Land Use Change, Air Pollution and Climate Change—Vegetation Response in Latin America

Chapter 19

Land Use Change, Air Pollution and Climate Change— Vegetation Response in Latin America Alessandra R. Kozovits*,1 and Mercedes M.C. Bustamante{ *

Departamento de Biodiversidade, Evoluc¸a˜o Meio Ambiente, Universidade Federal de Ouro Preto, Campus Morro do Cruzeiro, Ouro Preto, Brazil { Departamento de Ecologia, Instituto de Cieˆncias Biolo´gicas, Campus Universita´rio Darcy Ribeiro, Universidade de Brası´lia, Brası´lia, Brazil 1 Corresponding author:

Chapter Outline 19.1. Introduction 19.2. Latin America and Its Major Biomes 19.3. Land Use Change, Air Pollutant Emission and Regional Climate Change 19.4. Effects of Nitrogen Addition on Natural Savanna and Forest Ecosystems

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19.5. Ozone: A Growing Concern 19.6. Vegetation Responses to Global Change 19.7. Conclusions and Future Directions Acknowledgement References

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19.1 INTRODUCTION During the recent decades, Latin America has been facing a great challenge in combining unprecedented economic growth with environmental protection and sustainable development. Together with Caribbean countries, the region represents only 9% of the earth’s human population; however, it has the highest urbanization rates worldwide. Currently, about 80% of the population lives in urban areas. Besides the socio-environmental problems related to the Developments in Environmental Science, Vol. 13. http://dx.doi.org/10.1016/B978-0-08-098349-3.00019-0 © 2013 Elsevier Ltd. All rights reserved.

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rapidly growing urban and industrial centres, such as the decline in air quality, air pollution and climate change effects on natural and managed ecosystems are also causing concern. Harboring high biological diversity, countries such as Brazil bear a high responsibility not only for conserving native biomes but also for contributing significantly to agricultural productivity worldwide. At the same time, some research initiatives for long-term studies have been started to expand databases on emission sources of air pollutants and to understand the effects of climate change on forest types and their establishment. Substantial advances have become evident in research related to ecosystem responses of forests and crop plantations in the tropics and sub-tropics to climate change, in particular, as mediated through alterations in precipitation and temperature regimes, and as inherently interacting with air pollution. This chapter focuses on the major initiatives in the field in Latin America, with emphasis on Brazil, summarizing the results obtained to date. First, inventories of emitted air pollutants and their annual proportion in regional emission budgets are presented. Contributions by land use changes related to agriculture are highlighted. Extensive land areas are affected annually by the burning of grasslands and forests, deforestation and addition of fertilisers, these being events bearing long-distance and long-term consequences for ecosystem functioning and regional climate. The deposition of N, for example, results in significant changes in species diversity and ecosystem functions, such as nutrient cycling with litter turnover and greenhouse gas flux from soil to atmosphere under increasing N availability. In addition, changes in productivity, woody-plant diversity, C stocks and other parameters of tropical forest functioning under elevated CO2 concentration and altered water relations as, for example, imposed by prolonged drought in the Amazonian rainforest, have also been found to be causing some concern by regional long-term studies. Key findings from such studies are presented, after briefly featuring the Latin American biomes of relevance in this chapter in ecological terms. The chapter concludes by identifying major gaps in our knowledge of Latin America and suggesting potential strategies for strengthening research having socio-economic implications for policy-making and human welfare.

19.2 LATIN AMERICA AND ITS MAJOR BIOMES Approximately one quarter of the world’s forest area is located in Latin America and the Caribbean. In the region, biomes—together with others such as rainforests, savannas, mountainous and cloud forests, rocky outcrops, grasslands, dry forests, salt marshes, mangroves, xerophytic vegetation and wetlands—represent the largest reservoir of global biodiversity (Figure 19.1; Bovarnick et al., 2010; UNEP, 2010), being associated with an appreciable proportion of global aboveground carbon stock (Phillips et al., 2009). Tropical rain forests and savannas are the major native vegetation types within woody plant diversity in terms of coverage of terrestrial surface area.

