Positive Effect Of Species Richness Research Articles

Nitrogen addition strengthens the stabilizing effect of biodiversity on productivity by increasing plant trait diversity and species asynchrony in the artificial grassland communities.

Nitrogen (N) enrichment usually weakens the stabilizing effect of biodiversity on productivity. However, previous studies focused on plant species richness and thus largely ignored the potential contributions of plant functional traits to stability, even though evidence is increasing that functional traits are stronger predictors than species richness of ecosystem functions. We conducted a common garden experiment manipulating plant species richness and N addition levels to quantify effects of N addition on relations between species richness and functional trait identity and diversity underpinning the 'fast-slow' economics spectrum and community stability. Nitrogen addition had a minor effect on community stability but increased the positive effects of species richness on community stability. Increasing community stability was found in the species-rich communities dominated by fast species due to substantially increasing temporal mean productivity relative to its standard deviation. Furthermore, enhancement in 'fast-slow' functional diversity in species-rich communities dominated by fast species under N addition increased species asynchrony, resulting in a robust biodiversity-stability relationship under N addition the artificial grassland communities. The findings demonstrate mechanistic links between plant species richness, 'fast-slow' functional traits, and community stability under N addition, suggesting that dynamics of biodiversity-stability relations under global changes are the results of species-specific responses of 'fast-slow' traits on the plant economics spectrum.

Structural and taxonomic diversity predict above‐ground biomass better than functional measures of maximum height in mixed‐species forests

AbstractAims:Mixed‐species forests are known to be highly productive systems because of their high species diversity, including taxonomic diversity (species richness) and structural diversity. Recent empirical evidence also points to plant maximum height, as a functional trait that potentially drives forest above‐ground biomass (AGB). However, the interrelations between these biotic variables are complex, and it is not always predictable if structural diversity attributes or functional metrics of plant maximum height would act as the most important determinant of stand biomass. Here we evaluated the relative importance of structural diversity attributes and functional metrics of plant maximum height (Hmax) in predicting and mediating AGB response to variation in species richness in mixed‐species forests, while also accounting for fine‐scale environmental variation.Location:Northern Benin.Methods:We used forest inventory data from mixed‐species stands of native and exotic species. We quantified structural diversity as coefficient of variation of tree diameter at breast height (CVdbh) and of height (CVHt). For plant Hmax, we computed three metrics: functional range (FRHmax), functional divergence (FDHmax) and community‐weighted mean (CWMHmax). We used topographical variables such as elevation and slope to account for possible environmental effects. Simple and multiple mixed‐effects models, and structural equation models were performed to assess the direct and indirect links of AGB with species richness through structural diversity attributes and functional metrics of plant Hmax.Results:Species richness and CVdbh were positively related to AGB, while functional metrics of plant Hmax were not. Structural equation models revealed that species richness influenced AGB indirectly via CVdbh, which alone strongly promoted AGB. Elevation only had a positive direct effect on AGB. While increasing species richness enhanced CVdbh and functional measures of plant Hmax, there was no support for the latter mediating the effects of species richness on AGB.Conclusion:Structural diversity has a significant advantage in predicting and mediating the positive effect of species richness on AGB more so than functional measures of plant Hmax. We argue that structural diversity acts as a mechanism for the species richness–AGB relationship, and that maintaining high structural diversity would enhance biomass in mixed‐species forests.

Species Richness Promotes Productivity through Tree Crown Spatial Complementarity in a Species-Rich Natural Forest

