Inverse relationship between species competitiveness and intraspecific trait variability may enable species coexistence in experimental seedling communities.

Theory suggests that intraspecific trait variability may promote species coexistence when competitively inferior species have higher intraspecific trait variability than their superior competitors. Here, we provide empirical evidence for this phenomenon in tree seedlings. We evaluated intraspecific variability and plastic response of ten traits in 6750 seedlings of ten species in a three-year greenhouse experiment. While we observed no relationship between intraspecific trait variability and species competitiveness in competition-free homogeneous environments, an inverse relationship emerged under interspecific competition and in spatially heterogeneous environments. We showed that this relationship is driven by the plastic response of the competitively inferior species: Compared to their competitively superior counterparts, they exhibited a greater increase in trait variability, particularly in fine-root traits, in response to competition, environmental heterogeneity and their combination. Our findings contribute to understanding how interspecific competition and intraspecific trait variability together structure plant communities.

Responses of tree seedlings to understory filtering by the recalcitrant fern layer in a subtropical forest.

The recalcitrant understory fern layer is an important ecological filter for seedling regeneration, yet how the fern layer influences seedling regeneration dynamics remains unclear. Here we transplanted 576 seedlings of four dominant tree species, Castanopsis fargesii, Lithocarpus glaber, Schima superba and Hovenia acerba, to the treatments of Diplopterygium glaucum retention and removal under an evergreen broad-leaved forest in eastern China. We monitored the survival, growth and biomass data of these seedlings for 28 months, and then used generalized linear mixed models to evaluate the treatment effects on seedling survival, growth, biomass and root-shoot ratio. Our results showed that fern retention significantly inhibited the seedling establishment of all four species. During the seedling development stage, the seedling relative growth rate of L. glaber decreased under fern retention, which was not the case for the other three species. Root-shoot ratio of C. fargesii and L. glaber increased significantly under fern retention. Our findings provide new evidence of the filtering effect of a recalcitrant fern understory. Notably, we observed that the response of tree seedlings to the recalcitrant fern understory was more sensitive in the establishment stage. Finally, our work highlights that the filtering effect of the recalcitrant fern understory changes depending on the regeneration stages, and that shade-tolerant species, C. fargesii and L. glaber were even more affected by fern disturbed habitats, suggesting that effective management should attempt to curb forest fern outbreaks, thus unblocking forest recruitment.

Differences in Density Dependence among Tree Mycorrhizal Types Affect Tree Species Diversity and Relative Growth Rates.

Conspecific negative density dependence (CNDD) may vary by tree mycorrhizal type. However, whether arbuscular mycorrhizal (AM)-associated tree species suffer from stronger CNDD than ectomycorrhizal (EcM) and ericoid mycorrhizal (ErM)-associated tree species at different tree life stages, and whether EcM tree species can promote AM and ErM saplings and adults growth, remain to be studied. Based on the subtropical evergreen broad-leaved forest data in eastern China, the generalized linear mixed-effects model was used to analyze the effects of the conspecific density and heterospecific density grouped by symbiont mycorrhizal type on different tree life stages of different tree mycorrhizal types. The results showed that compared to other tree mycorrhizal types at the same growth stage, EcM saplings and AM adults experienced stronger CNDD. Heterospecific EcM density had a stronger positive effect on AM and ErM individuals. Species diversity and average relative growth rate (RGR) first increased and then decreased with increasing basal area (BA) ratios of EcM to AM tree species. These results suggested that the stronger CNDD of EcM saplings and AM adults favored local species diversity over other tree mycorrhizal types. The EcM tree species better facilitated the growth of AM and ErM tree species in the neighborhood, increasing the forest carbon sink rate. Interestingly, species diversity and average RGR decreased when EcM or AM tree species predominated. Therefore, our study highlights that manipulating the BA ratio of EcM to AM tree species will play a nonnegligible role in maintaining biodiversity and increasing forest carbon sink rates.

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.

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.

Large Underestimation of Intraspecific Trait Variation and Its Improvements.

