Are absorptive root traits good predictors of ecosystem functioning? A test in a natural temperate forest.

Trait-based approaches provide a useful framework to predict ecosystem functions under intensifying global change. However, our current understanding of trait-functioning relationships mainly relies on aboveground traits. Belowground traits (e.g. absorptive root traits) are rarely studied although these traits are related to important plant functions. We analyzed four pairs of analogous leaf and absorptive root traits of woody plants in a temperate forest and examined how these traits are coordinated at the community-level, and to what extent the trait covariation depends on local-scale environmental conditions. We then quantified the contributions of leaf and absorptive root traits and the environmental conditions in determining two important forest ecosystem functions, aboveground carbon storage, and woody biomass productivity. The results showed that both morphological trait pairs and chemical trait pairs exhibited positive correlations at the community level. Absorptive root traits show a strong response to environmental conditions compared to leaf traits. We also found that absorptive root traits were better predictors of the two forest ecosystem functions than leaf traits and environmental conditions. Our study confirms the important role of belowground traits in modulating ecosystem functions and deepens our understanding of belowground responses to changing environmental conditions.

Latitudinal patterns of forest ecosystem stability across spatial scales as affected by biodiversity and environmental heterogeneity.

Our planet is facing a variety of serious threats from climate change that are unfolding unevenly across the globe. Uncovering the spatial patterns of ecosystem stability is important for predicting the responses of ecological processes and biodiversity patterns to climate change. However, the understanding of the latitudinal pattern of ecosystem stability across scales and of the underlying ecological drivers is still very limited. Accordingly, this study examines the latitudinal patterns of ecosystem stability at the local and regional spatial scale using a natural assembly of forest metacommunities that are distributed over a large temperate forest region, considering a range of potential environmental drivers. We found that the stability of regional communities (regional stability) and asynchronous dynamics among local communities (spatial asynchrony) both decreased with increasing latitude, whereas the stability of local communities (local stability) did not. We tested a series of hypotheses that potentially drive the spatial patterns of ecosystem stability, and found that although the ecological drivers of biodiversity, climatic history, resource conditions, climatic stability, and environmental heterogeneity varied with latitude, latitudinal patterns of ecosystem stability at multiple scales were affected by biodiversity and environmental heterogeneity. In particular, α diversity is positively associated with local stability, while ß diversity is positively associated with spatial asynchrony, although both relationships are weak. Our study provides the first evidence that latitudinal patterns of the temporal stability of naturally assembled forest metacommunities across scales are driven by biodiversity and environmental heterogeneity. Our findings suggest that the preservation of plant biodiversity within and between forest communities and the maintenance of heterogeneous landscapes can be crucial to buffer forest ecosystems at higher latitudes from the faster and more intense negative impacts of climate change in the future.

Evaluation of the Site Form as a Site Productive Indicator in Temperate Uneven-Aged Multispecies Forests in Durango, Mexico.

Even though the site index is a popular method for describing forest productivity, its use is limited in uneven-aged multispecies forests. Accordingly, the site form (SF) is an alternative measure of productivity to the site index based on the tree height-diameter relationship. Our study aims to evaluate SF as a measure of productivity in the temperate uneven-aged multispecies forests of Durango, Mexico, applying three methods to estimate SF: (i) as the mean height of dominant trees at a reference diameter (SFH-D); (ii) as the expected mean height of dominant trees at a reference mean diameter (SFMH-MD), and (iii) as the expected height at a reference diameter for a given site (SFh-dbh). We assess the effectiveness of the SF based on two hypotheses: (i) the SF correlates to the total volume production, and (ii) the SF is independent of stand density. The SFH-D and the SFh-dbh showed a high correlation with productivity. However, they also did so with density. Contrary to this, the SFMH-MD had a weak correlation with density and productivity. We conclude that the SF is a suitable approach to describe site quality. Nonetheless, its effectiveness as a site quality indicator may be affected according to the method used.

Co-limitation towards lower latitudes shapes global forest diversity gradients.

