Temperature explains global variation in biomass among humid old‐growth forests

Aim To develop and test a simple climate‐based ecophysiological model of above‐ground biomass – an approach that can be applied directly to predicting the effects of climate change on forest carbon stores. Location Humid lowland forests world‐wide. Methods We developed a new approach to modelling the aboveground biomass of old‐growth forest (AGB_max) based on the influences of temperature on gross primary productivity (GPP) and what we call total maintenance cost (TMC), which includes autotrophic respiration as well as leaf, stem and other plant construction required to maintain biomass. We parameterized the models with measured carbon fluxes and tested them by comparing predicted AGB_max with measured AGB for another 109 old‐growth sites. Results Our models explained 57% of the variation in GPP across 95 sites and 79% of the variation in TMC across 17 sites. According to the best‐fit models, the ratio of GPP to maintenance cost per unit biomass (MCB) peaks at 16.5 °C, indicating that this is the air temperature leading to the highest possible AGB_max when temperatures are constant. Seasonal temperature variation generally reduces predicted AGB_max, and thus maritime temperate climates are predicted to have the highest AGB_max. The shift in temperatures from temperate maritime to tropical climates increases MCB more than GPP, and thus decreases AGB_max. Overall, our model explains exactly 50% of the variation in AGB among humid lowland old‐growth forests. Main conclusions Temperature plays an important role in explaining global variation in biomass among humid lowland old‐growth forests, a role that can be understood in terms of the dual effects of temperature on GPP and TMC. Our simple model captures these influences, and could be an important tool for predicting the effects of climate change on forest carbon stores.     

Allometric constraints and competition enable the simulation of size structure and carbon fluxes in a dynamic vegetation model of tropical forests (LM3PPA‐TV)

Tropical forests are a key determinant of the functioning of the Earth system, but remain a major source of uncertainty in carbon cycle models and climate change projections. In this study, we present an updated land model (LM3PPA‐TV) to improve the representation of tropical forest structure and dynamics in Earth system models (ESMs). The development and parameterization of LM3PPA‐TV drew on extensive datasets on tropical tree traits and long‐term field censuses from Barro Colorado Island (BCI), Panama. The model defines a new plant functional type (PFT) based on the characteristics of shade‐tolerant, tropical tree species, implements a new growth allocation scheme based on realistic tree allometries, incorporates hydraulic constraints on biomass accumulation, and features a new compartment for tree branches and branch fall dynamics. Simulation experiments reproduced observed diurnal and seasonal patterns in stand‐level carbon and water fluxes, as well as mean canopy and understory tree growth rates, tree size distributions, and stand‐level biomass on BCI. Simulations at multiple sites captured considerable variation in biomass and size structure across the tropical forest biome, including observed responses to precipitation and temperature. Model experiments suggested a major role of water limitation in controlling geographic variation forest biomass and structure. However, the failure to simulate tropical forests under extreme conditions and the systematic underestimation of forest biomass in Paleotropical locations highlighted the need to incorporate variation in hydraulic traits and multiple PFTs that capture the distinct floristic composition across tropical domains. The continued pressure on tropical forests from global change demands models which are able to simulate alternative successional pathways and their pace to recovery. LM3PPA‐TV provides a tool to investigate geographic variation in tropical forests and a benchmark to continue improving the representation of tropical forests dynamics and their carbon storage potential in ESMs.     

Roles of dominant understory Sasa bamboo in carbon and nitrogen dynamics following canopy tree removal in a cool‐temperate forest in northern Japan

To clarify the role of dense understory vegetation in the stand structure, and in carbon (C) and nitrogen (N) dynamics of forest ecosystems with various conditions of overstory trees, we: (i) quantified the above‐ and below‐ground biomasses of understory dwarf bamboo (Sasa senanensis) at the old canopy‐gap area and the closed‐canopy area and compared the stand‐level biomasses of S. senanensis with that of overstory trees; (ii) determined the N leaching, soil respiration rates, fine‐root dynamics, plant area index (PAI) of S. senanensis, and soil temperature and moisture at the tree‐cut patches (cut) and the intact closed‐canopy patches (control). The biomass of S. senanensis in the canopy‐gap area was twice that at the closed‐canopy area. It equated to 12% of total biomass above ground but 41% below ground in the stand. The concentrations of NO₃^? and NH₄⁺ in the soil solution and soil respiration rates did not significantly change between cut and control plots, indicating that gap creation did not affect the C or N dynamics in the soil. Root‐length density and PAI of S. senanensis were significantly greater at the cut plots, suggesting the promotion of S. senanensis growth following tree cutting. The levels of soil temperature and soil moisture were not changed following tree cutting. These results show that S. senanensis is a key component species in this cool‐temperate forest ecosystem and plays significant roles in mitigating the loss of N and C from the soil following tree cutting by increasing its leaf and root biomass and stabilizing the soil environment.     

