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The complex spatial structure and dynamics of ecological communities continue to defy explanation by simple principles despite decades of attention from ecologists and theoreticians. For example, the relationship between plant spatial distributions and species coexistence is often challenging to resolve in empirical settings. By analysing the spatial patterns of trees in 21 large forest plots we find a general and strong latitudinal gradient in the relationship between conspecific aggregation and abundance of tree species, with stronger negative abundance-dependency as latitude increases. To derive theoretical expectations for how interactions between multiple spatial pattern and processes can impact species coexistence, we incorporate the observed spatial patterns together with neighbourhood crowding competition into a mathematical model to estimate the local extinction risk of species. Strikingly, we find simple relationships that predict species local extinction risk from their demography and spatial distribution. Compared to a corresponding non-spatial analysis, accounting for spatial patterns reduces the 1000-year extinction risk on average by 52% when species invade from low abundances of 50 individuals. Additionally, based on their current abundances, only 8% of the species had an extinction risk greater than 5%. Our approach opens up new avenues for integrating observed spatial patterns with multiple ecological processes into mathematical theory. Our results demonstrate that emerging spatial patterns can contribute substantially to coexistence in species-rich forests, emphasizing the need to understand the interacting multiple processes underpinning spatial patterns in greater detail than has previously been appreciated.

Diversity–biomass relationships (DBRs) often vary with spatial scale in terrestrial ecosystems, but the mechanisms driving these scale-dependent patterns remain unclear, especially for highly heterogeneous forest ecosystems. This study explores how mutualistic associations between trees and different mycorrhizal fungi, i.e., arbuscular mycorrhizal (AM) vs. ectomycorrhizal (EM) association, modulate scale-dependent DBRs. We hypothesized that in soil-heterogeneous forests with a mixture of AM and EM tree species, (i) AM and EM tree species would respond in contrasting ways (i.e., positively vs. negatively, respectively) to increasing soil fertility, (ii) AM tree dominance would contribute to higher tree diversity and EM tree dominance to greater standing biomass, and that as a result (iii) mycorrhizal associations would exert an overall negative effect on DBRs across spatial scales. To empirically test these hypotheses, we collected detailed tree distribution and soil information (e.g., nitrogen, phosphorus, organic matter, pH) from seven temperate and subtropical AM–EM mixed forest megaplots (16–50 ha). Using a spatial codispersion null model and structural equation modeling, we identified the relationships among AM or EM tree dominance, soil fertility, tree species diversity, and biomass and, thus, DBRs across 0.01- to 1-ha scales. We found the first evidence overall supporting the three aforementioned hypotheses in these AM–EM mixed forests: (i) In most forests, with increasing soil fertility, tree communities changed from EM-dominated to AM-dominated; (ii) increasing AM tree dominance had an overall positive effect on tree diversity and a negative effect on biomass, even after controlling for soil fertility and number of trees. Together, (iii) the changes in mycorrhizal dominance along soil fertility gradients weakened the positive DBR observed at 0.01- to 0.04-ha scales in nearly all forests and drove negative DBRs at 0.25- to 1-ha scales in four out of seven forests. Hence, this study highlights a soil-related mycorrhizal dominance mechanism that could partly explain why, in many natural forests, biodiversity–ecosystem functioning (BEF) relationships shift from positive to negative with increasing spatial scale.

Ecosystem stability reveals how ecosystems respond to global change over time. Yet, the focus of past research on small spatial grains and extents overlooks scale dependence and how broad-scale environmental gradients shape stability. Here, we use forest inventory data covering a broad latitudinal gradient from the temperate to the tropical zone to examine cross-scale variation in stability of aboveground biomass and underlying stabilizing mechanisms. While stability did not shift systematically with latitude at either spatial grain, we found evidence that species asynchrony increased towards the tropics at the small spatial grain while species stability decreased at both spatial grains. Moreover, latitude stabilized forest communitiesviaits effects on both stabilizing mechanisms, which compensated for the weak and destabilizing effects of species richness. Yet, the trade-off in the relative importance of species stability and species asynchrony for stability was not mediated by latitude, suggesting that context-dependent factors - to a greater extent than macroecological ones - underlie large-scale patterns of stability. Our results highlight the crucial role of species asynchrony and species stability in determining ecosystem stability across broad-scale environmental gradients, suggesting that conserving biodiversity alone may not be sufficient for stabilizing naturally-assembled forest ecosystems.

Phylogenetic trees have been extensively used in community ecology. However, how the phylogeny construction affects ecological inferences is poorly understood. In this study, we constructed three different types of phylogenetic trees (a synthetic-tree generated using V.PhyloMaker, a barcode-tree generated using rbcL+matK+trnH-psbA, and a plastome-tree generated from plastid genomes) that represented an increasing level of phylogenetic resolution among 580 woody plant species from six forest dynamic plots in subtropical evergreen broadleaved forests of China. We then evaluated the performance of each phylogeny in estimations of community phylogenetic structure, turnover and phylogenetic signal in functional traits. As expected, the plastome-tree was most resolved and most supported for relationships among species. For local phylogenetic structure, the three trees showed consistent results with Faith's PD and MPD; however, only the synthetic-tree produced significant clustering patterns using MNTD for some plots. For phylogenetic turnover, contrasting results between the molecular trees and the synthetic-tree occurred only with nearest neighbor distance. The barcode-tree agreed more with the plastome-tree than the synthetic-tree for both phylogenetic structure and turnover. For functional traits, both the barcode-tree and plastome-tree detected phylogenetic signal in maximum height, but only the plastome-tree detected signal in leaf width. This is the first study that uses plastid genomes in large-scale community phylogenetics. Our results highlight the improvement of plastome-trees over barcode-trees and synthetic-trees for the analyses studied here. Our results also point to the possibility of type I and II errors in estimation of phylogenetic structure and turnover and detection of phylogenetic signal when using synthetic-trees.