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N

Legend Tropical rain broadleaf forest Atlantic rain forest Grassland/savanna/shrubland

0

625

1250

2500 km

FIGURE 19.1 Major biomes in Latin America. Modified from IPCC (2001).

Tropical rain forests are found in South and Central America, central Africa and South-East Asia, and are characterised by a high monthly mean temperature (above 18  C) and high precipitation (>100 mm per month). The dry season, if present, is short and with rare exceptions, rainfall always exceeds potential evapotranspiration (Dirzo, 2001). The Amazonian forest is the largest of all tropical rainforests worldwide. The Amazon biome extends across nine South American countries, covering about 30% of the continental area (Figure 19.1). It is distributed from the Andes’ base to the Atlantic coast and thus exhibits great structural heterogeneity because of variations in soil, topography, microclimate, disturbance regimes and proximity to, and influence of, rivers (Wittmann et al., 2006). The lowland forest, for example, grows in alluvium along the major rivers

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with sediment-rich water, whereas the Igapo´ (seasonally flooded forests) occurs along rivers of black and sediment-poor water (the dark colour of rivers results from organic acids released into the water through the decomposition of organic matter and the lack of terrestrial sediments). The ‘terra firme’ forest occurs outside the influence of rivers, and although such forests may vary in structure, they are commonly termed ‘rainforests’ (Lewis et al., 2009; Malhi et al., 2008). The plant communities of the Amazon basin are the most diverse on Earth, with more than 80,000 taxa of vascular plants described to date. In some areas such as in the upper Amazonia, it is possible to find more than 300 tree species (with individuals  10 cm in diameter at breast height) per hectare (Gentry, 1988), reaching heights of 45–60 m. Besides the diversity in soil and topography of the region across the Amazon basin, the gradient of precipitation varies from the extremely wet, continuously rainy northwest, to the wet-dry transition climate and to the prolonged dry-season climate of the south-eastern regions (Davidson et al., 2012). This climatic gradient is crucial in shaping forest structure and functioning, affecting resistance and resilience to natural and anthropogenic disturbances. The gradient is largely prone to land use change as well, as conversion to agricultural land is more dominant in the drier eastern and southern regions. Another type of rainforest, the Atlantic Forest, stretches along almost the entire coast of Brazil (Figure 19.1), extending inland in the south and into eastern Paraguay and northern Argentina. Besides its wide latitudinal distribution, this forest also reaches from sea level up to 1800 m high, generating evident altitudinal gradients of habitats and biodiversity. The Atlantic Forest domain includes different forest ecosystems such as the Atlantic ombrophilous dense forests, mixed ombrophilous forests, open ombrophilous forests, semi-deciduous stationary forests and deciduous stationary forests, besides the mangroves, restingas (a mosaic from herbaceous to woody vegetation that occurs on marine sand plains in the littoral), altitudinal grasslands, the countryside swamps, and the northeastern forest enclaves (Colombo and Joly, 2010; Marques et al., 2011). The original coverage area of the Atlantic Forest biome is home to 70% of the Brazilian population, and the use of this area accounts for 80% of Brazil’s gross domestic product. Despite the enormous population pressure, the Atlantic Forest has been identified as one of the 25 global biodiversity hotspots, containing between 1% and 8% of the world’s total species, a high number of them being endemic to the region (Myers et al., 2000). A recent assessment revealed appreciable numbers of endemic species in several groups, including 8000 species of trees (40% of the total number of Atlantic Forest trees), 200 species of birds (16%), 71 species of mammals (27%), 94 species of reptiles (31%), and 286 species of amphibians (60%), to mention only the best-known taxonomic groups (Myers et al., 2000). The IUCN Red List contains more than 110 Atlantic Forest species, 29 of which are critically endangered. The most recent estimates suggest that only 11.4–16% of the original 1.4 million