Linking canopy leaf area and light environments with tree size distributions to explain Amazon forest demography

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.     

Changes in grassland canopy structure across a precipitation gradient

Abstract. In temperate grasslands, the relative importance of above‐ground competition for light compared to below‐ground competition for water and nutrients is hypothesized to increase with increasing precipitation. Thus, competition for light is likely to exert an increasing influence on canopy structure and species composition as precipitation increases. We quantified canopy structure, light availability and changes in species composition at seven sites across the central grassland region of the United States to determine how these properties change across a precipitation gradient. Across the region, there was a disproportionate increase in leaf area and canopy height with increasing precipitation, indicating that plants become taller and leafier across the gradient. Leaf area index increased by a factor of 12 across the gradient while above‐ground net primary productivity increased by a factor of only 5.5. As precipitation increased, there was decreased light availability at the soil surface, increased seasonal variability in light transmission, increased biomass and leaf area at higher canopy layers and an increased proportion of tallstatured species. These observed changes in canopy structure support the prediction that competition for light increases in importance with increasing precipitation.     

Lidar remote sensing of above-ground biomass in three biomes

Estimation of the amount of carbon stored in forests is a key challenge for understanding the global carbon cycle, one which remote sensing is expected to help address. However, estimation of carbon storage in moderate to high biomass forests is difficult for conventional optical and radar sensors. Lidar (light detection and ranging) instruments measure the vertical structure of forests and thus hold great promise for remotely sensing the quantity and spatial organization of forest biomass. In this study, we compare the relationships between lidar‐measured canopy structure and coincident field measurements of above‐ground biomass at sites in the temperate deciduous, temperate coniferous, and boreal coniferous biomes. A single regression for all three sites is compared with equations derived for each site individually. The single equation explains 84% of variance in above‐ground biomass (P < 0.0001) and shows no statistically significant bias in its predictions for any individual site.     

Carbon balances of bioenergy systems using biomass from forests managed with long rotations: bridging the gap between stand and landscape assessments

Studies report different findings concerning the climate benefits of bioenergy, in part due to varying scope and use of different approaches to define spatial and temporal system boundaries. We quantify carbon balances for bioenergy systems that use biomass from forests managed with long rotations, employing different approaches and boundary conditions. Two approaches to represent landscapes and quantify their carbon balances – expanding vs. constant spatial boundaries – are compared. We show that for a conceptual forest landscape, constructed by combining a series of time‐shifted forest stands, the two approaches sometimes yield different results. We argue that the approach that uses constant spatial boundaries is preferable because it captures all carbon flows in the landscape throughout the accounting period. The approach that uses expanding system boundaries fails to accurately describe the carbon fluxes in the landscape due to incomplete coverage of carbon flows and influence of the stand‐level dynamics, which in turn arise from the way temporal system boundaries are defined on the stand level. Modelling of profit‐driven forest management using location‐specific forest data shows that the implications for carbon balance of management changes across the landscape (which are partly neglected when expanding system boundaries are used) depend on many factors such as forest structure and forest owners’ expectations of market development for bioenergy and other wood products. Assessments should not consider forest‐based bioenergy in isolation but should ideally consider all forest products and how forest management planning as a whole is affected by bioenergy incentives – and how this in turn affects carbon balances in forest landscapes and forest product pools. Due to uncertainties, we modelled several alternative scenarios for forest products markets. We recommend that future work consider alternative scenarios for other critical factors, such as policy options and energy technology pathways.     

