Vertical variation in leaf gas exchange parameters for a Southeast Asian tropical rainforest in Peninsular Malaysia

Vertical variation in leaf gas exchange characteristics of trees grown in a lowland dipterocarp forest in Peninsular Malaysia was investigated. Maximum net photosynthetic rate, stomatal conductance, and electron transport rate of leaves at the upper canopy, lower canopy, and forest floor were studied in situ with saturated condition photosynthetic photon flux density. The dark respiration rate of leaves at the various heights was also studied. Relationships among gas exchange characteristics, and also with nitrogen content per unit leaf area and leaf dry matter per area were clearly detected, forming general equations representing the vertical profile of several important parameters related to gas exchange. Numerical analysis revealed that the vertical distribution of gas exchange parameters was well determined showing both larger carbon gain for the whole canopy and at the same time positive carbon gain for the leaves of the lowest layer. For correct estimation of gas exchange at both leaf and canopy scales using multi-layer models, it is essential to consider the vertical distribution of gas exchange parameters with proper scaling coefficients.     

Coping with branch excision when measuring leaf net photosynthetic rates in a lowland tropical forest

Measuring leaf gas exchange from canopy leaves is fundamental for our understanding of photosynthesis and for a realistic representation of carbon uptake in vegetation models. Since canopy leaves are often difficult to reach, especially in tropical forests with emergent trees up to 60 m at remote places, canopy access techniques such as canopy cranes or towers have facilitated photosynthetic measurements. These structures are expensive and therefore not very common. As an alternative, branches are often cut to enable leaf gas exchange measurements. The effect of branch excision on leaf gas exchange rates should be minimized and quantified to evaluate possible bias. We compared light-saturated leaf net photosynthetic rates measured on excised and intact branches. We selected branches positioned at three canopy positions, estimated relative to the top of the canopy: upper sunlit foliage, middle canopy foliage, and lower canopy foliage. We studied the variation of the effects of branch excision and transport among branches at these different heights in the canopy. After excision and transport, light-saturated leaf net photosynthetic rates were close to zero for most leaves due to stomatal closure. However, when the branch had acclimated to its new environmental conditions—which took on average 20 min—light-saturated leaf net photosynthetic rates did not significantly differ between the excised and intact branches. We therefore conclude that branch excision does not affect the measurement of light-saturated leaf net photosynthesis, provided that the branch is recut under water and is allowed sufficient time to acclimate to its new environmental conditions.     

Remote sensing of forest gas exchange: Considerations derived from a tomographic perspective

The global exchange of gas (CO₂, H₂O) and energy (sensible and latent heat) between forest ecosystems and the atmosphere is often assessed using remote sensing (RS) products. Although these products are essential in quantifying the spatial variability of forest–atmosphere exchanges, large uncertainties remain from a measurement bias towards top of canopy fluxes since optical RS data are not sensitive for the vertically integrated forest canopy. We hypothesize that a tomographic perspective opens new pathways to advance upscaling gas exchange processes from leaf to forest stands and larger scales. We suggest a 3D modelling environment comprising principles of ecohydrology and radiative transfer modelling with measurements of micrometeorological variables, leaf optical properties and forest structure, and assess 3D fields of net CO₂ assimilation (A_n) and transpiration (T) in a Swiss temperate forest canopy. 3D simulations were used to quantify uncertainties in gas exchange estimates inherent to RS approaches and model assumptions (i.e. a big‐leaf approximation in modelling approaches). Our results reveal substantial 3D heterogeneity of forest gas exchange with top of canopy A_n and T being reduced by up to 98% at the bottom of the canopy. We show that a simplified use of RS causes uncertainties in estimated vertical gas exchange of up to 300% and that the spatial variation of gas exchange in the footprint of flux towers can exceed diurnal dynamics. We also demonstrate that big‐leaf assumptions can cause uncertainties up to a factor of 10 for estimates of A_n and T. Concluding, we acknowledge the large potential of 3D assessments of gas exchange to unravelling the role of vertical variability and canopy structure in regulating forest–atmosphere gas and energy exchange. Such information allows to systematically link canopy with global scale controls on forest functioning and eventually enables advanced understanding of forest responses to environmental change.     

