Unraveling the drivers and impacts of leaf phenological diversity in a subtropical forest: A fine-scale analysis using PlanetScope CubeSats.

Leaf phenology variations within plant communities shape community assemblages and influence ecosystem properties and services. However, questions remain regarding quantification, drivers, and productivity impacts of intra-site leaf phenological diversity. With a 50-ha subtropical forest plot in China's Heishiding Provincial Nature Reserve (part of the global ForestGEO network) as a testbed, we gathered a unique dataset combining ground-derived abiotic (topography, soil) and biotic (taxonomic diversity, functional diversity, functional traits) factors. We investigated drivers underlying leaf phenological diversity extracted from high-resolution PlanetScope data, and its influence on aboveground biomass (AGB) using structural equation modeling (SEM). Our results reveal considerable fine-scale leaf phenological diversity across the subtropical forest landscape. This diversity is directly and indirectly influenced by abiotic and biotic factors (e.g. slope, soil, traits, taxonomic diversity; r2 = 0.43). While a notable bivariate relationship between AGB and leaf phenological diversity was identified (r = -0.24, P < 0.05), this relationship did not hold in SEM analysis after considering interactions with other biotic and abiotic factors (P > 0.05). These findings unveil the underlying mechanism regulating intra-site leaf phenological diversity. While leaf phenology is known to be associated with ecosystem properties, our findings confirm that AGB is primarily influenced by functional trait composition and taxonomic diversity rather than leaf phenological diversity.

Effects of tree sex, maturity, local abiotic, and biotic neighborhoods on the growth of a subtropical dioecious tree species Diospyros morrisiana.

PREMISE: Understanding the drivers of the growth in long-lived woody trees is the key to predicting their responses to and maintaining their populations under global change. However, the role of tree sex and differential investment to reproduction are often not considered in models of individual tree growth, despite many gymnosperm and angiosperm species having separate male and female sexes. Thus, better models of tree growth should include tree sex and life stage along with the abiotic and biotic neighborhoods. METHODS: We used a sex-specific molecular marker to determine the sex of 2188 individual trees >1 cm DBH of the dioecious tree species Diospyros morrisiana in a 50-ha subtropical forest plot in China. We used long-term census data from about 300,000 trees, together with 625 soil samples and 2352 hemispherical photographs to characterize the spatially explicit biotic and abiotic neighborhoods. RESULTS: We found a male-biased effective sex ratio and a female-biased overall population sex ratio of D. morrisiana. No sex spatial segregation was detected for the overall population, mature, or immature trees. Immature trees grew faster than mature trees and females grew slower than males. Further, conspecific neighbors significantly decreased tree growth, while the abiotic neighborhood showed no significant effect. CONCLUSIONS: Our findings suggest that variation in resource allocation patterns within and across individual trees of different sexes and life-history stages should be more widely accounted for in models of tree growth. In addition, our study highlights the importance of sex-specific molecular markers for studying populations of long-lived dioecious tree species.

Interactions between all pairs of neighboring trees in 16 forests worldwide reveal details of unique ecological processes in each forest, and provide windows into their evolutionary histories.

When Darwin visited the Galapagos archipelago, he observed that, in spite of the islands' physical similarity, members of species that had dispersed to them recently were beginning to diverge from each other. He postulated that these divergences must have resulted primarily from interactions with sets of other species that had also diverged across these otherwise similar islands. By extrapolation, if Darwin is correct, such complex interactions must be driving species divergences across all ecosystems. However, many current general ecological theories that predict observed distributions of species in ecosystems do not take the details of between-species interactions into account. Here we quantify, in sixteen forest diversity plots (FDPs) worldwide, highly significant negative density-dependent (NDD) components of both conspecific and heterospecific between-tree interactions that affect the trees' distributions, growth, recruitment, and mortality. These interactions decline smoothly in significance with increasing physical distance between trees. They also tend to decline in significance with increasing phylogenetic distance between the trees, but each FDP exhibits its own unique pattern of exceptions to this overall decline. Unique patterns of between-species interactions in ecosystems, of the general type that Darwin postulated, are likely to have contributed to the exceptions. We test the power of our null-model method by using a deliberately modified data set, and show that the method easily identifies the modifications. We examine how some of the exceptions, at the Wind River (USA) FDP, reveal new details of a known allelopathic effect of one of the Wind River gymnosperm species. Finally, we explore how similar analyses can be used to investigate details of many types of interactions in these complex ecosystems, and can provide clues to the evolution of these interactions.

Species packing and the latitudinal gradient in 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 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.

Limited evidence of vertical fine-root segregation in a subtropical forest.