Ecological theory suggests that species with complementary architectural and physiological traits can optimize crown packing to improve resource efficiency and promote ecosystem productivity in forest communities. However, empirical evidence of this prediction is rare in species-rich natural forests, as little is known about how crown spatial complementarity regulates community species richness-productivity relationship (SRPR). In this study, we measured tree architectural traits (stem diameter, height, crown depth and width) for 11,337 trees, and quantified species richness, functional diversity, crown spatial complementarity, soil fertility and forest productivity for 44 quadrats (20 m × 20 m per quadrat) in the Badagongshan 25 ha forest plot, central China. We tested bivariate correlations between species richness, crown complementarity, functional diversity and forest productivity. We employed linear mixed effects models to predict crown complementarity and examined its relationship with functional diversity. Finally, we applied structural equation modeling to quantify the mediation effects of crown complementarity on SRPRs. Species richness promoted crown complementarity and forest productivity. Crown complementarity varied across quadrats, with increases driven primarily by changes in tree height. Crown complementarity was positively related to functional diversity and forest productivity. Species richness increased with soil total phosphorus, while functional diversity decreased with soil bulk density. Forest productivity increased with soil organic carbon and total nitrogen, but decreased with bulk density. Crown complementarity partially mediated the positive effect of species richness on forest productivity, and the mediation effect was mainly through functional diversity. Our results suggest that the crown complementarity index accurately reflects the niche complementarity through light utilization and carbon reallocation. Our study emphasizes that species richness can promote crown complementarity, leading to greater forest productivity, which provides greater insight into the mechanical understanding of the SRPRs.

What is the mechanism of submerged macrophyte biodiversity affecting biomass productivity along water depth: Niche complementarity or selection effect?

The impact of biodiversity on ecosystem functions and services remains a hot topic in ecology, especially under increasing biodiversity loss. Studies on the relationship between biodiversity and ecosystem functioning have identified two regulatory mechanisms: i.e., “niche complementarity” and “selection effect”. However, the relative importance of these mechanisms within communities and how they change along ecological gradients remain unclear. Moreover, most studies have focused on relatively stable ecosystems (e.g., forest, grassland), with limited attention paid to ecosystems with obvious seasonal changes, e.g., freshwater ecosystems. In this study we conducted a seasonal survey of submerged macrophyte communities to clarify the mechanisms of biodiversity on biomass productivity accounting for seasons and water depth. Our results showed that (1) seasonal variations of community indicators exhibited different trends along the water depth gradient. Biomass productivity showed high seasonal variation at intermediate depth while temporal beta diversity showed the opposite pattern. (2) Water depth not only inhibited biomass productivity directly but also indirectly by reducing species richness and enhancing temporal beta diversity. (3) The positive effect of species richness on biomass productivity was lower than the negative effect of evenness (path coefficients: 0.13 vs. 0.27), indicating that “selection effect” played a more important role in the relationship. (4) In shallow areas, the mechanism underlying the effects of biodiversity on biomass productivity was more from “selection effect”, whereas, in deep areas, the effects were related to both “niche complementarity” and “selection effect”. Our findings suggest that “niche complementarity” and “selection effect” are two complementary mechanisms underpinning the relationship between biodiversity and biomass productivity, and their relative importance varies with specific ecological gradients. Our results also provide a reference for studies on other freshwater organisms with analogous distribution patterns as submerged macrophytes with water depth. Scale dependence of the above relationship should be considered in future studies.

Evenness alters the positive effect of species richness on community drought resistance via changing complementarity

Biodiversity loss may decrease the resistance of ecosystems to disturbance. While many studies have tested the effects of species richness on the ability of communities to resist disturbance, few have considered the role of species evenness. We examined whether species evenness affects the relationship between species richness and the drought resistance of plant communities. We constructed experimental plant communities with four levels of species richness (1, 2-, 4- and 8-species) and three levels of species evenness (high, medium and low) and subjected them to two drought treatments (with vs. without drought). Species evenness significantly increased the drought resistance of the communities and also complementarity effects. Moreover, a positive relationship between species richness and the drought resistance of plant communities was found at high and medium evenness but not at low evenness. This result corresponded well with the fact that complementarity effects increased with species richness at high and medium evenness but not at low evenness. However, neither species evenness nor species richness significantly affected selection effects for the drought resistance of the communities. We conclude that species evenness can alter the relationships between species richness and the resistance to disturbance (e.g., drought) in plant communities by enhanced complementarity among species. Thus, in addition to species richness, evenness should also be considered an important indicator of the community resistance to future global changes, such as increased drought intensity and frequency.

