stomatal conductance (gs ) and photosynthetic capacity (Vcmax and Jmax ) contributed to the shift of ToptA by warming and EO3 . Neither Rdark measured at 25°C ( R dark 25 ) nor the temperature response of Rdark was affected by warming, EO3 , or their combination. The responses of Asat , Vcmax , and Jmax to warming and EO3 were closely correlated with changes in leaf nitrogen (N) content and N use efficiency. Overall, warming stimulated growth (leaf biomass and tree height), whereas EO3 reduced growth (leaf and woody biomass). The findings indicate that thermal acclimation of Asat may be overestimated if the impact of O3 pollution is not taken into account.

Ozone-induced effects on leaves in African crop species.

Tropospheric (ground-level) ozone is a harmful phytotoxic pollutant, and can have a negative impact on crop yield and quality in sensitive species. Ozone can also induce visible symptoms on leaves, appearing as tiny spots (stipples) between the veins on the upper leaf surface. There is little measured data on ozone concentrations in Africa and it can be labour-intensive and expensive to determine the direct impact of ozone on crop yield in the field. The identification of visible ozone symptoms is an easier, low cost method of determining if a crop species is being negatively affected by ozone pollution, potentially resulting in yield loss. In this study, thirteen staple African food crops (including wheat (Triticum aestivum), common bean (Phaseolus vulgaris), sorghum (Sorghum bicolor), pearl millet (Pennisetum glaucum) and finger millet (Eleusine coracana)) were exposed to an episodic ozone regime in a solardome system to monitor visible ozone symptoms. A more detailed examination of the progression of ozone symptoms with time was carried out for cultivars of P. vulgaris and T. aestivum, which showed early leaf loss (P. vulgaris) and an increased rate of senescence (T. aestivum) in response to ozone exposure. All of the crops tested showed visible ozone symptoms on their leaves in at least one cultivar, and ozone sensitivity varied between cultivars of the same crop. A guide to assist with identification of visible ozone symptoms (including photographs and a description of symptoms for each species) is presented.

Ozone critical levels for (semi-)natural vegetation dominated by perennial grassland species.

New critical levels for ozone based on accumulated flux through stomata (phytotoxic ozone dose, POD), for temperate perennial grassland (semi-)natural vegetation, have been agreed for use within the Convention on Long-Range Transboundary Air Pollution. These were based on data from several experiments conducted under naturally fluctuating environmental conditions that were combined and analysed to give linear dose-response relationships. Dose-response functions and flux-based critical levels were derived based on biomass and flower number. These parameters showed a statistically significant decline with increasing accumulated stomatal ozone flux. The functions and critical levels derived are based on sensitive species and can be used for risk assessments of the damaging effect of ozone on temperate vegetation communities dominated by perennial grassland species. The critical level based on flower number was lower than that for biomass, representing the greater sensitivity of flower number to ozone pollution.

Ozone affects plant, insect, and soil microbial communities: A threat to terrestrial ecosystems and biodiversity.

Elevated tropospheric ozone concentrations induce adverse effects in plants. We reviewed how ozone affects (i) the composition and diversity of plant communities by affecting key physiological traits; (ii) foliar chemistry and the emission of volatiles, thereby affecting plant-plant competition, plant-insect interactions, and the composition of insect communities; and (iii) plant-soil-microbe interactions and the composition of soil communities by disrupting plant litterfall and altering root exudation, soil enzymatic activities, decomposition, and nutrient cycling. The community composition of soil microbes is consequently changed, and alpha diversity is often reduced. The effects depend on the environment and vary across space and time. We suggest that Atlantic islands in the Northern Hemisphere, the Mediterranean Basin, equatorial Africa, Ethiopia, the Indian coastline, the Himalayan region, southern Asia, and Japan have high endemic richness at high ozone risk by 2100.

Melatonin enhances drought resistance by regulating leaf stomatal behaviour, root growth and catalase activity in two contrasting rapeseed (Brassica napus L.) genotypes.