Intraspecific trait variation (ITV) is common feature of natural communities and has gained increasing attention due to its significant ecological effects on community dynamics and ecosystem functioning. However, the estimation of ITV per se has yet to receive much attention, despite the need for accurate ITV estimation for trait-based ecological inferences. It remains unclear if, and to what extent, current estimations of ITV are biased. The most common method used to quantify ITV is the coefficient of variation (CV), which is dimensionless and can therefore be compared across traits, species, and studies. Here, we asked which CV estimator and data normalization method are optimal for quantifying ITV, and further identified the minimum sample size required for ±5% accuracy assuming a completely random sample scheme. To these ends, we compared the performance of four existing CV estimators, together with new simple composite estimators, across different data normalizations, and sample sizes using both a simulated and empirical trait datasets from local to regional scales. Our results consistently showed that the most commonly used ITV estimator (CV 1= σsample /µsample ), often underestimated ITV-in some cases by nearly 50%-and that underestimation varies largely among traits and species. The extent of this bias depends on the sample size, skewness and kurtosis of the trait value distribution. The bias in ITV can be substantially reduced by using log-transforming trait data and alternative CV estimators that take into consideration the above dependencies. We find that the CV4 estimator, also known as Bao's CV estimator, combined with log data normalization, exhibits the lowest bias and can reach ±5% accuracy with sample sizes greater than 20 for almost all examined traits and species. These results demonstrated that many previous ITV measurements may be substantially underestimated and, further, that these underestimations are not equal among species and traits even using the same sample size. These problems can be largely solved by log-transforming trait data first and then using the Bao's CV to quantify ITV. Together, our findings facilitate a more accurate understanding of ITV in community structures and dynamics, and may also benefit studies in other research areas that depend on accurate estimation of CV.

Distance-based methods for estimating density of nonrandomly distributed populations.

Population density is the most basic ecological parameter for understanding population dynamics and biological conservation. Distance-based methods (or plotless methods) are considered as a more efficient but less robust approach than quadrat-based counting methods in estimating plant population density. The low robustness of distance-based methods mainly arises from the oversimplistic assumption of completely spatially random (CSR) distribution of a population in the conventional distance-based methods for estimating density of non-CSR populations in natural communities. In this study we derived two methods to improve on density estimation for plant populations of non-CSR distribution. The first method modified an existing composite estimator to correct for the long-recognized bias associated with that estimator. The second method was derived from the negative binomial distribution (NBD) that directly deals with aggregation in the distribution of a species. The performance of these estimators was tested and compared against various distance-based estimators by both simulation and empirical data of three large-scale stem-mapped forests. Results showed that the NBD point-to-tree distance estimator has the best and most consistent performance across populations with vastly different spatial distributions. This estimator offers a simple, efficient and robust method for estimating density for empirical populations of plant species.

Distance-dependent seed‒seedling transition in the tree Castanopsis sclerophylla is altered by fragment size.

Negative distance-dependence of conspecific seedling mortality (NDisDM) is a crucial stabilizing force that regulates plant diversity, but it remains unclear whether and how fragment size shifts the strength of NDisDM. Here, we surveyed the seed‒seedling transition process for a total of 25,500 seeds of a local dominant tree species on islands of various sizes in a reservoir and on the nearby mainland. We found significant NDisDM on the mainland and large and medium islands, with significantly stronger NDisDM on medium islands. However, positive distance-dependent mortality was detected on small islands. Changes in distance-dependence were critically driven by both rodent attack and pathogen infestation, which were significantly affected by fragment size. Our results emphasize the necessity of incorporating the effects of fragment size on distance-dependent regeneration of dominant plant species into the existing frameworks for better predicting the consequences of habitat fragmentation.

Species with moderate intraspecific trait variability are locally abundant within an environmentally heterogeneous subtropical forest.

Species with large intraspecific trait variability (ITV) have larger niche breadth than species with low ITV and thus are expected to be more abundant at the local scale. However, whether the positive ITV-abundance relationship holds in heterogeneous local environments remains uncertain. Using an individual-based trait dataset encompassing three leaf traits (leaf area, specific leaf area, and leaf dry mass content) of 20,248 individuals across 80 species in an environmentally heterogeneous subtropical forest in eastern China, ITV for each trait of each species was estimated by rarefaction. Resource-based niche breadth and marginality (the absolute distance between the mean resource states used by a species and the mean plot-wise resource states) were estimated simultaneously by the K-S method and the outlying mean index, respectively. Species with moderate ITV were often locally abundant, while species with large or small ITV were locally rare. This unimodal relationship between ITV and species abundance persisted when traits were analyzed separately and for all tree size classes. There was also a hump-backed relationship between niche breadth and marginality, and ITV was positively associated with niche breadth. The combined results suggest either a trade-off between the benefit from expanding niche breadth to adapt to multiple habitats and the disadvantage of reducing competitive ability, or a scarcity of favorable resources. Our results do not support the traditional thought that ITV positively correlates with species abundance in heterogeneous local environments. Instead, our study suggests that moderate-rather than large-intraspecific trait variability increases species abundance at local scales.