Ecology cannot yet fully explain why so many tree species coexist in natural communities such as tropical forests. A major difficulty is linking individual-level processes to community dynamics. We propose a combination of tree spatial data, spatial statistics and dynamical theory to reveal the relationship between spatial patterns and population-level interaction coefficients and their consequences for multispecies dynamics and coexistence. Here we show that the emerging population-level interaction coefficients have, for a broad range of circumstances, a simpler structure than their individual-level counterparts, which allows for an analytical treatment of equilibrium and stability conditions. Mechanisms such as animal seed dispersal, which result in clustering of recruits that is decoupled from parent locations, lead to a rare-species advantage and coexistence of otherwise neutral competitors. Linking spatial statistics with theories of community dynamics offers new avenues for explaining species coexistence and calls for rethinking community ecology through a spatial lens., Tree spatial data, spatial statistics and dynamical theory reveal the relationship between spatial patterns and population-level interaction coefficients and their consequences for multispecies dynamics and coexistence.

Plant-associated microbes are essential for promoting plant well-being, maintaining biodiversity, and supporting ecosystem function. However, little is known about the geographic distribution of plant-microbe symbioses and how they are formed and change along latitudinal gradients. Here we identified leaf bacteria for 328 plant species sampled from 10 forests along a tropical to temperate gradient in China. We analyzed the diversity and composition of plant leaf-associated bacteria and quantified the contributions of hosts, habitats, and neighborhood plants to the plant-bacterial symbiosis. We found a strong latitudinal gradient in leaf bacterial diversity and composition. Bacterial assemblages on leaves were most strongly selected by host plants, and the selection pressure increased with latitude. In contrast, at low latitudes and at large geographical scales multiple factors were found to jointly regulate bacterial community composition. Our result also showed that plant-bacteria symbiotic networks were structured by network hub bacteria taxa with high co-occurrence network centrality, and the abundance of temperate hub taxa was more influenced by host plants than that in tropical forests. For the first time, we documented a previously unrecognized latitudinal gradient in plant-bacterial symbioses that was regulated by a joint effect of multiple factors at low latitudes but mostly by host selection at high latitudes, implying that leaf microbiomes are likely to respond differently to global change along the latitudinal gradient.

Plant functional traits are rarely used in tree species classification, and the impact of vertical canopy positions on collecting samples for classification also remains unclear. We aim to explore the feasibility and effectiveness of leaf traits in classification, as well as to detect the effect of vertical position on classification accuracy. This work will deepen our understanding of the ecological mechanism of natural forest structure and succession from new perspectives. In this study, we collected foliar samples from three canopy layers (upper, middle and lower) and measured their spectra, as well as eight well-known leaf traits. We used a leaf hyperspectral reflectance (LHR) dataset, leaf functional traits (LFT) dataset and LFT + LHR dataset to classify six dominant tree species in a subtropical evergreen broad-leaved forest. Our results showed that the LFT + LHR dataset achieved the highest classification results (overall accuracy (OA) = 77.65% and Kappa = 0.73), followed by the LFT dataset (OA = 74.26% and Kappa = 0.69) and the LHR dataset (OA = 69.06% and Kappa = 0.63). Along the vertical canopy, the OA and Kappa increased from the lower to the upper layers, and the combination data of the three canopy layers achieved the highest accuracy. For the individual tree species, the shade-tolerant species (including Machilus chinensis, Cryptocarya chinensis and Cryptocarya concinna) produced higher accuracies than the light-demanding species (including Schima superba and Castanopsis chinensis). Our results provide an approach for enhancing tree species recognition from the plant physiology and biochemistry perspective and emphasize the importance of vertical direction in forest community research.

Vertical stratification in trees may respond to selective pressures to enhance light interception and utilization; therefore, the vertical functional variation in leaf traits may indicate niche partitioning within forests. In this study, vertical variations in leaf and crown structure traits of seven common tree species were analysed with respect to differences between species in different height groups, within the same height range, in the same species across tree height, and different parts of the individual tree crown to reveal coexistence mechanisms in subtropical forest tree species. There were multiple levels of trait variation in the vertical dimension, validating the existence of vertical niche differentiation in subtropical forest species. The functional trait differences arose among different height groups, among species co-occurring within the same height range, in the same species across tree height, and among different parts of the individual tree crown. Variation in comparative advantages, which was characterised by those traits between species across different height ranges, was also one of the manifestations of niche differentiation in the vertical dimension. Moreover, contrasting results between lower height ranges and higher ranges in the relationship between species’ differences in functional traits and species’ difference of abundance were found, further confirming that there was obvious vertical niche separation in the community. This study emphasised the importance of vertical variation in species’ performances in elucidating the mechanisms of tree species coexistence in subtropical forests.

What causes variation in species abundance for a given site remains a central question in community ecology. Foundational to trait-based ecology is the expectation that functional traits determine species abundance. However, the relative success of using functional traits to predict relative abundance is questionable. One reason is that the diversity in plant function is greater than that characterized by the few most commonly and easily measurable traits. Here, we measured 10 functional traits and the stem density of 101 woody plant species in a 200,000 m2 permanent, mature, subtropical forest plot (high precipitation and high nitrogen, but generally light- and phosphorus-limited) in southern China to determine how well relative species abundance could be predicted by functional traits. We found that: (1) leaf phosphorus content, specific leaf area, maximum CO2 assimilation rate, maximum stomata conductance, and stem hydraulic conductivity were significantly and negatively associated with species abundance, (2) the ratio of leaf nitrogen content to leaf phosphorus content (N:P) and wood density were significantly positively correlated with species abundance; (3) neither leaf nitrogen content nor leaf turgor loss point were related to species abundance; (4) a combination of N:P and maximum stomata conductance accounted for 44% of the variation in species’ abundances. Taken together, our findings suggested that the combination of these functional traits are powerful predictors of species abundance. Species with a resource-conservative strategy that invest more in their tissues are dominant in the mature, subtropical, evergreen forest.