Two contrasting rapeseed (Brassica napus L.) genotypes, Qinyou 8 (drought-sensitive) and Q2 (drought-tolerant), were studied under drought stress with or without pretreatment with melatonin to (i) explore whether melatonin enhances drought resistance by regulating root growth and (ii) determine the relationship between the belowground and aboveground responses to melatonin under drought stress. Results show that the light-saturated rate of photosynthesis (Pn), stomatal conductance (gs), water use efficiency (WUE) and chlorophyll content were decreased by drought for Qinyou 8, whereas drought only decreased Pn and chlorophyll content for Q2. Drought decreased actual photochemical efficiency in saturated light (Fv'/Fm'), actual photochemical efficiency (PhiPSⅡ), quenching of photochemical efficiency (qL) and electron transport rate (ETR) in Qinyou 8. However drought only decreased Fv'/Fm' and qL in Q2. Drought increased malondialdehyde (MDA) and hydrogen peroxide (H2O2) contents in the roots of both genotypes. Melatonin had no significant additional effects on root guaiacol peroxidase (POD) and superoxide dismutase (SOD) activities, but enhanced root catalase (CAT) activity of droughted plants further. Melatonin promoted taproot and lateral root growth under drought stress. Melatonin also promoted stomatal opening resulting in enhanced photosynthesis in the two genotypes. The two mechanisms induced by melatonin synergistically enhance drought resistance of rapeseed as indicated by enhanced gas exchange parameters under melatonin pretreatment. The findings provide evidence for a physiological role of melatonin in improving drought resistance, especially in belowground parts.

Global change effects on plant communities are magnified by time and the number of global change factors imposed.

Global change drivers (GCDs) are expected to alter community structure and consequently, the services that ecosystems provide. Yet, few experimental investigations have examined effects of GCDs on plant community structure across multiple ecosystem types, and those that do exist present conflicting patterns. In an unprecedented global synthesis of over 100 experiments that manipulated factors linked to GCDs, we show that herbaceous plant community responses depend on experimental manipulation length and number of factors manipulated. We found that plant communities are fairly resistant to experimentally manipulated GCDs in the short term (<10 y). In contrast, long-term (≥10 y) experiments show increasing community divergence of treatments from control conditions. Surprisingly, these community responses occurred with similar frequency across the GCD types manipulated in our database. However, community responses were more common when 3 or more GCDs were simultaneously manipulated, suggesting the emergence of additive or synergistic effects of multiple drivers, particularly over long time periods. In half of the cases, GCD manipulations caused a difference in community composition without a corresponding species richness difference, indicating that species reordering or replacement is an important mechanism of community responses to GCDs and should be given greater consideration when examining consequences of GCDs for the biodiversity-ecosystem function relationship. Human activities are currently driving unparalleled global changes worldwide. Our analyses provide the most comprehensive evidence to date that these human activities may have widespread impacts on plant community composition globally, which will increase in frequency over time and be greater in areas where communities face multiple GCDs simultaneously.

Can Reduced Irrigation Mitigate Ozone Impacts on an Ozone-Sensitive African Wheat Variety?

Ground-level ozone (O3) pollution is known to adversely affect the production of O3-sensitive crops such as wheat. The magnitude of impact is dependent on the accumulated stomatal flux of O3 into the leaves. In well-irrigated plants, the leaf pores (stomata) tend to be wide open, which stimulates the stomatal flux and therefore the adverse impact of O3 on yield. To test whether reduced irrigation might mitigate O3 impacts on flag leaf photosynthesis and yield parameters, we exposed an O3-sensitive Kenyan wheat variety to peak concentrations of 30 and 80 ppb O3 for four weeks in solardomes and applied three irrigation regimes (well-watered, frequent deficit, and infrequent deficit irrigation) during the flowering and grain filling stage. Reduced irrigation stimulated 1000-grain weight and harvest index by 33% and 13%, respectively (when O3 treatments were pooled), which compensated for the O3-induced reductions observed in well-watered plants. Whilst full irrigation accelerated the O3-induced reduction in photosynthesis by a week, such an effect was not observed for the chlorophyll content index of the flag leaf. Further studies under field conditions are required to test whether reduced irrigation can be applied as a management tool to mitigate adverse impacts of O3 on wheat yield.

New Insights into Leaf Physiological Responses to Ozone for Use in Crop Modelling.

Estimating food production under future air pollution and climate conditions in scenario analysis depends on accurately modelling ozone (O3) effects on yield. This study tests several assumptions that form part of published approaches for modelling O3 effects on photosynthesis and leaf duration against experimental data. In 2015 and 2016, two wheat cultivars were exposed in eight hemispherical glasshouses to O3 ranging from 22 to 57 ppb (24 h mean), with profiles ranging from raised background to high peak treatments. The stomatal O3 flux (Phytotoxic Ozone Dose, POD) to leaves was simulated using a multiplicative stomatal conductance model. Leaf senescence occurred earlier as average POD increased according to a linear relationship, and the two cultivars showed very different senescence responses. Negative effects of O3 on photosynthesis were only observed alongside O3-induced leaf senescence, suggesting that O3 does not impair photosynthesis in un-senesced flag leaves at the realistic O3 concentrations applied here. Accelerated senescence is therefore likely to be the dominant O3 effect influencing yield in most agricultural environments. POD was better than 24 h mean concentration and AOT40 (accumulated O3 exceeding 40 ppb, daylight hours) at predicting physiological response to O3, and flux also accounted for the difference in exposure resulting from peak and high background treatments.