Ecosystem stability at the landscape scale is primarily associated with climatic history

Abstract There is an increasing interest in landscape‐scale perspectives of ecosystem functioning to inform policy and conservation decisions. However, we need a better understanding of the stability of ecosystem functioning (e.g. plant productivity) at the landscape scale to inform policy around topics such as global food security. We investigate the role of the ecological and environmental context on landscape‐scale stability of plant productivity in agricultural pasture using remotely sensed enhanced vegetation index data. We determine whether four measures of stability (variability, magnitude of extreme anomalies, recovery time and recovery rate) are predicted by (a) species richness of vascular plants, (b) regional land cover heterogeneity and (c) climatic history. Stability of plant productivity was primarily associated with climatic history, particularly a history of extreme events. These effects outweighed any positive effects of species richness in the agricultural landscape. A history of variable and extreme climates both increased and decreased contemporary ecosystem stability, suggesting both cumulative and legacy effects, whereas land cover heterogeneity had no effect on stability. The landscape scale is a relevant spatial scale for the management of an ecosystem's stability. At this scale, we find that past climate is a stronger driver of stability in plant productivity than species richness, differing from results at finer field scales. Management should take an integrated approach by incorporating the environmental context of the landscape, such as its climatic history, and consider multiple components of stability to maintain functioning in landscapes that are particularly vulnerable to environmental change. A free Plain Language Summary can be found within the Supporting Information of this article.

Structural diversity consistently mediates species richness effects on aboveground carbon along altitudinal gradients in northern Ethiopian grazing exclosures

Grazing exclosures have been promoted as an effective and low-cost land management strategy to recover vegetation and associated functions in degraded landscapes in the tropics. While grazing exclosures can be important reservoirs of biodiversity and carbon, their potential in playing a dual role of conservation of biodiversity and mitigation of climate change effects is not yet established. To address this gap, we assessed the effect of diversity on aboveground carbon (AGC) and the relative importance of the driving biotic (functional diversity, functional composition and structural diversity) and abiotic (climate, topography and soil) mechanisms. We used a dataset from 133 inventory plots across three altitudinal zones, i.e., highland, midland and lowland, in northern Ethiopia, which allowed local- (within altitudinal zone) and broad- (across altitudinal zones) environmental scale analysis of diversity-AGC relationships. We found that species richness-AGC relationship shifted from neutral in highlands to positive in mid- and lowlands as well as across the altitudinal zones. Structural diversity was consistently the strongest mediator of the positive effects of species richness on AGC within and across altitudinal zones, whereas functional composition linked species richness to AGC at the broad environmental scale only. Abiotic factors had direct and indirect effects via biotic factors on AGC, but their relative importance varied with altitudinal zones. Our results indicate that the effect of species diversity on AGC was altitude-dependent and operated more strongly through structural diversity (representing niche complementarity effect) than functional composition (representing selection effect). Our study suggests that maintaining high structural diversity and managing functionally important species while promoting favourable climatic and soil conditions can enhance carbon storage in grazing exclosures.

Climate reverses directionality in the richness\u2013abundance relationship across the World\u2019s main forest biomes

More tree species can increase the carbon storage capacity of forests (here referred to as the more species hypothesis) through increased tree productivity and tree abundance resulting from complementarity, but they can also be the consequence of increased tree abundance through increased available energy (more individuals hypothesis). To test these two contrasting hypotheses, we analyse the most plausible pathways in the richness-abundance relationship and its stability along global climatic gradients. We show that positive effect of species richness on tree abundance only prevails in eight of the twenty-three forest regions considered in this study. In the other forest regions, any benefit from having more species is just as likely (9 regions) or even less likely (6 regions) than the effects of having more individuals. We demonstrate that diversity effects prevail in the most productive environments, and abundance effects become dominant towards the most limiting conditions. These findings can contribute to refining cost-effective mitigation strategies based on fostering carbon storage through increased tree diversity. Specifically, in less productive environments, mitigation measures should promote abundance of locally adapted and stress tolerant tree species instead of increasing species richness.

