Extreme drought impacts have been underestimated in grasslands and shrublands globally.

Climate change is increasing the frequency and severity of short-term (~1 y) drought events-the most common duration of drought-globally. Yet the impact of this intensification of drought on ecosystem functioning remains poorly resolved. This is due in part to the widely disparate approaches ecologists have employed to study drought, variation in the severity and duration of drought studied, and differences among ecosystems in vegetation, edaphic and climatic attributes that can mediate drought impacts. To overcome these problems and better identify the factors that modulate drought responses, we used a coordinated distributed experiment to quantify the impact of short-term drought on grassland and shrubland ecosystems. With a standardized approach, we imposed ~a single year of drought at 100 sites on six continents. Here we show that loss of a foundational ecosystem function-aboveground net primary production (ANPP)-was 60% greater at sites that experienced statistically extreme drought (1-in-100-y event) vs. those sites where drought was nominal (historically more common) in magnitude (35% vs. 21%, respectively). This reduction in a key carbon cycle process with a single year of extreme drought greatly exceeds previously reported losses for grasslands and shrublands. Our global experiment also revealed high variability in drought response but that relative reductions in ANPP were greater in drier ecosystems and those with fewer plant species. Overall, our results demonstrate with unprecedented rigor that the global impacts of projected increases in drought severity have been significantly underestimated and that drier and less diverse sites are likely to be most vulnerable to extreme drought.

Dryland sensitivity to climate change and variability using nonlinear dynamics.

Primary productivity response to climatic drivers varies temporally, indicating state-dependent interactions between climate and productivity. Previous studies primarily employed equation-based approaches to clarify this relationship, ignoring the state-dependent nature of ecological dynamics. Here, using 40 y of climate and productivity data from 48 grassland sites across Mongolia, we applied an equation-free, nonlinear time-series analysis to reveal sensitivity patterns of productivity to climate change and variability and clarify underlying mechanisms. We showed that productivity responded positively to annual precipitation in mesic regions but negatively in arid regions, with the opposite pattern observed for annual mean temperature. Furthermore, productivity responded negatively to decreasing annual aridity that integrated precipitation and temperature across Mongolia. Productivity responded negatively to interannual variability in precipitation and aridity in mesic regions but positively in arid regions. Overall, interannual temperature variability enhanced productivity. These response patterns are largely unrecognized; however, two mechanisms are inferable. First, time-delayed climate effects modify annual productivity responses to annual climate conditions. Notably, our results suggest that the sensitivity of annual productivity to increasing annual precipitation and decreasing annual aridity can even be negative when the negative time-delayed effects of annual precipitation and aridity on productivity prevail across time. Second, the proportion of plant species resistant to water and temperature stresses at a site determines the sensitivity of productivity to climate variability. Thus, we highlight the importance of nonlinear, state-dependent sensitivity of productivity to climate change and variability, accurately forecasting potential biosphere feedback to the climate system.

Reintroducing bison results in long-running and resilient increases in grassland diversity.

The widespread extirpation of megafauna may have destabilized ecosystems and altered biodiversity globally. Most megafauna extinctions occurred before the modern record, leaving it unclear how their loss impacts current biodiversity. We report the long-term effects of reintroducing plains bison (Bison bison) in a tallgrass prairie versus two land uses that commonly occur in many North American grasslands: 1) no grazing and 2) intensive growing-season grazing by domesticated cattle (Bos taurus). Compared to ungrazed areas, reintroducing bison increased native plant species richness by 103% at local scales (10 m2) and 86% at the catchment scale. Gains in richness continued for 29 y and were resilient to the most extreme drought in four decades. These gains are now among the largest recorded increases in species richness due to grazing in grasslands globally. Grazing by domestic cattle also increased native plant species richness, but by less than half as much as bison. This study indicates that some ecosystems maintain a latent potential for increased native plant species richness following the reintroduction of native herbivores, which was unmatched by domesticated grazers. Native-grazer gains in richness were resilient to an extreme drought, a pressure likely to become more common under future global environmental change.

Heterogeneity promotes resilience in restored prairie: Implications for the environmental heterogeneity hypothesis.

