Development of TRIPLEX-Management model for simulating the response of forest growth to pre-commercial thinning

In order to simulate forest growth response to pre-commercial thinning (PCT), TRIPLEX1.0 - a process-based model designed to predict forest growth as well as carbon (C) and nitrogen (N) dynamics - was modified and improved to also simulate managed forest ecosystem thinning practices. A three-parameter Weibull distribution model was integrated to simulate thinning treatments within the newly developed TRIPLEX-Management model. The thinning intensity component within the model allows users to simulate thinning treatments by applying basal area, stand density and volume to quantify thinning intensity. Natural mortality decreased following thinning due to an increase in growing space for residual stems. Predicted litterfall pools also increased after thinning events took place. The TRIPLEX-Management model was tested against published observational data for Jack Pine (Pinus banksiana Lamb.) stands subjected to PCT in Northwestern Ontario, Canada. The coefficients of determination (R²) between the predicted and observed variables including stand density, mean DBH (diameter at breast height), the quadratic mean DBH, total volume and merchantable volume as well as belowground, aboveground, and total biomass ranged from 0.50 to 0.88 (n = 20, P < 0.001) with the exception of mean tree height (R² = 0.25, n = 20, P < 0.05). Overall, the Willmott index of agreement between predicted and observed variables ranged from 0.97 to 1.00. Results show that the TRIPLEX-Management model is generally capable of simulating growth response to PCT for Jack Pine stands.     

Growth response of Lebanon cedar (Cedrus libani) plantations to thinning intensity in Western Turkey

This paper presents the growth response of 25 yr old Lebanon cedar (Cedrus libani A. Rich.) plantation to thinnings of different intensities in Isparta in western Turkey. The thinning intensity was measured by using the residual basal area (%) as parameter. In spring of 2005, three treatments were tested; light, moderate and heavy thinning with respectively 10, 25 and 35% of basal area removed. The statistical design of the experiment was a randomized incomplete block with two blocks and three treatments. Variables such as diameter at breast height (diameter) and height were measured. Growth rate ratios of diameter in moderately thinned and heavily thinned stands were 1.02 and 1.03, respectively. Basal area growth rates in moderately thinned and heavily thinned plots were 0.93 and 1.05, respectively. The largest values for the mean tree were observed with the heaviest thinning treatment. Absolute diameter increment was positively correlated with initial diameter in all plots. Relative diameter growth was negatively correlated with initial diameter. Growth rate interpretations were supported by analysis of variance using Duncan's test of range multiple. The results obtained show significant differences between treatments for tree height growth, for the two inventories carried out (2005, 2008). However diameter basal area and volume were no found between treatments for tree.     

Dynamic growth model for Scots pine (Pinus sylvestris L.) plantations in Galicia (north-western Spain)

In this study we developed a dynamic growth model for Scots pine (Pinus sylvestris L.) plantations in Galicia (north-western Spain). The data used to develop the model were obtained from a network of permanent plots, of between 10 and 55-year-old, which the Unidade de Xestión Forestal Sostible (Sustainable Forest Management Unit) of the University of Santiago de Compostela has set up in pure plantations of this species of pine in its area of distribution in Galicia. In this model, the initial stand conditions at any point in time are defined by three state variables (number of trees per hectare, stand basal area and dominant height), and are used to estimate stand volume, classified by commercial classes, for a given projection age. The model uses three transition functions expressed as algebraic difference equations of the three corresponding state variables used to project the stand state at any point in time. In addition, the model incorporates a function for predicting initial stand basal area, which can be used to establish the starting point for the simulation. This alternative should only be used when the stand is not yet established or when no inventory data are available. Once the state variables are known for a specific moment, a distribution function is used to estimate the number of trees in each diameter class, by recovering the parameters of the Weibull function, using the moments of first and second order of the distribution (arithmetic mean diameter and variance, respectively). By using a generalized height–diameter function to estimate the height of the average tree in each diameter class, combined with a taper function that uses the above predicted diameter and height, it is then possible to estimate total or merchantable stand volume.     