不同邻体大头茶幼苗的适应特征

 在亚热带常绿阔叶林自然林窗下,以大头茶幼苗为基株,分别以马尾松、川灰木和大头茶幼苗为邻体。研究不同邻体如何改变可利用资源,影响大头茶幼苗基株生长动态和形态、树冠结构和生物量效应,并探讨影响基株生长的邻体的主导因素。结果表明,不同邻体的生长和截光效应差异明显;基株生长特征与邻体的树高、叶生物量、总生物量和树冠距地表10cm处截光量相关极小,却与邻体树冠距地表30cm处截光量相关极大,故影响基株生长的邻体主导因素是遮挡基株树冠的邻体的可利用光资源;基株生长动态,如树高、叶数、叶面积和叶大小,树冠结构和生物量显著受到不同邻体的影响,并且基株树高及增长量、叶面积、冠幅和生物量均为大头茶单株(GNO)>大头茶+川灰木(GNS)>大头茶+马尾松(GNP)>大头茶+大头茶(GNG)。大头茶幼苗种内竞争明显强于种间竞争。     

Biodiversity and climate determine the functioning of Neotropical forests

Aim

Tropical forests account for a quarter of the global carbon storage and a third of the terrestrial productivity. Few studies have teased apart the relative importance of environmental factors and forest attributes for ecosystem functioning, especially for the tropics. This study aims to relate aboveground biomass (AGB) and biomass dynamics (i.e., net biomass productivity and its underlying demographic drivers: biomass recruitment, growth and mortality) to forest attributes (tree diversity, community‐mean traits and stand basal area) and environmental conditions (water availability, soil fertility and disturbance).

Location

Neotropics.

Methods

We used data from 26 sites, 201 1‐ha plots and >92,000 trees distributed across the Neotropics. We quantified for each site water availability and soil total exchangeable bases and for each plot three key community‐weighted mean functional traits that are important for biomass stocks and productivity. We used structural equation models to test the hypothesis that all drivers have independent, positive effects on biomass stocks and dynamics.

Results

Of the relationships analysed, vegetation attributes were more frequently associated significantly with biomass stocks and dynamics than environmental conditions (in 67 vs. 33% of the relationships). High climatic water availability increased biomass growth and stocks, light disturbance increased biomass growth, and soil bases had no effect. Rarefied tree species richness had consistent positive relationships with biomass stocks and dynamics, probably because of niche complementarity, but was not related to net biomass productivity. Community‐mean traits were good predictors of biomass stocks and dynamics.

Main conclusions

Water availability has a strong positive effect on biomass stocks and growth, and a future predicted increase in (atmospheric) drought might, therefore, potentially reduce carbon storage. Forest attributes, including species diversity and community‐weighted mean traits, have independent and important relationships with AGB stocks, dynamics and ecosystem functioning, not only in relatively simple temperate systems, but also in structurally complex hyper‐diverse tropical forests.     

Lack of disturbance as an explanation for the additive basal area phenomenon in a stratified Afrotemperate forest

Abstract Explanations for the additive basal area (BA) phenomenon in forests are frequently given in the context of stratification and the avoidance of competition for space or resources (e.g. light, nutrients, moisture). Thus, large individuals avoid competition for light by emerging above the canopy and, in so doing, the BA of emergent individuals is often ‘additive’ to that of the rest of the individuals in a stand. The additive BA phenomenon was evident in a stratified Afrotemperate forest and was not confined to the emergent stratum but occurred also within the canopy stratum where gymnosperm BA appeared to be additive. However, there was no evidence that stand BA was at carrying capacity (i.e. presumably neither space nor nutrient resources were limiting) and there was no statistical evidence of competitive effects. We argue that avoiding competition is an insufficient explanation for how biomass accumulates in stratified forests and suggest that local variation in disturbance regime and tree life‐history provide a more plausible and general explanation for current forest stand structure and dynamics. We suggest that the additive effect is a result of long‐lived individuals persisting in the prolonged absence of disturbance to canopy and emergent strata.     

Leaf traits determine the growth-survival trade-off across rain forest tree species

A dominant hypothesis explaining tree species coexistence in tropical forest is that trade-offs in characters allow species to adapt to different light environments, but tests for this hypothesis are scarce. This study is the first that uses a theoretical plant growth model to link leaf trade-offs to whole-plant performances and to differential performances across species in different light environments. Using data of 50 sympatric tree species from a Bolivian rain forest, we observed that specific leaf area and photosynthetic capacity codetermined interspecific height growth variation in a forest gap; that leaf survival rate determined the variation in plant survival rate under a closed canopy; that predicted height growth and plant survival rate matched field observations; and that fast-growing species had low survival rates for both field and predicted values. These results show how leaf trade-offs influence differential tree performance and tree species' coexistence in a heterogeneous light environment.     

