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1.
Tropical forests vary substantially in the densities of trees of different sizes and thus in above-ground biomass and carbon stores. However, these tree size distributions show fundamental similarities suggestive of underlying general principles. The theory of metabolic ecology predicts that tree abundances will scale as the −2 power of diameter. Demographic equilibrium theory explains tree abundances in terms of the scaling of growth and mortality. We use demographic equilibrium theory to derive analytic predictions for tree size distributions corresponding to different growth and mortality functions. We test both sets of predictions using data from 14 large-scale tropical forest plots encompassing censuses of 473 ha and > 2 million trees. The data are uniformly inconsistent with the predictions of metabolic ecology. In most forests, size distributions are much closer to the predictions of demographic equilibrium, and thus, intersite variation in size distributions is explained partly by intersite variation in growth and mortality.  相似文献   

2.
Tree size distributions in an old-growth temperate forest   总被引:1,自引:0,他引:1  
Despite the wide variation in the structural characteristics in natural forests, tree size distribution show fundamental similarities that suggest general underlying principles. The metabolic ecology theory predicts the number of individual scales as the −2 power of tree diameter. The demographic equilibrium theory predicts tree size distribution starting from the relationship of size distributions with growth and mortality at demographic equilibrium. Several analytic predictions for tree size distributions are derived from the demographic equilibrium theory, based on different growth and mortality functions. In addition, some purely phenomenological functions, such as polynomial function, have been used to describe the tree size distributions. In this paper, we use the metabolic ecology theory, the demographic equilibrium theory and the polynomial function to predict the tree size distribution for both the whole community and each species in an old-growth temperate forest in northeastern China. The results show that metabolic ecology theory predictions for the scaling of tree abundance with diameter were unequivocally rejected in the studied forest. Although these predictions of demographic theory are the best models for most of the species in the temperate forest, the best models for some species ( Tilia amurensis , Quercus mongolica and Fraxinus mandshurica ) are compound curves (i.e. rotated sigmoid curves), best predicted by the polynomial function. Hence, the size distributions of natural forests were unlikely to be invariant and the predictive ability of general models was limited. As a result, developing a more sophisticated theory to predict tree size distributions remains a complex, yet tantalizing, challenge.  相似文献   

3.
Closed‐canopy forests are being rapidly fragmented across much of the tropical world. Determining the impacts of fragmentation on ecological processes enables better forest management and improves species‐conservation outcomes. Lianas are an integral part of tropical forests but can have detrimental and potentially complex interactions with their host trees. These effects can include reduced tree growth and fecundity, elevated tree mortality, alterations in tree‐species composition, degradation of forest succession, and a substantial decline in forest carbon storage. We examined the individual impacts of fragmentation and edge effects (0–100‐m transect from edge to forest interior) on the liana community and liana–host tree interactions in rainforests of the Atherton Tableland in north Queensland, Australia. We compared the liana and tree community, the traits of liana‐infested trees, and determinants of the rates of tree infestation within five forest fragments (23–58 ha in area) and five nearby intact‐forest sites. Fragmented forests experienced considerable disturbance‐induced degradation at their edges, resulting in a significant increase in liana abundance. This effect penetrated to significantly greater depths in forest fragments than in intact forests. The composition of the liana community in terms of climbing guilds was significantly different between fragmented and intact forests, likely because forest edges had more small‐sized trees favoring particular liana guilds which preferentially use these for climbing trellises. Sites that had higher liana abundances also exhibited higher infestation rates of trees, as did sites with the largest lianas. However, large lianas were associated with low‐disturbance forest sites. Our study shows that edge disturbance of forest fragments significantly altered the abundance and community composition of lianas and their ecological relationships with trees, with liana impacts on trees being elevated in fragments relative to intact forests. Consequently, effective control of lianas in forest fragments requires management practices which directly focus on minimizing forest edge disturbance.  相似文献   

4.
Forest biophysical structure – the arrangement and frequency of leaves and stems – emerges from growth, mortality and space filling dynamics, and may also influence those dynamics by structuring light environments. To investigate this interaction, we developed models that could use LiDAR remote sensing to link leaf area profiles with tree size distributions, comparing models which did not (metabolic scaling theory) and did allow light to influence this link. We found that a light environment‐to‐structure link was necessary to accurately simulate tree size distributions and canopy structure in two contrasting Amazon forests. Partitioning leaf area profiles into size‐class components, we found that demographic rates were related to variation in light absorption, with mortality increasing relative to growth in higher light, consistent with a light environment feedback to size distributions. Combining LiDAR with models linking forest structure and demography offers a high‐throughput approach to advance theory and investigate climate‐relevant tropical forest change.  相似文献   