Broad-scale spatial patterns of canopy cover and pond morphology affect the structure of a Neotropical amphibian metacommunity

Spatial and environmental processes influence species composition at distinct scales. Previous studies suggested that the distribution of larval anurans at the landscape-scale is influenced by environmental gradients related to adult breeding site selection, such as pond canopy cover, but not by water chemistry. However, the combined effects of spatial, pond morphology, and water chemistry variables on metacommunity structure of larval anurans have not been analyzed yet. We used a partial redundancy analysis with variation partitioning to analyze the relative influence of pond morphology (e.g., depth, area, and aquatic vegetation), water chemistry, and spatial variables on a tadpole metacommunity from southeastern Brazil. We predict that pond morphology and canopy cover will influence the metacommunity at broad spatial scales, while water chemistry would play a larger role at finer scales. We found that broad-scale spatial patterns of pond canopy cover and pond morphology strongly influenced metacommunity structure, with water chemistry being not significant. Additionally, species composition was spatially autocorrelated at short distances. We suggest that the reproductive behavior of adult anurans is driving tadpole metacommunity dynamics, since pond morphology, but not water chemistry affects breeding site selection by adults. Our results contribute to the understanding of amphibian species diversity in tropical wetlands.     

Sources of Canopy Chemical and Spectral Diversity in Lowland Bornean Forest

Sources of variation among the chemical and spectral properties of tropical forest canopies are poorly understood, yet chemical traits reveal potential ecosystem and phylogenetic controls, and spectral linkages to chemical traits are needed for remote sensing of functional and biological diversity. We analyzed 21 leaf traits in 395 fully sunlit canopies, representing 232 species and multiple growth forms, in a lowland mixed dipterocarp forest of Sarawak, Malaysia. Leaf traits related to light capture and growth (for example, photosynthetic pigments, nutrients) were up to 55% lower, and defense traits (for example, phenols, lignin) were 15–40% higher, in the dominant family Dipterocarpaceae and in its genus Shorea, as compared to all other canopy species. The chemical variation within Dipterocarpaceae and Shorea was equivalent to that of all other canopy species combined, highlighting the role that a single phylogenetic branch can play in creating canopy chemical diversity. Seventeen of 21 traits had more than 50% of their variation explained by taxonomic grouping, and at least 16 traits show a connection to remotely sensed spectroscopic signatures (RMSE < 15%). It is through these chemical-to-spectral linkages that studies of functional and biological diversity interactions become possible at larger spatial scales, thereby improving our understanding of the role of species in tropical forest ecosystem dynamics.     

Towards physiologically meaningful water‐use efficiency estimates from eddy covariance data

Intrinsic water‐use efficiency (iWUE) characterizes the physiological control on the simultaneous exchange of water and carbon dioxide in terrestrial ecosystems. Knowledge of iWUE is commonly gained from leaf‐level gas exchange measurements, which are inevitably restricted in their spatial and temporal coverage. Flux measurements based on the eddy covariance (EC) technique can overcome these limitations, as they provide continuous and long‐term records of carbon and water fluxes at the ecosystem scale. However, vegetation gas exchange parameters derived from EC data are subject to scale‐dependent and method‐specific uncertainties that compromise their ecophysiological interpretation as well as their comparability among ecosystems and across spatial scales. Here, we use estimates of canopy conductance and gross primary productivity (GPP) derived from EC data to calculate a measure of iWUE (G₁, “stomatal slope”) at the ecosystem level at six sites comprising tropical, Mediterranean, temperate, and boreal forests. We assess the following six mechanisms potentially causing discrepancies between leaf and ecosystem‐level estimates of G₁: (i) non‐transpirational water fluxes; (ii) aerodynamic conductance; (iii) meteorological deviations between measurement height and canopy surface; (iv) energy balance non‐closure; (v) uncertainties in net ecosystem exchange partitioning; and (vi) physiological within‐canopy gradients. Our results demonstrate that an unclosed energy balance caused the largest uncertainties, in particular if it was associated with erroneous latent heat flux estimates. The effect of aerodynamic conductance on G₁ was sufficiently captured with a simple representation. G₁ was found to be less sensitive to meteorological deviations between canopy surface and measurement height and, given that data are appropriately filtered, to non‐transpirational water fluxes. Uncertainties in the derived GPP and physiological within‐canopy gradients and their implications for parameter estimates at leaf and ecosystem level are discussed. Our results highlight the importance of adequately considering the sources of uncertainty outlined here when EC‐derived water‐use efficiency is interpreted in an ecophysiological context.     