Testing the diversity–biomass relationship in riverine fish communities

AbstractAimWe examined the diversity–biomass relationship in stream fish communities and quantified direct and indirect effects of abiotic variables on this relationship.LocationFrance.Time period1992–2012.Major taxa studiedStream fishes.MethodsWe analysed the relationship between biodiversity (species richness and functional diversity) and fish community biomass at 1,357 stream sites distributed throughout France. We used piecewise path analysis to quantify effects of abiotic and biodiversity variables on biomass and assess relative contributions of native and non‐native species on the diversity–biomass relationship.ResultsBiodiversity showed a direct positive relationship with biomass after controlling for sampling effort, and direct effects of biodiversity variables on community biomass exceeded those of climate and physical habitat variables. Our analysis indicates that positive effects of species richness on biomass exceeded those of functional diversity. Indirect effects of abiotic variables mediated by biodiversity metrics indicated that biomass increased in warmer, lowland habitats. Human impact had no effect on native biodiversity but had a positive effect on non‐native biodiversity.Main conclusionsWe provide evidence that direct effects of biodiversity on community biomass outweigh those of abiotic variables in riverine fish communities, but resource partitioning alone is unlikely to drive the effects of biodiversity on biomass in this system. Quantification of the relative roles of anthropogenic impacts, biodiversity and environmental context in the regulation of ecosystem functioning will be necessary to understand the potential consequences of ongoing global change.

Species richness both impedes and promotes alien plant invasions in the Brazilian Cerrado

Worldwide, alien plant invasions have been intensively studied in the past decades, but mechanisms controlling the invasibility of native communities are not fully understood yet. The stochastic niche hypothesis predicts that species-rich plant communities are less prone to alien plant invasions than species-poor communities, which is supported by some but not all field studies, with some very species-rich communities such as the Brazilian Cerrado becoming heavily invaded. However, species-rich communities potentially contain a greater variety of facilitative interactions in resource exploitation than species-poor communities, from which invasive plants might benefit. This alternative hypothetical mechanism might explain why nutrient-poor, species-rich ecosystems are prone to invasion. Here we show that a high species richness both impedes and promotes invasive plants in the Brazilian Cerrado, using structural equation modelling and data from 38 field sites. We found support for the stochastic niche hypothesis through an observed direct negative influence of species richness on abundance of alien invasive species, but an indirect positive effect of species richness on invasive alien plants through soil phosphatase activity that enhances P availability was also found. These field observations were supported with results from a mesocosm experiment. Root phosphatase activity of plants increased with species richness in the mesocosms, which was associated with greater community P and N uptake. The most prominent alien grass species of the region, Melinis minutiflora, benefited most from the higher N and P availability in the species mixtures. Hence, this study provides a novel explanation of why species-richness may sometimes promote rather than impede invasion, and highlights the need to perform facilitation experiments in multi-species communities.

Shifts in functional compositions predict desired multifunctionality along fragmentation intensities in an alpine grassland