5.
Over the last decade the field of tropical dendroecology has developed rapidly and major achievements have been made. We reviewed the advances in three main themes within the field. First, long chronologies for tropical tree species were constructed which allowed climate reconstructions, revealed sources of climatic variation and clarified climate–growth relations. Other studies combined tree-ring data and stable isotope (13C and 18O) measurements to evaluate the response of tropical trees to climatic variation and changes. A second set of studies assessed long-term growth patterns of individual trees throughout their life. These studies enhanced the understanding of growth trajectories to the canopy, quantified autocorrelated tree growth and yielded new estimates of tree ages. Such studies were also used to reconstruct the disturbance history of tropical forests. The last set of studies applied tree-ring data to growth models. Tree-ring data can replace diameter measurements from research plots, provide additional information to construct population models, improve timber yield models and validate model output. Based on our review, we propose two main directions for future research. (1) An evaluation of the causes and consequences of growth variation within and among trees and their relation to environmental variation. Studies evaluating this directly contribute to improved understanding of tropical tree ecology. (2) The simultaneous measurement of widths and stable isotope fractions in tree rings offers the potential to study responses of trees to climatic change. Given the major role of tropical forests in the global carbon cycle, knowing these responses is of high priority.  相似文献   

6.
Understory fires represent an accelerating threat to Amazonian tropical forests and can, during drought, affect larger areas than deforestation itself. These fires kill trees at rates varying from < 10 to c. 90% depending on fire intensity, forest disturbance history and tree functional traits. Here, we examine variation in bark thickness across the Amazon. Bark can protect trees from fires, but it is often assumed to be consistently thin across tropical forests. Here, we show that investment in bark varies, with thicker bark in dry forests and thinner in wetter forests. We also show that thinner bark translated into higher fire‐driven tree mortality in wetter forests, with between 0.67 and 5.86 gigatonnes CO2 lost in Amazon understory fires between 2001 and 2010. Trait‐enabled global vegetation models that explicitly include variation in bark thickness are likely to improve the predictions of fire effects on carbon cycling in tropical forests.  相似文献   

7.
The important role of tropical forests in the global carbon cycle makes it imperative to assess changes in their carbon dynamics for accurate projections of future climate–vegetation feedbacks. Forest monitoring studies conducted over the past decades have found evidence for both increasing and decreasing growth rates of tropical forest trees. The limited duration of these studies restrained analyses to decadal scales, and it is still unclear whether growth changes occurred over longer time scales, as would be expected if CO2‐fertilization stimulated tree growth. Furthermore, studies have so far dealt with changes in biomass gain at forest‐stand level, but insights into species‐specific growth changes – that ultimately determine community‐level responses – are lacking. Here, we analyse species‐specific growth changes on a centennial scale, using growth data from tree‐ring analysis for 13 tree species (~1300 trees), from three sites distributed across the tropics. We used an established (regional curve standardization) and a new (size‐class isolation) growth‐trend detection method and explicitly assessed the influence of biases on the trend detection. In addition, we assessed whether aggregated trends were present within and across study sites. We found evidence for decreasing growth rates over time for 8–10 species, whereas increases were noted for two species and one showed no trend. Additionally, we found evidence for weak aggregated growth decreases at the site in Thailand and when analysing all sites simultaneously. The observed growth reductions suggest deteriorating growth conditions, perhaps due to warming. However, other causes cannot be excluded, such as recovery from large‐scale disturbances or changing forest dynamics. Our findings contrast growth patterns that would be expected if elevated CO2 would stimulate tree growth. These results suggest that commonly assumed growth increases of tropical forests may not occur, which could lead to erroneous predictions of carbon dynamics of tropical forest under climate change.  相似文献   