Arbuscular mycorrhizal (AM) and ectomycorrhizal (EcM) associations are critical for host-tree performance. However, how mycorrhizal associations correlate with the latitudinal tree beta-diversity remains untested. Using a global dataset of 45 forest plots representing 2,804,270 trees across 3840 species, we test how AM and EcM trees contribute to total beta-diversity and its components (turnover and nestedness) of all trees. We find AM rather than EcM trees predominantly contribute to decreasing total beta-diversity and turnover and increasing nestedness with increasing latitude, probably because wide distributions of EcM trees do not generate strong compositional differences among localities. Environmental variables, especially temperature and precipitation, are strongly correlated with beta-diversity patterns for both AM trees and all trees rather than EcM trees. Results support our hypotheses that latitudinal beta-diversity patterns and environmental effects on these patterns are highly dependent on mycorrhizal types. Our findings highlight the importance of AM-dominated forests for conserving global forest biodiversity., The relationship of mycorrhizal associations with latitudinal gradients in tree beta-diversity is unexplored. Using a global dataset approach, this study examines how trees with arbuscular mycorrhizal and ectomycorrhizal associations contribute to latitudinal beta-diversity patterns and the environmental controls of these patterns.

The decline in species richness at higher latitudes is among the most fundamental patterns in ecology. Whether changes in species composition across space (beta-diversity) contribute to this gradient of overall species richness (gamma-diversity) remains hotly debated. Previous studies that failed to resolve the issue suffered from a well-known tendency for small samples in areas with high gamma-diversity to have inflated measures of beta-diversity. Here, we provide a novel analytical test, using beta-diversity metrics that correct the gamma-diversity and sampling biases, to compare beta-diversity and species packing across a latitudinal gradient in tree species richness of 21 large forest plots along a large environmental gradient in East Asia. We demonstrate that after accounting for topography and correcting the gamma-diversity bias, tropical forests still have higher beta-diversity than temperate analogues. This suggests that beta-diversity contributes to the latitudinal species richness gradient as a component of gamma-diversity. Moreover, both niche specialization and niche marginality (a measure of niche spacing along an environmental gradient) also increase towards the equator, after controlling for the effect of topographical heterogeneity. This supports the joint importance of tighter species packing and larger niche space in tropical forests while also demonstrating the importance of local processes in controlling beta-diversity.

Functional diversity is considered a key link between ecosystem functions and biodiversity, and forms the basis for making community diversity conservation strategies. Here, we chose a subtropical forest community in China as the research object, which is unique in that other regions of the world at the same latitude have almost no vegetation cover. We measured 17 functional traits of 100 plant species and calculated seven different functional diversity indices, based on functional richness, evenness, and divergence. We found that most functional diversity and species diversity indices significantly differed with plant habit. There was a significant positive correlation among functional richness indices. However, functional divergence indices, multidimensional functional divergence (FDiv), and Rao’s quadratic entropy index (RaoQ) were significantly negatively correlated, and RaoQ and functional divergence indices (FDis) were uncorrelated. The correlations between three types (richness, evenness, and divergence) of functional diversity indices and three species diversity indices were different. Lineage regression results generally showed that three functional richness indices (Average distance of functional traits (MFAD), Functional volume (FRic) and Posteriori functional group richness (FGR)) were increased with three species diversity indices (species richness (S), Shannon-Wiener index (H) and Pielou index (E)). The functional evenness index (FEve) decreased with species richness (S), Shannon-Wiener index (H) and increased with species evenness (Pielou index (E)), but the change trends were small. All three types of functional diversity indices declined with altitude, although altitude had a weak influence on them. Other environmental factors affected the functional diversity of the community. Here, soil total phosphorus (TP) was the most critical environmental factor and the convex had the least effect on functional diversity in our subtropical forest community. These results will contribute to our understanding of functional diversity in subtropical forests, and provide a basis for biodiversity conservation in this region.

The decline in species richness at higher latitudes is among the most fundamental patterns in ecology. Whether changes in species composition across space (beta-diversity) contribute to this gradient of overall species richness (gamma-diversity) remains hotly debated. Previous studies that failed to resolve the issue suffered from a well-known tendency for small samples in areas with high gamma-diversity to have inflated measures of beta-diversity. Here, we provide here a novel analytical test, using beta-diversity metrics that correct the gamma-diversity and sampling biases, to compare beta-diversity and species packing across a latitudinal gradient in tree species richness of 21 large forest plots along a large environmental gradient in East Asia. We demonstrate that after accounting for topography and correcting the gamma-diversity bias, tropical forests still have higher beta-diversity than temperate analogues. This suggests that beta-diversity contributes to the latitudinal species richness gradient as a component of gamma-diversity. Moreover, both niche specialization and niche marginality (a measure of niche spacing along an environmental gradient) also increase towards the equator, after controlling for the effect of topographic heterogeneity. This supports the joint importance of tighter species packing and larger niche space in tropical forests while also demonstrating the importance of local processes in controlling beta-diversity.