The latitudinal diversity gradient (LDG) is one of the most recognized global patterns of species richness exhibited across a wide range of taxa. Numerous hypotheses have been proposed in the past two centuries to explain LDG, but rigorous tests of the drivers of LDGs have been limited by a lack of high-quality global species richness data. Here we produce a high-resolution (0.025° × 0.025°) map of local tree species richness using a global forest inventory database with individual tree information and local biophysical characteristics from ~1.3 million sample plots. We then quantify drivers of local tree species richness patterns across latitudes. Generally, annual mean temperature was a dominant predictor of tree species richness, which is most consistent with the metabolic theory of biodiversity (MTB). However, MTB underestimated LDG in the tropics, where high species richness was also moderated by topographic, soil and anthropogenic factors operating at local scales. Given that local landscape variables operate synergistically with bioclimatic factors in shaping the global LDG pattern, we suggest that MTB be extended to account for co-limitation by subordinate drivers.

Assessing scale-dependent effects on Forest biomass productivity based on machine learning.

Estimating forest above-ground biomass (AGB) productivity constitutes one of the most fundamental topics in forest ecological research. Based on a 30-ha permanent field plot in Northeastern China, we modeled AGB productivity as output, and topography, species diversity, stand structure, and a stand density variable as input across a series of area scales using the Random Forest (RF) algorithm. As the grain size increased from 10 to 200 m, we found that the relative importance of explanatory variables that drove the variation of biomass productivity varied a lot, and the model accuracy was gradually improved. The minimum sampling area for biomass productivity modeling in this region was 140 × 140 m. Our study shows that the relationship of topography, species diversity, stand structure, and stand density variables with biomass productivity modeled using the RF algorithm changes when moving from scales typical of forest surveys (10 m) to larger scales (200 m) within a controlled methodology. These results should be of considerable interest to scientists concerned with forest assessment.

Decomposing Spatial ß-Diversity in the temperate forests of Northeastern China.

ß-Diversity, which describes the extent of change in species composition in a given region, has become a core issue in ecology in recent years. However, it is hard to understand the underlying mechanisms of ß-diversity by using indices that yield identical values under species replacement and nestedness pattern. Partitioning ß-diversity into turnover (caused by species replacement among plots) and nestedness components (caused by species loss or gain among plots) may provide improved understanding of the variation in species composition. Here, we collected presence-absence data of 456 one-tenth ha circular plots in the temperate forests of Northeastern China spanning a latitudinal range of 12° (41-53°N). We decomposed ß-diversity to assess the relative contribution of the turnover and nestedness components across latitudinal gradients. We used regression analysis to assess the relationship between spatial distance and ß-diversity. We applied variation partitioning to evaluate the importance of the measured environmental and spatial variables in explaining ß-diversity. We used the Tukey honest significant difference test to test the differences of ß-diversity along latitudinal gradients. Pearson correlations (r) and significance (p-value) were computed using the Mantel tests to verify the relationship between distance and ß-diversity. The ANOVA test was used to verify whether the variation of ß-diversity explained by the environment and distance was significant. Our results showed that (1) ß-diversity and the turnover component were higher at low latitudes (zones A and B) than at high latitudes (zones C and D), while there was no relationship between the nestedness component and latitude. (2) The turnover component was dominant. (3) The spatial distance explained more variation of ß-diversity than the measured environmental factors. Therefore, we conclude that ß-diversity is mainly a product of species turnover in our temperate forests, suggesting that different localities harbor different species. We find that decomposing ß-diversity into the turnover and nestedness components is a useful approach to explore the variation of community composition and their causes.

Scale-dependent effects of neighborhood biodiversity on individual tree productivity in a coniferous and broad-leaved mixed forest in China.

The relationship between biodiversity and productivity has stimulated an increasing body of research over the past decades, and this topic still occupies a central place in ecology. While most studies have focused on biomass production in quadrats or plots, few have investigated the scale-dependent relationship from an individual plant perspective. We present an analysis of the effects of biodiversity (species diversity and functional diversity) on individual tree growth with a data set of 16,060 growth records from a 30-ha temperate forest plot using spatially explicit individual tree-based methods. A significant relationship between species diversity and tree growth was found at the individual tree level in our study. The magnitude and direction of biodiversity effects varies with the spatial scale. We found positive effects of species diversity on tree growth at scales exceeding 9 m. Individual tree growth rates increased when there was a greater diversity of species in the neighborhood of the focal tree, which provides evidence of a niche complementarity effect. At small scales (3-5 m), species diversity had negative effects on tree growth, suggesting that competition is more prevalent than complementarity or facilitation in these close neighborhoods. The results also revealed many confounding factors which influence tree growth, such as elevation and available sun light. We conclude that the use of individual tree-based methods may lead to a better understanding of the biodiversity-productivity relationship in forest communities.

Assessing biotic and abiotic effects on forest productivity in three temperate forests.