Abstract Grassland ecosystems are experiencing increasing perturbations from climate change and anthropogenic activities, which often cause vegetation fragmentation and grassland degradation. Although individual ecosystem functions have been extensively investigated in the vegetation fragmentation and grassland degradation processes, the multiple ecosystem functions are rarely evaluated. Moreover, little is known on how species richness and functional compositions are associated with individual functions, and how such functions contribute to multifunctionality during vegetation fragmentation. In this study, 14 interconnected functions were derived from the seven fragmentation intensities of grassland on the Tibetan Plateau. According to vegetation continuity, three communities (an ungrazed grasses and Kobresia humilis community, a lightly-grazed Kobresia humilis and K. pygmaea community, and a medium-grazed K. pygmaea community) were treated as the pre-patchiness stage, and four communities (a heavily-grazed cracked K. pygmaea community, an eroded K. pygmaea community, a severely eroded K. pygmaea community, and a bare black-soil crust) were treated as the vegetation patchiness stage. Multivariate diversity interaction framework was used to test the effects of species richness and functional composition on multiple ecosystem functions. Desired multifunctionality was assessed based on three relatively independent functions: aboveground biomass, belowground biomass, and soil organic carbon (SOC) storage. The threshold was identified to signify the rapid decline of the multifunctionality. Our results showed that positive effects of species richness were observed among most of the functions in pre-patchiness communities. Such effects disappeared in vegetation patchiness communities. Shifts in functional composition explained the large variations in most of the functions in the two stages. Furthermore, a trade-off in abundance between grasses and sedges contributed to the negative correlation between the aboveground and belowground biomass in the pre-patchiness stage. Consistent decreases in the abundance of functional groups led to a rapid decline of the individual functions, and, as a consequence, an abrupt decline in the multifunctionality. Our results demonstrated that managing grassland for ecosystem multifunctionality and species conservation requires a detailed understanding of the effects of the various drivers, including species richness, functional composition, grazing intensity, and climate conditions.

Species richness influences the spatial distribution of trees in European forests

The functioning of plant communities is strongly influenced by the number of species in the community and their spatial arrangement. This is because plants interact with their nearest neighbors and this interaction is expected to be stronger when the interacting individuals are ecologically similar in terms of resource use. Recent evidence shows that species richness alters the balance of intra‐ versus interspecific competition, but the effect of species richness, and phylogenetic and functional diversity on the spatial pattern of the plant communities remain less studied. Even far, how forest stand structure derived from past management practices can influence the relationship between species richness and spatial pattern is still unknown. Here, we evaluate the spatial distribution of woody individuals (DBH >7.5 cm) in 209 forest stands (i.e. plots) with an increasing level of species richness (from 1 up to 10 species) in six forest types along a latitudinal gradient in Europe. We used completely mapped plots to investigate the spatial pattern in each forest stand with point pattern techniques. We fitted linear models to analyze the effect of species richness (positively correlated with phylogenetic diversity) and functional diversity on tree spatial arrangements. We also controled this relationship by forest type and stand structure as a proxy of the management legacy. Our results showed a generalized positive effect of species richness and functional diversity on the degree of spatial clustering of trees, and on the spatial independence of tree sizes regardless of the forest type. Moreover, current tree spatial arrangements were still conditioned by its history of management; however its effect was independent of the number of species in the community. Our study showed that species richness and functional diversity are relevant attributes of forests influencing the spatial pattern of plant communities, and consequently forest functioning.

Species asynchrony and response diversity determine multifunctional stability of natural grasslands

Abstract A growing body of empirical evidence has suggested that biodiversity affects the simultaneous performance of multiple ecosystem functions (i.e. ecosystem multifunctionality). Given increasing environmental variability and uncertainty under global change, an emerging question is how biodiversity influences the stability of multiple functions (i.e. multifunctional stability). We currently know little, however, about the determinants and mechanisms of multifunctional stability, which is of practical importance for ensuring the sustainable provision of multiple functions. Here, we examined mechanisms contributing to stability (quantified as the ratio of the mean to the standard deviation) of multiple functions related to ecosystem productivity and carbon sequestration, including plant above‐ground and below‐ground productivity, litter production, gross primary productivity and ecosystem respiration, in a large grassland biodiversity experiment in Inner Mongolia. We found that community‐wide species asynchrony was a strong driver to stabilize multiple functions. Community‐wide asynchrony mediated the positive effects of species richness and response diversity (describing how species with similar effects on ecosystem function respond differently to environmental change) on multifunctional stability. However, species richness had a negative direct effect on multifunctional stability because, although it increased the averaged temporal mean of multiple functions, it strongly increased the averaged temporal standard deviation of multiple functions. The overall effects of species richness on multifunctional stability were thus negative, whereas those of response diversity were positive. Synthesis. The studied ecosystem functions related to ecosystem productivity and carbon sequestration are important in natural grasslands across the world. We conclude that species asynchrony and response diversity, rather than species richness, are key to the ecosystem multifunctional stability. The loss of response diversity and compensatory mechanisms would likely reduce the long‐term sustainability of grasslands in the face of global change.