Dynamics of a human‐modified tropical peat swamp forest revealed by repeat lidar surveys

Tropical peat swamp forests (PSFs) are globally important carbon stores under threat. In Southeast Asia, 35% of peatlands had been drained and converted to plantations by 2010, and much of the remaining forest had been logged, contributing significantly to global carbon emissions. Yet, tropical forests have the capacity to regain biomass quickly and forests on drained peatlands may grow faster in response to soil aeration, so the net effect of humans on forest biomass remains poorly understood. In this study, two lidar surveys (made in 2011 and 2014) are compared to map forest biomass dynamics across 96 km² of PSF in Kalimantan, Indonesia. The peatland is now legally protected for conservation, but large expanses were logged under concessions until 1998 and illegal logging continues in accessible portions. It was hypothesized that historically logged areas would be recovering biomass while recently logged areas would be losing biomass. We found that historically logged forests were recovering biomass near old canals and railways used by the concessions. Lidar detected substantial illegal logging activity—579 km of logging canals were located beneath the canopy. Some patches close to these canals have been logged in the 2011–2104 period (i.e. substantial biomass loss) but, on aggregate, these illegally logged regions were also recovering. Unexpectedly, rapid growth was also observed in intact forest that had not been logged and was over a kilometre from the nearest known canal, perhaps in response to greater aeration of surface peat. Comparing these results with flux measurements taken at other nearby sites, we find that carbon sequestration in above‐ground biomass may have offset roughly half the carbon efflux from peat oxidation. This study demonstrates the power of repeat lidar survey to map fine‐scale forest dynamics in remote areas, revealing previously unrecognized impacts of anthropogenic global change.     

Above-ground biomass estimation in closed canopy Neotropical forests using lidar remote sensing: factors affecting the generality of relationships

Aim Previous studies have developed strong, site‐specific relationships between canopy metrics from lidar (light detecting and ranging) remote sensing data and forest structural characteristics such as above‐ground biomass (AGBM), but the generality of these relationships is unknown. In this study, we examine the generality of relationships between lidar metrics and forest structural characteristics, including AGBM, from two study areas in Central America with different precipitation patterns. Location A series of tropical moist forest sites in Panama and a tropical wet forest in Costa Rica. Methods Canopy metrics (e.g. canopy height) were calculated from airborne lidar data. Basal area, mean stem diameter and AGBM were calculated from measurements taken as a part of ongoing forest dynamics studies in both areas. We examined the generality of relationship between lidar metrics and forest structure, and possible environmental effects (e.g. leaf phenology). Results We found that lidar metrics were strongly correlated (R²: 0.65–0.92) with mean stem diameter, basal area and AGBM in both regions. We also show that the relationships differed between these regions. Deciduousness of canopy trees in the tropical moist forest area accounted for the differences in predictive equations for stem diameter and basal area. The relationships between lidar metrics and AGBM, however, remained significantly different between the two study areas even after adjusting for leaf drop. We attribute this to significant differences in the underlying allometric relationships between stem diameter and AGBM in tropical wet and moist forests. Conclusions Important forest structural characteristics can be estimated reliably across a variety of conditions sampled in these closed‐canopy tropical forests. Environmental factors such as drought deciduousness have an important influence on these relationships. Future efforts should continue to examine climatic factors that may influence the generality of the relationships between lidar metrics and forest structural characteristics and assess more rigorously the generality of field‐derived allometric relationships.     

