Functional diversity of leaf nitrogen concentrations drives grassland carbon fluxes

Little is known about the role of plant functional diversity for ecosystem‐level carbon (C) fluxes. To fill this knowledge gap, we translocated monoliths hosting communities with four and 16 sown species from a long‐term grassland biodiversity experiment (‘The Jena Experiment’) into a controlled environment facility for ecosystem research (Ecotron). This allowed quantifying the effects of plant diversity on ecosystem C fluxes as well as three parameters of C uptake efficiency (water and nitrogen use efficiencies and apparent quantum yield). By combining data on ecosystem C fluxes with vegetation structure and functional trait‐based predictors, we found that increasing plant species and functional diversity led to higher gross and net ecosystem C uptake rates. Path analyses and light response curves unravelled the diversity of leaf nitrogen concentration in the canopy as a key functional predictor of C fluxes, either directly or indirectly via LAI and aboveground biomass.     

Greater than the sum of the parts: how the species composition in different forest strata influence ecosystem function

The mechanisms underpinning forest biodiversity‐ecosystem function relationships remain unresolved. Yet, in heterogeneous forests, ecosystem function of different strata could be associated with traits or evolutionary relationships differently. Here, we integrate phylogenies and traits to evaluate the effects of elevational diversity on above‐ground biomass across forest strata and spatial scales. Community‐weighted means of height and leaf phosphorous concentration and functional diversity in specific leaf area exhibited positive correlations with tree biomass, suggesting that both positive selection effects and complementarity occur. However, high shrub biomass is associated with greater dissimilarity in seed mass and multidimensional trait space, while species richness or phylogenetic diversity is the most important predictor for herbaceous biomass, indicating that species complementarity is especially important for understory function. The strength of diversity‐biomass relationships increases at larger spatial scales. We conclude that strata‐ and scale‐ dependent assessments of community structure and function are needed to fully understand how biodiversity influences ecosystem function.     

Early subtropical forest growth is driven by community mean trait values and functional diversity rather than the abiotic environment

While functional diversity (FD) has been shown to be positively related to a number of ecosystem functions including biomass production, it may have a much less pronounced effect than that of environmental factors or species‐specific properties. Leaf and wood traits can be considered particularly relevant to tree growth, as they reflect a trade‐off between resources invested into growth and persistence. Our study focussed on the degree to which early forest growth was driven by FD, the environment (11 variables characterizing abiotic habitat conditions), and community‐weighted mean (CWM) values of species traits in the context of a large‐scale tree diversity experiment (BEF‐China). Growth rates of trees with respect to crown diameter were aggregated across 231 plots (hosting between one and 23 tree species) and related to environmental variables, FD, and CWM, the latter two of which were based on 41 plant functional traits. The effects of each of the three predictor groups were analyzed separately by mixed model optimization and jointly by variance partitioning. Numerous single traits predicted plot‐level tree growth, both in the models based on CWMs and FD, but none of the environmental variables was able to predict tree growth. In the best models, environment and FD explained only 4 and 31% of variation in crown growth rates, respectively, while CWM trait values explained 42%. In total, the best models accounted for 51% of crown growth. The marginal role of the selected environmental variables was unexpected, given the high topographic heterogeneity and large size of the experiment, as was the significant impact of FD, demonstrating that positive diversity effects already occur during the early stages in tree plantations.     

Dominance determines fish community biomass in a temperate seagrass ecosystem

Biodiversity and ecosystem function are often correlated, but there are multiple hypotheses about the mechanisms underlying this relationship. Ecosystem functions such as primary or secondary production may be maximized by species richness, evenness in species abundances, or the presence or dominance of species with certain traits. Here, we combine surveys of natural fish communities (conducted in July and August 2016) with morphological trait data to examine relationships between biodiversity and ecosystem function (quantified as fish community biomass) across 14 subtidal eelgrass meadows in the Northeast Pacific (54°N, 130°W). We employ both taxonomic and functional trait measures of diversity to investigate whether ecosystem function is best predicted by species diversity (complementarity hypothesis) or by the presence or dominance of species with particular trait values (selection or dominance hypotheses). After controlling for environmental variation, we find that fish community biomass is maximized when taxonomic richness and functional evenness are low, and in communities dominated by species with particular trait values, specifically those associated with benthic habitats and prey capture. While previous work on fish communities has found that species richness is often positively correlated with ecosystem function, our results instead highlight the capacity for regionally prevalent and locally dominant species to drive ecosystem function in moderately diverse communities. We discuss these alternate links between community composition and ecosystem function and consider their divergent implications for ecosystem valuation and conservation prioritization.     

