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.

Late-spring frost risk between 1959 and 2017 decreased in North America but increased in Europe and Asia.

Late-spring frosts (LSFs) affect the performance of plants and animals across the world's temperate and boreal zones, but despite their ecological and economic impact on agriculture and forestry, the geographic distribution and evolutionary impact of these frost events are poorly understood. Here, we analyze LSFs between 1959 and 2017 and the resistance strategies of Northern Hemisphere woody species to infer trees' adaptations for minimizing frost damage to their leaves and to forecast forest vulnerability under the ongoing changes in frost frequencies. Trait values on leaf-out and leaf-freezing resistance come from up to 1,500 temperate and boreal woody species cultivated in common gardens. We find that areas in which LSFs are common, such as eastern North America, harbor tree species with cautious (late-leafing) leaf-out strategies. Areas in which LSFs used to be unlikely, such as broad-leaved forests and shrublands in Europe and Asia, instead harbor opportunistic tree species (quickly reacting to warming air temperatures). LSFs in the latter regions are currently increasing, and given species' innate resistance strategies, we estimate that ∼35% of the European and ∼26% of the Asian temperate forest area, but only ∼10% of the North American, will experience increasing late-frost damage in the future. Our findings reveal region-specific changes in the spring-frost risk that can inform decision-making in land management, forestry, agriculture, and insurance policy.

Understory plant removal counteracts tree thinning effect on soil respiration in a temperate forest.

Elucidating the response mechanism of soil respiration (Rs) to silvicultural practices is pivotal to evaluating the effects of management practices on soil carbon cycling in planted forest ecosystems. However, as common management practices, how thinning, understory plant removal, and their interactions affect Rs and its autotrophic and heterotrophic components (Ra and Rh) remains unclear. Therefore, we investigated Rs, Ra and Rh by the trenching method from 2011 to 2015 in a Pinus tabuliformis plantation in northern China, subjecting to four treatments (intact control plots [CK], thinning [T], understory removal [UR], and thinning with understory removal [TUR]). Mean annual Rs was significantly increased by thinning (by 15.3%), whereas decreased by UR (by 17.4%), compared with CK. These variations in Rs were mainly attributed to changes in Ra. The increments of Ra were caused by the enhanced growth of fine root biomass after thinning. However, UR led to lower Ra compared with CK (p < .05), indicating that understory growth is inadequate to compensate for the decreased respiring root biomass induced by understory removal. Rs was unchanged between TUR and the intact control plot due to the opposite effects of thinning and UR on the Ra. Changes in Rh exhibited no significant differences among the treatments, partly because of the stable microbial biomass carbon (MBC) and forest floor mass (litter and fine woody debris). No interaction effect between thinning and understory removal was detected on Rs, Ra, and Rh. The lowest temperature sensitivity (Q10 ) value of Ra was found in CK. This study highlights the necessity of incorporating understory plant effects on soil CO2 efflux in assessing forest management practices on soil carbon cycling.

New forest biomass carbon stock estimates in Northeast Asia based on multisource data.

Forests play an important role in both regional and global C cycles. However, the spatial patterns of biomass C density and underlying factors in Northeast Asia remain unclear. Here, we characterized spatial patterns and important drivers of biomass C density for Northeast Asia, based on multisource data from in situ forest inventories, as well as remote sensing, bioclimatic, topographic, and human footprint data. We derived, for the first time, high-resolution (1 km × 1 km) maps of the current and future forest biomass C density for this region. Based on these maps, we estimated that current biomass C stock in northeastern China, the Democratic People's Republic of Korea, and Republic of Korea to be 2.53, 0.40, and 0.35 Pg C, respectively. Biomass C stock in Northeast Asia has increased by 20%-46% over the past 20 years, of which 40%-76% was contributed by planted forests. We estimated the biomass C stock in 2080 to be 6.13 and 6.50 Pg C under RCP4.5 and RCP8.5 scenarios, respectively, which exceeded the present region-wide C stock value by 2.85-3.22 Pg C, and were 8%-14% higher than the baseline C stock value (5.70 Pg C). The spatial patterns of biomass C densities were found to vary greatly across the Northeast Asia, and largely decided by mean diameter at breast height, dominant height, elevation, and human footprint. Our results suggest that reforestation and forest conservation in Northeast Asia have effectively expanded the size of the carbon sink in the region, and sustainable forest management practices such as precision forestry and close forest monitoring for fire and insect outbreaks would be important to maintain and improve this critical carbon sink for Northeast Asia.