Spatial scale and sampling resolution affect measures of gap disturbance in a lowland tropical forest: implications for understanding forest regeneration and carbon storage

Treefall gaps play an important role in tropical forest dynamics and in determining above-ground biomass (AGB). However, our understanding of gap disturbance regimes is largely based either on surveys of forest plots that are small relative to spatial variation in gap disturbance, or on satellite imagery, which cannot accurately detect small gaps. We used high-resolution light detection and ranging data from a 1500 ha forest in Panama to: (i) determine how gap disturbance parameters are influenced by study area size, and the criteria used to define gaps; and (ii) to evaluate how accurately previous ground-based canopy height sampling can determine the size and location of gaps. We found that plot-scale disturbance parameters frequently differed significantly from those measured at the landscape-level, and that canopy height thresholds used to define gaps strongly influenced the gap-size distribution, an important metric influencing AGB. Furthermore, simulated ground surveys of canopy height frequently misrepresented the true location of gaps, which may affect conclusions about how relatively small canopy gaps affect successional processes and contribute to the maintenance of diversity. Across site comparisons need to consider how gap definition, scale and spatial resolution affect characterizations of gap disturbance, and its inferred importance for carbon storage and community composition.     

Imaging spectroscopy links aspen genotype with below-ground processes at landscape scales

Fine-scale biodiversity is increasingly recognized as important to ecosystem-level processes. Remote sensing technologies have great potential to estimate both biodiversity and ecosystem function over large spatial scales. Here, we demonstrate the capacity of imaging spectroscopy to discriminate among genotypes of Populus tremuloides (trembling aspen), one of the most genetically diverse and widespread forest species in North America. We combine imaging spectroscopy (AVIRIS) data with genetic, phytochemical, microbial and biogeochemical data to determine how intraspecific plant genetic variation influences below-ground processes at landscape scales. We demonstrate that both canopy chemistry and below-ground processes vary over large spatial scales (continental) according to aspen genotype. Imaging spectrometer data distinguish aspen genotypes through variation in canopy spectral signature. In addition, foliar spectral variation correlates well with variation in canopy chemistry, especially condensed tannins. Variation in aspen canopy chemistry, in turn, is correlated with variation in below-ground processes. Variation in spectra also correlates well with variation in soil traits. These findings indicate that forest tree species can create spatial mosaics of ecosystem functioning across large spatial scales and that these patterns can be quantified via remote sensing techniques. Moreover, they demonstrate the utility of using optical properties as proxies for fine-scale measurements of biodiversity over large spatial scales.     