Water availability drives aboveground biomass and bird richness in forest restoration plantings to achieve carbon and biodiversity cobenefits

To combat global warming and biodiversity loss, we require effective forest restoration that encourages recovery of species diversity and ecosystem function to deliver essential ecosystem services, such as biomass accumulation. Further, understanding how and where to undertake restoration to achieve carbon sequestration and biodiversity conservation would provide an opportunity to finance ecosystem restoration under carbon markets. We surveyed 30 native mixed‐species plantings in subtropical forests and woodlands in Australia and used structural equation modeling to determine vegetation, soil, and climate variables most likely driving aboveground biomass accrual and bird richness and investigate the relationships between plant diversity, aboveground biomass accrual, and bird diversity. We focussed on woodland and forest‐dependent birds, and functional groups at risk of decline (insectivorous, understorey‐nesting, and small‐bodied birds). We found that mean moisture availability strongly limits aboveground biomass accrual and bird richness in restoration plantings, indicating potential synergies in choosing sites for carbon and biodiversity purposes. Counter to theory, woody plant richness was a poor direct predictor of aboveground biomass accrual, but was indirectly related via significant, positive effects of stand density. We also found no direct relationship between aboveground biomass accrual and bird richness, likely because of the strong effects of moisture availability on both variables. Instead, moisture availability and patch size strongly and positively influenced the richness of woodland and forest‐dependent birds. For understorey‐nesting birds, however, shrub cover and patch size predicted richness. Stand age or area of native vegetation surrounding the patch did not influence bird richness. Our results suggest that in subtropical biomes, planting larger patches to higher densities, ideally using a diversity of trees and shrubs (characteristics of ecological plantings) in more mesic locations will enhance the provision of carbon and biodiversity cobenefits. Further, ecological plantings will aid the rapid recovery of woodland and forest bird richness, with comparable aboveground biomass accrual to less diverse forestry plantations.     

Functional dominance rather than taxonomic diversity and functional diversity mainly affects community aboveground biomass in the Inner Mongolia grassland

The relationship between biodiversity and productivity has been a hot topic in ecology. However, the relative importance of taxonomic diversity and functional characteristics (including functional dominance and functional diversity) in maintaining community productivity and the underlying mechanisms (including selection and complementarity effects) of the relationship between diversity and community productivity have been widely controversial. In this study, 194 sites were surveyed in five grassland types along a precipitation gradient in the Inner Mongolia grassland of China. The relationships between taxonomic diversity (species richness and the Shannon–Weaver index), functional dominance (the community‐weighted mean of four plant traits), functional diversity (Rao's quadratic entropy), and community aboveground biomass were analyzed. The results showed that (1) taxonomic diversity, functional dominance, functional diversity, and community aboveground biomass all increased from low to high precipitation grassland types; (2) there were significant positive linear relationships between taxonomic diversity, functional dominance, functional diversity, and community aboveground biomass; (3) the effect of functional characteristics on community aboveground biomass is greater than that of taxonomic diversity; and (4) community aboveground biomass depends on the community‐weighted mean plant height, which explained 57.1% of the variation in the community aboveground biomass. Our results suggested that functional dominance rather than taxonomic diversity and functional diversity mainly determines community productivity and that the selection effect plays a dominant role in maintaining the relationship between biodiversity and community productivity in the Inner Mongolia grassland.     

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.     