The decline in species richness at higher latitudes is among the most fundamental patterns in ecology. Whether changes in species composition across space (beta-diversity) contribute to this gradient of overall species richness (gamma-diversity) remains hotly debated. Previous studies that failed to resolve the issue suffered from a well-known tendency for small samples in areas with high gamma-diversity to have inflated measures of beta-diversity. Here, we provide here a novel analytical test, using beta-diversity metrics that correct the gamma-diversity and sampling biases, to compare beta-diversity and species packing across a latitudinal gradient in tree species richness of 21 large forest plots along a large environmental gradient in East Asia. We demonstrate that after accounting for topography and correcting the gamma-diversity bias, tropical forests still have higher beta-diversity than temperate analogs. This suggests that beta-diversity contributes to the latitudinal species richness gradient as a component of gamma-diversity. Moreover, both niche specialization and niche marginality (a measure of niche spacing along an environmental gradient) also increase towards the equator, after controlling for the effect of topographic heterogeneity. This supports the joint importance of tighter species packing and larger niche space in tropical forests while also demonstrating the importance of local processes in controlling beta-diversity.

Legumes provide an essential service to ecosystems by capturing nitrogen from the atmosphere and delivering it to the soil, where it may then be available to other plants. However, this facilitation by legumes has not been widely studied in global tropical forests. Demographic data from 11 large forest plots (16–60 ha) ranging from 5.25° S to 29.25° N latitude show that within forests, leguminous trees have a larger effect on neighbor diversity than non-legumes. Where soil nitrogen is high, most legume species have higher neighbor diversity than non-legumes. Where soil nitrogen is low, most legumes have lower neighbor diversity than non-legumes. No facilitation effect on neighbor basal area was observed in either high or low soil N conditions. The legume–soil nitrogen positive feedback that promotes tree diversity has both theoretical implications for understanding species coexistence in diverse forests, and practical implications for the utilization of legumes in forest restoration., Xu et al. examine the effect of leguminous trees on neighbor diversity across 11 plots in tropical forests around the world, and find that in high soil nitrogen conditions, most legume species have higher neighbor diversity than non-legumes, and vice versa where soil nitrogen is low. Their results have practical implications for the utilization of legumes in forest restoration.

Environments in both biotic and abiotic ecosystems have been affected by the colonization of non-native flora. In this study, we examined the effect of Bidens alba invasion on different land-use types along a coastline in southern China. Bacterial communities in each site were determined using 16S rDNA sequencing, and soil physicochemical properties were analyzed using standard methods. Although our results indicated that B. alba invasion did not have a significant effect on the alpha diversity of bacteria, it caused significant differences in soil bacterial community composition between invaded and uninvaded soil across different land-use types. Beta diversity and several physicochemical properties in forest, orchard and waterfront environments were recorded to be more susceptible to B. alba invasion. A high proportion of the variation of bacterial communities can be explained by a combination of environmental variables, indicating that environmental selection rather than plant invasion is a more effective process in coastal microbial assemblages. By comparing topological roles of shared OTUs among invaded and uninvaded soil, keystone taxa in invaded soil were identified. Acidobacteria was the major phyla involved in the invasive process which could be driven by environmental selection. How key phyla react in our experiment should be verified by further studies.

Little attention has been paid to the impact of vertical canopy position on the leaf spectral properties of tall trees, and few studies have explored the ability of leaf spectra to characterize the variation of leaf traits across different canopy positions. Using a tower crane, we collected leaf samples from three canopy layers (lower, middle, and upper) and measured eight leaf traits (equivalent water thickness, specific leaf area, leaf carbon content, leaf nitrogen content, leaf phosphorus content, leaf chlorophyll content, flavonoid, and nitrogen balance index) in a subtropical evergreen broadleaved forest. We evaluated the variability of leaf traits and leaf spectral properties, as well as the ability of leaf spectra to track the variation of leaf traits among three canopy layers for six species within the entire reflectance spectrum. The results showed that the eight leaf traits that were moderately or highly correlated with each other showed significant differences along the vertical canopy profile. The three canopy layers of leaf spectra showed contrasting patterns for light-demanding (Castanopsis chinensis, Castanopsis fissa, Schima superba, and Machilus chinensis) and shade-tolerant species (Cryptocarya chinensis and Cryptocarya concinna) along the vertical canopy profile. The spectra at the lower and upper canopy layers were more sensitive than the middle layer for tracking the variation of leaf chlorophyll and flavonoid content. Our results revealed that it is important to choose an appropriate canopy layer for the field sampling of tall trees, and we suggest that flavonoid is an important leaf trait that can be used for mapping and monitoring plant growth with hyperspectral remote sensing.

Typhoons are having an increasingly severe and widespread impact on forest ecosystems around the world due to climate change. However, few studies have examined the impacts of strong typhoons on subtropical evergreen broadleaved forests, particularly in rare old-growth forests. Here, using a 20 ha permanent subtropical forest plot in Guangdong Province, southern China, the impact of super Typhoon Mangkhut on secondary and old-growth subtropical evergreen broadleaved forests was quantitatively investigated and compared. The old-growth forest was damaged more than twice as much as the secondary forest, as measured by the proportion of the basal area of damaged trees. Forest structure and composition played a significant role in determining the vulnerability of forests to powerful typhoons. This study advances knowledge about how old-growth forests respond to global climate change.