Hydrological Networks and Associated Topographic Variation as Templates for the Spatial Organization of Tropical Forest Vegetation

An understanding of the spatial variability in tropical forest structure and biomass, and the mechanisms that underpin this variability, is critical for designing, interpreting, and upscaling field studies for regional carbon inventories. We investigated the spatial structure of tropical forest vegetation and its relationship to the hydrological network and associated topographic structure across spatial scales of 10–1000 m using high-resolution maps of LiDAR-derived mean canopy profile height (MCH) and elevation for 4930 ha of tropical forest in central Panama. MCH was strongly associated with the hydrological network: canopy height was highest in areas of positive convexity (valleys, depressions) close to channels draining 1 ha or more. Average MCH declined strongly with decreasing convexity (transition to ridges, hilltops) and increasing distance from the nearest channel. Spectral analysis, performed with wavelet decomposition, showed that the variance in MCH had fractal similarity at scales of ∼30–600 m, and was strongly associated with variation in elevation, with peak correlations at scales of ∼250 m. Whereas previous studies of topographic correlates of tropical forest structure conducted analyses at just one or a few spatial grains, our study found that correlations were strongly scale-dependent. Multi-scale analyses of correlations of MCH with slope, aspect, curvature, and Laplacian convexity found that MCH was most strongly related to convexity measured at scales of 20–300 m, a topographic variable that is a good proxy for position with respect to the hydrological network. Overall, our results support the idea that, even in these mesic forests, hydrological networks and associated topographical variation serve as templates upon which vegetation is organized over specific ranges of scales. These findings constitute an important step towards a mechanistic understanding of these patterns, and can guide upscaling and downscaling.     

The phenology of leaf quality and its within‐canopy variation is essential for accurate modeling of photosynthesis in tropical evergreen forests

Leaf quantity (i.e., canopy leaf area index, LAI), quality (i.e., per‐area photosynthetic capacity), and longevity all influence the photosynthetic seasonality of tropical evergreen forests. However, these components of tropical leaf phenology are poorly represented in most terrestrial biosphere models (TBMs). Here, we explored alternative options for the representation of leaf phenology effects in TBMs that employ the Farquahar, von Caemmerer & Berry (FvCB) representation of CO₂ assimilation. We developed a two‐fraction leaf (sun and shade), two‐layer canopy (upper and lower) photosynthesis model to evaluate different modeling approaches and assessed three components of phenological variations (i.e., leaf quantity, quality, and within‐canopy variation in leaf longevity). Our model was driven by the prescribed seasonality of leaf quantity and quality derived from ground‐based measurements within an Amazonian evergreen forest. Modeled photosynthetic seasonality was not sensitive to leaf quantity, but was highly sensitive to leaf quality and its vertical distribution within the canopy, with markedly more sensitivity to upper canopy leaf quality. This is because light absorption in tropical canopies is near maximal for the entire year, implying that seasonal changes in LAI have little impact on total canopy light absorption; and because leaf quality has a greater effect on photosynthesis of sunlit leaves than light limited, shade leaves and sunlit foliage are more abundant in the upper canopy. Our two‐fraction leaf, two‐layer canopy model, which accounted for all three phenological components, was able to simulate photosynthetic seasonality, explaining ~90% of the average seasonal variation in eddy covariance‐derived CO₂ assimilation. This work identifies a parsimonious approach for representing tropical evergreen forest photosynthetic seasonality in TBMs that utilize the FvCB model of CO₂ assimilation and highlights the importance of incorporating more realistic phenological mechanisms in models that seek to improve the projection of future carbon dynamics in tropical evergreen forests.     

林冠结构是局域尺度木本植物功能性状beta多样性形成的重要驱动力

功能性状beta多样性反映了群落间功能性状组成的差异, 解析其形成机制是群落生态学研究的核心内容之一。本研究以云南西双版纳热带季节雨林20 ha动态监测样地为研究对象, 测定木本植物11个重要的功能性状, 采用多度加权的平均最近邻体性状距离度量不同取样尺度的功能性状beta多样性, 基于距离矩阵的多元回归方法解析林冠结构差异、环境异质性、空间距离在功能性状beta多样性格局形成中的相对作用。结果表明, 对于所有木本植物个体(DBH ≥ 1 cm)而言, 同时考虑林冠结构、环境和空间距离的模型为解释功能性状beta多样性格局的最优模型; 在3个不同取样尺度上, 林冠结构差异和环境距离都对功能性状beta多样性具有较大的解释力, 且随着取样尺度的增大而上升, 空间距离的作用基本可以忽略。本研究证实了林冠结构是局域尺度木本植物功能性状beta多样性格局形成的重要驱动力, 这一发现更新了环境异质性和空间距离是驱动功能性状beta多样性格局形成的主要因素的传统认知, 为将来研究功能性状beta多样性形成机制提供新的视角, 并证实了取样尺度在解析木本植物功能性状beta多样性格局形成机制中的重要性。     