8.
Forest degradation accounts for ~70% of total carbon losses from tropical forests. Substantial emissions are from selective logging, a land‐use activity that decreases forest carbon density. To maintain carbon values in selectively logged forests, climate change mitigation policies and government agencies promote the adoption of reduced‐impact logging (RIL) practices. However, whether RIL will maintain both carbon and timber values in managed tropical forests over time remains uncertain. In this study, we quantify the recovery of timber stocks and aboveground carbon at an experimental site where forests were subjected to different intensities of RIL (4, 8, and 16 trees/ha). Our census data span 20 years postlogging and 17 years after the liberation of future crop trees from competition in a tropical forest on the Guiana Shield, a globally important forest carbon reservoir. We model recovery of timber and carbon with a breakpoint regression that allowed us to capture elevated tree mortality immediately after logging. Recovery rates of timber and carbon were governed by the presence of residual trees (i.e., trees that persisted through the first harvest). The liberation treatment stimulated faster recovery of timber albeit at a carbon cost. Model results suggest a threshold logging intensity beyond which forests managed for timber and carbon derive few benefits from RIL, with recruitment and residual growth not sufficient to offset losses. Inclusion of the breakpoint at which carbon and timber gains outpaced postlogging mortality led to high predictive accuracy, including out‐of‐sample R2 values >90%, and enabled inference on demographic changes postlogging. Our modeling framework is broadly applicable to studies that aim to quantify impacts of logging on forest recovery. Overall, we demonstrate that initial mortality drives variation in recovery rates, that the second harvest depends on old growth wood, and that timber intensification lowers carbon stocks.  相似文献   

9.
《新西兰生态学杂志》2011,33(2):208-215
Large trees are a significant structural component of old-growth forests and are important as habitat for epiphytic biodiversity; as substantial stores of biomass, carbon and nutrient; as seed trees; and as engineers of large gap sites for regeneration. Their low density across the landscape is an impediment to accurately measuring growth and mortality, especially as infrequent tree deaths are rarely captured without long periods of monitoring. Here we present large-tree (≥ 30 cm in diameter at breast height) growth and mortality rates for six common New Zealand tree species over a 42-year period from 28 large permanent plots (0.4?0.8?ha) in the central North Island. Our goal was to examine how rates of growth and mortality varied with tree size and species. In total we sampled 1933 large trees across 11.6 ha, corresponding to a large-tree density of 167 trees?ha?1, of which we used 1542 as our six study species. Mean annual mortality rates varied more than 10-fold among species being least in Dacrydium cupressinum (0.16%) and greatest in Weinmannia racemosa (2.21%). Diameter growth rates were less variable among species and ranged from 1.8 mm?yr?1 in Ixerba brexioides to 3.3?mm?yr?1 in D.?cupressinum. Tree size influenced the rate of mortality in Beilschmiedia tawa, I. brexioides and W.?racemosa but there was no support for including tree size in models of the remaining three species. Likewise, tree size influenced growth rates in I.?brexioides and Nothofagus menziesii but not the remaining four species. These data provide robust size- and species-specific estimates of large-tree demographic rates that can be used as baselines for monitoring the impacts of management and global change in old-growth forests.  相似文献   

10.
Elevated atmospheric carbon dioxide (eCO2) is predicted to increase growth rates of forest trees. The extent to which increased growth translates to changes in biomass is dependent on the turnover time of the carbon, and thus tree mortality rates. Size‐ or age‐dependent mortality combined with increased growth rates could result in either decreased carbon turnover from a speeding up of tree life cycles, or increased biomass from trees reaching larger sizes, respectively. However, most vegetation models currently lack any representation of size‐ or age‐dependent mortality and the effect of eCO2 on changes in biomass and carbon turnover times is thus a major source of uncertainty in predictions of future vegetation dynamics. Using a reduced‐complexity form of the vegetation demographic model the Functionally Assembled Terrestrial Ecosystem Simulator to simulate an idealised tropical forest, we find increases in biomass despite reductions in carbon turnover time in both size‐ and age‐dependent mortality scenarios in response to a hypothetical eCO2‐driven 25% increase in woody net primary productivity (wNPP). Carbon turnover times decreased by 9.6% in size‐dependent mortality scenarios due to a speeding up of tree life cycles, but also by 2.0% when mortality was age‐dependent, as larger crowns led to increased light competition. Increases in aboveground biomass (AGB) were much larger when mortality was age‐dependent (24.3%) compared with size‐dependent (13.4%) as trees reached larger sizes before death. In simulations with a constant background mortality rate, carbon turnover time decreased by 2.1% and AGB increased by 24.0%, however, absolute values of AGB and carbon turnover were higher than in either size‐ or age‐dependent mortality scenario. The extent to which AGB increases and carbon turnover decreases will thus depend on the mechanisms of large tree mortality: if increased size itself results in elevated mortality rates, then this could reduce by about half the increase in AGB relative to the increase in wNPP.  相似文献   