Plant diversity loss reduces soil respiration across terrestrial ecosystems.

The rapid global biodiversity loss has led to the decline in ecosystem function. Despite the critical importance of soil respiration (Rs) in the global carbon and nutrient cycles, how plant diversity loss affects Rs remains uncertain. Here we present a meta-analysis using 446 paired observations from 95 published studies to evaluate the effects of plant and litter mixtures on Rs and its components. We found that total Rs and heterotrophic respiration (Rh) were, on average, greater in plant mixtures than expected from those of monocultures. These mixture effects increased with increasing species richness (SR) in both plant and litter mixtures. While the positive effects of species mixtures remained similar over time for total Rs, they increased over time for Rh in plant mixtures but decreased in litter mixtures. Despite the wide range of variations in mean annual temperature, annual aridity index, and ecosystem types, the plant mixture effects on total Rs and Rh did not change geographically, except for a more pronounced increase of total Rs in species mixtures with reduced water availability. Our structural equation models suggested that the positive effects of SR and stand age on total and Rh were driven by increased plant inputs and soil microbial biomass. Our results suggest that plant diversity loss has ubiquitous negative impacts on Rs, one of the fundamental carbon-cycle processes sustaining terrestrial element cycling and ecosystem function.

Covariation between tree size and shade tolerance modulates mixed-forest productivity

In tree communities, tree size inequality reduces productivity and interacts with tree shade tolerance to modulate stand productivity, with a higher productivity in stands where shade-intolerant species dominate shade-tolerant species in size. Positive diversity–productivity relationships have been reported in different plant communities, including tree communities. These effects may be strongly related to both structural diversity and functional diversity, but also to their interactions if there is a non-random distribution of species functional characteristics among canopy layers. We explore the relative effects on forest productivity of tree species diversity, tree size inequality, and species shade tolerance diversity, as well as the effect of the distribution of tree shade tolerance in the canopy. We used 11,054 mixed-species forest plots from the French Forest Inventory (IGN) distributed throughout France (2006–2011). We analyzed the effects of species richness, shade tolerance diversity, and height inequality on forest plot productivity, represented by basal area annual increment over a period of 5 years, while controlling for first-order structure characteristics (basal area and quadratic mean diameter) and environmental factors (soil water budget and sum of growing degree days). Using the covariance between tree height and shade tolerance in mixed species canopies, we also explored the effect of the distribution of species’ shade tolerance among canopy layers. The results showed a positive effect of species richness (effect size, 0.02) and a negative effect of height inequality (− 0.05) on mixed-forest productivity. We also showed that a negative covariance between shade tolerance and height (e.g., higher proportion of shade-tolerant species in lower height classes) increased productivity (0.01). Shade tolerance diversity did not affect productivity. In tree communities, as shown previously in monospecific forest stands, tree size inequality reduces productivity. This effect is modulated by the distribution of shade tolerance among canopy layers. Previous studies on species diversity effect have generally overlooked the importance of the size structure and the size hierarchy of functional characteristics. These effects are, however, crucial and deserve to be explored in greater detail.

Using root traits to understand temporal changes in biodiversity effects in grassland mixtures