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km2 of the Atlantic Forest has remained, with 32–40% of such remnants consisting of secondary forest and small forest fragments (<100 ha) (Ribeiro et al., 2009). The savannas, the second most extensive biome of tropical South America, are also dominant worldwide, being responsible for about 30% of the primary production of all terrestrial vegetation (Grace et al., 2006). In South America, the savannas cover an area of more than 2.821 million km2, mostly distributed across Brazil, Colombia, Venezuela and Bolivia (Gottsberger and SilberbauerGottsberger, 2009; Figure 19.1). The Brazilian savanna, locally known as Cerrado, covers about 22% of the country’s surface area, extending from the southern boundaries of the Amazonian region to the southern states of Sa˜o Paulo and Parana´ (Oliveira-Filho and Ratter, 2002). Cerrado is a also a biodiversity hotspot (Myers et al., 2000), comprising about 12,000 plant species, ca. 800 of them being large trees and shrubs, with a high proportion of endemism and rapid loss of habitats. The Cerrado, like other savanna-like vegetations, is characterised by a mosaic ranging from grassland to forest formations. Vegetation structure and diversity seem to be related to variation in soil physical and chemical parameters (fertility, pH, Al concentration, drainage), precipitation seasonality, and fire regime (Oliveira-Filho and Ratter, 2002). Across the region, the mean annual temperature varies between 18 and 28  C, and annual precipitation between 800 and 2000 mm, with a continuous 5–6-month dry season in the southern hemisphere winter. The typical Cerrado physiognomy arises on well-drained, deep, dystrophic and acidic soils, with low cation exchange capacity and high levels of Al saturation (Gottsberger and SilberbauerGottsberger, 2009). Besides being adapted to seasonal water limitation, Cerrado plants are also considered fire-tolerant and fire-dependent (Hoffmann and Moreira, 2002; Miranda et al., 2002).

19.3 LAND USE CHANGE, AIR POLLUTANT EMISSION AND REGIONAL CLIMATE CHANGE Although the agricultural population has declined over recent decades, now representing only 16% of labour in Latin America, agricultural activities still account for significant portions of gross national products in this region. The increasing global population and demand for resource consumption are directly related to the expansion of agricultural areas for both food and biofuel production (FAO, 2010). Related to the growth of the agricultural sector, especially because of expansion of pasture areas, forest cover in Latin America has been reduced by 8% since the 1990s. Aide et al. (2012) determined the extent and the spatial distribution of deforestation (and reforestation) between 2001 and 2010 within the 45 countries in Latin America and the Caribbean, estimating a net loss of 179,405 km2 of woody vegetation in the region. Most of the tropical rainforest deforestation was concentrated in the Brazilian

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Amazon, but Nicaragua and Guatemala also presented a negative balance between reforestation and deforestation. The dry forest and savanna/shrub biomes has also experienced extensive deforestation, so that about 200,000 km2 of woody vegetation has been lost altogether. Most losses occurred in northern Argentina, southeastern Bolivia, and western Paraguay, but Central Brazil exhibits a large loss of native areas as well. The conversion of rainforest and savanna habitats into agricultural and pasture lands, along with the abandonment of degraded areas, represent a substantial loss in aboveground biomass. Moreover, the fragmentation of forests and their conversion into pastures and croplands have been associated with changes in local and regional climate (Davidson et al., 2012). Depending on the local extent of deforestation and the new land use, annual rainfall may become reduced over the pasture area, while cloudiness along with instantaneous rainfall and thunderstorm intensity may increase (Avissar and Schmidt, 1998). At scales of hundreds to thousands of km2, however, rainfall over core clearings may be suppressed (Butt et al., 2011). Deforestation in another105 km2 is expected to significantly decrease the whole-basin precipitation (Coe et al., 2009). The conversion of native forests into agricultural areas in Latin America, especially into pastures, is closely associated with biomass burning and the emission of air pollutants. Vegetation fires are considered a major anthropogenic source of greenhouse gases (Cieslik et al., 2013; Sofiev, 2013), aerosols and pollutants to the atmosphere during the southern hemisphere dry season (July to October) over South America. Besides changing the microphysical processes within clouds, making droplets too small to precipitate (i.e. reducing local rainfall; Andreae et al., 2004), fires can cause additional plant stress by initiating ozone pollution (Grulke, 2009; Longo et al., 1999). Apart from high levels of O3 and its precursors occurring in the urban areas, deforestation and seasonal biomass burning are the most relevant pollution sources in Brazil, presumably enhancing O3 regimes also at rural landscape scales (Longo et al., 1999). Actually, 75% of the total amounts of CO2, CO and NOx released in Brazil are thought to originate from the conversion of native into pasture and cropland (MCT, 2009). Data from the Brazilian Inventory of Anthropogenic Greenhouse Gases Emissions and Removals demonstrate the contribution of changing land use to the emission balance in the country (MCT, 2009). The inventory takes into account five categories of economical sectors as considered by the IPCC: energy, industrial processes, agriculture, land use change and forestry, along with waste management. If only CO2, CH4 and N2O are considered as major greenhouse gases (Figure 19.2), unlike in many countries in other regions of the world, industrial activities are not their most relevant sources. About 76% of CO2 emission is derived from land use changes, compared with 1.6% and 22% from industrial and energy sectors, respectively. It is important to note that emission estimates related to land use change and forestry are based on all kinds of conversion, including that of native