Nitrogen availability does not affect ozone flux-effect relationships for biomass in birch (Betula pendula) saplings.

To investigate whether nitrogen (N) load affects the ozone (O3) stomatal flux-effect relationship for birch biomass, three-year old birch saplings were exposed to seven different O3 profiles (24 h mean of 35-66 ppb) and four different N loads (10, 30, 50 and 70 kg ha-1 yr-1) in precision-controlled hemispherical glasshouses (solardomes) in 2012 and 2013. Stomatal conductance (gs) under optimal growth conditions was stimulated by enhanced N supply but was not significantly affected by enhanced O3 exposure. Birch root, woody (stem + branches) and total biomass (root + woody) were not affected by the Phytotoxic Ozone Dose (POD1SPEC) after two seasons of O3 exposure, and enhanced N supply stimulated biomass production independent of POD1SPEC (i.e. there were no POD1SPEC × N interactions). There was a strong linear relationship between the stem cross-sectional area and tree biomass at the end of the experiment, which was not affected by O3 exposure or N load. Enhanced N supply stimulated the stem cross-sectional area at the end of season 2, but not at the end of season 1, which suggests a time lag before tree biomass responded to enhanced N supply. There was no significant effect of POD1SPEC on stem cross-sectional area after either the first or second growing season of the experiment. Contrasting results reported in the literature on the interactive impacts of O3 and N load on tree physiology and growth are likely due to species-specific responses, different duration of the experiments and/or a limitation of the number of O3 and N levels tested.

Increase of apoplastic ascorbate induced by ozone is insufficient to remove the negative effects in tobacco, soybean and poplar.

Apoplastic ascorbate (ASCapo) is an important contributor to the detoxification of ozone (O3). The objective of the study is to explore whether ASCapo is stimulated by elevated O3 concentrations. The detoxification of O3 by ASCapo was quantified in tobacco (Nicotiana L), soybean (Glycine max (L.) Merr.) and poplar (Populus L), which were exposed to charcoal-filtered air (CF) and elevated O3 treatments (E-O3). ASCapo in the three species were significantly increased by E-O3 compared with the values in the filtered treatment. For all three species, E-O3 significantly increased the malondialdehyde (MDA) content and decreased light-saturated rate of photosynthesis (Asat), suggesting that high O3 has induced injury/damage to plants. E-O3 significantly increased redox state in the apoplast (redox stateapo) for all species, whereas no effect on the apoplastic dehydroascorbate (DHAapo) was observed. In leaf tissues, E-O3 significantly enhanced reduced-ascorbate (ASC) and total ascorbate (ASC+DHA) in soybean and poplar, but significantly reduced these in tobacco, indicating different antioxidative capacity to the high O3 levels among the three species. Total antioxidant capacity in the apoplast (TACapo) was significantly increased by E-O3 in tobacco and poplar, but leaf tissue TAC was significantly enhanced only in tobacco. Leaf tissue superoxide anion (O2•-) in poplar and hydrogen peroxide (H2O2) in tobacco and soybean were significantly increased by E-O3. The diurnal variation of ASCapo, with maximum values occurring in the late morning and lower values experienced in the afternoon, appeared to play an important role in the harmful effects of O3 on tobacco, soybean and poplar.

Ozone pollution will compromise efforts to increase global wheat production.

