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.
Summary1. Biodiversity-ecosystem functioning (BEF) experiments address ecosystem-level consequences of species loss by comparing communities of high species richness with communities from which species have been gradually eliminated. BEF experiments originally started with microcosms in the laboratory and with grassland ecosystems. A new frontier in experimental BEF research is manipulating tree diversity in forest ecosystems, compelling researchers to think big and comprehensively. 2. We present and discuss some of the major issues to be considered in the design of BEF experiments with trees and illustrate these with a new forest biodiversity experiment established in subtropical China (Xingangshan, Jiangxi Province) in 2009/2010. Using a pool of 40 tree species, extinction scenarios were simulated with tree richness levels of 1, 2, 4, 8 and 16 species on a total of 566 plots of 25Á8 9 25Á8 m each. 3. The goal of this experiment is to estimate effects of tree and shrub species richness on carbon storage and soil erosion; therefore, the experiment was established on sloped terrain. The following important design choices were made: (i) establishing many small rather than fewer larger plots, (ii) using high planting density and random mixing of species rather than lower planting density and patchwise mixing of species, (iii) establishing a map of the initial 'ecoscape' to characterize site heterogeneity before the onset of biodiversity effects and (iv) manipulating tree species richness not only in random but also in trait-oriented extinction scenarios. 4. Data management and analysis are particularly challenging in BEF experiments with their hierarchical designs nesting individuals within-species populations within plots within-species compositions. Statistical analysis best proceeds by partitioning these random terms into fixed-term contrasts, for example, species composition into contrasts for species richness and the presence of particular functional groups, which can then be tested against the remaining random variation among compositions. 5. We conclude that forest BEF experiments provide exciting and timely research options. They especially require careful thinking to allow multiple disciplines to measure and analyse data jointly and effectively. Achiev- This is an open access article under the terms of the Creative Commons Attribution License, which permits use, distribution and reproduction in any medium, provided the original work is properly cited. 2014, 5, 74-89 doi: 10.1111/2041-210X.12126 ing specific research goals and synergy with previous experiments involves trade-offs between different designs and requires manifold design decisions. Methods in Ecology andEvolution
(2011). Community assembly during secondary forest succession in a Chinese subtropical forest. Ecological Monographs, 81(1): 25-41. Community assembly during secondary forest succession in a Chinese subtropical forest Abstract Subtropical broad-leaved forests in southeastern China support a high diversity of woody plants. Using a comparative study design with 30330 m plots (n=27) from five successional stages (<20, <40, <60, <80, and ≤80 yr), we investigated how the gradient in species composition reflects underlying processes of community assembly. In particular, we tested whether species richness of adult trees and shrubs decreased or increased and assessed to which degree this pattern was caused by negative density dependence or continuous immigration over time. Furthermore, we tested whether rare species were increasingly enriched and the species composition of adult trees and shrubs became more similar to species composition of seedlings during the course of succession. We counted the individuals of all adult species and shrubs >1 m in height in each plot and counted all woody recruits (bank of all seedlings ≤1 m in height) in each central 10310 m quadrant of each plot. In addition, we measured a number of environmental variables (elevation, slope, aspect, soil moisture, pH, C, N, and C/N ratio) and biotic structural variables (height and cover of layers). Adult species richness varied from 25 to 69 species per plot, and in total 148 woody species from 46 families were recorded. There was a clear successional gradient in species composition as revealed by nonmetric multidimensional scaling (NMDS), but only a poor differentiation of different successional stages with respect to particular species. Adult richness per 100 individuals (rarefaction method) increased with successional stage. None of the measured abiotic variables were significantly correlated with adult species richness. We found no evidence that rare species were responsible for the increasing adult species richness, as richness of rare species among both adults and recruits was independent of the successional stage. Furthermore, the similarity between established adults and recruits did not increase with successional stage. There was a constant number of recruit species and also of exclusive recruit species, i.e., those that had not been present as adult individuals, across all successional stages, suggesting a continuous random immigration over time. variables were significantly correlated with adult species richness. We found no evidence that 41 rare species were responsible for the increasing adult species richness, as richness of rare 42 species amongst both adults and recruits was independent of the successional stage.
Human-induced biodiversity change impairs ecosystem functions crucial to human well-being. However, the consequences of this change for ecosystem multifunctionality are poorly understood beyond effects of plant species loss, particularly in regions with high biodiversity across trophic levels. Here we adopt a multitrophic perspective to analyze how biodiversity affects multifunctionality in biodiverse subtropical forests. We consider 22 independent measurements of nine ecosystem functions central to energy and nutrient flow across trophic levels. We find that individual functions and multifunctionality are more strongly affected by the diversity of heterotrophs promoting decomposition and nutrient cycling, and by plant functional-trait diversity and composition, than by tree species richness. Moreover, cascading effects of higher trophic-level diversity on functions originating from lower trophic-level processes highlight that multitrophic biodiversity is key to understanding drivers of multifunctionality. A broader perspective on biodiversity-multifunctionality relationships is crucial for sustainable ecosystem management in light of non-random species loss and intensified biotic disturbances under future environmental change.
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