Martin Baruffol

Community assembly during secondary forest succession in a Chinese subtropical forest

Subtropical broad-leaved forests in southeastern China support a high diversity of woody plants. Using a comparative study design with 30 × 30 m plots (n = 27) from five successional stages ( 1 m in height in each plot and counted all woody recruits (bank of all seedlings ≤1 m in height) in each central 10 × 10 m quadrant of each plot. In addition, we measured a number of environmen...

Globally, functional traits are weak predictors of juvenile tree growth, and we do not know why

Additional co-authors: Herve Jactel, Xuefei Li, Kaoru Kitajima, Julia Koricheva, Cristina Martinez-Garza, Christian Messier, Alain Paquette, Christopher Philipson, Daniel Piotto, Lourens Poorter, Juan M. Posada, Catherine Potvin, Kalle Rainio, Sabrina E. Russo, Mariacarmen Ruiz-Jaen, Michael Scherer-Lorenzen, Campbell O. Webb, S. Joseph Wright, Rakan A. Zahawi, and Andy Hector

A guide to analyzing biodiversity experiments

Aims: The aim of this guide is to provide practical help for ecologists who analyze data from biodiversity–ecosystem functioning experiments. Our approach differs from others in the use of least squares-based linear models (LMs) together with restricted maximum likelihood-based mixed models (MMs) for the analysis of hierarchical data. An original data set containing diameter and height of young trees grown in monocultures, 2- or 4-species mixtures under ambient light or shade is used as an example. Methods: Starting with a simple LM, basic features of model fitting and the subsequent analysis of variance (ANOVA) for significance tests are summarized. From this, more complex models are developed. We use the statistical software R for model fitting and to demonstrate similarities and complementarities between LMs and MMs. The formation of contrasts and the use of error (LMs) or random-effects (MMs) terms to account for hierarchical data structure in ANOVAs are explained. Important Findings: Data from biodiversity experiments can be analyzed at the level of entire plant communities (plots) and plant individuals. The basic explanatory term is species composition, which can be divided into contrasts in many ways depending on specific biological hypotheses. Typically, these contrasts code for aspects of species richness or the presence of particular species. For significance tests in ANOVAs, contrast terms generally are compared with remaining variation of the explanatory terms from which they have been ‘carved out’. Once a final model has been selected, parameters (e.g. means or slopes for fixed-effects terms and variance components for error or random-effects terms) can be estimated to indicate the direction and size of effects.

Kinetic energy of Throughfall in subtropical forests of SE China - effects of tree canopy structure, functional traits, and biodiversity.

Throughfall kinetic energy (TKE) plays an important role in soil erosion in forests. We studied TKE as a function of biodiversity, functional diversity as well as structural stand variables in a secondary subtropical broad-leaved forest in the Gutianshan National Nature Reserve (GNNR) in south-east China, a biodiversity hotspot in the northern hemisphere with more than 250 woody species present. Using a mixed model approach we could identify significant effects of all these variables on TKE: TKE increased with rarefied tree species richness and decreased with increasing proportion of needle-leaved species and increasing leaf area index (LAI). Furthermore, for average rainfall amounts TKE was decreasing with tree canopy height whereas for high rainfall amounts this was not the case. The spatial pattern of throughfall was stable across several rain events. The temporal variation of TKE decreased with rainfall intensity and increased with tree diversity. Our results show that more diverse forest stands over the season have to cope with higher cumulative raindrop energy than less diverse stands. However, the kinetic energy (KE) of one single raindrop is less predictable in diverse stands since the variability in KE is higher. This paper is the first to contribute to the understanding of the ecosystem function of soil erosion prevention in diverse subtropical forests.

Woody plant phylogenetic diversity mediates bottom–up control of arthropod biomass in species-rich forests

Global change is predicted to cause non-random species loss in plant communities, with consequences for ecosystem functioning. However, beyond the simple effects of plant species richness, little is known about how plant diversity and its loss influence higher trophic levels, which are crucial to the functioning of many species-rich ecosystems. We analyzed to what extent woody plant phylogenetic diversity and species richness contribute to explaining the biomass and abundance of herbivorous and predatory arthropods in a species-rich forest in subtropical China. The biomass and abundance of leaf-chewing herbivores, and the biomass dispersion of herbivores within plots, increased with woody plant phylogenetic diversity. Woody plant species richness had much weaker effects on arthropods, but interacted with plant phylogenetic diversity to negatively affect the ratio of predator to herbivore biomass. Overall, our results point to a strong bottom–up control of functionally important herbivores mediated particularly by plant phylogenetic diversity, but do not support the general expectation that top–down predator effects increase with plant diversity. The observed effects appear to be driven primarily by increasing resource diversity rather than diversity-dependent primary productivity, as the latter did not affect arthropods. The strong effects of plant phylogenetic diversity and the overall weaker effects of plant species richness show that the diversity-dependence of ecosystem processes and interactions across trophic levels can depend fundamentally on non-random species associations. This has important implications for the regulation of ecosystem functions via trophic interaction pathways and for the way species loss may impact these pathways in species-rich forests.