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.

Spatial scale changes the relationship between beta diversity, species richness and latitude.

The relationship between ß-diversity and latitude still remains to be a core question in ecology because of the lack of consensus between studies. One hypothesis for the lack of consensus between studies is that spatial scale changes the relationship between latitude and ß-diversity. Here, we test this hypothesis using tree data from 15 large-scale forest plots (greater than or equal to 15 ha, diameter at breast height ≥ 1 cm) across a latitudinal gradient (3-30o) in the Asia-Pacific region. We found that the observed ß-diversity decreased with increasing latitude when sampling local tree communities at small spatial scale (grain size ≤0.1 ha), but the observed ß-diversity did not change with latitude when sampling at large spatial scales (greater than or equal to 0.25 ha). Differences in latitudinal ß-diversity gradients across spatial scales were caused by pooled species richness (γ-diversity), which influenced observed ß-diversity values at small spatial scales, but not at large spatial scales. Therefore, spatial scale changes the relationship between ß-diversity, γ-diversity and latitude, and improving sample representativeness avoids the γ-dependence of ß-diversity.

Conspecific Leaf Litter-Mediated Effect of Conspecific Adult Neighborhood on Early-Stage Seedling Survival in A Subtropical Forest.

Conspecific adults have strong negative effect on the survival of nearby early-stage seedlings and thus can promote species coexistence by providing space for the regeneration of heterospecifics. The leaf litter fall from the conspecific adults, and it could mediate this conspecific negative adult effect. However, field evidence for such effect of conspecific leaf litter remains absent. In this study, we used generalized linear mixed models to assess the effects of conspecific leaf litter on the early-stage seedling survival of four dominant species (Machilus leptophylla, Litsea elongate, Acer pubinerve and Distylium myricoides) in early-stage seedlings in a subtropical evergreen broad-leaved forest in eastern China. Our results consistently showed that the conspecific leaf litter of three species negatively affected the seedling survival. Meanwhile, the traditional conspecific adult neighborhood indices failed to detect this negative conspecific adult effect. Our study revealed that the accumulation of conspecific leaf litter around adults can largely reduce the survival rate of nearby seedlings. Ignoring it could result in underestimation of the importance of negative density dependence and negative species interactions in the natural forest communities.

Detangling the Effects of Environmental Filtering and Dispersal Limitation on Aggregated Distributions of Tree and Shrub Species: Life Stage Matters.

The pervasive pattern of aggregated tree distributions in natural communities is commonly explained by the joint effect of two clustering processes: environmental filtering and dispersal limitation, yet little consensus remains on the relative importance of the two clustering processes on tree aggregations. Different life stages of examined species were thought to be one possible explanation of this disagreement, because the effect of environmental filtering and dispersal limitation are expected to increase and decrease with tree life stages, respectively. However, few studies have explicitly tested these expectations. In this study, we evaluated these expectations by three different methods (species-habitat association test based on Poisson Clustering model and spatial point pattern analyses based on Heterogeneous Poisson model and the jointly modeling approach) using 36 species in a 20-ha subtropical forest plot. Our results showed that the percentage of species with significant habitat association increased with life stages, and there were fewer species affected by dispersal limitation in later life stages compared with those in earlier stages. Percentage of variance explained by the environmental filtering and dispersal limitation also increases and decreases with life stages. These results provided a promising alternative explanation on the existing mixed results about the relative importance of the two clustering processes. These findings also highlighted the importance of plant life stages for fully understanding species distributions and species coexistence.

Quantifying effects of habitat heterogeneity and other clustering processes on spatial distributions of tree species.