Vertical root segregation and the resulting niche partitioning can be a key underpinning of species coexistence. This could result from substantial interspecific variations in root profiles and rooting plasticity in response to soil heterogeneity and neighbours, but they remain largely untested in forest communities. In a diverse forest in subtropical China, we randomly sampled > 4000 root samples from 625 0-30 cm soil profiles. Using morphological and DNA-based methods, we identified 109 woody plant species, determined the degree of vertical fine-root segregation, and examined rooting plasticity in response to soil heterogeneity and neighbour structure. We found no evidence of vertical fine-root segregation among cooccurring species. By contrast, root abundance of different species tended to be positively correlated within soil zones. Underlying these findings was a lack of interspecific variation in fine-root profiles with over 90% of species concentrated in the 0-10 cm soil zone with only one species dominating in the 10-20 cm soil zone. Root profiles exhibited low responsiveness to root neighbours but tended to be shallow in soils with low phosphorus and copper content. These findings suggest that if there is niche differentiation leading to coexistence in this diverse forest, it would be occurring by mechanisms other than vertical fine-root segregation.

The complexity of trait-environment performance landscapes in a local subtropical forest.

That functional traits should affect individual performance and, in turn, determine fitness and population growth, is a foundational assumption of trait-based ecology. This assumption is, however, not supported by a strong empirical base. Here, we measured simultaneously two individual performance metrics (survival and growth), seven traits and 10 environmental properties for each of 3981 individuals of 205 species in a 50-ha stem-mapped subtropical forest. We then modelled survival/growth as a function of traits, environments and trait × environment interactions, and quantified their relative importance at both the species and individual levels. We found evidence of alternative functional designs and multiple performance peaks along environmental gradients, indicating the presence of complicated trait × environment interactions. However, such interactions were relatively unimportant in our site, which had relatively low environmental variations. Moreover, individual performance was not better predicted, and trait × environment interactions were not more likely detected, at the individual level than at the species level. Although the trait × environment interactions might be safely ignored in relatively homogeneous environments, we encourage future studies to test the interactive effects of traits and environments on individual performances and lifelong fitness at larger spatial scales or along experimentally manipulated environmental gradients.

Direct effects of selection on aboveground biomass contrast with indirect structure-mediated effects of complementarity in a subtropical forest.

Understanding the multiple biotic and abiotic controls of aboveground biomass (AGB) is important for projecting the consequences of global change and to effectively manage carbon storage. Although large-scale studies have identified the major environmental and biological controls of AGB, drivers of local-scale variation are less well known. Additionally, involvement of multiple causal paths and scale dependence in effect sizes potentially confounds comparisons among studies differing in methodology and sampling grain. We tested for scale dependence in evidence supporting selection, complementarity and environmental factors as the main determinants of AGB variation over a 50 ha study extent in subtropical China, modelling this at four sampling grains (0.01, 0.04, 0.25 and 1 ha). At each grain, we used piecewise structural equation models to quantify the direct and indirect effects of environmental (topographic and edaphic properties) and forest attributes (structure, diversity and functional traits) on AGB, while controlling for spatial autocorrelation. Direct scale-invariant effects on AGB were evident for structure and community-mean traits, supporting dominance of selection effects. However, diversity had strong indirect effects on AGB via forest structure, particularly at larger sampling grains (≥ 0.25 ha), while direct effects only emerged at the smallest grain size (0.01 ha). The direct and indirect effects of edaphic and topographic factors were also important for explaining both forest attributes and AGB across all scales. Although selection effects appeared to be more influential on ecosystem function, ignoring indirect causal pathways for diversity via structural attributes risks overlooking the importance of complementarity on ecosystem functioning, particularly as sampling grain increases.

Direct and indirect effects of climate on richness drive the latitudinal diversity gradient in forest trees.

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.

The effects of habitat filtering and non-habitat processes on species spatial distribution vary across life stages.

PREMISE OF THE STUDY: Habitat filtering and non-habitat processes are two major processes affecting spatial distributions of species. Because trees at different life stages perform differently, the life stage of tree species could play an important role in shaping the spatial distribution of species and community assembly. Here, we examined the possible changes of spatial distributions of species and evaluated the shifts in the relative importance of habitat filtering and non-habitat processes across life stages in a 50-ha subtropical forest plot in China. METHODS: We modeled species distribution with and without life stages using three point process models. The performance of these models, with and without considering life stages, was evaluated by comparing the species-area curve and the degree of clustering. The relative effects of habitat filtering and non-habitat processes across life stages were quantified using a spatial variance decomposition method. KEY RESULTS: The incorporation of life stage considerably improved the goodness-of-fit of these point process models at both the community and species levels. Non-habitat processes explained about 90% of the total variation in spatial distribution, while habitat filtering explained about 10%. The relative importance of habitat filtering only increased slightly from sapling to adult stages. CONCLUSIONS: Point process models performed better when life stages are included, indicating the importance of considering life stage when modeling spatial distributions for understanding community assembly. The finding that habitat acts weakly and non-habitat processes act dominantly in determining spatial distributions of species suggests a strong dependence of spatial patterns on non-habitat processes.