Ormosia formosana is an important hardwood species and its seeds are popular as decorative jewelry. Currently, this species is threatened in the natural forests due to habitat destruction. Here, we first report the chloroplast genome of O. formosana for future studies in ecology, phylogeny, and conservation. The chloroplast genome of O. formosana is 173,587 bp in length with a GC content of 35.80%. It includes a large single-copy region of 73,550 bp, a small single-copy region of 18,683 bp, and two inverted repeat regions of 40,696 bp and 40,658 bp, respectively. The genome was totally annotated with 135 genes, including 90 protein-coding genes, eight ribosomal RNA genes, and 37 transfer RNA genes. Phylogenetic analysis indicated that O. formosana is most genetically similar to O. boluoensis.

Community assembly in natural communities is commonly explained by stochastic and niche-based processes such as environmental filtering and biotic interactions. Many studies have inferred the importance of these processes using a trait-based approach, however, there are still unknowns around what factors affect the importance of different assembly processes in natural communities. In this study, the trait dispersion patterns of 134 species were examined across different functional traits, habitat types, ontogenetic stages and spatial scales from a 20-ha Dinghushan Forest Dynamic Plot in China. The results showed that (1) functional traits related to productivity such as specific leaf area and leaf area mainly showed functional clustering, indicating these two functional traits were more affected by environmental filtering. However, trait dispersion patterns depended on more than the ecological significances of functional traits. For example, trait dispersions of leaf dry matter content, leaf thickness and maximum height did not show consistent patterns across habitat types and ontogenetic stages, suggesting more complex mechanisms may operate on these traits, (2) the trait dispersion varied with the habitat types and ontogenetic stages. Specifically, we found that habitat types only affected the strength of trait dispersions for all the five traits, but ontogenetic stages influenced both the strength and direction of trait dispersions, which depended on the traits selected, (3) the relative importance of soil, topography and space to trait dispersion varied with ontogenetic stages. Topography and space were more important for trait dispersion of saplings but soil was more important for trait dispersion of adults, (4) biotic interactions dominated community assembly at smaller spatial scales but environmental filtering dominated community assembly at larger spatial scales. Overall, the results highlight the importance of functional traits, habitat types, ontogenetic stages and spatial scales to community assembly in natural communities.

The competition-colonization trade-off, by which species can partition spatial niches, is a potentially important mechanism allowing the maintenance of species diversity in plant communities. We examined whether there was evidence for this trade-off among tree species in a subtropical forest and how it correlated with eight functional traits. We developed and estimated a metric for colonization ability that incorporates both fecundity and seed dispersal based on seed trap data and the sizes and distributions of adult trees. Competitive ability was estimated as survival probability under high crowding conditions based on neighborhood models. Although we found no significant relationship between colonization and competitive abilities, there was a significant negative correlation between long distance dispersal ability and competitive ability at the 5 cm size class. Colonizers had traits associated with faster growth, such as large leaves and low leaf lamina density, whereas competitors had traits associated with higher survival, such as dense wood. Our results imply that any trade-off between competition and colonization may be more determined by dispersal ability than by fecundity, suggesting that seed dispersal is an important contributor to diversity maintenance. Future work should test how competitive ability covaries with the components of colonization ability, as we did here.

Soil microbes re-establish plant diversity and ecosystem functions after disturbance events. Deterministic and stochastic processes are expected to contribute to microbial community assembly during long-term ecosystem recovery. We characterized soil prokaryotic and fungal communities, to determine their assembly patterns, along two chronosequences with early to later successional subtropical forests. Prokaryotic and fungal community composition was more variable in early successional forests but converged in the later successional forests. The community composition was governed by deterministic processes in the early stages, while the relative influence of stochasticity increased in the later stages. Environmental factors that predicted the shift in deterministic and stochastic balance varied within and across successional stages. In particular, the compositional dissimilarity of plant communities strongly predicted the relative influences of the two processes during succession. These findings suggest that the relative influences of deterministic and stochastic processes in shaping microbial community assembly shift during secondary succession. Consequently, plant communities are important predictors of assembly processes in soil microbial communities during long-term ecosystem recovery.

Environmental filtering and competitive interactions are important ecological processes in community assembly. The contribution of the two processes to community assembly can be evaluated by shifts in functional diversity patterns. We examined the correlations between functional diversity of six traits (leaf chlorophyll concentration, dry matter content, size, specific leaf area, thickness and wood density) and environmental gradients (topography and soil) for 92 species in the 20-ha Dinghushan forest plot in China. A partial Mantel test showed that most of the community-weighted mean trait values changed with terrain convexity and soil fertility, which implied that environmental filtering was occurring. Functional diversity of many traits significantly increased with increasing terrain convexity and soil fertility, which was associated with increased light and below-ground resources respectively. These results suggest that co-occurring species are functionally convergent in regions of strong abiotic stress under the environmental filtering, but functionally divergent in more benign environments due to resource partitioning and competitive interactions. Single-trait diversity and multivariate functional diversity had different relationships with environmental factors, indicating that traits were related to different niche axes, and associated with different ecological processes, which demonstrated the importance of focusing niche axes in traits selection. Between 9% and 41% of variation in functional diversity of different traits was explained by environmental factors in stepwise multiple regression models. Terrain convexity and soil fertility were the best predictors of functional diversity, which contributed 30.5% and 29.0% of total R2to the model. These provided essential evidence that different environmental factors had distinguishing impacts on regulating diversity of traits.