Linking above-ground and below-ground effects in autotrophic microcosms: effects of shading and defoliation on plant and soil properties

Although factors affecting plant growth and plant carbon/nutrient balance – e.g., light availability and defoliation by herbivores – may also propagate changes in below‐ground food webs, few studies have aimed at linking the above‐ground and below‐ground effects. We established a 29‐week laboratory experiment (～one growing season) using autotrophic microcosms to study the effects of light and defoliation on plant growth, plant carbon/nutrient balance, soil inorganic N content, and microbial activity and biomass in soil. Each microcosm contained three substrate layers – mineral soil, humus and plant litter – and one Nothofagus solandri var. cliffortioides seedling. The experiment constituted of the presence or absence of two treatments in a full factorial design: shading (50% decrease in light) and artificial defoliation (approximately 50% decrease in leaf area in the beginning of the growing season). At the end of the experiment a range of above‐ground and below‐ground properties were measured. The shading treatment reduced root and shoot mass, root/shoot ratio and leaf production of the seedlings, while the defoliation treatment significantly decreased leaf mass only. Leaf C and N content were not affected by either treatment. Shading increased NO ₃–N concentration and decreased microbial biomass in humus, while defoliation did not significantly affect inorganic N or microbes in humus. The results show that plant responses to above‐ground treatments have effects which propagate below ground, and that rather straightforward mechanisms may link above‐ground and below‐ground effects. The shading treatment, which reduced overall seedling growth and thus below‐ground N use and C allocation, also led to changes in humus N content and microbial biomass, whereas defoliation, which did not affect overall growth, did not influence these below‐ground properties. The study also shows the carbon/nutrient balance of N. solandri var. cliffortioides seedlings to be highly invariant to both shading and defoliation.     

Pantropical dynamics of 'intact' rain forest canopy texture

Aside from deforestation and selective logging, mature tropical rain forests appear to be in a state of near‐term flux. At plot scales over a 20‐year period, forest inventory data (representing areas < 750 ha) have shown stands to be accumulating above‐ground biomass pantropically while displaying signs of increasing turnover. To assess whether or not such disequilibrium is manifest at broader spatial scales (i.e. coarser grain sizes and larger extents), we performed a change detection analysis of landscape texture (i.e. the organization of reflective properties), in satellite images of closed‐canopy tropical rain forests considered to be anthropically undisturbed. Here we show that fractal properties of pixel spectral values depicting low and high levels of photosynthetic activity underwent significant shifts from the 1970s to the 1980s. Following expectations for aggrading forests, canopy texture became more random throughout the tropics. Although subject to periodic disturbance events such as natural exogenic perturbations and/or synchronous die‐off, which should produce no consistent trend, these forested landscapes across the globe exhibited similar dynamics at fine temporal (decadal) intervals. Such biophysical changes (representing areas > 1000 000 ha) directly affect atmospheric boundary layer conditions and could have implications with respect to biodiversity and carbon cycling in these systems.     

Topography shapes the structure,composition and function of tropical forest landscapes

Topography is a key driver of tropical forest structure and composition, as it constrains local nutrient and hydraulic conditions within which trees grow. Yet, we do not fully understand how changes in forest physiognomy driven by topography impact other emergent properties of forests, such as their aboveground carbon density (ACD). Working in Borneo – at a site where 70‐m‐tall forests in alluvial valleys rapidly transition to stunted heath forests on nutrient‐depleted dip slopes – we combined field data with airborne laser scanning and hyperspectral imaging to characterise how topography shapes the vertical structure, wood density, diversity and ACD of nearly 15 km² of old‐growth forest. We found that subtle differences in elevation – which control soil chemistry and hydrology – profoundly influenced the structure, composition and diversity of the canopy. Capturing these processes was critical to explaining landscape‐scale heterogeneity in ACD, highlighting how emerging remote sensing technologies can provide new insights into long‐standing ecological questions.     

No evidence for consistent long‐term growth stimulation of 13 tropical tree species: results from tree‐ring analysis

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.     

