Designing forest biodiversity experiments : general considerations illustrated by a new large experiment in subtropical China

Funded by German Research Foundation. Grant Number: DFG FOR 891/1 and 2 National Natural Science Foundation of China. Grant Numbers: NSFC 30710103907, 30930005, 31170457 , 31210103910 Swiss National Science Foundation (SNSF) Sino-German Centre for Research Promotion in BeijingPeer reviewedPublisher PD

Towards a methodical framework for comprehensively assessing forest multifunctionality

Funded by Deutsche Forschungsgemeinschaft. Grant Number: DFG FOR 891/1-3 National Natural Science Foundation of China. Grant Numbers: 30710103907, 30930005, 31170457, 31210103910 Swiss National Science Foundation (SNSF) Sino-German Centre for Research Promotion in Beijing. Grant Number: GZ 986Peer reviewedPublisher PD

Early positive effects of tree species richness on herbivory in a large-scale forest biodiversity experiment influence tree growth

Despite the importance of herbivory for the structure and functioning of species-rich forests, little is known about how herbivory is affected by tree species richness, and more specifically by random vs. non-random species loss. We assessed herbivore damage and its effects on tree growth in the early stage of a large-scale forest biodiversity experiment in subtropical China that features random and non-random extinction scenarios of tree mixtures numbering between one and 24 species. In contrast to random species loss, the non-random extinction scenarios were based on the tree species' local rarity and specific leaf area - traits that may strongly influence the way herbivory is affected by plant species richness. Herbivory increased with tree species richness across all scenarios and was unaffected by the different species compositions in the random and non-random extinction scenarios. Whereas tree growth rates were positively related to herbivory on plots with smaller trees, growth rates significantly declined with increasing herbivory on plots with larger trees. Our results suggest that the effects of herbivory on growth rates increase from monocultures to the most species-rich plant communities and that negative effects with increasing tree species richness become more pronounced with time as trees grow larger.Synthesis. Our results indicate that key trophic interactions can be quick to become established in forest plantations (i.e. already 2.5years after tree planting). Stronger herbivory effects on tree growth with increasing tree species richness suggest a potentially important role of herbivory in regulating ecosystem functions and the structural development of species-rich forests from the very start of secondary forest succession. The lack of significant differences between the extinction scenarios, however, contrasts with findings from natural forests of higher successional age, where rarity had negative effects on herbivory. This indicates that the effects of non-random species loss could change with forest succession

Mixed afforestation of young subtropical trees promotes nitrogen acquisition and retention

Afforestation is globally increasing to produce timber and pulp wood, but also to enhance ecosystem services such as carbon sequestration, nutrient retention or groundwater recharge. In China, large areas have been and will be afforested in order to compensate for the negative impacts of former clear-cuttings and to make use of the ecosystem services associated with afforestation. In order to further optimize these services with regard to balanced nutrient (particularly nitrogen) cycles, it is important to know whether the use of mixtures of native tree species in afforestation projects promotes the acquisition and retention of nitrogen compared with the currently established large-scale monocultures. To test the effect of species richness on system N retention and tree sapling N uptake, we conducted a N-15 tracer experiment in a young tree plantation. To this end, saplings of four abundant early successional tree species were planted in monocultures, in two- and four-species mixtures and as single trees. Nitrogen retention increased with higher species richness due to enhanced N pools in sapling biomass. These species richness effects strengthened over time. Species-specific differences in N-15 recoveries over time revealed below-ground niche differentiation with regard to N uptake, which is likely to result in complementary resource use among coexisting species. Synthesis and applications. This study provides evidence that mixed afforestation promotes N retention from the sapling stage. To further improve ecosystem services associated with afforestation, we strongly suggest the use of mixtures of native tree species instead of monocultures. Mixtures of four species may reduce system N losses and thus may lessen groundwater contamination due to N leaching. We encourage further investigations to find optimal species combinations that promote a wide range of ecosystem services related to more closed nutrient cycles and minimized soil erosion. In our study, the plantations' capability to retain N could be optimized by means of both increasing tree species richness and by choosing the optimal species combinations. This study provides evidence that mixed afforestation promotes N retention from the sapling stage. To further improve ecosystem services associated with afforestation, we strongly suggest the use of mixtures of native tree species instead of monocultures. Mixtures of four species may reduce system N losses and thus may lessen groundwater contamination due to N leaching. We encourage further investigations to find optimal species combinations that promote a wide range of ecosystem services related to more closed nutrient cycles and minimized soil erosion. In our study, the plantations' capability to retain N could be optimized by means of both increasing tree species richness and by choosing the optimal species combinations

Interspecific and intraspecific variation in specific root length drives aboveground biodiversity effects in young experimental forest stands