11.
The size distribution of trees in natural forests is a fundamental attribute of forest structure. Previous attempts to model tree size distributions using simple functions (such as power or Weibull functions) have had limited success, typically overestimating the number of large stems observed. We describe a model which assumes that the dominant mortality process is asymmetric competition when trees are smaller, and size‐independent processes (e.g. disturbance) when trees are larger. This combination of processes leads to a size distribution which takes the form of a power distribution in the small tree phase and a Weibull distribution in the large tree phase. Analyses of data from four large‐scale (≥ 24 ha each) subtropical and temperate forest plots totalling 99 ha and approximately 0.4 million trees provide support for this model in two respects: (a) the combined function provided unbiased predictions and (b) power‐law functions fitted to small trees had exponents that deviated from the universal exponent of –2 predicted by metabolic scaling theory, gradually decreasing from subtropical evergreen to temperate deciduous forests along the latitudinal gradient.  相似文献   

12.
Aim Insect assemblages associated with lianas in tropical forests are poorly studied compared with those associated with trees. The importance of lianas for the maintenance of local species richness of insect herbivores in tropical forests is therefore poorly understood. With this in mind, a comparative study of the relative importance of trees and lianas as hosts for phytophagous beetles was carried out. Location The study area was located in the canopy of a dry tropical forest in Parque Natural Metropolitano, Panama province, Republic of Panama. Methods A crane system was utilized to access the canopy. The number of species and host specialization of adult phytophagous beetles associated with twenty‐six liana species of ten different families, and twenty‐four tree species of twelve different families were compared. Results A total of 2561 host associations of 697 species of beetles were determined (1339 for trees and 1222 for lianas). On average 55.8 ± 6.8 beetle species were found to be associated with each tree species while the comparable number for lianas was 47.0 ± 6.1. The pooled numbers of phytophagous beetle species associated with trees and lianas, respectively, were not significantly different. However, there were significantly more species feeding on green plant parts on lianas than on trees, and there were significantly more wood eaters on trees than on lianas. Phytophagous beetles associated with lianas were significantly more specialized than the tree associates due to a higher degree of specialization among the species feeding on green plant parts of lianas. Wood eaters and flower visitors showed no differences in host specialization on different growth forms. Main conclusion The present study shows that lianas are at least as important as trees for the maintenance of local species diversity of phytophagous beetles at this site. The mechanisms that drive the patterns can only be hypothesized. Plant architecture, size, and length of growing season are probably involved. Further studies, should include measurements of plant traits to elucidate experimentally what mechanisms that drive the patterns. Additional insight would come from similar studies in other forest types, and also studies of other major taxonomic groups of arthropod herbivores.  相似文献   

13.
Tropical forests are carbon rich ecosystems and small changes in tropical forest tree growth substantially influence the global carbon cycle. Forest monitoring studies report inconsistent growth changes in tropical forest trees over the past decades. Most of the studies highlighted changes in the forest level carbon gain, neglecting the species-specific growth changes which ultimately determine community-level responses. Tree-ring analysis can provide historical data on species-specific tree growth with annual resolution. Such studies are inadequate in Bangladesh, which is one of the most climate sensitive regions in the tropics. In this study, we investigated long-term growth rates of Toona ciliata in a moist tropical forest of Bangladesh by using tree-ring analysis. We sampled 50 trees of varying size, obtained increment cores from these trees and measured tree-ring width. Analyses of growth patterns revealed size-dependent growth increments. After correcting for the effect of tree size on tree growth (ontogenetic changes) by two different methods we found declining growth rates in T. ciliata from 1960 to 2013. Standardized ring-width index (RWI) was strongly negatively correlated with annual mean and maximum temperatures suggesting that rising temperature might cause the observed growth decline in T. ciliata. Assuming that global temperatures will rise at the current rate, the observed growth decline is assumed to continue. The analysis of stable carbon and oxygen isotopes may reveal more insight on the physiological response of this species to future climatic changes.  相似文献   