A landscape‐scale assessment of the relationship between grassland functioning,community diversity,and functional traits

Livestock farmers rely on a high and stable grassland productivity for fodder production to sustain their livelihoods. Future drought events related to climate change, however, threaten grassland functionality in many regions across the globe. The introduction of sustainable grassland management could buffer these negative effects. According to the biodiversity–productivity hypothesis, productivity positively associates with local biodiversity. The biodiversity–insurance hypothesis states that higher biodiversity enhances the temporal stability of productivity. To date, these hypotheses have mostly been tested through experimental studies under restricted environmental conditions, hereby neglecting climatic variations at a landscape‐scale. Here, we provide a landscape‐scale assessment of the contribution of species richness, functional composition, temperature, and precipitation on grassland productivity. We found that the variation in grassland productivity during the growing season was best explained by functional trait composition. The community mean of plant preference for nutrients explained 24.8% of the variation in productivity and the community mean of specific leaf area explained 18.6%, while species richness explained only 2.4%. Temperature and precipitation explained an additional 22.1% of the variation in productivity. Our results indicate that functional trait composition is an important predictor of landscape‐scale grassland productivity.     

Partitioning the effect of composition and diversity of tree communities on leaf litter decomposition and soil respiration

The decomposition of plant material is an important ecosystem process influencing both carbon cycling and soil nutrient availability. Quantifying how plant diversity affects decomposition is thus crucial for predicting the effect of the global decline in plant diversity on ecosystem functioning. Plant diversity could affect the decomposition process both directly through the diversity of the litter, and/or indirectly through the diversity of the host plant community and its affect on the decomposition environment. Using a biodiversity experiment with trees in which both functional and taxonomic diversity were explicitly manipulated independently, we tested the effects of the functional diversity and identity of the living trees separately and in combination with the functional diversity and identity of the decomposing litter on rates of litter decomposition and soil respiration. Plant traits, predominantly leaf chemical and physical traits, were correlated with both litter decomposition and soil respiration rates. Surface litter decomposition, quantified by mass loss in litterbags, was best explained by abundance‐weighted mean trait values of tree species from which the litter was assembled (functional identity). In contrast, soil respiration, which includes decomposition of dissolved organic carbon and root respiration, was best explained by the variance in trait values of the host trees (functional diversity). This research provides insight into the effect of loss of tree diversity in forests on soil processes. Such understanding is essential to predicting changes in the global carbon budget brought on by biodiversity loss.     

A functional trait-based approach to understand community assembly and diversity–productivity relationships over 7 years in experimental grasslands

Several multi-year biodiversity experiments have shown positive species richness–productivity relationships which strengthen over time, but the mechanisms which control productivity are not well understood. We used experimental grasslands (Jena Experiment) with mixtures containing different numbers of species (4, 8, 16 and 60) and plant functional groups (1–4; grasses, legumes, small herbs, tall herbs) to explore patterns of variation in functional trait composition as well as climatic variables as predictors for community biomass production across several years (from 2003 to 2009). Over this time span, high community mean trait values shifted from the dominance of trait values associated with fast growth to trait values suggesting a conservation of growth-related resources and successful reproduction. Increasing between-community convergence in means of several productivity-related traits indicated that environmental filtering and exclusion of competitively weaker species played a role during community assembly. A general trend for increasing functional trait diversity within and convergence among communities suggested niche differentiation through limiting similarity in the longer term and that similar mechanisms operated in communities sown with different diversity. Community biomass production was primarily explained by a few key mean traits (tall growth, large seed mass and leaf nitrogen concentration) and to a smaller extent by functional diversity in nitrogen acquisition strategies, functional richness in multiple traits and functional evenness in light-acquisition traits. Increasing species richness, presence of an exceptionally productive legume species (Onobrychis viciifolia) and climatic variables explained an additional proportion of variation in community biomass. In general, community biomass production decreased through time, but communities with higher functional richness in multiple traits had high productivities over several years. Our results suggest that assembly processes within communities with an artificially maintained species composition maximize functional diversity through niche differentiation and exclusion of weaker competitors, thereby maintaining their potential for high productivity.     