Root-centric ß diversity reveals functional homogeneity while phylogenetic heterogeneity in a subtropical forest.

Root-centric studies have revealed fast taxonomic turnover across root neighborhoods, but how such turnover is accompanied by changes in species functions and phylogeny (i.e., ß diversity) remains largely unknown. As ß diversity can reflect the degree of community-wide biotic homogenization, such information is crucial for better inference of below-ground assembly rules, community structuring, and ecosystem processes. We collected 2480 root segments from 625 0-30 cm soil profiles in a subtropical forest in China. Root segments were identified into 138 species with DNA-barcoding with six root morphological and architectural traits measured per species. By using the mean pairwise (Dpw ) and mean nearest neighbor distance (Dnn ) to quantify species ecological differences, we first tested the non-random functional and phylogenetic turnover of root neighborhoods that would lend more support to deterministic over stochastic community assembly processes. Additionally, we examined the distance-decay pattern of ß diversity, and finally partitioned ß diversity into geographical and environmental components to infer their potential drivers of environmental filtering, dispersal limitation, and biotic interactions. We found that functional turnover was often lower than expected given the taxonomic turnover, whereas phylogenetic turnover was often higher than expected. Phylogenetic Dpw (e.g., interfamily species) turnover exhibited a distance-decay pattern, likely reflecting limited dispersal or abiotic filtering that leads to the spatial aggregation of specific plant lineages. Conversely, both functional and phylogenetic Dnn (e.g., intrageneric species) exhibited an inverted distance-decay pattern, likely reflecting strong biotic interactions among spatially and phylogenetically close species leading to phylogenetic and functional divergence. While the spatial distance was generally a better predictor of ß diversity than environmental distance, the joint effect of environmental and spatial distance usually overrode their respective pure effects. These findings suggest that root neighborhood functional homogeneity may somewhat increase forest resilience after disturbance by exhibiting an insurance effect. Likewise, root neighborhood phylogenetic heterogeneity may enhance plant fitness by hindering the transmission of host-specific pathogens through root networks or by promoting interspecific niche complementarity not captured by species functions. Our study highlights the potential role of root-centric ß diversity in mediating community structures and functions largely ignored in previous studies.

Does local soil factor drive functional leaf trait variation? A test on Neilingding Island, South China.

Leaf traits were affected by soil factors and displayed varietal differences in forest. However, few examples have been reported on the Island ecosystems. We comprehensively investigated 9 leaf traits (leaf length, leaf width, leaf area, SLA, leaf fresh weight, leaf C content, leaf N content, leaf K content, leaf C:N ratio) of 54 main subtropical woody species and soil parameters (soil pH, total C content, total N content, total K content, available N content, available P content, available K content and soil moisture) in Neilingding Island, Shenzhen, southern China. Intra-and interspecific variation of leaf traits were measured and their correlations with soil parameters were explored. The interspecific variations of leaf C:N ratio, leaf N content and leaf fresh weight were higher than their intraspecific variations. The intraspecific variation of leaf K content was larger than that of interspecific one, accounting for 80.69% of the total variance. Positive correlations were found among intraspecific coefficients of variations in leaf morphological traits. The correlation analysis between the variation of intraspecific traits and the variation of soil parameters showed that changes in soil factors affected leaf morphology and stoichiometry. The interaction between soil moisture and soil available P content was the key factor on intraspecific variations of leaf traits including leaf area, leaf fresh weight, leaf C and leaf K content. We concluded that leaf traits of plants in the island were tightly related to soil parameters. Soil parameters, especially soil moisture and available P content, affected plant leaf morphology and stoichiometry at the local scale.

The unimodal intransitivity-fertility relationship is not mediated by demographic trade-offs in a subtropical forest.