14.
Recent studies have suggested that tropical forests may not be resilient against climate change in the long term, primarily owing to predicted reductions in rainfall and forest productivity, increased tree mortality, and declining forest biomass carbon sinks. These changes will be caused by drought‐induced water stress and ecosystem disturbances. Several recent studies have reported that climate change has increased tree mortality in temperate and boreal forests, or both mortality and recruitment rates in tropical forests. However, no study has yet examined these changes in the subtropical forests that account for the majority of China's forested land. In this study, we describe how the monsoon evergreen broad‐leaved forest has responded to global warming and drought stress using 32 years of data from forest observation plots. Due to an imbalance in mortality and recruitment, and changes in diameter growth rates between larger and smaller trees and among different functional groups, the average DBH of trees and forest biomass have decreased. Sap flow measurements also showed that larger trees were more stressed than smaller trees by the warming and drying environment. As a result, the monsoon evergreen broad‐leaved forest community is undergoing a transition from a forest dominated by a cohort of fewer and larger individuals to a forest dominated by a cohort of more and smaller individuals, with a different species composition, suggesting that subtropical forests are threatened by their lack of resilience against long‐term climate change.  相似文献   

15.
Aim   We seek to determine the factors which control the success of lianas across macroecological gradients. Lianas have a strong impact on the growth, mortality and biomass of tropical trees, and are reported to be increasing in dominance, so understanding their behaviour is important from the perspectives of both ecological and global change.
Location   Lowland and montane Neotropical forests.
Methods   Using 65 standardized samples of lianas (≥ 2.5 cm diameter) from across the Neotropics, we attempted to account for characteristics of both the environment and the forest in explaining macroecological variation in liana success in Neotropical forests, using regression analyses and structural equation modelling.
Results   We found that both liana density and basal area were unrelated to mean annual precipitation, dry season length or soil variables, except for a weak effect of mean annual precipitation on liana basal area. Structural characteristics of the forest explained more of the variation in liana density and basal area than the physical environment. More disturbed forests generally tended to have a higher liana density. Liana basal area, however, was highest in undisturbed forests.
Main conclusions   The availability of host trees and their characteristics may be more important than the direct effects of the physical environment in controlling the success of lianas in Neotropical forests. Changes to the tropical climate in the coming century may not strongly affect lianas directly, but could have very substantial indirect effects via changes in tree community structure and dynamics.  相似文献   

16.
Understanding the mechanisms generating species distributions remains a challenge, especially in hyperdiverse tropical forests. We evaluated the role of rainfall variation, soil gradients and herbivory on seedling mortality, and how variation in seedling performance along these gradients contributes to habitat specialisation. In a 4‐year experiment, replicated at the two extremes of the Amazon basin, we reciprocally transplanted 4638 tree seedlings of 41 habitat‐specialist species from seven phylogenetic lineages among the three most important forest habitats of lowland Amazonia. Rainfall variation, flooding and soil gradients strongly influenced seedling mortality, whereas herbivory had negligible impact. Seedling mortality varied strongly among habitats, consistent with predictions for habitat specialists in most lineages. This suggests that seedling performance is a primary determinant of the habitat associations of adult trees across Amazonia. It further suggests that tree diversity, currently mostly harboured in terra firme forests, may be strongly impacted by the predicted climate changes in Amazonia.  相似文献   

17.
Forest succession depends strongly on the life history strategies of individual trees. An important strategic element is the ability to survive unfavourable environmental conditions that result in strongly reduced tree growth. In this study, we investigated whether the relationship between growth and mortality differs among tree species and site conditions. We analysed 10 329 trees of nine tree species (Picea abies , Taxus baccata , Fagus sylvatica , Tilia cordata , Carpinus betulus , Fraxinus excelsior , Quercus robur , Betula spp. and Alnus glutinosa ) from unmanaged forests of Europe: the continental Białowieża forest (Poland) and several oceanically influenced Swiss forest reserves. For each species, we calculated a set of flexible logistic regression models with the explanatory variables growth (as measured by relative basal area increment), tree size and site. We selected the species-specific model with the highest goodness-of-fit and calculated its discriminatory power (area under the receiver operating characteristic curve, AUC) and calibration measures. Most models achieved at least a good discriminatory power (AUC>0.7) and the AUC ranged from 0.62 to 0.87; calibration curves did not indicate any overfitting. Almost all growth–mortality relationships differed among species and sites, i.e. there is no universal growth–mortality relationship. Some species such as F. excelsior showed reduced survival probabilities for both unfavourable and very good growth conditions. We conclude that the growth–mortality relationships presented here can contribute to the life-history classification of trees and that they should also help to improve projections of forest succession models.  相似文献   