Emergent interactions influence functional traits and success of dune building ecosystem engineers

Aims Dune building processes are affected by interactions between the growth of ecosystem engineering dune grasses and environmental factors associated with disturbance such as sand burial and sea spray. Research investigating how species interactions influence dune community structure and functional trait responses in high abiotic stress environments is minimal. We investigated how species interactions influence the functional trait responses of three dominant dune grasses to common abiotic stressors.Methods We performed a multi-factorial greenhouse experiment by planting three common dune grasses (Ammophila breviligulata Fern., Uniola paniculata L. and Spartina patens Muhl.) in different interspecific combinations, using sand burial and sea spray as abiotic stressors. Sand burial was applied once at the beginning of the study. Sea spray was applied three times per week using a calibrated spray bottle. Morphological functional trait measurements (leaf elongation, maximum root length, aboveground biomass and belowground biomass) were collected at the end of the study. The experiment continued from May 2015 to August 2015.Important findings Species interactions between A. breviligulata and U. paniculata negatively affected dune building function traits of A. breviligulata, indicating that interactions with U. paniculata could alter dune community structure. Furthermore, A. breviligulata had a negative interaction with S. patens, which decreased S. patens functional trait responses to abiotic stress. When all species occurred together, the interactions among species brought about coexistence of all three species. Our data suggest that species interactions can change traditional functional trait responses of dominant species to abiotic stress.     

Do temperate tree species diversity and identity influence soil microbial community function and composition?

Studies of biodiversity–ecosystem function in treed ecosystems have generally focused on aboveground functions. This study investigates intertrophic links between tree diversity and soil microbial community function and composition. We examined how microbial communities in surface mineral soil responded to experimental gradients of tree species richness (SR ), functional diversity (FD ), community‐weighted mean trait value (CWM ), and tree identity. The site was a 4‐year‐old common garden experiment near Montreal, Canada, consisting of deciduous and evergreen tree species mixtures. Microbial community composition, community‐level physiological profiles, and respiration were evaluated using phospholipid fatty acid (PLFA ) analysis and the MicroResp^? system, respectively. The relationship between tree species richness and glucose‐induced respiration (GIR ), basal respiration (BR ), metabolic quotient (qCO ₂) followed a positive but saturating shape. Microbial communities associated with species mixtures were more active (basal respiration [BR ]), with higher biomass (glucose‐induced respiration [GIR ]), and used a greater number of carbon sources than monocultures. Communities associated with deciduous tree species used a greater number of carbon sources than those associated with evergreen species, suggesting a greater soil carbon storage capacity. There were no differences in microbial composition (PLFA ) between monocultures and SR mixtures. The FD and the CWM of several functional traits affected both BR and GIR . In general, the CWM of traits had stronger effects than did FD , suggesting that certain traits of dominant species have more effect on ecosystem processes than does FD . Both the functions of GIR and BR were positively related to aboveground tree community productivity. Both tree diversity (SR ) and identity (species and functional identity—leaf habit) affected soil microbial community respiration, biomass, and composition. For the first time, we identified functional traits related to life‐history strategy, as well as root traits that influence another trophic level, soil microbial community function, via effects on BR and GIR .     

Is the relationship between algal diversity and biomass in North American lakes consistent with biodiversity experiments?

