There is mounting evidence that species diversity increases the temporal stability of forest growth. This stabilising effect of diversity has mainly been attributed to species differences in their response to fluctuating environmental conditions. Interactions among individuals could also contribute to the stabilising effect of diversity by increasing the mean and reducing the variance of tree growth, however, this has never been directly demonstrated. We used tree-ring width chronologies from temperate and boreal mixed stands of Eastern Canada to identify the role of interactions among individuals in the stabilising effect of diversity on forest growth. Using neighbourhood competition index and a mixed model, we compared the effect of interspecific and intraspecific interactions on the mean and the variance of tree growth. We found that interspecific interactions are less detrimental to tree growth than intraspecific interactions. We also found that interspecific interactions buffer tree response to drought and thereby reduce the variance of tree growth. Our results indicate diversity may increase the mean and reduce the variance of tree growth through interactions among individuals. Thus, we demonstrate interactions among individuals play a role in the stabilising effect of diversity on forest growth, and in doing so, we bring to light other mechanisms of the insurance hypothesis.

Forest diversity-productivity relationships have been intensively investigated in recent decades. However, few studies have considered the interplay between species and structural diversity in driving productivity. We analyzed these factors using data from 52 permanent plots in southwestern Germany with more than 53,000 repeated tree measurements. We used basal area increment as a proxy for productivity and hypothesized that: (1) structural diversity would increase tree and stand productivity, (2) diversity-productivity relationships would be weaker for species diversity than for structural diversity, and (3) species diversity would also indirectly impact stand productivity via changes in size structure. We measured diversity using distance-independent indices. We fitted separate linear mixed-effects models for fir, spruce and beech at the tree level, whereas at the stand level we pooled all available data. We tested our third hypothesis using structural equation modeling. Structural and species diversity acted as direct and independent drivers of stand productivity, with structural diversity being a slightly better predictor. Structural diversity, but not species diversity, had a significant, albeit asymmetric, effect on tree productivity. The functioning of structurally diverse, mixed forests is influenced by both structural and species diversity. These sources of trait diversity contribute to increased vertical stratification and crown plasticity, which in turn diminish competitive interferences and lead to more densely packed canopies per unit area. Our research highlights the positive effects of species diversity and structural diversity on forest productivity and ecosystem dynamics.

As one of Earth's most carbon-dense regions, tropical forests are central to climate change mitigation efforts. Their unparalleled species richness also makes them vital for safeguarding biodiversity. However, because research has not been conducted at management-relevant scales and has often not accounted for forest disturbance, the biodiversity implications of carbon conservation strategies remain poorly understood. We investigated tropical carbon-biodiversity relationships and trade-offs along a forest-disturbance gradient, using detailed and extensive carbon and biodiversity datasets. Biodiversity was positively associated with carbon in secondary and highly disturbed primary forests. Positive carbon-biodiversity relationships dissipated at around 100 MgC ha(-1), meaning that in less disturbed forests more carbon did not equal more biodiversity. Simulated carbon conservation schemes therefore failed to protect many species in the most species-rich forests. These biodiversity shortfalls were sensitive to opportunity costs and could be decreased for small carbon penalties. To ensure that the most ecologically valuable forests are protected, biodiversity needs to be incorporated into carbon conservation planning.

Stand structure can strongly influence forest growth and other processes, such as the water balance, carbon partitioning, nutrient cycling and light dynamics. However, individual structural variables can be positively or negatively correlated with growth. This is the case for variables such as size inequality and those that describe resource partitioning, such as the degree of symmetric/asymmetric competition and growth dominance. Several contrasting growth-structure correlations are reviewed and linked to forest processes by considering the different types of tree interactions they are associated with. Contrasting growth-structure correlations appear to converge when they are examined using a simple framework where stand growth is a function of three variables as opposed to any one of the variables alone; stand density, size distributions and tree size-growth relationships. The size distributions quantify how the stand density is distributed between the different sizes while the size growth relationships quantify how growth is partitioned between different sizes. Size inequality may not often be a useful explanatory variable and instead it appears to sometimes correlate with growth because it can be correlated with other variables that influence growth. The spatial and temporal dynamics of the effects of structure on growth have received little attention and a long-term growth and yield data set from central Europe was used to examine how the effects of structure can change along climatic gradients. The simple framework of three variables could be used to separate the effects of structure and functioning when comparing mixed and monospecific forests, as well as to design silvicultural interventions or to determine whether past management interventions have achieved their goals. The implications for selecting which structural variables to use and when scaling up to the stand level, are also discussed.