Spatially explicit consideration of species distribution can significantly add to our understanding of species coexistence. In this paper, we evaluated the relative importance of habitat heterogeneity and other clustering processes (e.g., dispersal limitation, collectively called the non-habitat clustering process) in explaining the spatial distribution patterns of 341 tree species in three stem-mapped 25-50 ha plots of tropical, subtropical, and temperate forests. Their relative importance was estimated by a method that can take one mechanism into account when estimating the effects of the other mechanism and vice versa. Our results demonstrated that habitat heterogeneity was less important in explaining the observed species patterns than other clustering processes in plots with flat topography but was more important in one of the three plots that had a complex topography. Meanwhile, both types of clustering mechanisms (habitat or non-habitat) were pervasive among species at the 50-ha scale across the studied plots. Our analyses also revealed considerable variation among species in the relative importance of the two types of mechanism within each plot and showed that this species-level variation can be partially explained by differences in dispersal mode and growth form of species in a highly heterogeneous environment. Our findings provide new perspectives on the formation of species clustering. One important finding is that a significant species-habitat association does not necessarily mean that the habitat heterogeneity has a decisive influence on species distribution. The second insight is that the large species-level variation in the relative importance of the two types of clustering mechanisms should not be ignored. Non-habitat clustering processes can play an important role on species distribution.

[Dynamics of soil arthropod community structure and its responses to forest fragmentation during the decomposition of Castanopsis eyrei leaf litter].

Five evergreen broad-leaved forests (one continuous forest and four fragmented forests) in the mountain areas in the juncture of Zhejiang, Fujian, and Jiangxi Provinces, East China were selected as test objects to study the dynamics of soil arthropod community structure and its responses to forest fragmentation during the decomposition of dominant tree species Castanopsis eyrei leaf litter. A total of 899 soil arthropods were collected, belonging to 9 classes and 25 orders. Lepidoptera was the dominant taxon, accounting for 10% of the individual, while Hymenoptera, Collembola, Diptera, Prostigmata, and Geophilomorpha were the common taxa. The decomposition rate of C. eyrei leaf litter was the highest in August and lower in April-June and December, which was in accordance with the seasonal dynamics of the taxa number and individual number of soil arthropods. Meanwhile, the taxa number, individual number, and species diversity of soil arthropods differed between continuous forest and fragmented forests, suggesting that both area effect and edge effect affected the dynamics of soil arthropod community structure during the decomposition of C. eyrei leaf litter.

[Decomposition of Schima superba leaf litter and dynamics change of soil meso-micro arthropods community structure in evergreen broad-leaved forest fragments].

Taking four evergreen broad-leaved forest fragments within the adjacent mountainous region of Zhejiang, Fujian, and Jiangxi provinces as study objects, and the continuous forest in Gutianshan National Natural Reserve as the control, an investigation was made by litter bag method from June 2004 to April 2006, aimed to understand the community structure and its dynamics change of soil meso-micro arthropods during the decomposition of Schima superba leaf litter. A total of 1050 soil meso-micro arthropods belonging to 8 classes and 23 orders were collected, among which, Lepidoptera, Hymenoptera, Collembola and Diptera were the dominant taxa. The impact of habitat fragmentation on the community composition of soil arthropods was mainly manifested in the differences of rare taxa, and the dominance of different taxa at different decomposition stages of S. superba leaf litter varied with the functions of the taxa in litter decomposition. After two-year decomposition, the mass loss of S. superba leaf litter was 60%-70%, and the species diversity indices of soil arthropods showed certain changes, being different between forest fragments and continuous forest.

Species-area relationships explained by the joint effects of dispersal limitation and habitat heterogeneity.

Species-area relationships (SARs) characterize the spatial distribution of species diversity in community ecology, but the biological mechanisms underlying the SARs have not been fully explored. Here, we examined the roles of dispersal limitation and habitat heterogeneity in shaping SARs in two large-scale forest plots. One is a 24-ha subtropical forest in Gutianshan National Nature Reserve, China. The other is a 50-ha tropical rain forest in Barro Colorado Island, Panama. Spatial point pattern models were applied to investigate the contributions of dispersal and habitat heterogeneity and their interactions to the formation of the SARs in the two sites. The results showed that, although dispersal and habitat heterogeneity each could significantly contribute to the SARs, each alone was insufficient to explain the SARs. Their joint effects sufficiently explained the real SARs, suggesting that heterogeneous habitat and dispersal limitation are two predominant mechanisms for maintaining the spatial distributions of the species in these two forests. These results add to our understanding of the ecological processes underlying the spatial variation of SARs in natural forest communities.