Validation of a multispecies forest dynamics model using 50-year growth from Eucalyptus forests in eastern Australia

One of the key problems confronting ecological forecasting is the validation of computer models. Here we report successful validation of a forest dynamics model Ecosystem Dynamics Simulator (EDS), adapted from the JABOWA-II forest succession model. This model and many variants derived from it have successfully simulated growth dynamics of uneven-aged mixed forests under changing environment with a moderate amount of input data. But rarely are adequate time-series data available for quantitative model validation. This study tested the performance of EDS in projecting the tree density, tree diameter at breast height (dbh), tree height, basal area and aboveground biomass of uneven-aged, mixed species sclerophyll forests in St. Mary state forests of eastern Australia. The test data were collected between 1951 and 2005. Every tree was uniquely numbered, tagged and measured in consecutive re-measurements. Projected growth attributes were compared with those observed in an independent validation dataset. The model produced satisfactory projections of tree density (91.7%), dbh (92.3%), total tree height (82.8%), basal area (89.3%) and aboveground biomass (87.6%) compared to the observed attributes. These results suggest that the EDS model can provide reasonable capability in projecting growth dynamics of uneven-aged, mixed species sclerophyll forests.     

Response of tree growth to a changing climate in boreal central Canada: A comparison of empirical,process-based,and hybrid modelling approaches

The impact of 2 × CO₂ driven climate change on radial growth of boreal tree species Pinus banksiana Lamb., Populus tremuloides Michx. and Picea mariana (Mill.) BSP growing in the Duck Mountain Provincial Forest of Manitoba (DMPF), Canada, is simulated using empirical and process-based model approaches. First, empirical relationships between growth and climate are developed. Stepwise multiple-regression models are conducted between tree-ring growth increments (TRGI) and monthly drought, precipitation and temperature series. Predictive skills are tested using a calibration–verification scheme. The established relationships are then transferred to climates driven by 1× and 2 × CO₂ scenarios using outputs from the Canadian second-generation coupled global climate model. Second, empirical results are contrasted with process-based projections of net primary productivity allocated to stem development (NPP_s). At the finest scale, a leaf-level model of photosynthesis is used to simulate canopy properties per species and their interaction with the variability in radiation, temperature and vapour pressure deficit. Then, a top-down plot-level model of forest productivity is used to simulate landscape-level productivity by capturing the between-stand variability in forest cover. Results show that the predicted TRGI from the empirical models account for up to 56.3% of the variance in the observed TRGI over the period 1912–1999. Under a 2 × CO₂ scenario, the predicted impact of climate change is a radial growth decline for all three species under study. However, projections obtained from the process-based model suggest that an increasing growing season length in a changing climate could counteract and potentially overwhelm the negative influence of increased drought stress. The divergence between TRGI and NPP_s simulations likely resulted, among others, from assumptions about soil water holding capacity and from calibration of variables affecting gross primary productivity. An attempt was therefore made to bridge the gap between the two modelling approaches by using physiological variables as TRGI predictors. Results obtained in this manner are similar to those obtained using climate variables, and suggest that the positive effect of increasing growing season length would be counteracted by increasing summer temperatures. Notwithstanding uncertainties in these simulations (CO₂ fertilization effect, feedback from disturbance regimes, phenology of species, and uncertainties in future CO₂ emissions), a decrease in forest productivity with climate change should be considered as a plausible scenario in sustainable forest management planning of the DMPF.     