It is well understood that biotic and abiotic variables influence forest productivity. However, in regard to temperate forests, the relative contributions of the aforementioned drivers to biomass demographic processes (i.e., the growth rates of the survivors and recruits) have not received a great deal of attention. Thus, this study focused on the identification of the relative influencing effects of biotic and abiotic variables in the demographic biomass processes of temperate forests.This study was conducted in the Changbai Mountain Nature Reserve, in northeastern China. Based on the observational data collected from three 5.2-hectare forest plots, the annual above-ground biomass (AGB) increment (productivity) of the surviving trees, recruits, and the total tree community (survivors + recruits) were estimated. Then, the changes in the forest productivity in response to biotic variables (including species diversity, structural diversity, and density variables) along with abiotic variables (including topographic and soil variables) were evaluated using linear mixed-effect models.This study determined that the biotic variables regulated the variabilities in productivity. Density variables were the most critical drivers of the annual AGB increments of the surviving trees and total tree community. Structural diversity enhanced the annual AGB increments of the recruits, but diminished the annual AGB increments of the surviving trees and the total tree community. Species diversity and abiotic variables did not have impacts on the productivity in the examined forest plots.The results highlighted the important roles of forest density and structural diversity in the biomass demographic processes of temperate forests. The surviving and recruit trees were found to respond differently to the biotic variables, which suggested that the asymmetric competition had shaped the productivity dynamics in forests. Therefore, the findings emphasized the need to consider the demographic processes of forest productivity to better understand the functions of forests.

Functional and phylogenetic diversity determine woody productivity in a temperate forest.

Understanding the relationships between biodiversity and ecosystem productivity has become a central issue in ecology and conservation biology studies, particularly when these relationships are connected with global climate change and species extinction. However, which facets of biodiversity (i.e. taxonomic, functional, and phylogenetic diversity) account most for variations in productivity are still not understood very well. This is especially true with regard to temperate forest ecosystems. In this study, we used a dataset from a stem-mapped permanent forest plot in northeastern China exploring the relationships between biodiversity and productivity at different spatial scales (20 × 20 m; 40 × 40 m; and 60 × 60 m). The influence of specific environmental conditions (topographic conditions) and stand maturity (expressed by initial stand volume and biomass) were taken into account using the multivariate approach known as structural equation models. The variable "Biodiversity" includes taxonomic (Shannon), functional (FDis), and phylogenetic diversity (PD). Biodiversity-productivity relationships varied with the spatial scales. At the scale of 20 × 20 m, PD and FDis significantly affected forest biomass productivity, while Shannon had only indirect effects. At the 40 × 40 m and 60 × 60 m scales, biodiversity and productivity were weakly correlated. The initial stand volume and biomass were the most important drivers of forest productivity. The local environmental conditions significantly influenced the stand volume, biomass, biodiversity, and productivity. The results highlight the scale dependency of the relationships between forest biodiversity and productivity. The positive role of biodiversity in facilitating forest productivity was confirmed at the smaller scales. Our findings emphasize the fundamental role of environmental conditions in determining forest ecosystem performances. The results of this study provide a better understanding of the underlying ecological processes that influence specific forest biodiversity and productivity relationships.

How beta diversity and the underlying causes vary with sampling scales in the Changbai mountain forests.

This study aims to establish a relationship between the sampling scale and tree species beta diversity temperate forests and to identify the underlying causes of beta diversity at different sampling scales. The data were obtained from three large observational study areas in the Changbai mountain region in northeastern China. All trees with a dbh ≥1 cm were stem-mapped and measured. The beta diversity was calculated for four different grain sizes, and the associated variances were partitioned into components explained by environmental and spatial variables to determine the contributions of environmental filtering and dispersal limitation to beta diversity. The results showed that both beta diversity and the causes of beta diversity were dependent on the sampling scale. Beta diversity decreased with increasing scales. The best-explained beta diversity variation was up to about 60% which was discovered in the secondary conifer and broad-leaved mixed forest (CBF) study area at the 40 × 40 m scale. The variation partitioning result indicated that environmental filtering showed greater effects at bigger grain sizes, while dispersal limitation was found to be more important at smaller grain sizes. What is more, the result showed an increasing explanatory ability of environmental effects with increasing sampling grains but no clearly trend of spatial effects. The study emphasized that the underlying causes of beta diversity variation may be quite different within the same region depending on varying sampling scales. Therefore, scale effects should be taken into account in future studies on beta diversity, which is critical in identifying different relative importance of spatial and environmental drivers on species composition variation.