Understanding the role of dominant species in structuring the distribution of neighbor species is an important part of understanding community assembly, a central goal of ecology. Phylogenetic information helps resolve the multitude of processes driving community assembly and the importance of evolution in the assembly process. In this study, we classified species in a 20-ha subtropical forest in southern China into groups with different degrees of phylogenetic relatedness to the dominant species Castanopsis chinensis. Species surrounding individuals of C. chinensis were sampled in an equal area annulus at six spatial scales, counting the percent of relatives and comparing this to permutation tests of a null model and variance among species groups. The results demonstrated that dominant species affected their relatives depending on community successional stage. Theory would predict that competitive exclusion and density-dependence mechanisms should lead to neighbors that are more distant in phylogeny from C. chinensis. However, in mature forests distant relatives were subjected to competitive repulsion by C. chinensis, while environment filtering led to fewer distant species, regardless of scale. A variety of biological and non-biological factors appear to result in a U-shaped quantitative distribution determined by the dominant species C. chinensis. Scale effects also influenced the dominant species. As a dominant species, C. chinensis played an important role in structuring the species distributions and coexistence of neighbor species in a subtropical forest.

Climate is widely recognised as an important determinant of the latitudinal diversity gradient. However, most existing studies make no distinction between direct and indirect effects of climate, which substantially hinders our understanding of how climate constrains biodiversity globally. Using data from 35 large forest plots, we test hypothesised relationships amongst climate, topography, forest structural attributes (stem abundance, tree size variation and stand basal area) and tree species richness to better understand drivers of latitudinal tree diversity patterns. Climate influences tree richness both directly, with more species in warm, moist, aseasonal climates and indirectly, with more species at higher stem abundance. These results imply direct limitation of species diversity by climatic stress and more rapid (co-)evolution and narrower niche partitioning in warm climates. They also support the idea that increased numbers of individuals associated with high primary productivity are partitioned to support a greater number of species.

Topography is an important environmental feature that affects the spatial genetic structures of plant species. However, its influence on such structures at small spatial scales has hardly been investigated for forest tree species even though many of them are located in mountains with complex topographic features. Here we report the genetic structures of a pioneer species, Castanopsis chinensis (Fagaceae), in a 20-ha forest plot in Dinghushan in lower subtropical China, which has complex topographic features, such as valleys and ridges. C. chinensis typically undergoes wind pollination, but its seeds are dispersed by gravity and animals. Therefore, the mechanisms of transportation of its seeds may result in topography-dependent genetic structures, whereas wind-mediated pollen flow of C. chinensis may reduce such structures through counteracting the influence of topography. Our results indicate that most of the genetic patterns in C. chinensis in our study plot were attributable to wind-mediated pollen flow. However, we also found both seed and pollen flows could be impeded by ridges, causing some varied genetic patterns even between locations separated by only small distances. We observed that owing to topographic positions where they grew, the 13 old individuals that were probably the oldest in the southeast corner of the plot had not made major genetic contributions to the young individuals that had recently colonised previously clear-cut land in the rest of the plot. Therefore, our results indicate that we should consider both species life-history traits and topography when studying the genetic structures of plant populations in order to design sound conservation and management programs for the areas with complex topography.

The accurate characterization of aboveground biomass (AGB) dynamics and their driving forces in sub-tropical forests is important to evaluate AGB's contribution to global carbon stocks. In order to estimate the effects of local variables on temporal and spatial patterns of AGB we investigated the variation of AGB in a 20-ha sub-tropical permanent plot. We found that the AGB was 153.7 Mg ha-1 in 2005, and 149.3 Mg ha-1 in 2010. During the five-year period, AGB changed due to tree mortality, growth (diameter at breast height, DBH) and recruitment. Medium-sized trees (DBH: 30–50 cm) were the largest contributor to total AGB. Both the tree species diversity index and the number of individuals within medium (DBH: 30–50 cm) and large (DBH: >50 cm) DBH ranges had significant positive effects on AGB. In addition, the abiotic factors such as elevation, slope, and convexity had negative significant relationships with AGB. The results indicate the importance of including forest community characteristics and environmental variables in forest carbon studies. This study also provides important data that can be used to develop and validate carbon cycling models for old-growth sub-tropical forests.

The relationship between species richness and ecosystem function, as measured by productivity or biomass, is of long-standing theoretical and practical interest in ecology. This is especially true for forests, which represent a majority of global biomass, productivity and biodiversity. Here, we conduct an analysis of relationships between tree species richness, biomass and productivity in 25 forest plots of area 8–50 ha from across the world. The data were collected using standardized protocols, obviating the need to correct for methodological differences that plague many studies on this topic. We found that at very small spatial grains (0.04 ha) species richness was generally positively related to productivity and biomass within plots, with a doubling of species richness corresponding to an average 48% increase in productivity and 53% increase in biomass. At larger spatial grains (0.25 ha, 1 ha), results were mixed, with negative relationships becoming more common. The results were qualitatively similar but much weaker when we controlled for stem density: at the 0.04 ha spatial grain, a doubling of species richness corresponded to a 5% increase in productivity and 7% increase in biomass. Productivity and biomass were themselves almost always positively related at all spatial grains. Synthesis. This is the first cross-site study of the effect of tree species richness on forest biomass and productivity that systematically varies spatial grain within a controlled methodology. The scale-dependent results are consistent with theoretical models in which sampling effects and niche complementarity dominate at small scales, while environmental gradients drive patterns at large scales. Our study shows that the relationship of tree species richness with biomass and productivity changes qualitatively when moving from scales typical of forest surveys (0.04 ha) to slightly larger scales (0.25 and 1 ha). This needs to be recognized in forest conservation policy and management.