Tree Diversity Explains Variation in Ecosystem Function in a Neotropical Forest in Panama

Many experimental studies show that a decline in species number has a negative effect on ecosystem function, however less is known about this pattern in natural communities. We examined the relative importance of environment, space, and diversity on ecosystem function, specifically tree carbon storage in four plant types (understory/canopy; trees/palms), in a tropical forest in central Panama. The objectives of this study were to detect the relationship between tree diversity and carbon storage given the environmental and spatial variation that occur in natural forests and to determine which species diversity measure is more important to tree carbon storage: richness or dominance. We used redundancy analyses to partition the effect of these sources of variation on tree carbon storage. We showed that together, environment, space, and diversity accounted for 43 percent of tree carbon storage, where diversity (19%) alone is the most important source of variation and explained more variation than space (13%) and environment (1%) together. Therefore, even in natural forests where substantial environment and spatial variation can be found, it is still possible to detect the effect of diversity on ecosystem function at scales relevant to conservation. Moreover, both richness and dominance are important to explain the variation on tree carbon storage in natural forests suggesting that these two diversity measures are complementary. Thus, tree diversity is important to predict tree carbon storage in hyperdiverse forests.     

Do edge effects increase the susceptibility of rainforest fragments to structural damage resulting from a severe tropical cyclone?

Abstract If changes in the structural characteristics of rainforest at edges are caused by wind, then physical damage from a tropical cyclone might be greatest at edges or in small fragments that have a high proportion of edge. We tested whether this was true of a fragmented rainforest landscape impacted by a category 4 severe tropical cyclone in March 2006. Six structural variables (canopy cover, canopy height, cover of ground vegetation, leaf litter, stem density and counts of woody debris) were surveyed at 18 rainforest sites (six small linear remnants, and both edges and interiors of six large remnants) on the Atherton Tableland in north‐eastern Queensland, Australia. Data collected 7 and 12 months after the passage of Cyclone Larry were compared with an identical survey conducted 4 years prior to the cyclone. The cyclone had large effects across many components of forest structure. However, sites within 30 m of forest edges in small and large remnants were not impacted more than the interiors of large remnants. It is likely that the high wind intensity from severe tropical cyclones overrides the modest wind protection provided by surrounding forest. The cyclone's effects were highly patchy at local scales (0.5–1.0 km), leading to an increase in among‐site variation in forest structure and the disappearance of significant spatial autocorrelation among large remnant edge‐interior site pairs which had existed prior to the cyclone. The main effect of Cyclone Larry at these study sites was to increase the spatial heterogeneity of forest structure at local scales.     

海南岛热带天然针叶林主要树种的空间格局及关联性

天然针叶林在热带地区虽较为少见, 但其对维持热带地区的生物多样性和生境异质性具有特殊意义。在我国热带天然针叶林集中分布面积最大的海南霸王岭林区, 作者选择伴生阔叶树种优势度不同的两种典型南亚松(Pinus latteri)天然林(简称纯林和混交林), 采用点格局法分析了其林冠层、亚林层和林下层主要树种的空间分布格局及其关联性。结果表明: (1)纯林中林冠层的南亚松主要为聚集分布, 混交林中在较小尺度上为聚集分布, 在较大尺度上为随机分布。(2)纯林中亚林层树种在较小尺度上为聚集分布, 在较大尺度上为随机分布, 在混交林中主要为聚集分布。(3)纯林中林下层树种主要呈现为随机分布, 而在混交林中主要为聚集分布。(4)随着尺度的增加, 林冠层与其他两个层次的树种, 在纯林中表现出从空间无关联到正关联的变化趋势, 而在混交林中则表现出从空间无关联到负关联的变化趋势。(5)亚林层与林下层树种在各个尺度上都表现为空间正关联。由此可见, 热带天然针叶林中优势种南亚松对伴生阔叶树种的分布格局具有重要影响。     