Light-related interactions can increase productivity in tree-species mixtures compared with monocultures due to higher stand-level absorption of photosynthetically active radiation (APAR) or light-use efficiency (LUE). However, the effects of different light-related interactions, and their relative importance, have rarely been quantified. Here, measurements of vertical leaf-area distributions, tree sizes, and stand density were combined with a tree-level light model (Maestra) to examine how crown architecture and vertical or horizontal canopy structure influence the APAR of 16 monocultures and eight different two-species mixtures with 16 different species in a Chinese subtropical tree diversity experiment. A higher proportion of crown leaf area occurred in the upper crowns of species with higher specific leaf areas. Tree-level APAR depended largely on tree leaf area and also, but to a lesser extent, on relative height (i.e., tree dominance) and leaf-area index (LAI). Stand-level APAR depended on LAI and canopy volume, but not on the vertical stratification or canopy leaf-area density. The mixing effects, in terms of relative differences between mixtures and monocultures, on stand-level APAR were correlated with the mixing effects on basal area growth, indicating that light-related interactions may have been responsible for part of the mixing effects on basal area growth. While species identity influences the vertical distributions of leaf area within tree crowns, this can have a relatively small effect on tree and stand APAR compared with the size and vertical positioning of the crowns, or the LAI and canopy volume.

The insurance hypothesis, stating that biodiversity can increase ecosystem stability, has received wide research and political attention. Recent experiments suggest that climate change can impact how plant diversity influences ecosystem stability, but most evidence of the biodiversity-stability relationship obtained to date comes from local studies performed under a limited set of climatic conditions. Here, we investigate how climate mediates the relationships between plant (taxonomical and functional) diversity and ecosystem stability across the globe. To do so, we coupled 14 years of temporal remote sensing measurements of plant biomass with field surveys of diversity in 123 dryland ecosystems from all continents except Antarctica. Across a wide range of climatic and soil conditions, plant species pools, and locations, we were able to explain 73% of variation in ecosystem stability, measured as the ratio of the temporal mean biomass to the SD. The positive role of plant diversity on ecosystem stability was as important as that of climatic and soil factors. However, we also found a strong climate dependency of the biodiversity-ecosystem stability relationship across our global aridity gradient. Our findings suggest that the diversity of leaf traits may drive ecosystem stability at low aridity levels, whereas species richness may have a greater stabilizing role under the most arid conditions evaluated. Our study highlights that to minimize variations in the temporal delivery of ecosystem services related to plant biomass, functional and taxonomic plant diversity should be particularly promoted under low and high aridity conditions, respectively.

Insurance effects of biodiversity can stabilize the functioning of multispecies ecosystems against environmental variability when differential species' responses lead to asynchronous population dynamics. When responses are not perfectly positively correlated, declines in some populations are compensated by increases in others, smoothing variability in ecosystem productivity. This variance reduction effect of biodiversity is analogous to the risk-spreading benefits of diverse investment portfolios in financial markets. We use data from the BIODEPTH network of grassland biodiversity experiments to perform a general test for stabilizing effects of plant diversity on the temporal variability of individual species, functional groups, and aggregate communities. We tested three potential mechanisms: reduction of temporal variability through population asynchrony; enhancement of long-term average performance through positive selection effects; and increases in the temporal mean due to overyielding. Our results support a stabilizing effect of diversity on the temporal variability of grassland aboveground annual net primary production through two mechanisms. Two-species communities with greater population asynchrony were more stable in their average production over time due to compensatory fluctuations. Overyielding also stabilized productivity by increasing levels of average biomass production relative to temporal variability. However, there was no evidence for a performance-enhancing effect on the temporal mean through positive selection effects. In combination with previous work, our results suggest that stabilizing effects of diversity on community productivity through population asynchrony and overyielding appear to be general in grassland ecosystems.