Structure and demographics in many tropical forests is changing, but the causes of these changes remain unclear. We studied 5 y (2005–2010) of species turnover, recruitment, mortality and population change data from a 20-ha subtropical forest plot in Dinghushan, China, to identify trends in forest change, and to test whether tree mortality is associated with intraspecific or interspecific competition. We found the Dinghushan forest to be more dynamic than one temperate and two tropical forests in a comparison of large, long-term forest dynamics plots. Within Dinghushan, size-class distributions were bell-shaped only for the three most dominant species and reverse J-shaped for other species. Bell-shaped population distributions can indicate a population in decline, but our data suggest that these large and long-lived species are not in decline because the pattern is driven by increasing probabilities of transition to larger size class with increasing size and fast growth in saplings. Spatially aggregated tree species distributions were common for surviving and dead individuals. Competitive associations were more frequently intraspecific than interspecific. The competition that induced tree mortality was more associated with intraspecific than interspecific interactions. Intraspecific competitive exclusion and density-dependence appear to play important roles in tree mortality in this subtropical forest.

Tree mortality and recruitment are key factors influencing forest dynamics, but the driving mechanisms of these processes remain unclear. To better understand these driving mechanisms, we studied forest dynamics over a 5-year period in a 20-ha sub-tropical forest in the Dinghushan Nature Reserve, South China. The goal was to identify determinants of tree mortality/recruitment at the local scale using neighborhood analyses on some locally dominant tree species. Results show that the study plot was more dynamic than some temperate and tropical forests in a comparison to large, long-term forest dynamics plots. Over the 5-year period, mortality rates ranged from 1.67 to 12.33% per year while recruitment rates ranged from 0 to 20.26% per year. Tree size had the most consistent effect on mortality across species. Recruitment into the ≥1-cm size class consistently occurred where local con-specific density was high. This suggests that recruitment may be limited by seed dispersal. Hetero-specific individuals also influenced recruitment significantly for some species. Canopy species had low recruitment into the ≥1-cm size class over the 5-year period. In conclusion, tree mortality and recruitment for sixteen species in this plot was likely limited by seed dispersal and density-dependence.

In most plants, the contributions of pollen and seed flow to their genetic structures are generally difficult to disentangle. For typical wind-pollinated and wind-dispersed species Engelhardia roxburghiana in a 20-ha natural forest plot in lower subtropic China, because the prevailing wind directions change during its pollen release and seed dispersal seasons, we could compare its genetic structures in different directions, which could result primarily from pollen or seed flow. Furthermore, because the plot has undergone from an open to a closed canopy stage historically, we also examined forest canopy effects on gene flow in different generations and different directions. Using 522 E. roxburghiana individuals mapped in the plot, our results revealed that greater pollen flow led to biased gene flow in the pollen dispersal-predominant direction (pollen direction), while greater seed flow generated less spatial genetic structure in the seed dispersal-predominant direction (seed direction). The results predicted from generalized additive models indicated that canopy closure enhanced resistance to gene flow from the old generation to the new generation. Analyses by landscape genetic models for the new generation revealed that gene flow associated with pollen direction was more strongly affected by canopy than with seed direction. Our study is new by proposing an alternative way to separate effects of the pollen and seed flow on spatial variation patterns in E. roxburghiana. To our knowledge, our study is also the first attempt to use landscape genetic models to represent canopy effects for different dispersal vectors in spatial scales only up to a few hundred meters.

In general, even within a local area, many common plant species are found in diff erent types of environment. We propose that if the association of a common plant species with diff erent types of environment is investigated, by analysing all individuals in a given population as a single entity, the results might be misleading or incomplete owing to intraspecifi c variation. To test this hypothesis, we used molecular markers to classify mature Castanopsis chinensis individuals with a diameter at breast height � 40 cm into diff erent genetic groups and analysed the associations of these groups with topographic features and habitats within a 20-ha Dinghushan forest plot, South China. Our results indicated that the diff erent groups had diff erent topographical associations, and that the spatial distributions and genetic structures of individuals varied among the groups. Th erefore, if signifi cant genetic structure exists in the population of a common species within a community, to understand the relationship between the spatial distributions of individuals in the population and the environment, it is necessary to classify the individuals into genetic groups and analyse the data for these groups, rather than for a combined group of all individuals.

Community assembly has long been an important issue in community ecology. The study of community phylogenetic structure, which applies phylogeny to community ecology studies, has provided an effective way to disentangle the most important ecological processes that drive community assembly. Study- ing the phylogenetic structure of a community involves firstly the construction of a supertree representing the species pool of the community, then a calculation of phylogenetic distances between all species within the community, and finally an inference of phylogenetic structure (e.g., clustering, overdispersion) obtained by statistically testing whether the obtained phylogenetic distances are different from those expected under ran- dom model, hence revealing key ecological processes involved in community assembly (e.g., habitat filter- ing, competition exclusion). Community phylogenetic structure is different when studied at different taxo- nomic, spatial or temporal scales. At small spatial scales, community phylogenetic pattern tends to change from clustering to overdispersion with decreasing taxonomical scale or increasing tree age class, while the pattern tends to be tighter clustering at larger spatial scales. Phylogenetic information also indicates the in

Functional redundancy is an important tool for justifying and prioritizing species protection in forest ecosystem, but it is a scale-dependent. If functional redundancy really exists, functional trait composition tends to have higher predictive ability of community assembly than species composition. Thus, comparing the differences in the predictive ability of community assembly between species and functional trait compositions across spatial scale represents a useful tool to quantify how functional redundancy varies across spatial scales. Here, we used variation partitioning in combination with distance-based Moran’s eigenvector maps to compare the differences in the predictive ability of community assembly between species composition and functional trait composition across spatial scales (20, 30, 40, 50, and 100 m) in a 20-ha subtropical forest plot. We found that functional trait composition possessed higher predictive ability of niche-based abiotic filtering process than species composition within 40 m. At 50 and 100 m scales, both species and functional trait compositions had approximately equal predictive ability of dispersal limitation processes. Thus, functional redundancy can only exist within 40 m scale but not 50 and 100 m scales. As a result, priority species loss protection should be performed at 50 and 100 m scales.

