Haibao Ren

Partitioning beta diversity in a subtropical broad-leaved forest of China.

The classical environmental control model assumes that species distribution is determined by the spatial variation of underlying habitat conditions. This niche-based model has recently been challenged by the neutral theory of biodiversity which assumes that ecological drift is a key process regulating species coexistence. Understanding the mechanisms that maintain biodiversity in communities critically depends on our ability to decompose the variation of diversity into the contributions of different processes affecting it. Here we investigated the effects of pure habitat, pure spatial, and spatially structured habitat processes on the distributions of species richness and species composition in a recently established 24-ha stem-mapping plot in the subtropical evergreen broad-leaved forest of Gutianshan National Nature Reserve in East China. We used the new spatial analysis method of principal coordinates of neighbor matrices (PCNM) to disentangle the contributions of these processes. The results showed that (1) habitat and space jointly explained approximately 53% of the variation in richness and approximately 65% of the variation in species composition, depending on the scale (sampling unit size); (2) tree diversity (richness and composition) in the Gutianshan forest was dominantly controlled by spatially structured habitat (24%) and habitat-independent spatial component (29%); the spatially independent habitat contributed a negligible effect (6%); (3) distributions of richness and species composition were strongly affected by altitude and terrain convexity, while the effects of slope and aspect were weak; (4) the spatial distribution of diversity in the forest was dominated by broad-scaled spatial variation; (5) environmental control on the one hand and unexplained spatial variation on the other (unmeasured environmental variables and neutral processes) corresponded to spatial structures with different scales in the Gutianshan forest plot; and (6) five habitat types were recognized; a few species were statistically significant indicators of three of these habitats, whereas two habitats had no significant indicator species. The results suggest that the diversity of the forest is equally governed by environmental control (30%) and neutral processes (29%). In the fine-scale analysis (10 x 10 m cells), neutral processes dominated (43%) over environmental control (20%).

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.

Community-level consequences of density dependence and habitat association in a subtropical broad-leaved forest

How extraordinary numbers of species can coexist in hyper-diverse communities remains unresolved. While numerous hypotheses have been proposed based on observational and theoretical investigations, little is known about which mechanisms are truly active in forest communities and less is known about their relative contributions to community assembly. In this study, generalized linear mixed models with crossed random effects were used to assess the relative contributions of density dependence and habitat association to community-level diversity maintenance. Species habitat associations were classified based on soil nutrients, topography and species composition. Local neighbourhood effects were also addressed with spatially explicit models of seedling survival. The results shown here reveal that local- and community-level seedling dynamics were consistent with density-dependent predictions, although habitat association played a more important role in shaping short-term seedling survival. We conclude that density dependence could promote species coexistence on the premise of habitat partitioning.

The use of the Mexican Hat and the Morlet wavelets for detection of ecological patterns

In this paper, we compare the relationship between scale and period in ecological pattern analysis and wavelet analysis. We also adapt a commonly used wavelet, the Morlet, to ecological pattern analysis. Using Monte Carlo assessments, we apply methods of statistical significance test to wavelet analysis for pattern analysis. In order to understand the inherent strength and weakness of the Morlet and the Mexican Hat wavelets, we also investigate and compare the properties of two frequently used wavelets by testing with field data and four artificial transects of different typical patterns which is often encountered in ecological research. It is shown that the Mexican Hat provides better detection and localization of patch and gap events over the Morlet, whereas the Morlet offers improved detection and localization of scale over the Mexican Hat. There is always a trade-off between the detection and localization of scale versus patch and gap events. Therefore, the best composite analysis is the combination of their advantages. The properties of wavelet in dealing with ecological data may be affected by characteristics intrinsic to wavelet itself. The peaks of different scales in isograms of wavelet power spectrum from the Mexican Hat may overlap with each other. Alternatively, these peaks of different scales in isograms of wavelet power spectrum may combine with each other unless the size of the analyzed scales is significantly different. These overlapping or combining lead to combining of peaks for different scales, or the masking of trough between peaks of different scales in the scalogram. Ecologists should combine all the information in scalogram and isograms of wavelet coefficient and wavelet power spectrum from different wavelets, which can provide us a broader view and precise pattern information.

The Contribution of Rare Species to Community Phylogenetic Diversity across a Global Network of Forest Plots

Niche differentiation has been proposed as an explanation for rarity in species assemblages. To test this hypothesis requires quantifying the ecological similarity of species. This similarity can potentially be estimated by using phylogenetic relatedness. In this study, we predicted that if niche differentiation does explain the co-occurrence of rare and common species, then rare species should contribute greatly to the overall community phylogenetic diversity (PD), abundance will have phylogenetic signal, and common and rare species will be phylogenetically dissimilar. We tested these predictions by developing a novel method that integrates species rank abundance distributions with phylogenetic trees and trend analyses, to examine the relative contribution of individual species to the overall community PD. We then supplement this approach with analyses of phylogenetic signal in abundances and measures of phylogenetic similarity within and between rare and common species groups. We applied this analytical approach to 15 long-term temperate and tropical forest dynamics plots from around the world. We show that the niche differentiation hypothesis is supported in six of the nine gap-dominated forests but is rejected in the six disturbance-dominated and three gap-dominated forests. We also show that the three metrics utilized in this study each provide unique but corroborating information regarding the phylogenetic distribution of rarity in communities.