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.

Contrasting Patterns of Fungal and Bacterial Endophytes Inhabiting Temperate Tree Leaves in Response to Thinning.

The phyllosphere is an important but underestimated habitat for a variety of microorganisms, with limited knowledge about leaf endophytes as a crucial component of the phyllosphere microbiome. In this study, we investigated the mechanisms of communities and co-occurrence networks of leaf endophytes in response to forest thinning in a temperate forest. As we expected, contrasting responses of fungal and bacterial endophytes were observed. Specifically, the diversity of leaf endophytic fungi and the complexity of their co-occurrence networks increased significantly with thinning intensity, whereas the complexity of endophytic bacterial co-occurrence networks decreased. In particular, microbiota inhabiting damaged leaves seem to be more intensively interacting, showing an evident fungi-bacteria trade-off under forest thinning. In damaged leaves, besides the direct effects of thinning, thinning-induced changes in neighbor tree diversity indirectly altered the diversity of leaf fungal and bacterial endophytes via modifying leaf functional traits such as leaf dry matter content and specific leaf area. These findings provide new experimental evidence for the trade-offs between leaf endophytic fungi and bacteria under the different magnitudes of deforestation, highlighting their dependence on the presence or absence of leaf damage.

Unravelling Trait-Environment Relationships at Local and Regional Scales in Temperate Forests.

Understanding the trait-environment relationships has been a core ecological research topic in the face of global climate change. However, the strength of trait-environment relationships at the local and regional scales in temperate forests remains poorly known. In this study, we investigated the local and regional scale forest plots of the natural broad-leaved temperate forest in northeastern China, to assess what extent community-level trait composition depends on environmental drivers across spatial scales. We measured five key functional traits (leaf area, specific leaf area, leaf carbon content, leaf nitrogen content, and wood density) of woody plant, and quantified functional compositions of communities by calculating the "specific" community-weighted mean (CWM) traits. The sum of squares decomposition method was used to quantify the relative contribution of intraspecific trait variation to total trait variation among communities. Multiple linear regression model was then used to explore the community-level trait-environment relationships. We found that (i) intraspecific trait variation contributed considerably to total trait variation and decreased with the spatial scale from local to regional; (ii) functional composition was mainly affected by soil and topography factors at the local scale and climate factor at the regional scale, while explaining that variance of environment factors were decreased with increasing spatial scale; and (iii) the main environment driver of functional composition was varied depending on the traits and spatial scale. This work is one of the few multi-scale analyses to investigate the environmental drivers of community functional compositions. The extent of intraspecific trait variation and the strength of trait-environment relationship showed consistent trends with increasing spatial scale. Our findings demonstrate the influence of environmental filtering on both local- and regional-scale temperate forest communities, and contribute to a comprehensive understanding of trait-environment relationships across spatial scales.

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.

A global synthesis reveals increases in soil greenhouse gas emissions under forest thinning.

Forest thinning is a major forest management practice worldwide and may lead to profound alterations in the fluxes of soil greenhouse gases (GHGs). However, the global patterns and underlying mechanisms of soil GHG fluxes in response to forest thinning remain poorly understood. Here, we conducted a global meta-analysis of 106 studies to assess the effects of forest thinning on soil GHG fluxes and the underpinning mechanisms. The results showed that forest thinning significantly increased soil CO2 emission (mean lnRR: 0.07, 95% CI: 0.03-0.11), N2O emission (mean lnRR: 0.39, 95% CI: 0.16-0.61) and decreased CH4 uptake (mean Hedges' d: 0.98, 95% CI: 0.32-1.64). Furthermore, the negative response of soil CH4 uptake was amplified by thinning intensity, and the positive response of soil N2O emission decreased with recovery time after thinning. The response of soil CO2 emission was mainly correlated with changes in fine root biomass and soil nitrogen content, and the response of soil CH4 uptake was related to the changes in soil moisture and litterfall. Moreover, the response of soil N2O emission was associated with changes in soil temperature and soil nitrate nitrogen content. Thinning also increased the total balance of the three greenhouse gas fluxes in combination, which decreased with recovery time. Our findings highlight that thinning significantly increases soil GHG emissions, which is crucial to understanding and predicting ecosystem-climate feedbacks in managed forests.