We assessed the effects of topographic heterogeneity and stem density on species composition between grains of different sizes (20 × 20, 50 × 50, and 100 × 100 m), based on partial Mantel tests. Similarity in species composition was measured by the abundance-based Jaccard index (C_J) and by an index that incorporates phylogenetic information into C_J (pC_J). Plants were divided into five groups, arbor, subarbor, and shrub according to life form and two other groups: species that produce dry fruits (PDF) and that produce fleshy fruits (PFF). C_J and pC_J between any two grains at each grain size were calculated separately for these groups and for all species combined. In order to examine what influences C_J and pC_J, we analyzed their correlations with topographic heterogeneity variables and two dispersal limitation-related variables (stem and topographic resistance). Our data indicate that at all three grain sizes, C_J and pC_J decrease with increasing distance for all plant groups. Dispersal limitation and topographic heterogeneity were both important at 20 × 20 and 50 × 50 m grain sizes for C_J and pC_J of all plant groups; and at 100 × 100 m grain size, topographic heterogeneity dominates over dispersal limitation for some plant groups. C_J and pC_J of PDFs are less negatively correlated with stem resistance than those of PFFs. We conclude that both beta diversity and phylobetadiversity are dependent on plant groups and grain sizes.

Habitat specialization has been considered as a primary factor in determining the distribution of species. In this study, we investigated species–habitat associations while controlling for spatial neighbourhood effects in a large-scale (20 ha) stem-mapping plot in a species-rich subtropical forest of China. Habitat specialization was measured by topographic variation and its effects on species distributions were modelled at three different spatial scales (10×10, 20×20 and 25×25 m2) using log-linear regression models and randomization tests. Our results showed: (1) 83% of the species were related to at least one or more topographic variables. Among them, 66%, 60%, 65% and 70% were closely dependent on slope, aspect, elevation and convexity, respectively. (2) Topographic variables have much stronger non-linear (quadratic and cubic) effects on species distributions than linear effects. (3) The effects of topographic heterogeneity on the distribution of shrubs species are smaller than on the distribution of canopy species, and smaller effects were also found in less abundant species. (4) There was a strong neighbourhood effect on species distribution: In 85% of the species, abundance in a focal quadrat was significantly correlated with abundance in the neighbour quadrats. We conclude that habitat specialization plays an important role in maintaining the diversity of this species-rich subtropical forest.

Characterizing the spatial distribution patterns of soil microorganisms is helpful in understanding the biogeochemical processes they perform, but has been less studied relative to those of macroorganisms. In this study, we investigated and compared the spatially explicit distribution patterns of ammonia-oxidizing archaea (AOA) abundance and the influential factors between an early (ES) and a late successional (LS) subtropical forest stand. The average AOA abundance, vegetational attributes and soil nutrient contents were mostly greater in the LS than the ES stand (P = 0.085 or smaller), but their spatial variations were more pronounced in the ES than the LS stand. The spatial distribution patches of AOA abundance were smaller and more irregular in the ES stand (patch size

Tropical forests play a disproportionately important role in the global carbon (C) cycle, but it remains unclear how local environments and functional diversity regulate tree aboveground C storage. We examined how three components (environments, functional dominance and diversity) affected C storage in Dinghushan 20-ha plot in China. There was large fine-scale variation in C storage. The three components significantly contributed to regulate C storage, but dominance and diversity of traits were associated with C storage in different directions. Structural equation models (SEMs) of dominance and diversity explained 34% and 32% of variation in C storage. Environments explained 26–44% of variation in dominance and diversity. Similar proportions of variation in C storage were explained by dominance and diversity in regression models, they were improved after adding environments. Diversity of maximum diameter was the best predictor of C storage. Complementarity and selection effects contributed to C storage simultaneously, and had similar importance. The SEMs disengaged the complex relationships among the three components and C storage, and established a framework to show the direct and indirect effects (via dominance and diversity) of local environments on C storage. We concluded that local environments are important for regulating functional diversity and C storage.

BACKGROUND Ecologists have been monitoring community dynamics with the purpose of understanding the rates and causes of community change. However, there is a lack of monitoring of community dynamics from the perspective of phylogeny. METHODS/PRINCIPLE FINDINGS We attempted to understand temporal phylogenetic turnover in a 50 ha tropical forest (Barro Colorado Island, BCI) and a 20 ha subtropical forest (Dinghushan in southern China, DHS). To obtain temporal phylogenetic turnover under random conditions, two null models were used. The first shuffled names of species that are widely used in community phylogenetic analyses. The second simulated demographic processes with careful consideration on the variation in dispersal ability among species and the variations in mortality both among species and among size classes. With the two models, we tested the relationships between temporal phylogenetic turnover and phylogenetic similarity at different spatial scales in the two forests. Results were more consistent with previous findings using the second null model suggesting that the second null model is more appropriate for our purposes. With the second null model, a significantly positive relationship was detected between phylogenetic turnover and phylogenetic similarity in BCI at a 10 m×10 m scale, potentially indicating phylogenetic density dependence. This relationship in DHS was significantly negative at three of five spatial scales. This could indicate abiotic filtering processes for community assembly. Using variation partitioning, we found phylogenetic similarity contributed to variation in temporal phylogenetic turnover in the DHS plot but not in BCI plot. CONCLUSIONS/SIGNIFICANCE The mechanisms for community assembly in BCI and DHS vary from phylogenetic perspective. Only the second null model detected this difference indicating the importance of choosing a proper null model.