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Elevated CO2 (eCO2) experiments provide critical information to quantify the effects of rising CO2 on vegetation1–6. Many eCO2 experiments suggest that nutrient limitations modulate the local magnitude of the eCO2 effect on plant biomass1,3,5, but the global extent of these limitations has not been empirically quantified, complicating projections of the capacity of plants to take up CO27,8. Here, we present a data-driven global quantification of the eCO2 effect on biomass based on 138 eCO2 experiments. The strength of CO2 fertilization is primarily driven by nitrogen (N) in ~65% of global vegetation and by phosphorus (P) in ~25% of global vegetation, with N- or P-limitation modulated by mycorrhizal association. Our approach suggests that CO2 levels expected by 2100 can potentially enhance plant biomass by 12 ± 3% above current values, equivalent to 59 ± 13 PgC. The global-scale response to eCO2 we derive from experiments is similar to past changes in greenness9 and biomass10 with rising CO2, suggesting that CO2 will continue to stimulate plant biomass in the future despite the constraining effect of soil nutrients. Our research reconciles conflicting evidence on CO2 fertilization across scales and provides an empirical estimate of the biomass sensitivity to eCO2 that may help to constrain climate projections.

Active restoration often aims to accelerate ecosystem recovery. However, active restoration may not be worthwhile if its effects are overwhelmed by changes that occur passively. Moreover, it can be challenging to separate the effects of passive processes, such as dispersal and natural succession, from active restoration efforts. We assess the 24‐year impact of actively restoring a Minnesota old‐field grassland via seed addition of native tallgrass prairie species. We compared the abundance of four functional plant groups in actively restored plots against abundances in three reference classes: (1) unrestored plots undergoing passive recovery within the same old field, (2) passively recovering plots in two nearby old fields of similar age and (3) a chronosequence of 21 old fields within the same landscape. Active restoration led to a higher abundance of native grasses and forbs in the 36 m2 treatment plots. Seed addition was more effective if the original vegetation was first removed using herbicide, burning and tilling. However, long‐term conclusions about the efficacy of active restoration varied widely depending on the choice of reference class. In our small‐scale restoration experiment, native abundance was similarly high in both the actively restored and reference plots after 24 years, suggesting either (1) passive recovery or (2) local dispersal of native species from nearby treatment plots (i.e. cross‐contamination). In contrast, a comparison with two nearby reference fields suggested active restoration resulted in much higher native abundance relative to passive recovery. A smaller, positive effect was detected when we compared actively restored plots to the chronosequence of old fields. In the chronosequence, many passively recovering old fields had transitioned to native grass dominance naturally, although active restoration appeared to increase native forb abundance. Synthesis and applications: Our findings highlight the importance of using scale‐appropriate references for assessing the efficacy and need for active restoration. Comparing actively restored plots with the surrounding landscape, we found that active restoration and passive recovery led to similar plant communities after 24 years. Because local dispersal from actively restored sites can nearby references, caution should be exercised when evaluating long‐term restoration projects using only small‐scale experiments. [ABSTRACT FROM AUTHOR]

Despite decades of research documenting the consequences of naturalized and invasive plant species on ecosystem functions, our understanding of the functional underpinnings of these changes remains rudimentary. This is partially due to ineffective scaling of trait differences between native and naturalized species to whole plant communities. Working with data from over 75,000 plots and over 5,500 species from across the United States, we show that changes in the functional composition of communities associated with increasing abundance of naturalized species mirror the differences in traits between native and naturalized plants. We find that communities with greater abundance of naturalized species are more resource acquisitive aboveground and belowground, shorter, more shallowly rooted, and increasingly aligned with an independent strategy for belowground resource acquisition via thin fine roots with high specific root length. We observe shifts toward herbaceous-dominated communities but shifts within both woody and herbaceous functional groups follow community-level patterns for most traits. Patterns are remarkably similar across desert, grassland, and forest ecosystems. Our results demonstrate that the establishment and spread of naturalized species, likely in combination with underlying environmental shifts, leads to predictable and consistent changes in community-level traits that can alter ecosystem functions. [ABSTRACT FROM AUTHOR]