Over the past few decades, a large body of research has examined how biodiversity loss influences the functioning of ecosystems, as well as the cascading impacts on the goods and services ecosystems provide to humanity. The relationship between biodiversity and ecosystem functions quantified in prior experiments suggests that initial losses of biodiversity have relatively small impacts on properties like community biomass production; however, beyond some threshold, increasing losses lead to accelerating declines in function. Some have questioned whether a saturating relationship between diversity and community biomass production is an artifact of overly simplified experiments that manipulate diversity in homogeneous conditions over short time‐scales in which niche differences may not be realized. Others have questioned whether even the modest effects of biodiversity observed in experiments would be discernible in natural systems where they could be over‐ridden by the stronger influence of abiotic factors. Here, we used a biogeographic dataset to assess how the taxonomic richness of aquatic primary producers relates to community biomass in unmanipulated lake ecosystems in the US, and then compared these findings to prior experiments. We used structural equation modeling to evaluate hypotheses about the effects of algal richness on community biomass while accounting for covariance with environmental parameters measured in the USEPA's National Lakes Assessment (NLA), which sampled 1157 freshwater lakes. These analyses converged on a single best‐fit model (χ²= 0.31, p = 0.58) wherein community algal biomass was a function of three explanatory variables – nitrogen, phosphorus, and algal richness. The quantitative magnitude of the algal diversity (x) – biomass (y) relationship in the NLA dataset is statistically greater than that documented in the average biodiversity experiment. It did, however, lie at approximately the 75th percentile of experimental relationships, indicating the diversity–biomass relationship in unmanipulated lakes is within the range that has been characterized experimentally.     

Leaf‐trait plasticity and species vulnerability to climate change in a Mongolian steppe

Climate change is expected to modify plant assemblages in ways that will have major consequences for ecosystem functions. How climate change will affect community composition will depend on how individual species respond, which is likely related to interspecific differences in functional traits. The extraordinary plasticity of some plant traits is typically neglected in assessing how climate change will affect different species. In the Mongolian steppe, we examined whether leaf functional traits under ambient conditions and whether plasticity in these traits under altered climate could explain climate‐induced biomass responses in 12 co‐occurring plant species. We experimentally created three probable climate change scenarios and used a model selection procedure to determine the set of baseline traits or plasticity values that best explained biomass response. Under all climate change scenarios, plasticity for at least one leaf trait correlated with change in species performance, while functional leaf‐trait values in ambient conditions did not. We demonstrate that trait plasticity could play a critical role in vulnerability of species to a rapidly changing environment. Plasticity should be considered when examining how climate change will affect plant performance, species' niche spaces, and ecological processes that depend on plant community composition.     

草甸和沼泽植物群落功能多样性与生产力的关系

功能多样性-生产力关系研究结果支持质量比假说和多样性假说, 但对于这两种假说的适用条件尚有争议。通过对吉林省西部草甸和沼泽植物群落的地上生物量、2个物种多样性指标(物种丰富度和Shannon-Weaver指数)、7种植物性状的两类功能多样性指标(群落权重均值和Rao二次熵), 以及土壤环境因子进行调查测量, 研究了群落功能多样性与生产力的关系。结果表明: 1)功能多样性与生产力的关系比物种多样性与生产力的关系更为密切; 2)功能群落权重均值解释生产力变异的能力好于Rao二次熵, 即优势物种对群落生产力的影响作用更大; 3)水淹条件影响着功能多样性与生产力的关系, 以群落权重均值为基础的质量比假说适于解释草甸群落功能多样性与生产力的关系, 而以Rao二次熵为基础的多样性假说适于解释有强烈环境筛(水淹)的沼泽群落功能多样性与生产力的关系。     

Relationship between functional diversity and productivity in meadow and marsh plant communities

功能多样性-生产力关系研究结果支持质量比假说和多样性假说, 但对于这两种假说的适用条件尚有争议。通过对吉林省西部草甸和沼泽植物群落的地上生物量、2个物种多样性指标(物种丰富度和Shannon-Weaver指数)、7种植物性状的两类功能多样性指标(群落权重均值和Rao二次熵), 以及土壤环境因子进行调查测量, 研究了群落功能多样性与生产力的关系。结果表明: 1)功能多样性与生产力的关系比物种多样性与生产力的关系更为密切; 2)功能群落权重均值解释生产力变异的能力好于Rao二次熵, 即优势物种对群落生产力的影响作用更大; 3)水淹条件影响着功能多样性与生产力的关系, 以群落权重均值为基础的质量比假说适于解释草甸群落功能多样性与生产力的关系, 而以Rao二次熵为基础的多样性假说适于解释有强烈环境筛(水淹)的沼泽群落功能多样性与生产力的关系。     

The unseen invaders: introduced earthworms as drivers of change in plant communities in North American forests (a meta‐analysis)