Analysing taxonomic structures and local ecological processes in temperate forests in North Eastern China.

BACKGROUND: One of the core issues of forest community ecology is the exploration of how ecological processes affect community structure. The relative importance of different processes is still under debate. This study addresses four questions: (1) how is the taxonomic structure of a forest community affected by spatial scale? (2) does the taxonomic structure reveal effects of local processes such as environmental filtering, dispersal limitation or interspecific competition at a local scale? (3) does the effect of local processes on the taxonomic structure vary with the spatial scale? (4) does the analysis based on taxonomic structures provide similar insights when compared with the use of phylogenetic information? Based on the data collected in two large forest observational field studies, the taxonomic structures of the plant communities were analyzed at different sampling scales using taxonomic ratios (number of genera/number of species, number of families/number of species), and the relationship between the number of higher taxa and the number of species. Two random null models were used and the "standardized effect size" (SES) of taxonomic ratios was calculated, to assess possible differences between the observed and simulated taxonomic structures, which may be caused by specific ecological processes. We further applied a phylogeny-based method to compare results with those of the taxonomic approach. RESULTS: As expected, the taxonomic ratios decline with increasing grain size. The quantitative relationship between genera/families and species, described by a linearized power function, showed a good fit. With the exception of the family-species relationship in the Jiaohe study area, the exponents of the genus/family-species relationships did not show any scale dependent effects. The taxonomic ratios of the observed communities had significantly lower values than those of the simulated random community under the test of two null models at almost all scales. Null Model 2 which considered the spatial dispersion of species generated a taxonomic structure which proved to be more consistent with that in the observed community. As sampling sizes increased from 20 m × 20 m to 50 m × 50 m, the magnitudes of SESs of taxonomic ratios increased. Based on the phylogenetic analysis, we found that the Jiaohe plot was phylogenetically clustered at almost all scales. We detected significant phylogenetically overdispersion at the 20 m × 20 m and 30 m × 30 m scales in the Liangshui plot. CONCLUSIONS: The results suggest that the effect of abiotic filtering is greater than the effects of interspecific competition in shaping the local community at almost all scales. Local processes influence the taxonomic structures, but their combined effects vary with the spatial scale. The taxonomic approach provides similar insights as the phylogenetic approach, especially when we applied a more conservative null model. Analysing taxonomic structure may be a useful tool for communities where well-resolved phylogenetic data are not available.

Analysis of the Global Warming Potential of Biogenic CO2 Emission in Life Cycle Assessments.

Biomass is generally believed to be carbon neutral. However, recent studies have challenged the carbon neutrality hypothesis by introducing metric indicators to assess the global warming potential of biogenic CO2 (GWPbio). In this study we calculated the GWPbio factors using a forest growth model and radiative forcing effects with a time horizon of 100 years and applied the factors to five life cycle assessment (LCA) case studies of bioproducts. The forest carbon change was also accounted for in the LCA studies. GWPbio factors ranged from 0.13-0.32, indicating that biomass could be an attractive energy resource when compared with fossil fuels. As expected, short rotation and fast-growing biomass plantations produced low GWPbio. Long-lived wood products also allowed more regrowth of biomass to be accounted as absorption of the CO2 emission from biomass combustion. The LCA case studies showed that the total life cycle GHG emissions were closely related to GWPbio and energy conversion efficiency. By considering the GWPbio factors and the forest carbon change, the production of ethanol and bio-power appeared to have higher GHG emissions than petroleum-derived diesel at the highest GWPbio.

Stoichiometry patterns in the androdioecious Acer tegmentosum.

This study evaluates stoichiometry patterns in the androdioecious Acer tegmentosum, a species characterized by a rare reproductive system where males and hermaphrodites coexist. Altogether 31 hermaphrodites and 29 male plants were harvested and samples of leaves, current-year shoots, branches and coarse roots were analyzed to explore gender differences in biomass, C, N and P concentrations of these four components. The nitrogen to phosphorus relationship of each component was examined using SMA estimates. Males had significantly greater amounts of leaf and coarse root dry matter content than hermaphrodites. C, N and P stoichiometry differed significantly between genders, especially in the newly emerging vegetative components (leaves and shoots). Males had higher C/N and C/P ratios in current-year shoots and lower C/P ratios in leaves and branches. Hermaphrodites had higher N/P ratios in the leaves and branches. Males had higher rates of increase in leaf P content than hermaphrodites. This study suggests that stoichiometry patterns may be significantly affected by gender.