Introduction of high-performing crop cultivars and crop/soil water management practices that increase the stomatal uptake of carbon dioxide and photosynthesis will be instrumental in realizing the United Nations Sustainable Development Goal (SDG) of achieving food security. To date, however, global assessments of how to increase crop yield have failed to consider the negative effects of tropospheric ozone, a gaseous pollutant that enters the leaf stomatal pores of plants along with carbon dioxide, and is increasing in concentration globally, particularly in rapidly developing countries. Earlier studies have simply estimated that the largest effects are in the areas with the highest ozone concentrations. Using a modelling method that accounts for the effects of soil moisture deficit and meteorological factors on the stomatal uptake of ozone, we show for the first time that ozone impacts on wheat yield are particularly large in humid rain-fed and irrigated areas of major wheat-producing countries (e.g. United States, France, India, China and Russia). Averaged over 2010-2012, we estimate that ozone reduces wheat yields by a mean 9.9% in the northern hemisphere and 6.2% in the southern hemisphere, corresponding to some 85 Tg (million tonnes) of lost grain. Total production losses in developing countries receiving Official Development Assistance are 50% higher than those in developed countries, potentially reducing the possibility of achieving UN SDG2. Crucially, our analysis shows that ozone could reduce the potential yield benefits of increasing irrigation usage in response to climate change because added irrigation increases the uptake and subsequent negative effects of the pollutant. We show that mitigation of air pollution in a changing climate could play a vital role in achieving the above-mentioned UN SDG, while also contributing to other SDGs related to human health and well-being, ecosystems and climate change.

Modelling spatial patterns of correlations between concentrations of heavy metals in mosses and atmospheric deposition in 2010 across Europe.

BACKGROUND: This paper aims to investigate the correlations between the concentrations of nine heavy metals in moss and atmospheric deposition within ecological land classes covering Europe. Additionally, it is examined to what extent the statistical relations are affected by the land use around the moss sampling sites. Based on moss data collected in 2010/2011 throughout Europe and data on total atmospheric deposition modelled by two chemical transport models (EMEP MSC-E, LOTOS-EUROS), correlation coefficients between concentrations of heavy metals in moss and in modelled atmospheric deposition were specified for spatial subsamples defined by ecological land classes of Europe (ELCE) as a spatial reference system. Linear discriminant analysis (LDA) and logistic regression (LR) were then used to separate moss sampling sites regarding their contribution to the strength of correlation considering the areal percentage of urban, agricultural and forestry land use around the sampling location. After verification LDA models by LR, LDA models were used to transform spatial information on the land use to maps of potential correlation levels, applicable for future network planning in the European Moss Survey. RESULTS: Correlations between concentrations of heavy metals in moss and in modelled atmospheric deposition were found to be specific for elements and ELCE units. Land use around the sampling sites mainly influences the correlation level. Small radiuses around the sampling sites examined (5 km) are more relevant for Cd, Cu, Ni, and Zn, while the areal percentage of urban and agricultural land use within large radiuses (75-100 km) is more relevant for As, Cr, Hg, Pb, and V. Most valid LDA models pattern with error rates of < 40% were found for As, Cr, Cu, Hg, Pb, and V. Land use-dependent predictions of spatial patterns split up Europe into investigation areas revealing potentially high (= above-average) or low (= below-average) correlation coefficients. CONCLUSIONS: LDA is an eligible method identifying and ranking boundary conditions of correlations between atmospheric deposition and respective concentrations of heavy metals in moss and related mapping considering the influence of the land use around moss sampling sites.

Origin and spatial distribution of metals in moss samples in Albania: A hotspot of heavy metal contamination in Europe.

This study presents the spatial distribution of 37 elements in 48 moss samples collected over the whole territory of Albania and provides information on sources and factors controlling the concentrations of elements in the moss. High variations of trace metals indicate that the concentrations of elements are affected by different factors. Relations between the elements in moss, geochemical interpretation of the data, and secondary effects such as redox conditions generated from local soil and/or long distance atmospheric transport of the pollutants are discussed. Zr normalized data, and the ratios of different elements are calculated to assess the origin of elements present in the current moss samples with respect to different geogenic and anthropogenic inputs. Factor analysis (FA) is used to identify the most probable sources of the elements. Four dominant factors are identified, i.e. natural contamination; dust emission from local mining operations; atmospheric transport of contaminants from local and long distance sources; and contributions from air borne marine salts. Mineral particle dust from local emission sources is classified as the most important factor affecting the atmospheric deposition of elements accumulated in the current moss samples. The open slag dumps of mining operation in Albania is probably the main factor contributing to high contents of Cr, Ni, Fe, Ti and Al in the moss. Enrichment factors (EF) were calculated to clarify whether the elements in the present moss samples mainly originate from atmospheric deposition and/or local substrate materials.

Leaf traits and photosynthetic responses of Betula pendula saplings to a range of ground-level ozone concentrations at a range of nitrogen loads.