18.
Stegen JC  White EP 《Ecology letters》2008,11(12):1287-1293
It has been suggested that frequency distributions of individual tree masses in natural stands are characterized by power-law distributions with exponents near -3/4, and that therefore tree communities exhibit energetic equivalence among size classes. Because the mass of trees is not measured directly, but estimated from diameter, this supposition is based on the fact that the observed distribution of tree diameters is approximately characterized by a power-law with an exponent approximately -2. Here we show that diameter distributions of this form are not equivalent to mass distributions with exponents of -3/4, but actually to mass distributions with exponents of -11/8. We discuss the implications of this result for the metabolic theory of ecology and for understanding energetic equivalence and the processes structuring tree communities.  相似文献   

19.
Most North American forests are at some stage of post‐disturbance regrowth, subject to a changing climate, and exhibit growth and mortality patterns that may not be closely coupled to annual environmental conditions. Distinguishing the possibly interacting effects of these processes is necessary to put short‐term studies in a longer term context, and particularly important for the carbon‐dense, fire‐prone boreal forest. The goals of this study were to combine dendrochronological sampling, inventory records, and machine‐learning algorithms to understand how tree growth and death have changed at one highly studied site (Northern Old Black Spruce, NOBS) in the central Canadian boreal forest. Over the 1999–2012 inventory period, mean tree diameter increased even as stand density and basal area declined significantly. Tree mortality averaged 1.4 ± 0.6% yr?1, with most mortality occurring in medium‐sized trees; new recruitment was minimal. There have been at least two, and probably three, significant influxes of new trees since stand initiation, but none in recent decades. A combined tree ring chronology constructed from sampling in 2001, 2004, and 2012 showed several periods of extreme growth depression, with increased mortality lagging depressed growth by ~5 years. Higher minimum and maximum air temperatures exerted a negative influence on tree growth, while precipitation and climate moisture index had a positive effect; both current‐ and previous‐year data exerted significant effects. Models based on these variables explained 23–44% of the ring‐width variability. We suggest that past climate extremes led to significant mortality still visible in the current forest structure, with decadal dynamics superimposed on slower patterns of fire and succession. These results have significant implications for our understanding of previous work at NOBS, the carbon sequestration capability of old‐growth stands in a disturbance‐prone landscape, and the sustainable management of regional forests in a changing climate.  相似文献   

20.
Lianas (woody vines) are increasing in neotropical forests, representing one of the first large-scale structural changes documented for these important ecosystems. The potential ramifications of increasing lianas are huge, as lianas alter both tropical forest diversity and ecosystem functioning. At the community level, lianas affect tree species co-existence and diversity by competing more intensely with some tree species than others, and thus will likely alter tree species composition. At the ecosystem level, lianas affect forest carbon and nutrient storage and fluxes. A decrease in forest carbon storage and sequestration may be the most important ramification of liana increases. Lianas reduce tree growth and increase tree mortality—thus reducing forest-level carbon storage. The increase in lianas, which have much less wood than trees, compensates only partially for the amount of carbon lost in the displaced trees. Because tropical forests contribute approximately one-third of global terrestrial carbon stocks and net primary productivity, the effect of increasing lianas for tropical forest carbon cycles may have serious repercussions at the global scale.Key words: carbon cycle, CO2, disturbance, global change, land use change, liana increases, structural changes, tropical forestsTropical forests contain most of the earth''s plant species and contribute more to carbon storage in the form of above ground biomass than any other terrestrial ecosystem. Temperate and boreal forests are changing rapidly in response to global anthropogenic drivers. Similar large-scale changes are now being detected in tropical forests. One of the largest contemporary changes in tropical forests is an increase in lianas (woody vines),1 which could have serious consequences for tree species diversity and composition, as well as the reducing capacity of tropical forests to store carbon.13  相似文献   

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