Intransitive competition has long been acknowledged as a potential mechanism favoring species coexistence. However, its prevalence, variance along environmental gradients, and possible underlying mechanisms (trade-offs) in plant communities (especially in forests) has seldom been examined. A recently developed "reverse-engineering" approach based on Markov Chain allowed us to estimate the competitive transition matrices and competitive intransitivity from observational abundance data. Using this approach, we estimated competitive intransitivity of five dominant species in a subtropical forest and then related it to soil fertility (soil organic matter and soil pH) and demographic trade-offs (growth-survival and stature-recruitment trade-offs). In our forest plot, intransitive competition was common among the dominant species and peaked at the intermediate level of soil organic matter. Neither the growth-survival trade-off nor the stature-recruitment trade-off was positively related to competitive intransitivity. Our study for the first time empirically supported the unimodal intransitivity-fertility relationship in forests, which, however, was not mediated by the two demographic trade-offs in our plot.

Response of leaf functional traits to soil nutrients in the wet and dry seasons in a subtropical forest on an island.

Introduction: Island ecosystems often have a disproportionate number of endemic species and unique and fragile functional characteristics. However, few examples of this type of ecosystem have been reported. Methods: We conducted a comprehensive field study on Neilingding Island, southern China. The leaf samples of 79 subtropical forest tree species were obtained and their functional traits were studied in the dry and wet seasons to explain the relationships between plant functional traits and soil nutrients. Results: We found a greater availability of soil moisture content (SMC) and nutrients in the wet season than in the dry season. The values of wet season soil available phosphorus (5.97 mg·kg-1), SMC (17.67%), and soil available potassium (SAK, 266.96 mg·kg-1) were significantly higher than those of the dry season. The leaf dry matter content, specific leaf weight, leaf density, leaf total carbon, leaf total nitrogen, leaf total calcium, and the N/P and C/P ratios of leaves were all significantly higher in the dry season than in the wet season, being 18.06%, 12.90%, 12.00%, 0.17%, 3.41%, 9.02%, 26.80%, and 24.14% higher, respectively. In contrast, the leaf area (51.01 cm2), specific leaf area (152.76 cm2·g-1), leaf water content (0.59%), leaf total nitrogen (1.31%), leaf total phosphorus (0.14%), and leaf total magnesium (0.33%) were much lower in the dry season than in the wet one. There were significant pairwise correlations between leaf functional traits, but the number and strength of correlations were significantly different in the dry and wet seasons. The SAK, soil total phosphorus (STP), and pH impacted plant leaf functional traits in the dry season, whereas in the wet season, they were affected by SAK, STP, pH, and NO3- (nitrate). Discussion: Both soil nutrients and water availability varied seasonally and could cause variation in a number of leaf traits.

A DNA barcode library for woody plants in tropical and subtropical China.

The application of DNA barcoding has been significantly limited by the scarcity of reliable specimens and inadequate coverage and replication across all species. The deficiency of DNA barcode reference coverage is particularly striking for highly biodiverse subtropical and tropical regions. In this study, we present a comprehensive barcode library for woody plants in tropical and subtropical China. Our dataset includes a standard barcode library comprising the four most widely used barcodes (rbcL, matK, ITS, and ITS2) for 2,520 species from 4,654 samples across 49 orders, 144 families, and 693 genera, along with 79 samples identified at the genus level. This dataset also provides a super-barcode library consisting of 1,239 samples from 1,139 species, 411 genera, 113 families, and 40 orders. This newly developed library will serve as a valuable resource for DNA barcoding research in tropical and subtropical China and bordering countries, enable more accurate species identification, and contribute to the conservation and management of tropical and subtropical forests.

RecruitNet: A global database of plant recruitment networks.

Plant recruitment interactions (i.e., what recruits under what) shape the composition, diversity, and structure of plant communities. Despite the huge body of knowledge on the mechanisms underlying recruitment interactions among species, we still know little about the structure of the recruitment networks emerging in ecological communities. Modeling and analyzing the community-level structure of plant recruitment interactions as a complex network can provide relevant information on ecological and evolutionary processes acting both at the species and ecosystem levels. We report a data set containing 143 plant recruitment networks in 23 countries across five continents, including temperate and tropical ecosystems. Each network identifies the species under which another species recruits. All networks report the number of recruits (i.e., individuals) per species. The data set includes >850,000 recruiting individuals involved in 118,411 paired interactions among 3318 vascular plant species across the globe. The cover of canopy species and open ground is also provided. Three sampling protocols were used: (1) The Recruitment Network (RN) protocol (106 networks) focuses on interactions among established plants ("canopy species") and plants in their early stages of recruitment ("recruit species"). A series of plots was delimited within a locality, and all the individuals recruiting and their canopy species were identified; (2) The paired Canopy-Open (pCO) protocol (26 networks) consists in locating a potential canopy plant and identifying recruiting individuals under the canopy and in a nearby open space of the same area; (3) The Georeferenced plot (GP) protocol (11 networks) consists in using information from georeferenced individual plants in large plots to infer canopy-recruit interactions. Some networks incorporate data for both herbs and woody species, whereas others focus exclusively on woody species. The location of each study site, geographical coordinates, country, locality, responsible author, sampling dates, sampling method, and life habits of both canopy and recruit species are provided. This database will allow researchers to test ecological, biogeographical, and evolutionary hypotheses related to plant recruitment interactions. There are no copyright restrictions on the data set; please cite this data paper when using these data in publications.