Modelling the response of tree growth to temperature and CO2 elevation as related to the fertility and current temperature sum of a site

A methodology for simulating climate change impacts on tree growth was introduced into a statistical growth and yield model in relation to variations in site fertility and location implemented with current temperature sum. This was based on a procedure in which the relative enhancement in stem volume growth was calculated from short-term runs of a physiological simulation model for Scots pine (Pinus sylvestris L.), Norway spruce (Picea abies (L.) Karst.) and silver birch (Betula pendula Roth.) stands. These simulations were made for a set of stands with species-specific variations in stand characteristics, location and fertility type first in current climatic conditions and then in different combinations of CO₂ and temperature elevations. Based on these simulations, the relative enhancement of volume growth induced by the climate change (relative scenario effect, RSEv) was calculated and modelled in relation to: (i) CO₂ and temperature elevation, stand density and the competition status of the tree in its stand, and (ii) variations in site fertility type and current temperature sum of a stand. Finally, these transfer functions for RSEv were applied to adapt the stem volume growth in the statistical growth and yield model to reflect the response to climate change.     

3 forest simulation model

The 3 forest simulation model is a process model of tree growth, carbon and nitrogen dynamics in a single-species, even-aged forest stand. It is based on the model. Major changes include the computation of sun angle and radiation as a function of latitude and day of the year, the closed-form integration of canopy production as a function of day and hour, the introduction of tree number, height, and diameter as separate state variables, and different growth strategies, mortalities, and resulting self-thinning as function of crowding competition.The tree/soil system is described by a set of nonlinear ordinary differential equations for the state variables: tree number, base diameter, tree height, wood biomass, nitrogen in wood, leaf mass, fine root mass, fruit biomass, assimilate, carbon and nitrogen in litter, carbon and nitrogen in soil organic matter, and plant-available nitrogen. The model includes explicit formulations of all relevant ecophysiological processes such as: computation of radiation as a function of seasonal time, daytime and cloudiness, light attenuation in the canopy, and canopy photosynthesis as function of latitude, seasonal time, and daytime, respiration of all parts, assimilate allocation, increment formation, nitrogen fixation, mineralization, humification and leaching, forest management (thinning, felling, litter removal, fertilization etc.), temperature effects on respiration and decomposition, and environmental effects (pollution damage to photosynthesis, leaves, and fine roots). Only ecophysiological parameters which can be either directly measured or estimated with reasonable certainty are used. 3 is a generic process model which requires species- and site-specific parametrization. It can be applied to deciduous and coniferous forests under tropical, as well as temperate or boreal conditions.The paper presents a full documentation of the mathematical model as well as representative simulation results for spruce and acacia.     

treedyn3 forest simulation model

The treedyn3 forest simulation model is a process model of tree growth, carbon and nitrogen dynamics in a single-species, even-aged forest stand. It is based on the treedyn model. Major changes include the computation of sun angle and radiation as a function of latitude and day of the year, the closed-form integration of canopy production as a function of day and hour, the introduction of tree number, height, and diameter as separate state variables, and different growth strategies, mortalities, and resulting self-thinning as function of crowding competition.The tree/soil system is described by a set of nonlinear ordinary differential equations for the state variables: tree number, base diameter, tree height, wood biomass, nitrogen in wood, leaf mass, fine root mass, fruit biomass, assimilate, carbon and nitrogen in litter, carbon and nitrogen in soil organic matter, and plant-available nitrogen. The model includes explicit formulations of all relevant ecophysiological processes such as: computation of radiation as a function of seasonal time, daytime and cloudiness, light attenuation in the canopy, and canopy photosynthesis as function of latitude, seasonal time, and daytime, respiration of all parts, assimilate allocation, increment formation, nitrogen fixation, mineralization, humification and leaching, forest management (thinning, felling, litter removal, fertilization etc.), temperature effects on respiration and decomposition, and environmental effects (pollution damage to photosynthesis, leaves, and fine roots). Only ecophysiological parameters which can be either directly measured or estimated with reasonable certainty are used. treedyn3 is a generic process model which requires species- and site-specific parametrization. It can be applied to deciduous and coniferous forests under tropical, as well as temperate or boreal conditions.The paper presents a full documentation of the mathematical model as well as representative simulation results for spruce and acacia.     