Variation in tropical forest growth rates: combined effects of functional group composition and resource availability

Rates of tree growth in tropical forests reflect variation in life history strategies, contribute to the determination of species' distributional limits, set limits to timber harvesting and control the carbon balance of the stands. Here, we review the resources that limit tree growth at different temporal and spatial scales, and the different growth rates and responses of functional groups defined on the basis of regeneration strategy, maximum size, and species' associations with particular edaphic and climatic conditions.Variation in soil water availability determines intra- and inter-annual patterns of growth within seasonal forests, whereas irradiance may have a more important role in aseasonal forests. Nutrient supply limits growth rates in montane forests and may determine spatial variation in growth of individual species in lowland forests. However, its role in determining spatial variation in stand-level growth rates is unclear. In terms of growth rate, we propose a functional classification of tropical tree species which contrasts inherently fast-growing, responsive species (pioneer, large-statured species), from slow-growing species that are less responsive to increasing resource availability (shade-bearers, small-statured species). In a semi-deciduous forest in Ghana, pioneers associated with high-rainfall forests with less fertile soils, had significantly lower growth rates than pioneers that are more abundant in low-rainfall forests with more fertile soils. These results match patterns found in seedling trials and suggest for pioneers that species' associations with particular environmental conditions are useful indicators of maximum growth rate.The effects of variation in resource availability and of inherent differences between species on stand-level patterns of growth will not be independent if the functional group composition of tropical forests varies along resource gradients. We find that there is increasing evidence of such spatial shifts at both small and large scales in tropical forests. Quantifying these gradients is important for understanding spatial patterns in forest growth rates.     

Leaf litter inputs reinforce islands of nitrogen fertility in a lowland tropical forest

The role of lowland tropical forest tree communities in shaping soil nutrient cycling has been challenging to elucidate in the face of high species diversity. Previously, we showed that differences in tree species composition and canopy foliar nitrogen (N) concentrations correlated with differences in soil N availability in a mature Costa Rican rainforest. Here, we investigate potential mechanisms explaining this correlation. We used imaging spectroscopy to identify study plots containing 10–20 canopy trees with either high or low mean canopy N relative to the landscape mean. Plots were restricted to an uplifted terrace with relatively uniform parent material and climate. In order to assess whether canopy and soil N could be linked by litterfall inputs, we tracked litter production in the plots and measured rates of litter decay and the carbon and N content of leaf litter and leaf litter leachate. We also compared the abundance of putative N fixing trees and rates of free-living N fixation as well as soil pH, texture, cation exchange capacity, and topographic curvature to assess whether biological N fixation and/or soil properties could account for differences in soil N that were, in turn, imprinted on the canopy. We found no evidence of differences in legume communities, free-living N fixation, or abiotic properties. However, soils beneath high canopy N assemblages received ~ 60% more N via leaf litterfall due to variability in litter N content between plot types. The correlation of N in canopy leaves, leaf litter, and soil suggests that, under similar abiotic conditions, litterfall-mediated feedbacks can help maintain soil N differences among tropical tree assemblages in this diverse tropical forest.