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N2 O 2.7%

3.1%

Energy

11.3%

Industrial processes Agriculture/Husbandry

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Change in land use and forests Waste treatment

FIGURE 19.2 Contribution of the main sectors (Energy, Industrial processes, Agriculture/Husbandry, Change in land use and forests and Waste treatment) to the Brazilian annual emissions of CO2, CH4 and N2O. Data extracted from MCT (2009).

vegetation to agricultural and pasture land, besides expansion of urban areas, construction of water reservoirs and succession towards secondary forests or reforestation. Thus, considering the expansion of pasture areas during recent years, for example, to the detriment of forest and savanna ecosystems, one can attribute a large portion of CO2 emissions indirectly to the agricultural sector. Focusing on the Amazon and Cerrado biomes of Brazil, Bustamante et al. (2012b) estimated the proportion of greenhouse gas emissions that can be attributed to land cover conversion, in particular, to cattle ranching. The authors considered the deforestation and subsequent biomass burning for the establishment of new pasture areas, the burning related to pasture maintenance, and the enteric fermentation and wastes from cattle. The full set of emissions derived from cattle farming was responsible for approximately half of all Brazilian emissions, estimated to be approximately 1055 Mt CO2eq in 2005, for example. Agriculture is also directly responsible for 71% of the methane and 91% of nitrous oxide emission budgets (Figure 19.2). Similar data can be also be observed in India (Pandey et al., 2013).These amounts should be added to the indirect contribution upon land use change, accounting for 15% and 3% of the total Brazilian emissions of CH4 and N2O, respectively. Savanna and rainforests, as pointed out earlier, appear to be adapted to the natural variation in climate and soil conditions, as well as to natural cycles of climatic variation and extreme events, showing, for example, considerable resilience to moderate annual drought (Davidson et al., 2012 for a revision). However, what kind of responses would savanna and rainforest vegetation show to the effects of increasing land use change, accompanied by a distinct natural or anthropogenic enhancement of greenhouse-gas emissions, nitrogen deposition and variation in frequency and intensity of drought?

19.4 EFFECTS OF NITROGEN ADDITION ON NATURAL SAVANNA AND FOREST ECOSYSTEMS Common practices related to agricultural activities in Latin America such as biomass burning, deforestation, fertiliser additions and the spreading cultivation of N-fixing crops, together with the burning of fossil fuels and