Aims Although the net biodiversity effect (NE) can be statistically partitioned into complementarity and selection effects (CE and SE), there are different underlying mechanisms that can cause a certain partitioning. Our objective was to assess the role of resource partitioning and species interactions as two important mechanisms that can bring about CEs by interspecific and intraspecific trait variation. Methods We measured tree height of 2493 living individuals in 57 plots and specific root length (SRL) on first-order roots of 368 of these individuals across different species richness levels (1, 2, 4, 8 species) in a large-scale forest biodiversity and ecosystem functioning experiment in subtropical China (BEF-China) established in 2009. We describe the effects of resource partitioning between species by a fixed component of interspecific functional diversity (RaoQ) and further effects of species interactions by variable components of interspecific and intraspecific functional diversity (community weighted trait similarity and trait dissimilarity, CWS and CWD). Finally, we examined the relationships between biodiversity effects on stand-level tree height and functional diversity (RaoQ, CWS and CWD) in SRL using linear regression and assessed the relative importance of these three components of functional diversity in explaining the diversity effects. Important Findings Our results show that species richness significantly affected SRL in five and tree height in ten out of 16 species. A positive NE was generally brought about by a positive CE on stand-level tree height and related to high values of RaoQ and CWS in SRL. A positive CE was related to high values of all three components of root functional diversity (RaoQ, CWS and CWD). Our study suggests that both resource partitioning and species interactions are the underlying mechanisms of biodiversity effects on stand-level tree growth in subtropical forest

Impact of tree diversity and environmental conditions on the survival of shrub species in a forest biodiversity experiment in subtropical China

Aims Although shrubs are an important component of forests, their role has not yet been considered in forest biodiversity experiments. In the biodiversity-ecosystem functioning (BEF) experiment with subtropical tree species in south-east China (BEF-China), we factorially combined tree with shrub species-diversity treatments. Here, we tested the hypotheses that shrub survival differs between the 10 planted shrub species, with lower survival rates of late-than early-successional species and is affected by environmental conditions, such as topography and top soil characteristics, as well as by biotic factors, represented by tree, shrub and herb layer characteristics. Methods We analyzed the survival of 42 000 shrub individuals in 105 plots varying in tree and shrub species richness of the BEF-China project four years after planting. Shrub survival was analyzed with generalized linear mixed effects models at the level of individuals and with variance partitioning at the plot level. Random intercept and random slope models of different explanatory variables were compared with respect to the Bayesian Information Criterion (BIC). Important Findings Survival rates differed largely between the 10 shrub species, ranging from 26% to 91% for Ardisia crenata and Distylium buxifolium, respectively. Irrespective of species identity, single abiotic factors explained up to 5% of species survival, with a negative effect of altitude and slope inclination and a positive effect of the topsoil carbon to nitrogen ratio, which pointed to drought as the major cause of shrub mortality. In contrast, neither tree nor shrub richness affected shrub survival at this early stage of the experiment. Among the biotic predictors, only herb layer species richness and cover of the dominant fern species (Dicranopteris pedata) affected shrub survival. Overall, our models that included all variables could explain about 65% in shrub survival, with environmental variables being most influential, followed by shrub species identity, while tree species diversity (species richness and identity) and herb layer characteristics contributed much less. Thus, in this early stage of the experiment the biotic interactions among shrubs and between shrubs and trees have not yet overruled the impact of abiotic environmental factors

Multiple components of plant diversity loss determine herbivore phylogenetic diversity in a subtropical forest experiment

1. Plant diversity loss can alter higher trophic-level communities via non-random species interactions, which in turn may cascade to affect key ecosystem functions. These non-random linkages might be best captured by patterns of phylogenetic diversity, which take into account co-evolutionary dependencies. However, lack of adequate phylogenetic data of higher trophic levels hampers our mechanistic understanding of biodiversity relationships in species-rich ecosystems. 2. We used DNA barcoding to generate data on the phylogenetic diversity of lepidopteran caterpillars in a large-scale forest biodiversity experiment in subtropical China. We analysed how different metrics of lepidopteran phylogenetic diversity (Faith's PD, MPD, MNTD) and taxonomic diversity were influenced by multiple components of tree diversity (taxonomic, functional, phylogenetic). 3. Our data from six sampling periods represent 7,204 mitochondrial cytochrome c oxidase subunit I (COI) sequences of lepidopteran larvae, clustered into 461 molecular operational taxonomic units. Lepidopteran abundance, the effective number of species (irrespective of the focus on rare or common species) and Faith's PD and MPD (reflecting basal evolutionary splits), but not MNTD (reflecting recent evolutionary splits), significantly increased with experimentally manipulated tree species richness. Lepidopteran MNTD decreased with increasing tree MNTD. Path analyses showed that tree phylogenetic and functional diversity explained part, but not all of the effects of tree species richness on lepidopteran diversity. Importantly, tree diversity effects on lepidopteran diversity were to a large extent indirect, operating via changes in lepidopteran abundance. 4. Synthesis. Our study shows that evolutionary dependencies determine the response of herbivore communities to changes in host plant diversity. Incorporating a wider range of diversity metrics both at the level of producers and consumers can thus help to develop a more comprehensive understanding of the functional consequences of biodiversity change across trophic levels. Moreover, the dependence of trophic linkages on herbivore abundances underlines the need to address the consequences of current declines in insect abundances for ecosystem structure and functioning