     

Environmental drivers of tree community turnover in western Amazonian forests

A more comprehensive understanding of the factors governing tropical tree community turnover at different spatial scales is needed to support land‐management and biodiversity conservation. We used new forest inventory data from 263 permanent plots in the Carnegie Biodiversity‐Biomass Forest Plot Network spanning the eastern Andes to the western Amazonian lowlands of Peru to examine environmental factors driving genus‐level canopy tree compositional variation at regional and landscape scales. Across the full plot network, constrained ordination analysis indicated that all environmental variables together explained 23.8% of the variation in community composition, while soil, topographic, and climatic variables each explained 15.2, 10.9, and 17.0%, respectively. A satellite‐derived metric of cloudiness was the single strongest predictor of community turnover, and constrained ordination revealed a primary gradient of environmentally‐driven community turnover spanning from cloudy, high elevation sites to warm, wet, lowland sites. For three focal landscapes within the region, local environmental variation explained 13.4–30.8% of compositional variation. Community turnover at the landscape scale was strongly driven by topo‐edaphic factors in the two lowland landscapes examined and strongly driven by potential insolation and topography in the montane landscape. At the regional scale, we found that the portion of compositional variation that was uniquely explained by spatial variation was relatively small (2.7%), and was effectively zero within the three focal landscapes. Overall, our results show strong canopy tree compositional turnover in response to environmental gradients at both regional and landscape scales, though the most important environmental drivers differed between scales and among landscapes. Our results also highlight the usefulness of key satellite‐derived environmental covariates that should be considered when conducting biodiversity analyses in tropical forests.     

Using Light-Use and Production Efficiency Models to Predict Photosynthesis and Net Carbon Exchange During Forest Canopy Disturbance

Vegetation growth models are used with remotely sensed and meteorological data to monitor terrestrial carbon dynamics at a range of spatial and temporal scales. Many of these models are based on a light-use efficiency equation and two-component model of whole-plant growth and maintenance respiration that have been parameterized for distinct vegetation types and biomes. This study was designed to assess the robustness of these parameters for predicting interannual plant growth and carbon exchange, and more specifically to address inconsistencies that may arise during forest disturbances and the loss of canopy foliage. A model based on the MODIS MOD17 algorithm was parameterized for a mature upland hardwood forest by inverting CO₂ flux tower observations during years when the canopy was not disturbed. This model was used to make predictions during a year when the canopy was 37% defoliated by forest tent caterpillars. Predictions improved after algorithms were modified to scale for the effects of diffuse radiation and loss of leaf area. Photosynthesis and respiration model parameters were found to be robust at daily and annual time scales regardless of canopy disturbance, and differences between modeled net ecosystem production and tower net ecosystem exchange were only approximately 2 g C m⁻² d⁻¹ and less than 23 g C m⁻² y⁻¹. Canopy disturbance events such as insect defoliations are common in temperate forests of North America, and failure to account for cyclical outbreaks of forest tent caterpillars in this stand could add an uncertainty of approximately 4–13% in long-term predictions of carbon sequestration.     

Scale matters: fire–vegetation feedbacks are needed to explain tropical tree cover at the local scale

At a broad (regional to global) spatial scale, tropical vegetation is controlled by climate; at the local scale, it is believed to be determined by interactions between disturbance, vegetation and local conditions (soil and topography) through feedback processes. It has recently been suggested that strong fire–vegetation feedback processes may not be needed to explain tree‐cover patterns in tropical ecosystems and that climate–fire determinism is an alternative possibility. This conclusion was based on the fact that it is possible to reproduce observed patterns in tropical regions (e.g. a trimodal frequency distribution of tree cover) using a simple model that does not explicitly incorporate fire–vegetation feedback processes. We argue that these two mechanisms (feedbacks versus fire–climate control) operate at different spatial and temporal scales; it is not possible to evaluate the role of a process acting at fine scales (e.g. fire–vegetation feedbacks) using a model designed to reproduce regional‐scale pattern (scale mismatch). While the distributions of forest and savannas are partially determined by climate, many studies are providing evidence that the most parsimonious explanation for their environmental overlaps is the existence of feedback processes. Climate is unlikely to be an alternative to feedback processes; rather, climate and fire–vegetation feedbacks are complementary processes at different spatial and temporal scales.