Effects of density dependence in a temperate forest in northeastern China.

Negative density dependence may cause reduced clustering among individuals of the same species, and evidence is accumulating that conspecific density-dependent self-thinning is an important mechanism regulating the spatial structure of plant populations. This study evaluates that specific density dependence in three very large observational studies representing three successional stages in a temperate forest in northeastern China. The methods include standard spatial point pattern analysis and a heterogeneous Poisson process as the null model to eliminate the effects of habitat heterogeneity. The results show that most of the species exhibit conspecific density-dependent self-thinning. In the early successional stage 11 of the 16 species, in the intermediate successional stage 18 of the 21 species and in the old growth stage all 21 species exhibited density dependence after removing the effects of habitat heterogeneity. The prevalence of density dependence thus varies among the three successional stages and exhibits an increase with increasing successional stage. The proportion of species showing density dependence varied depending on whether habitat heterogeneity was removed or not. Furthermore, the strength of density dependence is closely related with species abundance. Abundant species with high conspecific aggregation tend to exhibit greater density dependence than rare species.

Drivers of seedling survival in a temperate forest and their relative importance at three stages of succession.

Negative density dependence (NDD) and niche partitioning have been perceived as important mechanisms for the maintenance of species diversity. However, little is known about their relative contributions to seedling survival. We examined the effects of biotic and abiotic neighborhoods and the variations of biotic neighborhoods among species using survival data for 7503 seedlings belonging to 22 woody species over a period of 2 years in three different forest types, a half-mature forest (HF), a mature forest (MF), and an old-growth forest (OGF), each of these representing a specific successional stage in a temperate forest ecosystem in northeastern China. We found a convincing evidence for the existence of NDD in temperate forest ecosystems. The biotic and abiotic variables affecting seedlings survival change with successional stage, seedling size, and age. The strength of NDD for the smaller (<20 cm in height) and younger seedlings (1-2 years) as well as all seedlings combined varies significantly among species. We found no evidence that a community compensatory trend (CCT) existed in our study area. The results of this study demonstrate that the relative importance of NDD and habitat niche partitioning in driving seedling survival varies with seedling size and age and that the biotic and abiotic factors affecting seedlings survival change with successional stage.

Spatial characteristics of tree diameter distributions in a temperate old-growth forest.

This contribution identifies spatial characteristics of tree diameter in a temperate forest in north-eastern China, based on a fully censused observational study area covering 500×600 m. Mark correlation analysis with three null hypothesis models was used to determine departure from expectations at different neighborhood distances. Tree positions are clumped at all investigated scales in all 37 studied species, while the diameters of most species are spatially negatively correlated, especially at short distances. Interestingly, all three cases showing short-distance attraction of dbh marks are associated with light-demanding shrub species. The short-distance attraction of dbh marks indicates spatially aggregated cohorts of stems of similar size. The percentage of species showing significant dbh suppression peaked at a 4 m distance under the heterogeneous Poisson model. At scales exceeding the peak distance, the percentage of species showing significant dbh suppression decreases sharply with increasing distances. The evidence from this large observational study shows that some of the variation of the spatial characteristics of tree diameters is related variations of topography and soil chemistry. However, an obvious interpretation of this result is still lacking. Thus, removing competitors surrounding the target trees is an effective way to avoid neighboring competition effects reducing the growth of valuable target trees in forest management practice.

A case study of forest management planning using a new heuristic algorithm.

We present an approach to generate and evaluate different silvicultural development paths and to optimize the development of a Norway spruce stand, using a long-term planning horizon. To generate a silvicultural path, the maximum stand density was applied. At each thinning event, three possible thinning intensities (10, 20, 30% of the stem number per ha) were randomly chosen. A search algorithm known as modified Accelerated Simulated Annealing (mASA) was used to estimate the optimum combination of stand paths for a given forest as a whole. Production and economic management objectives were considered and then compared. The economic criterion was the Expected Stand Value (ESV) with a 4% discount rate. The generated data set of 38 Norway spruce stands (comprising a total of 123.8 ha) was used in the case study. The result with the best combination of paths was presented in a digitized forest map. Forest management simulation was performed using a specially developed computer program, for a planning horizon of 20 years. The mASA proved to be an effective search method for identifying optimum paths.