Biodiversity–ecosystem functioning (BEF) studies typically show that species richness enhances community biomass, but the underlying mechanisms remain debated. Here, we combine metrics from BEF research that distinguish the contribution of dominant species (selection effects, SE) from those due to positive interactions such as resource partitioning (complementarity effects, CE) with a functional trait approach in an attempt to reveal the functional characteristics of species that drive community biomass in species mixtures. In a biodiversity experiment with 16 plant species in monocultures, 4‐species and 16‐species mixtures, we used aboveground biomass to determine the relative contributions of CE and SE to biomass production in mixtures in the second, dry year of the experiment. We also measured root traits (specific root length, root length density, root tissue density and the deep root fraction) of each species in monocultures and linked the calculated community weighted mean (CWM) trait values and trait diversity of mixtures to CE and SE. In the second year of the experiment, community biomass, CE and SE increased compared to the first year. The contribution of SE to this positive effect was greater than that of CE. The increased contribution of SE was associated with root traits: SE increased most in communities with high abundance of species with deep, thick and dense roots. In contrast, changes in CE were not related to trait diversity or CWM trait values. Together, these results suggest that increased positive effects of species richness on community biomass in a dry year were mainly driven by increased dominance of deep‐rooting species, supporting the insurance hypothesis of biodiversity. Positive CE indicates that other positive interactions did occur, but we could not find evidence that belowground resource partitioning or facilitation via root trait diversity was important for community productivity in our biodiversity experiment.

Tree diversity mitigates defoliation after a drought-induced tipping point.

Understanding the processes that underlie drought-related tree vitality loss is essential for anticipating future forest dynamics, and for developing management plans aiming at increasing the resilience of forests to climate change. Forest vitality has been continuously monitored in Europe since the acid rain alert in the 1980s, and the intensive monitoring plots of ICP Forests offer the opportunity to investigate the effects of air pollution and climate change on forest condition. By making use of over 100 long-term monitoring plots, where crown defoliation has been assessed extensively since 1990, we discovered a progressive shift from a negative to a positive effect of species richness on forest health. The observed tipping point in the balance of net interactions, from competition to facilitation, has never been reported from real ecosystems outside experimental conditions; and the strong temporal consistency of our observations with increasing drought stress emphasizes its climate change relevance. Furthermore, we show that higher species diversity has reduced the severity of defoliation in the long term. Our results confirm the greater resilience of diverse forests to future climate change-induced stress. More generally, they add to an accumulating body of evidence on the large potential of tree species mixtures to face manifold disturbances in a changing world.

Stabilizing effects in temporal fluctuations: management, traits, and species richness in high‐diversity communities

The loss of biodiversity is thought to have adverse effects on multiple ecosystem functions, including the decline of community stability. Decreased diversity reduces the strength of the portfolio effect, a mechanism stabilizing community temporal fluctuations. Community stability is also expected to decrease with greater variability in individual species populations and with synchrony of their fluctuations. In semi-natural meadows, eutrophication is one of the most important drivers of diversity decline; it is expected to increase species fluctuations and synchrony among them, all effects leading to lower community stability. With a 16-year time series of biomass data from a temperate species-rich meadow with fertilization and removal of the dominant species, we assessed population biomass temporal (co)variation under different management types and competition intensity, and in relation to species functional traits and to species diversity. Whereas the effect of dominant removal was relatively small (with a tendency toward lower stability), fertilization markedly decreased community stability (i.e., increased coefficient of variation in the total biomass) and species diversity. On average, the fluctuations of individual populations were mutually independent, with a slight tendency toward synchrony in unfertilized plots, and a tendency toward compensatory dynamics in fertilized plots and no effects of removal. The marked decrease of synchrony with fertilization, contrary to the majority of the results reported previously, follows the predictions of increased compensatory dynamics with increased asymmetric competition for light in a more productive environment. Synchrony increased also with species functional similarity stressing the importance of shared ecological strategies in driving similar species responses to weather fluctuations. As expected, the decrease of temporal stability of total biomass was mainly related to the decrease of species richness, with its effect remaining significant also after accounting for fertilization. The weakening of the portfolio effect with species richness decline is a crucial driver of community destabilization. However, the positive effect of species richness on temporal stability of total biomass was not due to increased compensatory dynamics, since synchrony increased with species richness. This shows that the negative effect of eutrophication on community stability does not operate through increasing synchrony, but through the reduction of diversity.