Ground-level ozone (O3) concentrations and atmospheric nitrogen (N) deposition rates have increased strongly since the 1950s. Rising ground-level O3 concentrations and atmospheric N deposition both affect plant physiology and growth, however, impacts have often been studied in isolation rather than in combination. In addition, studies are often limited to a control treatment and one or two elevated levels of ozone and/or nitrogen supply. In the current study, three-year old Betula pendula saplings were exposed to seven different O3 profiles (24h mean O3 concentration of 36-68ppb in 2013, with peaks up to an average of 105ppb) in precision-controlled hemispherical glasshouses (solardomes) and four different N loads (10, 30, 50 or 70kgNha-1y-1) in 2012 and 2013. Here we report on the effects of enhanced O3 concentrations and N load on leaf traits and gas exchange in leaves of varying age and developmental stage in 2013. The response of leaf traits to O3 (but not N) vary with leaf developmental stage. For example, elevated O3 did not affect the chlorophyll content of the youngest fully expanded leaf, but it reduced the chlorophyll content and photosynthetic parameters in aging leaves, relatively more so later than earlier in the growing season. Elevated O3 enhanced the N content of senesced leaves prior to leaf fall, potentially affecting subsequent N cycling in the soil. Enhanced N generally stimulated the chlorophyll content and photosynthetic capacity. Whilst elevated O3 reduced the light-saturated rate of photosynthesis (Asat) in aging leaves, it did not affect stomatal conductance (gs). This suggests that photosynthesis and gs are not closely coupled at elevated O3 under-light saturating conditions. We did not observe any interactions between O3 and N regarding photosynthetic parameters (Vc,max, Jmax, Asat), chlorophyll content, gs, N content in senesced leaves and leaf number. Hence, the sensitivity of these leaf traits to O3 in young silver birch trees is neither reduced nor enhanced by N load.

Current and future ozone risks to global terrestrial biodiversity and ecosystem processes.

Risks associated with exposure of individual plant species to ozone (O3) are well documented, but implications for terrestrial biodiversity and ecosystem processes have received insufficient attention. This is an important gap because feedbacks to the atmosphere may change as future O3 levels increase or decrease, depending on air quality and climate policies. Global simulation of O3 using the Community Earth System Model (CESM) revealed that in 2000, about 40% of the Global 200 terrestrial ecoregions (ER) were exposed to O3 above thresholds for ecological risks, with highest exposures in North America and Southern Europe, where there is field evidence of adverse effects of O3, and in central Asia. Experimental studies show that O3 can adversely affect the growth and flowering of plants and alter species composition and richness, although some communities can be resilient. Additional effects include changes in water flux regulation, pollination efficiency, and plant pathogen development. Recent research is unraveling a range of effects belowground, including changes in soil invertebrates, plant litter quantity and quality, decomposition, and nutrient cycling and carbon pools. Changes are likely slow and may take decades to become detectable. CESM simulations for 2050 show that O3 exposure under emission scenario RCP8.5 increases in all major biomes and that policies represented in scenario RCP4.5 do not lead to a general reduction in O3 risks; rather, 50% of ERs still show an increase in exposure. Although a conceptual model is lacking to extrapolate documented effects to ERs with limited or no local information, and there is uncertainty about interactions with nitrogen input and climate change, the analysis suggests that in many ERs, O3 risks will persist for biodiversity at different trophic levels, and for a range of ecosystem processes and feedbacks, which deserves more attention when assessing ecological implications of future atmospheric pollution and climate change.

Spatially valid data of atmospheric deposition of heavy metals and nitrogen derived by moss surveys for pollution risk assessments of ecosystems.

For analysing element input into ecosystems and associated risks due to atmospheric deposition, element concentrations in moss provide complementary and time-integrated data at high spatial resolution every 5 years since 1990. The paper reviews (1) minimum sample sizes needed for reliable, statistical estimation of mean values at four different spatial scales (European and national level as well as landscape-specific level covering Europe and single countries); (2) trends of heavy metal (HM) and nitrogen (N) concentrations in moss in Europe (1990-2010); (3) correlations between concentrations of HM in moss and soil specimens collected across Norway (1990-2010); and (4) canopy drip-induced site-specific variation of N concentration in moss sampled in seven European countries (1990-2013). While the minimum sample sizes on the European and national level were achieved without exception, for some ecological land classes and elements, the coverage with sampling sites should be improved. The decline in emission and subsequent atmospheric deposition of HM across Europe has resulted in decreasing HM concentrations in moss between 1990 and 2010. In contrast, hardly any changes were observed for N in moss between 2005, when N was included into the survey for the first time, and 2010. In Norway, both, the moss and the soil survey data sets, were correlated, indicating a decrease of HM concentrations in moss and soil. At the site level, the average N deposition inside of forests was almost three times higher than the average N deposition outside of forests.