The composition of communities is strongly altered by anthropogenic manipulations of biogeochemical cycles, abiotic conditions, and trophic structure in all major ecosystems. Whereas the effects of species loss on ecosystem processes have received broad attention, the consequences of altered species dominance for emergent properties of communities and ecosystems are poorly investigated. Here we propose a framework guiding our understanding of how dominance affects species interactions within communities, processes within ecosystems, and dynamics on regional scales. Dominance (or the complementary term, evenness) reflects the distribution of traits in a community, which in turn affects the strength and sign of both intraspecifc and interspecific interactions. Consequently, dominance also mediates the effect of such interactions on species coexistence. We review the evidence for the fact that dominance directly affects ecosystem functions such as process rates via species identity (the dominant trait) and evenness (the frequency distribution of traits), and indirectly alters the relationship between process rates and species richness. Dominance also influences the temporal and spatial variability of aggregate community properties and compositional stability (invasibility). Finally, we propose that dominance affects regional species coexistence by altering metacommunity dynamics. Local dominance leads to high beta diversity, and rare species can persist because of source-sink dynamics, but anthropogenically induced environmental changes result in regional dominance and low beta diversity, reducing regional coexistence. Given the rapid anthropogenic alterations of dominance in many ecosystems and the strong implications of these changes, dominance should be considered explicitly in the analysis of consequences of altered biodiversity.

Estimates of recent biodiversity change remain inconsistent, debated, and infrequently assessed for their functional implications. Here, we report that spatial scale and type of biodiversity measurement influence evidence of temporal biodiversity change. We show a pervasive scale dependence of temporal trends in taxonomic (TD) and functional (FD) diversity for an similar to 50-year record of avian assemblages from North American Breeding Bird Survey and a record of global extinctions. Average TD and FD increased at all but the global scale. Change in TD exceeded change in FD toward large scales, signaling functional resilience. Assemblage temporal dissimilarity and turnover (replacement of species or functions) declined, while nestedness (tendency of assemblages to be subsets of one another) increased with scale. Patterns of FD change varied strongly among diet and foraging guilds. We suggest that monitoring, policy, and conservation require a scale-explicit framework to account for the pervasive effect that scale has on perceived biodiversity change.

Topography is an important driver that determines diversity patterns and ecosystem functioning in tropical forests. However, there are few studies analyzing contrasting topographical conditions on the relative importance of species to ecosystem functioning, mainly on those who have a greater contribution (i.e., hyperdominant species). We aimed to evaluate whether contrasting topographical conditions determine changes in tree species richness, community composition, and the number of stem and biomass hyperdominant species in a Brazilian Atlantic remnant forest fragment. We selected two areas on distinct hillsides with contrasting topographic conditions, at the biological reserve of the Federal University of Vicosa, Minas Gerais state, southeastern Brazil. Each area (100x100 m) was sub-divided into 100 plots of 10x10 m. From each plot, all trees having diameter at breast height (DBH) >= 10 cm were identified to the species level and tagged for measurement. We measured three topographic variables (slope, elevation, and convexity) in each plot, based on the assumption that these variables may affect tree species diversity, species composition, and ecosystem function (aboveground biomass). The AGB of individual stems was calculated in all plots. We performed a multivariate regression tree for estimating topographical heterogeneity in each area. We found that species richness differed significantly between areas. Species richness in the Northeast area (the more topographically heterogeneous one) was 48% higher than that in the Southeast area, which is less topographically heterogeneous. The tree species composition varied considerably between areas, with similar AGB patterns being registered among plots. The number of stem hyperdominants varied significantly between areas. In the Southeast area, only two species out of the 85 recorded (2.38%) accounted for 50% of the number of stems hyperdominants, while in the Northeast area 10 species (7.94%) accounted for 50% of stems hyperdominants. Our results showed that high topographic heterogeneity induces high species richness and that the number of stem and biomass hyperdominant species increase along with richness on a local scale. Based on our results, we presume that biomass hyperdominance can also strongly influence forest ecosystem functioning on a local scale.