Covariation in Plant Functional Traits and Soil Fertility within Two Species-Rich Forests

The distribution of plant species along environmental gradients is expected to be predictable based on organismal function. Plant functional trait research has shown that trait values generally vary predictably along broad-scale climatic and soil gradients. This work has also demonstrated that at any one point along these gradients there is a large amount of interspecific trait variation. The present research proposes that this variation may be explained by the local-scale sorting of traits along soil fertility and acidity axes. Specifically, we predicted that trait values associated with high resource acquisition and growth rates would be found on soils that are more fertile and less acidic. We tested the expected relationships at the species-level and quadrat-level (20×20 m) using two large forest plots in Panama and China that contain over 450 species combined. Predicted relationships between leaf area and wood density and soil fertility were supported in some instances, but the majority of the predicted relationships were rejected. Alternative resource axes, such as light gradients, therefore likely play a larger role in determining the interspecific variability in plant functional traits in the two forests studied.

The Relative Importance of Janzen-Connell Effects in Influencing the Spatial Patterns at the Gutianshan Subtropical Forest

The Janzen-Connell hypothesis is among the most important theories put forward to explain species coexistence in species-rich communities. However, the relative importance of Janzen-Connell effects with respect to other prominent mechanisms of community assembly, such as dispersal limitation, self-thinning due to competition, or habitat association, is largely unresolved. Here we use data from a 24-ha Gutianshan subtropical forest to address it. First we tested for significant associations of adults, juveniles, and saplings with environmental variables. Second we evaluated if aggregation decreased with life stage. In a third analysis we approximately factored out the effect of habitat association and comprehensively analyzed the spatial associations of intraspecific adults and offspring (saplings, juveniles) of 46 common species at continuous neighborhood distances. We found i) that, except for one, all species were associated with at least one environmental variable during at least one of their life stages, but the frequency of significant habitat associations declined with increasing life stage; ii) a decline in aggregation with increasing life stage that was strongest from juveniles to adults; and iii) intraspecific adult-offspring associations were dominated by positive relationships at neighborhood distances up to 10 m. Our results suggest that Janzen-Connell effects were not the dominant mechanisms in structuring the spatial patterns of established trees in the subtropical Gutianshan forest. The spatial patterns may rather reflect the joint effects of size-dependent self-thinning, dispersal limitation and habitat associations. Our findings contribute to a more comprehensive understanding of the relative importance of Janzen-Connell effects in influencing plant community structure under strong topographic heterogeneity.

Geographical Range and Local Abundance of Tree Species in China

Most studies on the geographical distribution of species have utilized a few well-known taxa in Europe and North America, with little research in China and its wide range of climate and forest types. We assembled large datasets to quantify the geographic ranges of tree species in China and to test several biogeographic hypotheses: 1) whether locally abundant species tend to be geographically widespread; 2) whether species are more abundant towards their range-centers; and 3) how abundances are correlated between sites. Local abundances of 651 species were derived from four tree plots of 20–25 ha where all individuals ≥1 cm in stem diameter were mapped and identified taxonomically. Range sizes of these species across China were then estimated from over 460,000 geo-referenced records; a Bayesian approach was used, allowing careful measures of error of each range estimate. The log-transformed range sizes had a bell-shaped distribution with a median of 703,000 km2, and >90% of 651 species had ranges >105 km2. There was no relationship between local abundance and range size, and no evidence for species being more abundant towards their range-centers. Finally, species’ abundances were positively correlated between sites. The widespread nature of most tree species in China suggests few are vulnerable to global extinction, and there is no indication of the double-peril that would result if rare species also had narrow ranges.

Stochastic assembly in a subtropical forest chronosequence: evidence from contrasting changes of species, phylogenetic and functional dissimilarity over succession.

Deterministic and stochastic processes jointly determine the community dynamics of forest succession. However, it has been widely held in previous studies that deterministic processes dominate forest succession. Furthermore, inference of mechanisms for community assembly may be misleading if based on a single axis of diversity alone. In this study, we evaluated the relative roles of deterministic and stochastic processes along a disturbance gradient by integrating species, functional, and phylogenetic beta diversity in a subtropical forest chronosequence in Southeastern China. We found a general pattern of increasing species turnover, but little-to-no change in phylogenetic and functional turnover over succession at two spatial scales. Meanwhile, the phylogenetic and functional beta diversity were not significantly different from random expectation. This result suggested a dominance of stochastic assembly, contrary to the general expectation that deterministic processes dominate forest succession. On the other hand, we found significant interactions of environment and disturbance and limited evidence for significant deviations of phylogenetic or functional turnover from random expectations for different size classes. This result provided weak evidence of deterministic processes over succession. Stochastic assembly of forest succession suggests that post-disturbance restoration may be largely unpredictable and difficult to control in subtropical forests.

Effects of historical logging on soil microbial communities in a subtropical forest in southern China

Background and AimsGaining a better understanding of the legacy effects of logging and forest restoration on soil microbial communities could improve our ability to conserve biodiversity and promote ecosystem sustainability. Herein, we investigated how soil microbial community is linked to natural, restored, and planted forests and the legacies of historical forest.MethodsSoil microbial biomass and composition were measured in four forest types (i.e., primary forest, once-clearcut forest, twice-logged forest, and plantation forest) and related to physico-chemical soil properties and forest community structure data by using analysis of covariance.ResultsFungal, bacterial, and total microbial biomass measured by phospholipid fatty acid profiles were significantly lower in the two secondary forests and the plantation than in the primary forest. The conversion of vegetation and soil regimes due to forest logging altered microbial communities.ConclusionsOur findings elucidate the correlation of plant communities and soil characteristics to soil microbial communities in the context of subtropical forest management. Naturally restored and planted forests may affect soil microorganisms largely by directly modifying the soil labile C and N fractions of organic matter.