Invasive alien species (IAS) threaten human livelihoods and biodiversity globally. Increasing globalization facilitates IAS arrival, and environmental changes, including climate change, facilitate IAS establishment. Here we provide the first global, spatial analysis of the terrestrial threat from IAS in light of twenty-first century globalization and environmental change, and evaluate national capacities to prevent and manage species invasions. We find that one-sixth of the global land surface is highly vulnerable to invasion, including substantial areas in developing economies and biodiversity hotspots. The dominant invasion vectors differ between high-income countries (imports, particularly of plants and pets) and low-income countries (air travel). Uniting data on the causes of introduction and establishment can improve early-warning and eradication schemes. Most countries have limited capacity to act against invasions. In particular, we reveal a clear need for proactive invasion strategies in areas with high poverty levels, high biodiversity and low historical levels of invasion.

Exotic species dominate many communities; however the functional significance of species' biogeographic origin remains highly contentious. This debate is fuelled in part by the lack of globally replicated, systematic data assessing the relationship between species provenance, function and response to perturbations. We examined the abundance of native and exotic plant species at 64 grasslands in 13 countries, and at a subset of the sites we experimentally tested native and exotic species responses to two fundamental drivers of invasion, mineral nutrient supplies and vertebrate herbivory. Exotic species are six times more likely to dominate communities than native species. Furthermore, while experimental nutrient addition increases the cover and richness of exotic species, nutrients decrease native diversity and cover. Native and exotic species also differ in their response to vertebrate consumer exclusion. These results suggest that species origin has functional significance, and that eutrophication will lead to increased exotic dominance in grasslands.

Terrestrial ecosystem productivity is widely accepted to be nutrient limited(1). Although nitrogen (N) is deemed a key determinant of aboveground net primary production (ANPP)(2,3), the prevalence of co-limitation by N and phosphorus (P) is increasingly recognized(4-8). However, the extent to which terrestrial productivity is co-limited by nutrients other than N and P has remained unclear. Here, we report results from a standardized factorial nutrient addition experiment, in which we added N, P and potassium (K) combined with a selection of micronutrients (K+μ), alone or in concert, to 42 grassland sites spanning five continents, and monitored ANPP. Nutrient availability limited productivity at 31 of the 42 grassland sites. And pairwise combinations of N, P, and K+μ co-limited ANPP at 29 of the sites. Nitrogen limitation peaked in cool, high latitude sites. Our findings highlight the importance of less studied nutrients, such as K and micronutrients, for grassland productivity, and point to significant variations in the type and degree of nutrient limitation. We suggest that multiple-nutrient constraints must be considered when assessing the ecosystem-scale consequences of nutrient enrichment.

Summary: Uncertainty persists within trait‐based ecology, partly because few studies assess multiple axes of functional variation and their effect on plant performance.For 55 species from two semiarid grasslands, we quantified: (1) covariation between economic traits of leaves and absorptive roots, (2) covariation among economic traits, plant height, leaf size, and seed mass, and (3) relationships between these traits and species' abundance.Pairs of analogous leaf and root traits were at least weakly positively correlated (e.g. specific leaf area (SLA) and specific root length (SRL)). Two pairs of such traits, N content and DMC of leaves and roots, were at least moderately correlated (r > 0.5) whether species were grouped by site, taxonomic group and growth form, or life history. Root diameter was positively correlated with seed mass for all groups of species except annuals and monocots. Species with higher leaf dry matter content (LDMC) tended to be more abundant (r = 0.63). Annuals with larger seeds were more abundant (r = 0.69).Compared with global‐scale syntheses with many observations from mesic ecosystems, we observed stronger correlations between analogous leaf and root traits, weaker correlations between SLA and leaf N, and stronger correlations between SRL and root N. In dry grasslands, plant persistence may require coordination of above‐ and belowground traits, and dense tissues may facilitate dominance. [ABSTRACT FROM AUTHOR]