Introducing effects of temperature and CO2 elevation on tree growth into a statistical growth and yield model

Impacts of elevated temperature and CO₂ on tree growth were introduced into a statistical growth and yield model for Finnish conditions based on corresponding predictions obtained from a physiological growth model. This one-way link between models was made by means of species-specific transfer functions describing the increase in stem volume growth of trees as a function of elevated temperature and CO₂, stand density and the tree's competition status in a stand of Scots pine (Pinus sylvestris), silver birch (Betula pendula) and Norway spruce (Picea abies). This method allows the inner dynamics of the statistical model to be followed when the impacts of temperature and CO₂ elevation on tree growth are introduced into the calculation of volume growth and further allocated between diameter and height growth. In this way compatibility with previous predictions of tree growth by means of statistical models and related model systems under current climatic conditions could be retained.The performance of the statistical model with species-specific transfer functions was evaluated by comparing its predictions with corresponding predictions given by a physiological model under conditions of elevated temperature and CO₂. These calculations revealed that the growth response of individual trees to elevated temperature and CO₂ can be introduced into the statistical model from a physiological growth model with an outcome that results in fairly satisfactory growth responses at the stand level as well.     

Combining a generic process-based productivity model and a statistical classification method to predict the presence and absence of tree species in the Pacific Northwest,U.S.A.

Although long-lived tree species experience considerable environmental variation over their life spans, their geographical distributions reflect sensitivity mainly to mean monthly climatic conditions. We introduce an approach that incorporates a physiologically based growth model to illustrate how a half-dozen tree species differ in their responses to monthly variation in four climatic-related variables: water availability, deviations from an optimum temperature, atmospheric humidity deficits, and the frequency of frost. Rather than use climatic data directly to correlate with a species’ distribution, we assess the relative constraints of each of the four variables as they affect predicted monthly photosynthesis for Douglas-fir, the most widely distributed species in the region. We apply an automated regression-tree analysis to create a suite of rules, which differentially rank the relative importance of the four climatic modifiers for each species, and provide a basis for predicting a species’ presence or absence on 3737 uniformly distributed U.S. Forest Services’ Forest Inventory and Analysis (FIA) field survey plots. Results of this generalized rule-based approach were encouraging, with weighted accuracy, which combines the correct prediction of both presence and absence on FIA survey plots, averaging 87%. A wider sampling of climatic conditions throughout the full range of a species’ distribution should improve the basis for creating rules and the possibility of predicting future shifts in the geographic distribution of species.     

Modelling coupled peach tree-aphid population dynamics and their control by winter pruning and nitrogen fertilization

Nitrogen fertilization and winter pruning are commonly used to control crop production in peach [Prunus persica (L.) Batsch] orchards. They are also known to affect the dynamics of Myzus persicae (Sulzer) (Homoptera: Aphididae) aphid populations via bottom-up regulation processes. Interactions between crops and pests can cause complex system behaviour in response to management practices. An integrated approach will therefore improve the understanding of the effects of these two cultural practices on aphid and peach performances.We developed a simulation model that describes the cultural control of interacting peach tree and aphid population dynamics. It uses the principles of common trophic models while gathering available knowledge and explicit assumptions on peach and aphid functioning and the effects of cultural practices.The model was able to qualitatively reproduce the system behaviour observed in the field. It accounted for actions and feedback such as stimulation of foliar growth by winter pruning, consecutive aphid population increase, subsequent damage to foliage, and partial compensatory growth of foliage. The model also reproduced low losses in fruit production due to aphid infestations. However, it called for further integration of ‘long-term’ effects. Analysis of the model showed the complexity of peach tree and aphid responses to leaf N × winter pruning interactions. Simulations indicated that fruit production losses remained low within a range of realistic values of leaf N and pruning intensity, whereas manipulating peach and aphid dynamics, their interactions and their relationships to practices could result in higher losses.The model is useful to evaluate the relevance of cultural practices for a bottom-up regulation of aphid dynamics in crop-pest management. After considering other control methods and fruit quality, it can be used to find a combination of practices that optimises trade-offs between fruit production and environmental conservation goals. A modelling approach that links crop growth and pest population dynamics and integrates management practice effects has strong potential for improving crop-pest management in an integrated crop production context.     