There is increasing evidence that mixed-species forests can provide multiple ecosystem services at a higher level than their monospecific counterparts. However, most studies concerning tree diversity and ecosystem functioning relationships use data from forest inventories (under noncontrolled conditions) or from very young plantation experiments. Here, we investigated temporal dynamics of diversity-productivity relationships and diversity-stability relationships in the oldest tropical tree diversity experiment. Sardinilla was established in Panama in 2001, with 22 plots that form a gradient in native tree species richness of one-, two-, three- and five-species communities. Using annual data describing tree diameters and heights, we calculated basal area increment as the proxy of tree productivity. We combined tree neighbourhood- and community-level analyses and tested the effects of both species diversity and structural diversity on productivity and its temporal stability. General patterns were consistent across both scales indicating that tree-tree interactions in neighbourhoods drive observed diversity effects. From 2006 to 2016, mean overyielding (higher productivity in mixtures than in monocultures) was 25%-30% in two- and three-species mixtures and 50% in five-species stands. Tree neighbourhood diversity enhanced community productivity but the effect of species diversity was stronger and increased over time, whereas the effect of structural diversity declined. Temporal stability of community productivity increased with species diversity via two principle mechanisms: asynchronous responses of species to environmental variability and overyielding. Overyielding in mixtures was highest during a strong El Niño-related drought. Overall, positive diversity-productivity and diversity-stability relationships predominated, with the highest productivity and stability at the highest levels of diversity. These results provide new insights into mixing effects in diverse, tropical plantations and highlight the importance of analyses of temporal dynamics for our understanding of the complex relationships between diversity, productivity and stability. Under climate change, mixed-species forests may provide both high levels and high stability of production.© 2019 The Authors. Global Change Biology published by John Wiley & Sons Ltd.

A major ecosystem effect of biodiversity is to stabilise assemblages that perform particular functions. However, diversity-stability relationships (DSRs) are analysed using a variety of different population and community properties, most of which are adopted from theory that makes several restrictive assumptions that are unlikely to be reflected in nature. Here, we construct a simple synthesis and generalisation of previous theory for the DSR. We show that community stability is a product of two quantities: the synchrony of population fluctuations, and an average species-level population stability that is weighted by relative abundance. Weighted average population stability can be decomposed to consider effects of the mean-variance scaling of abundance, changes in mean abundance with diversity and differences in species' mean abundance in monoculture. Our framework makes explicit how unevenness in the abundances of species in real communities influences the DSR, which occurs both through effects on community synchrony, and effects on weighted average population variability. This theory provides a more robust framework for analysing the results of empirical studies of the DSR, and facilitates the integration of findings from real and model communities.© 2012 Blackwell Publishing Ltd/CNRS.

The stability of ecological communities is critical for the stable provisioning of ecosystem services, such as food and forage production, carbon sequestration, and soil fertility. Greater biodiversity is expected to enhance stability across years by decreasing synchrony among species, but the drivers of stability in nature remain poorly resolved. Our analysis of time series from 79 datasets across the world showed that stability was associated more strongly with the degree of synchrony among dominant species than with species richness. The relatively weak influence of species richness is consistent with theory predicting that the effect of richness on stability weakens when synchrony is higher than expected under random fluctuations, which was the case in most communities. Land management, nutrient addition, and climate change treatments had relatively weak and varying effects on stability, modifying how species richness, synchrony, and stability interact. Our results demonstrate the prevalence of biotic drivers on ecosystem stability, with the potential for environmental drivers to alter the intricate relationship among richness, synchrony, and stability.

The term "size hierarchy" has been used frequently by plant population biologists but it has not been defined. Positive skewness of the size distribution, which has been used to evaluate size hierarchies, is inappropriate. We suggest that size hierarchy is equivalent to size inequality. Methods developed by economists to evaluate inequalities in wealth and income, the Lorenz curve and Gini Coefficient, provide a useful quantification of inequality and allow us to compare populations. A measure of inequality such as the Gini Coefficient will usually be more appropriate than a measure of skewness for addressing questions concerning plant population structure.