On the combined effect of soil fertility and topography on tree growth in subtropical forest ecosystems-a study from SE China

Aims The aim of our research was to understand small-scale effects of topography and soil fertility on tree growth in a forest biodiversity and ecosystem functioning (BEF) experiment in subtropical SE China. Methods Geomorphometric terrain analyses were carried out at a spatial resolution of 5 x 5 m. Soil samples of different depth increments and data on tree height were collected from a total of 566 plots (667 m(2) each). The soils were analyzed for carbon (soil organic carbon [SOC]), nitrogen, acidity, cation exchange capacity (CEC), exchangeable cations and base saturation as soil fertility attributes. All plots were classified into geomorphological units. Analyses of variance and linear regressions were applied to all terrain, soil fertility and tree growth attributes. Important Findings In general, young and shallow soils and relatively small differences in stable soil properties suggest that soil erosion has truncated the soils to a large extent over the whole area of the experiment. This explains the concurrently increasing CEC and SOC stocks downslope, in hollows and in valleys. However, colluvial, carbon-rich sediments are missing widely due to the convexity of the foot-slopes caused by uplift and removal of eroded sediments by adjacent waterways. The results showed that soil fertility is mainly influenced by topography. Monte-Carlo flow accumulation (MCCA), curvature, slope and aspect significantly affected soil fertility. Furthermore, soil fertility was affected by the different geomorphological positions on the experimental sites with ridge and spur positions showing lower exchangeable base cation contents, especially potassium (K), due to leaching. This geomorphological effect of soil fertility is most pronounced in the topsoil and decreases when considering the subsoil down to 50 cm depth. Few soil fertility attributes affect tree height after 1-2 years of growth, among which C stocks proved to be most important while pH(KCl) and CEC only played minor roles. Nevertheless, soil acidity and a high proportion of Al on the exchange complex affected tree height even after only 1-2 years growth. Hence, our study showed that forest nutrition is coupled to a recycling of litter nutrients, and does not only depend on subsequent supply of nutrients from the mineral soil. Besides soil fertility, topography affected tree height. We found that especially MCCA as indicator of water availability affected tree growth at small-scale, as well as aspect. Overall, our synthesis on the interrelation between fertility, topography and tree growth in a subtropical forest ecosystem in SE China showed that topographic heterogeneity lead to ecological gradients across geomorphological positions. In this respect, small-scale soil-plant interactions in a young forest can serve as a driver for the future development of vegetation and biodiversity control on soil fertility. In addition, it shows that terrain attributes should be accounted for in ecological research

Evaluating structural and compositional canopy characteristics to predict the light-demand signature of the forest understorey in mixed, semi-natural temperate forests

International audienceQuestions Light availability at the forest floor affects many forest ecosystem processes, and is often quantified indirectly through easy-to-measure stand characteristics. We investigated how three such characteristics, basal area, canopy cover and canopy closure, were related to each other in structurally complex mixed forests. We also asked how well they can predict the light-demand signature of the forest understorey (estimated as the mean Ellenberg indicator value for light [''EIVLIGHT''] and the proportion of ``forest specialists'' [''%FS''] within the plots). Furthermore, we asked whether accounting for the shade-casting ability of individual canopy species could improve predictions ofEIV(LIGHT)and %FS. Location A total of 192 study plots from nineteen temperate forest regions across Europe. Methods In each plot, we measured stand basal area (all stems >7.5 cm diameter), canopy closure (with a densiometer) and visually estimated the percentage cover of all plant species in the herb (7 m). We used linear mixed-effect models to assess the relationships between basal area, canopy cover and canopy closure. We performed model comparisons, based onR(2)and the Akaike Information Criterion (AIC), to assess which stand characteristics can predictEIV(LIGHT)and %FSbest, and to assess whether canopy shade-casting ability can significantly improve model fit. Results Canopy closure and cover were weakly related to each other, but showed no relation with basal area. For bothEIV(LIGHT)and %FS, canopy cover was the best predictor. Including the share of high-shade-casting species in both the basal-area and cover models improved the model fit forEIV(LIGHT), but not for %FS. Conclusions The typically expected relationships between basal area, canopy cover and canopy closure were weak or even absent in structurally complex mixed forests. In these forests, easy-to-measure structural canopy characteristics were poor predictors of the understorey light-demand signature, but accounting for compositional characteristics could improve predictions