Globally, biological invasions can have strong impacts on biodiversity as well as ecosystem functioning. While less conspicuous than introduced aboveground organisms, introduced belowground organisms may have similarly strong effects. Here, we synthesize for the first time the impacts of introduced earthworms on plant diversity and community composition in North American forests. We conducted a meta‐analysis using a total of 645 observations to quantify mean effect sizes of associations between introduced earthworm communities and plant diversity, cover of plant functional groups, and cover of native and non‐native plants. We found that plant diversity significantly declined with increasing richness of introduced earthworm ecological groups. While plant species richness or evenness did not change with earthworm invasion, our results indicate clear changes in plant community composition: cover of graminoids and non‐native plant species significantly increased, and cover of native plant species (of all functional groups) tended to decrease, with increasing earthworm biomass. Overall, these findings support the hypothesis that introduced earthworms facilitate particular plant species adapted to the abiotic conditions of earthworm‐invaded forests. Further, our study provides evidence that introduced earthworms are associated with declines in plant diversity in North American forests. Changing plant functional composition in these forests may have long‐lasting effects on ecosystem functioning.     

Predicting individual plant performance in grasslands

Plant functional traits are widely used to predict community productivity. However, they are rarely used to predict individual plant performance in grasslands. To assess the relative importance of traits compared to environment, we planted seedlings of 20 common grassland species as phytometers into existing grassland communities varying in land‐use intensity. After 1 year, we dug out the plants and assessed root, leaf, and aboveground biomass, to measure plant performance. Furthermore, we determined the functional traits of the phytometers and of all plants growing in their local neighborhood. Neighborhood impacts were analyzed by calculating community‐weighted means (CWM) and functional diversity (FD) of every measured trait. We used model selection to identify the most important predictors of individual plant performance, which included phytometer traits, environmental conditions (climate, soil conditions, and land‐use intensity), as well as CWM and FD of the local neighborhood. Using variance partitioning, we found that most variation in individual plant performance was explained by the traits of the individual phytometer plant, ranging between 19.30% and 44.73% for leaf and aboveground dry mass, respectively. Similarly, in a linear mixed effects model across all species, performance was best predicted by phytometer traits. Among all environmental variables, only including land‐use intensity improved model quality. The models were also improved by functional characteristics of the local neighborhood, such as CWM of leaf dry matter content, root calcium concentration, and root mass per volume as well as FD of leaf potassium and root magnesium concentration and shoot dry matter content. However, their relative effect sizes were much lower than those of the phytometer traits. Our study clearly showed that under realistic field conditions, the performance of an individual plant can be predicted satisfyingly by its functional traits, presumably because traits also capture most of environmental and neighborhood conditions.     

物种多样性和功能性状驱动高寒草原地上生物量对长期禁牧的响应

围封对草地生物多样性和初级生产力的影响是草地生态学研究的热点问题之一。基于2013—2021年在青藏高原东北部紫花针茅(Stipa purpurea)高寒草原围栏内外植物群落长期调查数据,从物种多样性、功能性状的角度解析了高寒草原地上生物量对长期围封的生态响应过程。结果表明：(1)围封处理对高寒草原物种多样性的负效应具有强烈的时间依赖性。围封处理显著提高地上生物量,但也显著降低了生物量稳定性和异步性,意味着高寒草原稳定的、可持续的生态系统服务功能被长期围封处理削弱。(2)植物功能性状对长期围封处理表现出差异性响应模式;与叶绿素性状相比,叶形态性状对长期围封处理表现出更强的敏感性。(3)物种多样性和功能性状与地上生物量之间均存在显著相关关系,并且物种多样性的影响被功能性状调控进而对地上生物量发挥间接效应,群落加权性状和功能分异度共同对草地生物量发挥直接的主导效应。研究结果证明了植物功能性状通过介导物种多样性与其共同驱动高寒草原地上生物量对长期围封的响应。因此,在未来草地管理过程中,同步研究植物物种和功能属性对于全面揭示生态系统的响应机制至关重要。