Tree Diversity Enhances Stand Carbon Storage but Not Leaf Area in a Subtropical Forest

Research about biodiversity–productivity relationships has focused on herbaceous ecosystems, with results from tree field studies only recently beginning to emerge. Also, the latter are concentrated largely in the temperate zone. Tree species diversity generally is much higher in subtropical and tropical than in temperate or boreal forests, with reasons not fully understood. Niche overlap and thus complementarity in the use of resources that support productivity may be lower in forests than in herbaceous ecosystems, suggesting weaker productivity responses to diversity change in forests. We studied stand basal area, vertical structure, leaf area, and their relationship with tree species richness in a subtropical forest in south-east China. Permanent forest plots of 30 x 30 m were selected to span largely independent gradients in tree species richness and secondary successional age. Plots with higher tree species richness had a higher stand basal area. Also, stand basal area increases over a 4-year census interval were larger at high than at low diversity. These effects translated into increased carbon stocks in aboveground phytomass (estimated using allometric equations). A higher variability in tree height in more diverse plots suggested that these effects were facilitated by denser canopy packing due to architectural complementarity between species. In contrast, leaf area was not or even negatively affected by tree diversity, indicating a decoupling of carbon accumulation from leaf area. Alternatively, the same community leaf area might have assimilated more C per time interval in more than in less diverse plots because of differences in leaf turnover and productivity or because of differences in the display of leaves in vertical and horizontal space. Overall, our study suggests that in species-rich forests niche-based processes support a positive diversity–productivity relationship and that this translates into increased carbon storage in long-lived woody structures. Given the high growth rates of these forests during secondary succession, our results further indicate that a forest management promoting tree diversity after disturbance may accelerate CO2 sequestration from the atmosphere and thus be relevant in a climate-change context.

Tree species traits but not diversity mitigate stem breakage in a subtropical forest following a rare and extreme ice storm

Future climates are likely to include extreme events, which in turn have great impacts on ecological systems. In this study, we investigated possible effects that could mitigate stem breakage caused by a rare and extreme ice storm in a Chinese subtropical forest across a gradient of forest diversity. We used Bayesian modeling to correct stem breakage for tree size and variance components analysis to quantify the influence of taxon, leaf and wood functional traits, and stand level properties on the probability of stem breakage. We show that the taxon explained four times more variance in individual stem breakage than did stand level properties; trees with higher specific leaf area (SLA) were less susceptible to breakage. However, a large part of the variation at the taxon scale remained unexplained, implying that unmeasured or undefined traits could be used to predict damage caused by ice storms. When aggregated at the plot level, functional diversity and wood density increased after the ice storm. We suggest that for the adaption of forest management to climate change, much can still be learned from looking at functional traits at the taxon level.

Strong positive biodiversity-productivity relationships in a subtropical forest experiment

Forest ecosystems contribute substantially to global terrestrial primary productivity and climate regulation, but, in contrast to grasslands, experimental evidence for a positive biodiversity-productivity relationship in highly diverse forests is still lacking1. Here, we provide such evidence from a large forest biodiversity experiment with a novel design2 in subtropical China. Productivity (stand-level tree basal area, aboveground volume and carbon and their annual increment) increased linearly with the logarithm of tree species richness. Additive partitioning3 showed that increasing positive complementarity effects combined with weakening negative selection effects caused a strengthening of the relationship over time. In 2-species mixed stands, complementary effects increased with functional distance and selection effects with vertical crown dissimilarity between species. Understorey shrubs reduced stand-level tree productivity, but this effect of competition was attenuated by shrub species richness, indicating that a diverse understorey may facilitate overall ecosystem functioning. Identical biodiversity-productivity relationships were found in plots of different size, suggesting that extrapolation to larger scales is possible. Our results highlight the potential of multi-species afforestation strategies to simultaneously contribute to mitigation of climate change and biodiversity restoration.

Impacts of species richness on productivity in a large-scale subtropical forest experiment

Tree diversity improves forest productivity Experimental studies in grasslands have shown that the loss of species has negative consequences for ecosystem functioning. Is the same true for forests? Huang et al. report the first results from a large biodiversity experiment in a subtropical forest in China. The study combines many replicates, realistic tree densities, and large plot sizes with a wide range of species richness levels. After 8 years of the experiment, the findings suggest strong positive effects of tree diversity on forest productivity and carbon accumulation. Thus, changing from monocultures to more mixed forests could benefit both restoration of biodiversity and mitigation of climate change. Science, this issue p. 80 In a replicated experiment in a subtropical forest, higher tree species diversity promoted productivity and carbon storage. Biodiversity experiments have shown that species loss reduces ecosystem functioning in grassland. To test whether this result can be extrapolated to forests, the main contributors to terrestrial primary productivity, requires large-scale experiments. We manipulated tree species richness by planting more than 150,000 trees in plots with 1 to 16 species. Simulating multiple extinction scenarios, we found that richness strongly increased stand-level productivity. After 8 years, 16-species mixtures had accumulated over twice the amount of carbon found in average monocultures and similar amounts as those of two commercial monocultures. Species richness effects were strongly associated with functional and phylogenetic diversity. A shrub addition treatment reduced tree productivity, but this reduction was smaller at high shrub species richness. Our results encourage multispecies afforestation strategies to restore biodiversity and mitigate climate change.