Increased nutrient inputs due to anthropogenic activity are expected to increase primary productivity across terrestrial ecosystems, but changes in allocation aboveground versus belowground with nutrient addition have different implications for soil carbon (C) storage. Thus, given that roots are major contributors to soil C storage, understanding belowground net primary productivity (BNPP) and biomass responses to changes in nutrient availability is essential to predicting carbon–climate feedbacks in the context of interacting global environmental changes. To address this knowledge gap, we tested whether a decade of nitrogen (N) and phosphorus (P) fertilization consistently influenced aboveground and belowground biomass and productivity at nine grassland sites spanning a wide range of climatic and edaphic conditions in the continental United States. Fertilization effects were strong aboveground, with both N and P addition stimulating aboveground biomass at nearly all sites (by 30% and 36%, respectively, on average). P addition consistently increased root production (by 15% on average), whereas other belowground responses to fertilization were more variable, ranging from positive to negative across sites. Site‐specific responses to P were not predicted by the measured covariates. Atmospheric N deposition mediated the effect of N fertilization on root biomass and turnover. Specifically, atmospheric N deposition was positively correlated with root turnover rates, and this relationship was amplified with N addition. Nitrogen addition increased root biomass at sites with low N deposition but decreased it at sites with high N deposition. Overall, these results suggest that the effects of nutrient supply on belowground plant properties are context dependent, particularly with regard to background N supply rates, demonstrating that site conditions must be considered when predicting how grassland ecosystems will respond to increased nutrient loading from anthropogenic activity. [ABSTRACT FROM AUTHOR]

Climate change, disturbance, and plant invasion threaten grassland ecosystems, but their combined and interactive effects are poorly understood. Here, we examine how the combination of disturbance and plant invasion influences the sensitivity of mixed‐grass prairie to elevated carbon dioxide (eCO2) and warming. We established subplots of intact prairie and disturbed/invaded prairie within a free‐air CO2 enrichment (to 600 ppmv) by infrared warming (+1.5°C day, 3°C night) experiment and followed plant and soil responses for 5 years. Elevated CO2 initially led to moderate increases in biomass and plant diversity in both intact and disturbed/invaded prairie, but these effects shifted due to strong eCO2 responses of the invasive forb Centaurea diffusa. In the final 3 years, biomass responses to eCO2 in disturbed/invaded prairie were 10 times as large as those in intact prairie (+186% vs. +18%), resulting in reduced rather than increased plant diversity (−17% vs. +10%). At the same time, warming interacted with disturbance/invasion and year, reducing the rate of topsoil carbon recovery following disturbance. The strength of these interactions demonstrates the need to incorporate disturbance into predictions of climate change effects. In contrast to expectations from studies in intact ecosystems, eCO2 may threaten plant diversity in ecosystems subject to soil disturbance and invasion. [ABSTRACT FROM AUTHOR]

The effects of climate change on plants and ecosystems are mediated by plant hydraulic traits, including interspecific and intraspecific variability of trait phenotypes. Yet, integrative and realistic studies of hydraulic traits and climate change are rare. In a semiarid grassland, we assessed the response of several plant hydraulic traits to elevated CO2 (+200 ppm) and warming (+1.5 to 3°C; day to night). For leaves of five dominant species (three graminoids and two forbs), and in replicated plots exposed to 7 years of elevated CO2, warming, or ambient climate, we measured: stomatal density and size, xylem vessel size, turgor loss point, and water potential (pre‐dawn). Interspecific differences in hydraulic traits were larger than intraspecific shifts induced by elevated CO2 and/or warming. Effects of elevated CO2 were greater than effects of warming, and interactions between treatments were weak or not detected. The forbs showed little phenotypic plasticity. The graminoids had leaf water potentials and turgor loss points that were 10% to 50% less negative under elevated CO2; thus, climate change might cause these species to adjust their drought resistance strategy away from tolerance and toward avoidance. The C4 grass also reduced allocation of leaf area to stomata under elevated CO2, which helps explain observations of higher soil moisture. The shifts in hydraulic traits under elevated CO2 were not, however, simply due to higher soil moisture. Integration of our results with others' indicates that common species in this grassland are more likely to adjust stomatal aperture in response to near‐term climate change, rather than anatomical traits; this contrasts with apparent effects of changing CO2 on plant anatomy over evolutionary time. Future studies should assess how plant responses to drought may be constrained by the apparent shift from tolerance (via low turgor loss point) to avoidance (via stomatal regulation and/or access to deeper soil moisture). [ABSTRACT FROM AUTHOR]