Variation in potential for isoprene emissions among Neotropical forest sites

As part of the Large Scale Biosphere–Atmosphere Experiment in Amazônia (LBA), we have developed a bottom‐up approach for estimating canopy‐scale fluxes of isoprene. Estimating isoprene fluxes for a given forest ecosystem requires knowledge of foliar biomass, segregated by species, and the isoprene emission characteristics of the individual tree species comprising the forest. In this study, approximately 38% of 125 tree species examined at six sites in the Brazilian Amazon emitted isoprene. Given logistical difficulties and extremely high species diversity, it was possible to screen only a small percentage of tree species, and we propose a protocol for estimating the emission capacity of unmeasured taxa using a taxonomic approach, in which we assign to an unmeasured genus a value based on the percentage of genera within its plant family which have been shown to emit isoprene. Combining this information with data obtained from 14 tree censuses at four Neotropical forest sites, we have estimated the percentage of isoprene‐emitting biomass at each site. The relative contribution of each genus of tree is estimated as the basal area of all trees of that genus divided by the total basal area of the plot. Using this technique, the percentage of isoprene‐emitting biomass varied from 20% to 42% (mean=31%; SD=8%). Responses of isoprene emission to varying light and temperature, measured on a sun‐adapted leaf of mango (Mangifera indica L.), suggest that existing algorithms developed for temperate species are adequate for tropical species as well. Incorporating these algorithms, estimates of isoprene‐emitting biomass, isoprene emission capacity, and site foliar biomass into a canopy flux model, canopy‐scale fluxes of isoprene were predicted and compared with the above‐canopy fluxes measured at two sites. Our bottom‐up approach overestimates fluxes by about 50%, but variations in measured fluxes between the two sites are largely explained by observed variation in the amount of isoprene‐emitting biomass.     

Forest biomass,productivity and carbon cycling along a rainfall gradient in West Africa

Net Primary Productivity (NPP) is one of the most important parameters in describing the functioning of any ecosystem and yet it arguably remains a poorly quantified and understood component of carbon cycling in tropical forests, especially outside of the Americas. We provide the first comprehensive analysis of NPP and its carbon allocation to woody, canopy and root growth components at contrasting lowland West African forests spanning a rainfall gradient. Using a standardized methodology to study evergreen (EF), semi‐deciduous (SDF), dry forests (DF) and woody savanna (WS), we find that (i) climate is more closely related with above and belowground C stocks than with NPP (ii) total NPP is highest in the SDF site, then the EF followed by the DF and WS and that (iii) different forest types have distinct carbon allocation patterns whereby SDF allocate in excess of 50% to canopy production and the DF and WS sites allocate 40%–50% to woody production. Furthermore, we find that (iv) compared with canopy and root growth rates the woody growth rate of these forests is a poor proxy for their overall productivity and that (v) residence time is the primary driver in the productivity‐allocation‐turnover chain for the observed spatial differences in woody, leaf and root biomass across the rainfall gradient. Through a systematic assessment of forest productivity we demonstrate the importance of directly measuring the main components of above and belowground NPP and encourage the establishment of more permanent carbon intensive monitoring plots across the tropics.     

Developing a Multilayered Integrated Numerical Model of Surface Physics – Growing Plants Interaction (MINoSGI)

This paper describes a Multilayered Integrated Numerical Model of Surface Physics – Growing Plants Interaction (MINoSGI), which represents interactions between the dynamics of forest ecosystems and microclimate. Aiming at a large‐scale study in the future, we describe forest dynamics by using area‐averaged prognostic equations for thedistributions of plant density and plant weight with respect to plant height classes, instead of individual‐based treatments for small‐scale forest patches. Growth and mortality of plants are modelled based on the carbon balance of each plant height class. The area‐averaged microclimate (e.g., light, wind speed, temperature, humidity, CO₂ concentration) within the forest canopy is simulated by a one‐dimensional multilayer canopy model, which includes most of the physical and physiological processes that control the forest microclimate. Owing to its multilayered framework, a direct specification is possible for the difference in the growing environment among plants of different size and species. Given hourly meteorological conditions, the model outputs energy, water, CO₂ and momentum fluxes to and from a forest, of which the structure changes through competition among plants. The model's performance was tested by comparing its outputs with observed data on the development of plant size distribution taken over a 5‐year period in an evergreen coniferous (Cryptomeria japonica) forest. The model produced realistic estimates of the total biomass increments during the period. The ratio of net primary production to gross primary production (=0.45) was consistent with previous estimates for temperate forests. The bimodal seasonal pattern in net ecosystem production was similar to the seasonal trend in the CO₂ flux measured over a forest of the same species. Although some limitations due to the one‐dimensional representation of microclimate were noticeable, the model adequately simulated distributions of annual growth rate, plant weight and diameter across plant height classes. Since the basic equations can be extended to include the effect of spatial variability with marginal increase of computational costs, the present model framework is feasible for large‐scale studies.