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urbanization, can also act indirectly on biodiversity and ecosystem functioning. As described in the previous section, many of these activities transfer large amounts of nitrogen species (NOx, NH3, NO3) from the land surface into the atmosphere. Besides the effects on the cycling and formation of other gaseous pollutants such as O3 that act as greenhouse gases, part of the released N returns to the terrestrial ecosystems through dry and wet deposition, considerably affecting the N availability to plants and, consequently, the terrestrial and aquatic ecosystems’ productivity. The N deposition over some biodiversity hotspots in Latin America may achieve levels higher than 10 kg ha 1 year 1 (Phoenix et al., 2006), especially in regions of intensive agricultural activity (Dentener et al., 2006). The Brazilian Atlantic Forest is presumed to suffer the highest N deposition of about 22 kg ha 1 year 1, followed by the Madrean and Magdalena regions in Colombia and the tropical Andes, which receive between 10 and 15 kg ha 1 year 1. In addition, to enhance productivity, huge amounts of phosphorus and other elements are being applied to and lost from croplands in Latin America. Falkowski et al. (2000) estimated that anthropogenic introduction of nutrients has amplified global N and P cycles by 100% and 400%, respectively, since the Industrial Revolution. The effects of the increasing availability of limiting elements such as N and P on plant diversity and functioning, as already widely observed in several ecosystems of the Northern hemisphere (Bobbink et al., 2010, see Eugster and Ha¨ni, 2013 and Tuovinen et al., 2013), have also been evaluated in some tropical biomes. A 10-year-long N and P fertilising experiment has been conducted in the Cerrado, the Brazilian kind of savanna (Bustamante et al., 2012a). Developing under conditions of seasonal rainfall distribution and highly weathered soils with low nutrient availability, plant species have been evolutionarily selected for mechanisms of high nutrient use, efficiency and retention. In response to long-term N and P additions, Cerrado plots exhibited visible changes in plant density as well as species dominance, richness and diversity, both within the herbaceous and the tree-shrub layers. Considering the woody species, plots fertilised with N alone or N plus P showed significantly lower levels of diversity and evenness indices compared to unfertilised plots (Jacobson et al., 2011). Patterns of species density and dominance in both the N and N plus P treatment deviated from those in unfertilised plots. In general, species abundant in control plots displayed a lowered density in N and N plus P plots. N and P concentrations in leaf litter were also modified by N and P additions, changing the rate of litter decomposition. Enhanced N and P levels in leaf litter were found, especially upon the combined addition of these nutrients, indicating a reduced proficiency in element re-translocation during senescence, and accelerating litter mass losses and nutrient release in comparison to the control plots (Kozovits et al., 2007). Changes in species diversity, vegetation cover and biomass production were more evident in the herbaceous than the woody layer (Bustamante

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et al., 2012a). The highest and lowest species richness occurred in control and N plus P plots, respectively. The addition of P alone or in combination with N induced invasion by Melinis minutiflora (exotic, African C4 grass), resulting in negative impacts on native grass species. Besides changes in aboveground biomass, addition of N and P also led, although to a lesser extent, to changes in the root morphology and biomass, which was reduced under simultaneous N and P additions. The effects of a long-term N and P addition experiment on vegetation structure and tree species biomass and diversity have also been measured in the Amazon rainforest areas undergoing secondary succession (Siddique et al., 2010). In response to 6 years of repeated N and P additions, tree assemblage evenness was reduced and tree species biomass accumulation over time was delayed. While N stimulated the growth of two pioneer and one early secondary tree species, P stimulated the growth of only the dominant pioneer tree, Rollinia exsucca. Absolute tree growth rates were enhanced within 2 years of nutrient addition. Conversely, nutrient-induced shifts in relative tree species growth and reduced assemblage evenness were observed even for more than 3 years after nutrient addition, favouring the nutrient-responsive species.

19.5 OZONE: A GROWING CONCERN Although the focus of this chapter is on available long-term studies that have assessed the effects of climate change and nitrogen deposition on the vegetation in Latin America, one plant stressor intrinsically associated with and intensified through such changes is tropospheric O3 as occurring at enhanced ground-level concentrations. There is no doubt today that ground-level O3 regimes enhanced above pre-industrial levels must be regarded as an inherent component of climate and global change (Matyssek et al., 2013a,b). However, in comparison to other agents of such scenarios, long-term studies on O3 action are lacking in the southern hemisphere. Surface O3 levels in urban areas of Brazil and other South-American countries range above 40 nl l 1 (Muramoto et al., 2003; Olcese and Toselli, 1998; Sa´nchez-Ccoyllo et al., 2006). Maximum levels typically occur during spring (August to November) with means between 40 and 60 nl l 1 (even up to more than 90 nl l 1), mainly because of intense biomass burning. A similar situation may also occur in certain regions in Africa (Laakso et al., 2013). During seasonal vegetation fires in South America, several hundreds of fire spots can occur every day (Langmann et al., 2009), and the smoke-generated aerosols and ozone, transported by the atmospheric motions, produce a huge plume of continental scale covering areas of around 4–5  106 km2 (Freitas et al., 2005). Long-range O3 transport has been measured in Brazil throughout latitudes of 35 to 5 S (Boian and Kirchhoff, 2005). In Parana´ State of Southern Brazil, high O3 events with maximum