Differential use of shelter holes by sympatric species of blennies (Blennidae)

Comparative use of shelter use by three sympatric species of combtooth blenny (Ecsenius stictus, Glyptoparus delicatulus, and Salarias patzneri) was studied among micro-atolls in the lagoon at Lizard Island (14°42′S, 145°30′E), northern Great Barrier Reef, Australia. Blenny species used different sized holes; however, the average diameter and depth of holes used by the smallest and largest species differed by only 4 and 25 mm, respectively, indicating interspecific differences in suitable refuge can be very subtle. Both hole diameter and depth were positively related to total length of fish, suggesting use of holes relates to interspecific differences in body size. Total abundance of blennies was best explained by a general linear model that included either the number of holes or total habitat area on individual micro-atolls, predictor variables that were positively correlated with each other. However, the relative importance of variables differed among the three species, feeding area being most important for S. patzneri, feeding area and number of holes for E. stictus, and variance in hole diameter being the best explanatory variable for G. delicatulus abundance. The number of blenny species on a micro-atoll was best explained by variance in hole diameter, emphasizing the influence of refuge size variety in fish diversity. It is likely that subtle habitat partitioning, which relates to interspecific differences in body size, contributes to the co-existence of blenny species within the same microhabitat, but presence of holes is unlikely to regulate abundance of these fish.     

Calibration of the forest vegetation simulator (FVS) model for the main forest species of Ontario,Canada

The forest vegetation simulator (FVS) model was calibrated for use in Ontario, Canada, to predict the growth of forest stands. Using data from permanent sample plots originating from different regions of Ontario, new models were derived for dbh growth rate, survival rate, stem height and species group density index for large trees and height and dbh growth rate for small trees. The dataset included black spruce (Picea mariana (Mill.) B.S.P.) and jack pine (Pinus banksiana Lamb.) for the boreal region, sugar maple (Acer saccharum Marsh.), white pine (Pinus strobus L.), red pine (Pinus resinosa Ait.) and yellow birch (Betula alleghaniensis Britton) for the Great Lakes-St. Lawrence region, and balsam fir (Abies balsamea (L.) Mill.) and trembling aspen (Populus tremuloides Michx.) for both regions. These new models were validated against an independent dataset that consisted of permanent sample plots located in Quebec. The new models predicted biologically consistent growth patterns whereas some of the original models from the Lake States version of FVS occasionally did not. The new models also fitted the calibration (Ontario) data better than the original FVS models. The validation against independent data from Quebec showed that the new models generally had a lower prediction error than the original FVS models.     

Carbon sequestration potential of Rhizophora mucronata and Avicennia marina as influenced by age, season, growth and sediment characteristics in southeast coast of India

This work analysed the carbon sequestration potential in two species of mangroves (Rhizophora mucronata and Avicennia marina) along with their growth, biomass, sediment characteristics for four seasons of the year 2009–2010, in planted stands of different age (1–17.5 years) in the Vellar-Coleroon estuarine complex, India. The mangroves were recorded to store significant amount of biomass. Avicennia marina performed better to display 75 % higher rate of carbon sequestration than that in Rhizophora mucronata. This could be attributed to growth efficiency and high biomass production. For instance, Avicennia marina exhibited 2.7 fold higher girth, 24 % higher net canopy photosynthesis, 2 fold aboveground biomass (AGB), 40 % more belowground biomass (BGB) and 77.3 % higher total biomass, than R. mucronata did. Seasonally the rate of carbon sequestration was 7.3 fold higher in post-monsoon, 3.4 fold in monsoon, 73 % more in summer than that in pre-monsoon. The rate of carbon sequestration was positively correlated with age of planted site, tree height, tree diameter, net canopy photosynthesis, AGB, BGB, total biomass, carbon stock, growth efficiency, AGB/tree height tree girth, leaf area index, silt content (p?<?0.01). The carbon sequestration was negatively corrected with soil temperature and clay content (p?<?0.05). Mangroves were found to be a productive system and important sink of carbon in the tropical coastal zone, but increasing soil temperature due to global warming would have a negative impact on carbon sequestration potential of the mangroves.     