Effects of local biotic neighbors and habitat heterogeneity on tree and shrub seedling survival in an old-growth temperate forest.

Seedling dynamics play a crucial role in determining species distributions and coexistence. Exploring causes of variation in seedling dynamics can therefore provide key insights into the factors affecting these phenomena. We examined the relative importance of biotic neighborhood processes and habitat heterogeneity using survival data for 5,827 seedlings in 39 tree and shrub species over 2 years from an old-growth temperate forest in northeastern China. We found significant negative density-dependence effects on survival of tree seedlings, and limited effects of habitat heterogeneity (edaphic and topographic variables) on survival of shrub seedlings. The importance of negative density dependence on young tree seedling survival was replaced by habitat in tree seedlings ≥ 4 years old. As expected, negative density dependence was more apparent in gravity-dispersed species compared to wind-dispersed and animal-dispersed species. Moreover, we found that a community compensatory trend existed for trees. Therefore, although negative density dependence was not as pervasive as in other forest communities, it is an important mechanism for the maintenance of community diversity in this temperate forest. We conclude that both negative density dependence and habitat heterogeneity drive seedling survival, but their relative importance varies with seedling age classes and species traits.

Highly spatial variation of soil microbial respiration and temperature sensitivity in a subtropical forest.

Quantifying the spatial variation and drivers of microbe-driven soil carbon (C) decomposition (also called soil microbial respiration, MR) and its temperature sensitivity (Q10) is crucial for reducing the uncertainty in modelling the terrestrial C cycle under global warming. To this end, most previous studies sampled soils from multiple sites at regional scales and incubated them at the same temperature level in the laboratory. However, this unified incubation temperature is too warm to the cold sites, and too cold to the warm sites, thus causing a large bias in the MR and Q10 estimations. Here, we conducted fine scale intensive sampling (194 soil samples) and measurements within a 4-ha subtropical forest plot to examine the underlying mechanisms driving the spatial pattern of MR and Q10. Our results showed that both MR and Q10 varied spatially within subtropical forests. The fine scale variation of MR was dominated by soil nitrogen concentration and slope position, and Q10 was dominated by soil fungi abundance. Overall, the 35 investigated biotic and abiotic factors explained 38% of the spatial variation of MR and 9% of the spatial variation of Q10 in the subtropical forest. This suggests that the fine scale variation of soil C dynamics is much more complex than that at the regional scale reported in previous studies, which should be considered in the assessments of terrestrial soil C cycles.

Plastome-based phylogeny improves community phylogenetics of subtropical forests in China.

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.

Arbuscular mycorrhizal trees influence the latitudinal beta-diversity gradient of tree communities in forests worldwide.

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.

Experimental Evidence for the Importance of Light on Understory Grass Communities in a Subtropical Forest.

Light is one of the most important environmental filters for forest understory grass communities. It is predicted that light can select species with the same light requirements, resulting in a decrease in species compositional dissimilarity among grass communities experiencing the same light intensity, and an increase in community dissimilarity under variable light intensities. However, these predictions have been questioned recently in light of modern coexistence theories, and evidence for them in natural communities is often indistinguishable from patterns created by dispersal limitation and biotic interactions. To help fill this gap, we sampled 48 understory grass communities that had regenerated from the same soil seed bank in Southern China. Plots were established under a light intensity gradient. Changes in species composition and neighborhood densities were monitored over a growing season. Our experimental setup controls for bias from dispersal limitation and is useful for detecting the effects of biotic interactions at different intensities of light. As expected, (1) compositional dissimilarity of grass communities increased between communities with different light intensities. The extent to which communities became more dissimilar was positively correlated with the difference in the light intensity. (2) No significant change in compositional dissimilarity was observed among communities experiencing the same light intensity. (3) Finally, relative neighborhood density significantly decreased in communities with moderate to high shading treatments. Our results clearly show that light can drive compositional divergence among communities under different light densities. However, the light may not lead to convergence among communities experiencing the same low light intensity, because intense competition induced by low light might enlarge species compositional differences, as shown with the neighborhood density analysis. Therefore, our study provides more convincing evidence for the importance of light on understory grass communities in subtropical forests and highlights the need to jointly consider biotic interactions when testing for evidence for environmental filtering.