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levels of 93–173 nl l 1 (average around 89 nl l 1) were observed during the dry season. Such episodes were a consequence of transport from regions with intense biomass burning, as is the case in Para (Amazonian Rain Forest region), Mato Grosso, Mato Grosso do Sul and Tocantins (savanna-rain forest transition regions). Thus, the air quality of areas neighbouring fire zones is often reduced, and in some cases, may turn even worse than that measured in some South American megacities, such as Sa˜o Paulo (Brazil Health, 2006). Although ozone should obviously be conceived as a factor that can cause serious impacts on the vegetation of tropical and subtropical ecosystems, there are few studies, especially on a long-term scale, which are devoted to measuring O3 effects on plants in Latin America. In Brazil, responses of tree species of the Atlantic Rain Forest to elevated O3 levels have been based on macroscopic foliar injury and physiological parameters (e.g. Bulbovas et al., 2010; Furlan et al., 2007, 2008; Moraes et al., 2006; Pina and Moraes, 2010; Rezende and Furlan, 2009). In such surveys, native tree species have proven to be sensitive to current and predicted O3 levels in SE Brazil. Recent studies have also concentrated efforts on finding suitable species for acting as bio-indicators of O3 impact in the tropics (Dafre´-Martinelli et al., 2011; Ferreira et al., 2012).

19.6 VEGETATION RESPONSES TO GLOBAL CHANGE Given the currently predicted scenarios of global change in South America, relevant modifications in tropical and subtropical forest dynamics are expected to occur (Marengo et al., 2009). Scenarios such as SRES A2 and B2 (IPCC, 2007) presume a reduction in rainfall in Amazonia and in tropical South America east of the Andes, warm nights across the entirety of South America and an increase in rainfall extremes in the SE of South America from 2071 through 2100. Variation in rainfall amount and distribution and increase in atmospheric CO2 are considered to be major drivers of vegetation changes (Bytnerowicz et al., 2013; Lewis et al., 2009; Malhi et al., 2008). Increases in temperature and humidity evidentially generate changes in the energy balance at the earth’s surface, enhancing the seasonality in tropical regions, with shortened but intense rainy seasons and accentuated, prolonged dry seasons (Cox et al., 2008; Malhi et al., 2008; Phillips et al., 2009). The dry season that occurred in the western Amazon in 2005 has been the most severe over the past 100 years (Fisher et al., 2007), being considered similar to what is expected to occur with increasing frequency during the twenty-first century. Several changes resulted in the Amazon forest. A 100 mm deficit on average in rainfall caused losses of 5.3 t in above-ground biomass production per hectare (1.2–1.6  109 t throughout the Amazon rainforest). The group of softwood trees, including pioneer and some successionally intermediate trees species, proved to be sensitive to drought, exhibiting enhanced mortality rates. In the long term, such selective death might alter