Response of plant species to coal-mine soil materials

A two-year experiment was conducted on the Black Mesa Coal Mine near Kayenta, Arizona to investigate the growth and establishment of seven plant species in unmined soil (undisturbed soil) and coal-mine soil (spoils). Natural rainfall (20 cm/yr) and natural rainfull plus sprinkler irrigation (50 cm/yr) were the irrigation treatments applied to each soil material. Plant species grew better in unmined soil than in coal-mine soil. Supplemental irrigation water resulted in more plant growth than did natural rainfall alone in both soil materials; however, there were highly significant differences among species. Alfalfa (Medicago sativa L.) and all of the annual perennial grasses used, excluding Indian ricegrass (Oryzopsis hymenoides Ricker), produced effective ground cover on both soil materials when they received supplemental irrigation water. Fourwing saltbush (Atriplex canescens Pursh.) had low germination (emergence), seedling establishment, and stem production during the first year of growth; however, in the second year of growth, this species produced a dense ground cover on coal-mine soil with natural rainfall plus irrigation. In revegetating the Black Mesa Coal Mine, Harlan II barley (Hordeum vulgare L.) and Super X wheat (Triticum aestivum L.) provided initial protective cover until adapted perennial species could be established for permanent stabilisation. The Black Mesa Research Study indicated that irrigation water during seedling establishment was essential for the effective stabilisation of coal-mine soils in a semiarid environment.     

Efficient thinning regimes for Eucalyptus fastigata: Multi-objective stand-level optimisation using the island model genetic algorithm

A stand-level optimisation problem formulated to determine a set of efficient thinning regimes satisfying two objectives, i.e. value production for sawlog harvesting and volume production for a pulpwood market, was demonstrated for a Eucalyptus fastigata trial in Kaingaroa Forest, New Zealand. Genetic algorithms were used to estimate the set of efficient thinning regimes (i.e. regimes that occur when it is not possible to increase the achievement of one objective without reducing another) known as a Pareto frontier. Each thinning regime specified a unique combination of initial planting density; frequency, timing and intensity of thinning; final crop number; and rotation length. Specifications for the “best” regime in the Pareto set (i.e. the one that satisfied a balanced trade-off between value and volume production) were similar to those recommended through professional judgment based on pooled long-term field observations from different eucalypt species grown throughout New Zealand. The advantage of Pareto optimality was the ability of not only identifying a unique thinning regime, but equally efficient regimes each providing a different combination of value and volume production. Research on this approach has the potential of being applied to other forest sites, providing there is sufficient re-measurement data to reflect stand growth dynamics.     

Simulation of interannual responses of trembling aspen stands to climatic variation and insect defoliation in western Canada

A carbon-based model has been developed to simulate responses of trembling aspen (Populus tremuloides Michx.) stands to interannual climatic variation and insect defoliation. The model is designed for medium time scale (10–100 years) simulations and requires only daily maximum and minimum temperature and precipitation as meteorological inputs. The modelling approach is similar to FOREST-BGC but includes additional processes known to be important in deciduous forests. These include removal of leaf area during outbreaks of forest tent caterpillar (Malacosoma disstria Hbn.), phenological changes in leaf area index, storage and allocation of non-structural carbohydrate and the contribution of understorey vegetation to evapotranspiration. The model was used for simulations of growth and mortality of biomass carbon in two mature aspen forests located in the climatically dry transition zone between the boreal forest and prairie grassland regions of Saskatchewan, Canada. Model inputs of annual defoliation intensity were based on historic records of insect defoliation and the incidence of light-coloured tree rings in disks or cores collected from aspen at each of the two sites. At both sites, moderately good correlations (r²=0.47–0.54) were obtained between modelled interannual changes in stem carbon growth and observed interannual changes in stem basal area increment obtained from tree-ring analysis. Model outputs of stem biomass carbon were found to be highly sensitive to parameters describing seasonal leaf area duration, insect defoliation intensity, photosynthesis and root respiration and carbohydrate allocation to growth versus storage.     