The mediation roles of intraspecific and interspecific functional trait diversity for linking the response of aboveground biomass to species richness across forest strata in a subtropical forest

Intraspecific functional trait diversity (FTD) has improved our understandings about the key mechanisms of species coexistence in plant communities. Yet, little is known about whether and how intraspecific and interspecific FTD mediate the response of aboveground biomass to species richness across forest strata (i.e. overstorey and understorey) and at whole-community in forests. To address this question, we tested the direct and indirect responses of aboveground biomass to species richness via intraspecific and interspecific FTD based on specific leaf area (FTDSLA) and leaf dry matter content (FTDLDMC) using structural equation modeling, in addition to the effects of soil nutrients, across 125 plots in a 5-ha subtropical forest in Eastern China. Results showed that intraspecific FTD mediates the response of aboveground biomass to species richness at overstorey and understorey strata, and whole-community level, while interspecific FTD did so at understorey strata only. At overstorey strata, 14% of the variation in aboveground biomass was accounted by the strong direct positive effect of species richness only. At understorey strata, soil nutrients had a strong negative direct effect followed by positive effects of species richness and FTDLDMC on aboveground biomass with 44–45% of the variation in both intraspecific and interspecific FTD models. At whole community level, 14% of the variation in aboveground biomass was explained by the strong positive direct effect of species richness followed by negative direct effect of intraspecific FTDSLA. Intraspecific and interspecific FTDLDMC had positively mediated the response of aboveground biomass to species richness at understorey strata through niche differentiation. Intraspecific FTD had negligible mediation role, whereas interspecific FTD had no role, for linking the response of aboveground biomass to species richness at overstorey strata, indicating that only dominant species with a specific functional strategy may largely determine community trait space. Intraspecific FTDSLA had negative relationship with aboveground biomass at the whole-community, probably due to the presence of a few large trees occupying larger niche space in a community. We conclude that intraspecific versus interspecific FTD plays a central role for linking the direct and indirect responses of aboveground biomass to species richness, but these relationships depend on the forest strata of a community.

The positive relationships between plant coverage, species richness, and aboveground biomass are ubiquitous across plant growth forms in semi-steppe rangelands

The positive relationships between biodiversity and aboveground biomass are important for biodiversity conservation and greater ecosystem functioning and services that humans depend on. However, the interaction effects of plant coverage and biodiversity on aboveground biomass across plant growth forms (shrubs, forbs and grasses) in natural rangelands are poorly studied. Here, we hypothesized that, while accounting for environmental factors and disturbance intensities, the positive relationships between plant coverage, biodiversity, and aboveground biomass are ubiquitous across plant growth forms in natural rangelands. We applied structural equation models (SEMs) using data from 735 quadrats across 35 study sites in semi-steppe rangelands in Iran. The combination of plant coverage and species richness rather than Shannon's diversity or species diversity (a latent variable of species richness and evenness) substantially enhance aboveground biomass across plant growth forms. In all selected SEMs, plant coverage had a strong positive direct effect on aboveground biomass (β = 0.72 for shrubs, 0.84 for forbs and 0.80 for grasses), followed by a positive effect of species richness (β = 0.26 for shrubs, 0.05 for forbs and 0.09 for grasses), and topographic factors. Disturbance intensity had a negative effect on plant coverage, whereas it had a variable effect on species richness across plant growth forms. Plant coverage had a strong positive total effect on aboveground biomass (β = 0.84 for shrubs, 0.88 for forbs, and 0.85 for grasses), followed by a positive effect of species richness, and a negative effect of disturbance intensity across plant growth forms. Our results shed light on the management of rangelands that is high plant coverage can significantly improve species richness and aboveground biomass across plant growth forms. We also found that high disturbance intensity due to heavy grazing has a strong negative effect on plant coverage rather than species richness in semi-steppe rangelands. This study suggests that proper grazing systems (e.g. rotational system) based on carrying capacity and stocking rate of a rangeland may be helpful for biodiversity conservation, better grazing of livestock, improvement of plant coverage and enhancement of aboveground biomass.