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.

Allometric biomass equations for 12 tree species in coniferous and broadleaved mixed forests, Northeastern China.

Understanding forest carbon budget and dynamics for sustainable resource management and ecosystem functions requires quantification of above- and below-ground biomass at individual tree species and stand levels. In this study, a total of 122 trees (9-12 per species) were destructively sampled to determine above- and below-ground biomass of 12 tree species (Acer mandshuricum, Acer mono, Betula platyphylla, Carpinus cordata, Fraxinus mandshurica, Juglans mandshurica, Maackia amurensis, P. koraiensis, Populus ussuriensis, Quercus mongolica, Tilia amurensis and Ulmus japonica) in coniferous and broadleaved mixed forests of Northeastern China, an area of the largest natural forest in the country. Biomass allocation was examined and biomass models were developed using diameter as independent variable for individual tree species and all species combined. The results showed that the largest biomass allocation of all species combined was on stems (57.1%), followed by coarse root (21.3%), branch (18.7%), and foliage (2.9%). The log-transformed model was statistically significant for all biomass components, although predicting power was higher for species-specific models than for all species combined, general biomass models, and higher for stems, roots, above-ground biomass, and total tree biomass than for branch and foliage biomass. These findings supplement the previous studies on this forest type by additional sample trees, species and locations, and support biomass research on forest carbon budget and dynamics by management activities such as thinning and harvesting in the northeastern part of China.

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.

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.

Combined effects of nitrogen addition and organic matter manipulation on soil respiration in a Chinese pine forest.

The response of soil respiration (Rs) to nitrogen (N) addition is one of the uncertainties in modelling ecosystem carbon (C). We reported on a long-term nitrogen (N) addition experiment using urea (CO(NH2)2) fertilizer in which Rs was continuously measured after N addition during the growing season in a Chinese pine forest. Four levels of N addition, i.e. no added N (N0: 0 g N m-2 year-1), low-N (N1: 5 g N m-2 year-1), medium-N (N2: 10 g N m-2 year-1), and high-N (N3: 15 g N m-2 year-1), and three organic matter treatments, i.e. both aboveground litter and belowground root removal (LRE), only aboveground litter removal (LE), and intact soil (CK), were examined. The Rs was measured continuously for 3 days following each N addition application and was measured approximately 3-5 times during the rest of each month from July to October 2012. N addition inhibited microbial heterotrophic respiration by suppressing soil microbial biomass, but stimulated root respiration and CO2 release from litter decomposition by increasing either root biomass or microbial biomass. When litter and/or root were removed, the "priming" effect of N addition on the Rs disappeared more quickly than intact soil. This is likely to provide a point of view for why Rs varies so much in response to exogenous N and also has implications for future determination of sampling interval of Rs measurement.

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.

Reproduction and vegetative growth in the dioecious shrub Acer barbinerve in temperate forests of Northeast China.

KEY MESSAGE: Trade-off in dioecious plant. The trade-off between reproduction, vegetative growth and maintenance is a major issue in the life history of an organism and a record of the process which is producing the largest possible number of living offspring by natural selection. Dioecious species afford an excellent opportunity for detecting such possible trade-offs in resource allocation. In this study, we selected the dioecious shrub Acer barbinerve to examine possible trade-offs between reproduction and vegetative growth in both genders at different modular levels during three successive years. Reproductive and vegetative biomass values were assessed during successive years to evaluate their intra-annual and inter-annual trade-offs. These trade-offs were examined at shoot, branch and shrub modular levels in Acer barbinerve shrubs. An intra-annual trade-off was detected at the shoot level for both genders in 2011 and 2012. Both males and females showed a negative correlation between reproduction and vegetative growth, but this was more prominent in males. For the females of the species, inter-annual trade-offs were only found at branch and shrub levels. Slightly negative correlations in females were detected between the reproduction in 2012 and the reproduction in the two previous years. The gender ratio was significantly male biased during the three successive years of our investigation. Females had higher mortality rates in the larger diameter classes, both in 2011 and 2012. This study revealed a clear trade-off between reproduction and vegetative growth in Acer barbinerve, but results varied between males and females. The degree of autonomy of the different modular levels may affect the ability to detect such trade-offs.