Incorporating Lindenmayer systems for architectural development in a functional-structural tree model

LIGNUM is a functional-structural tree model that represents coniferous and broad-leaved trees with modelling units corresponding to the real structure of trees. The units are tree segments, axes, branching points and buds. Metabolic processes are explicitly related to the structural units in which they take place.This paper enhances the modelling capabilities of LIGNUM with the possibility to formally describe the architectural development of trees with Lindenmayer systems. This is achieved by presenting an algorithm to convert tree structures generated by Lindenmayer systems to the LIGNUM representation of trees with feedback of results of events or processes from LIGNUM to Lindenmayer system. We then give two example applications that model the development of Scots pine (Pinus sylvestris L.) and the dwarf shrub bearberry (Arctostaphylos uva-ursi L.). Finally we discuss our approach and its consequences for the future development of LIGNUM.     

More asymmetric tree competition brings about more evapotranspiration and less runoff from the forest ecosystems: A simulation study

The present paper reports how stand size-structure dynamics due to competition between different-sized trees affect long-term forested water balance in Japanese cool-temperate planted stands (evergreen coniferous Cryptomeria japonica and deciduous coniferous Larix kaempferi stands) using a fully coupled multi-layered meteorological surface physics—terrestrial ecosystems model. The simulation captured the well-known annual variation in leaf area index (LAI) accurately with stand age in monocultured and even-aged stands; the occurrence of maximum LAI during the early growth stage and then a gradual decline followed by a steady state after the maximum LAI. The simulations also detected a high dependency of annual evapotranspiration (AETr) on LAI with stand age that is well known by prior observational researches. In the C. japonica (shade-tolerant late-successional species) stand, the relationship between annual net primary productivity of an individual tree (NPP_ind) and individual tree mass (w) changed from linear to a convex curve during self-thinning, indicating that the degree of asymmetric tree competition intensified with forest stand development. The higher degree of competitive asymmetry characterized by the convex-shaped NPPind-w relationship produced greater size inequality, i.e., the formation of trees stratified by height. Under such conditions, AETr and annual transpiration (ATr) were mainly regulated by larger trees. On the other hand, the NPPind-w relationships in the L. kaempferi (shade-intolerant early-successional species) stand were linear throughout the simulated period, indicating the lower degree of competitive asymmetry. Under such conditions, the growth of intermediate-sized trees was enhanced and these trees became a dominant source of AETr (and also ATr) during self-thinning. Furthermore, the sensitivity analysis of the effects of ecophysiological parameters such as foliage profile (i.e., vertical distribution of leaf area density) of an individual tree (distribution pattern is described by the parameter η), the maximum carboxylation velocity (V_cmax0) and biomass allocation pattern of individual plant growth (μ₁) on AETr, ATr and annual runoff (AR_off) showed that the temporal trends of AETr, ATr, AR_off and NPPind-w relationships were completely the same as those in the control simulations. However, the NPPind-w relationship during self-thinning indicated higher degrees of competitive asymmetry when η or V_cmax0 were greater than those in the control simulation and generated greater AETr and ATr and thus smaller AR_off. We found that more asymmetric tree competition brings about greater size inequality between different-sized trees and thus more evapotranspiration and less runoff in a forest stand. Overall, our simulation approach revealed that not only LAI dynamics but also plant competition, and thus size-structure dynamics, in a forest ecosystem are essential to long-term future projections of forested water balance.