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forest composition, biodiversity and functioning. Also in 2005, the third highest atmospheric CO2 concentration since the beginning of recording was registered. Part of this increase was suggested to relate to the drought-induced reduction in carbon stocks of neotropical rain forests, caused by tree death and reduction in net primary productivity (Phillips et al., 2009). For clarification of the effects of prolonged drought in the Amazon Forest, a throughfall exclusion experiment is being performed in Tapajo´s, eastern Brazil, since 2000. The understory of one hectare of evergreen tropical seasonal forest remained covered with plastic panels (installed at 1–2 m height) throughout the rainy seasons (Nepstad et al., 2002). About 40% of the annual rainfall has experimentally been intercepted, simulating rainfall regimes similar to those of the savanna climate. Over the last decade, the decline in net primary productivity and a 38% increase in mortality were the main effects of the water deficit in the Amazon forest, reducing the total biomass by about 46.5 t ha 1 and thus, the stock of forest carbon (Brando et al., 2008; Fisher et al., 2008; Nepstad et al., 2007). Besides the effects of drought, the increasing atmospheric CO2 concentration is considered to be another strong factor that drives the dynamics and changes in tropical forests (Lloyd and Farquhar, 2008). The Amazon Forest Inventory Network, known as ‘RAINFOR’ (Malhi et al., 2002), has measured significant increases of about 0.62  0.23 t C ha 1 year 1 in above-ground biomass during the 1980s and 1990s (Baker et al., 2004). Such an increase sufficiently balanced the carbon lost through deforestation in the region. Rates of mortality, growth and recruitment were accelerated, as well as forest productivity, as a response to the largest supply of limiting resources for growth, especially CO2 (Phillips et al., 1998). Studies in tropical forests have demonstrated that under elevated CO2 concentrations, seedlings and understory trees lowered light compensation points, exhibiting a growth increase of up to 50% with less than 0.5% of the daylight (Wu¨rth et al., 1998). RAINFOR data also suggest an overall imbalance amongst forest functional groups, with a substantial increase in density and relative dominance of lianas (see above, Phillips et al., 2002). Ko¨rner (2004) found that seedling growth of the tropical liana Gonolobus viridiflorus was stimulated by 130% under CO2 concentrations near the current (420 ml l 1) as compared to the pre-industrial level (280 ml l 1). Lianas are structural parasites competing with trees for light, exerting a major ecological impact on forest functioning. If highly abundant, they intensify shading, suppress tree growth and increase tree mortality. Such effects can cause forest biomass loss as well as changes in forest species composition and structure.

19.7 CONCLUSIONS AND FUTURE DIRECTIONS Examples of long-term studies presented here indicate that the large biomes in Latin America, that is, both the rainforest and savanna ecosystems, are highly

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sensitive to the anthropogenic increase in the availabilities of N, P and CO2, which typically limit plant productivity, and potentially in O3 levels, which exacerbate the limitation. Notwithstanding the limiting O3 effects, productivity may increase, although shifts may occur between plant functional groups. Alien species may become profiteers, if they gain in competitiveness under a less limiting environment. Conversely, limitation in rainfall or by enhanced O3 levels can result in productivity loss and selective death of functional groups. How may plant functional groups respond to the combined effects of increased N and CO2 availabilities and shortage of water supply during the dry season? Can increased CO2 concentrations counteract drought or O3 effects? In the long term, changes in plant species composition and vegetation structure, reductions in biodiversity, and changes in C stocks and fluxes, including that of water and nutrients, are to be expected (see also Cudlı´n et al., 2013). Additional factors must be considered when predicting vegetation responses and ecosystem functioning under future climate and air pollution scenarios characterised by enhanced heat and drought. Such factors are, for example, fire incidences (Malhi et al., 2008) and herbivory rates (Bale et al., 2002), which are expected to increase. The quantification of such changes and their effects on ecosystems are current challenges for ecological research in Latin America. Further studies should seek to analyse the joint effects of all such factorial drivers that are crucial modulators of forest dynamics. Moreover, with an economy in transition and ascending, Latin America should also further modify the land use in response to regional and global demands for food, biofuels and forest resources. Thus, there is urgent need for an improved understanding of the trade-offs between land cover, air pollution, carbon stocks, climate change, habitat conservation and human health under future scenarios of economic development (Davidson et al., 2012). Continued augmentation of scientific knowledge and technological capacity, as well as improvements in promoting the training of new scientists and technologists will be required in the Latin American region to provide adequate support to policy-making.

ACKNOWLEDGEMENT The authors thank Dr. Mariangela G.P. Leite for designing the figures.

REFERENCES Aide, T.M., Clark, M.L., Grau, H.R., Lopez-Carr, D., Levy, M.A., Redo, D., Bonilla-Moheno, M., Riner, G., Andrade-Nunez, M.J., Muniz, M., 2012. Deforestation and reforestation of Latin America and the Caribbean (2001–2010). Biotropica 45, 1–10. Andreae, M.O., Rosenfeld, D., Artaxo, P., Costa, A.A., Frank, G.P., Longo, K.M., Silva-Dias, M. A.F., 2004. Smoking rain clouds over the Amazon. Science 303, 1337–1342.

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