Enhancing resilience in formerly degraded ecosystems is an important goal of restoration ecology. However, evidence for the recovery of resilience and its underlying mechanisms require long-term experiments and comparison with reference ecosystems. We used data from an experimental prairie restoration that featured long-term soil heterogeneity manipulations and data from two long-term experiments located in a comparable remnant (reference) prairie to (1) quantify the recovery of ecosystem functioning (i.e., productivity) relative to remnant prairie, (2) compare the resilience of restored and remnant prairies to a natural drought, and (3) test whether soil heterogeneity enhances resilience of restored prairie. We compared sensitivity and legacy effects between prairie types (remnant and restored) and among four prairie sites that included two remnant prairie sites and prairie restored under homogeneous and heterogeneous soil conditions. We measured sensitivity and resilience as the proportional change in aboveground net primary productivity (ANPP) during and following drought (sensitivity and legacy effects, respectively) relative to average ANPP based on 4 pre-drought years (2014-2017). In nondrought years, total ANPP was similar between remnant and restored prairie, but remnant prairie had higher grass productivity and lower forb productivity compared with restored prairie. These ANPP patterns generally persisted during drought. The sensitivity of total ANPP to drought was similar between restored and remnant prairie, but grasses in the restored prairie were more sensitive to drought. Post-drought legacy effects were more positive in the restored prairie, and we attributed this to the more positive and less variable legacy response of forb ANPP in the restored prairie, especially in the heterogeneous soil treatment. Our results suggest that productivity recovers in restored prairie and exhibits similar sensitivity to drought as in remnant prairie. Furthermore, creating heterogeneity promotes forb productivity and enhances restored prairie resilience to drought.

As above, not so below: Long-term dynamics of net primary production across a dryland transition zone.

Drylands are key contributors to interannual variation in the terrestrial carbon sink, which has been attributed primarily to broad-scale climatic anomalies that disproportionately affect net primary production (NPP) in these ecosystems. Current knowledge around the patterns and controls of NPP is based largely on measurements of aboveground net primary production (ANPP), particularly in the context of altered precipitation regimes. Limited evidence suggests belowground net primary production (BNPP), a major input to the terrestrial carbon pool, may respond differently than ANPP to precipitation, as well as other drivers of environmental change, such as nitrogen deposition and fire. Yet long-term measurements of BNPP are rare, contributing to uncertainty in carbon cycle assessments. Here, we used 16 years of annual NPP measurements to investigate responses of ANPP and BNPP to several environmental change drivers across a grassland-shrubland transition zone in the northern Chihuahuan Desert. ANPP was positively correlated with annual precipitation across this landscape; however, this relationship was weaker within sites. BNPP, on the other hand, was weakly correlated with precipitation only in Chihuahuan Desert shrubland. Although NPP generally exhibited similar trends among sites, temporal correlations between ANPP and BNPP within sites were weak. We found chronic nitrogen enrichment stimulated ANPP, whereas a one-time prescribed burn reduced ANPP for nearly a decade. Surprisingly, BNPP was largely unaffected by these factors. Together, our results suggest that BNPP is driven by a different set of controls than ANPP. Furthermore, our findings imply belowground production cannot be inferred from aboveground measurements in dryland ecosystems. Improving understanding around the patterns and controls of dryland NPP at interannual to decadal scales is fundamentally important because of their measurable impact on the global carbon cycle. This study underscores the need for more long-term measurements of BNPP to improve assessments of the terrestrial carbon sink, particularly in the context of ongoing environmental change.

Soil nematode assemblages respond to interacting environmental changes.

Multi-factor experiments suggest that interactions among environmental changes commonly influence biodiversity and community composition. However, most field experiments manipulate only single factors. Soil food webs are critical to ecosystem health and may be particularly sensitive to interactions among environmental changes that include soil warming, eutrophication, and altered precipitation. Here, we asked how environmental changes interacted to alter soil nematode communities in a northern Chihuahuan Desert grassland. Factorial manipulations of nitrogen, winter rainfall, and nighttime warming matched predictions for regional environmental change. Warming reduced nematode diversity by 25% and genus-level richness by 32%, but declines dissipated with additional winter rain, suggesting that warming effects occurred via drying. Interactions between precipitation and nitrogen also altered nematode community composition, but only weakly affected total nematode abundance, indicating that most change involved reordering of species abundances. Specifically, under ambient precipitation, nitrogen fertilizer reduced bacterivores by 68% and herbivores by 73%, but did not affect fungivores. In contrast, under winter rain addition, nitrogen fertilization increased bacterivores by 95%, did not affect herbivores, and doubled fungivore abundance. Rain can reduce soil nitrogen availability and increase turnover in the microbial loop, potentially promoting the recovery of nematode populations overwhelmed by nitrogen eutrophication. Nematode communities were not tightly coupled to plant community composition and may instead track microbes, including biocrusts or decomposers. Our results highlight the importance of interactions among environmental change stressors for shaping the composition and function of soil food webs in drylands.

Resolving the Dust Bowl paradox of grassland responses to extreme drought.

During the 1930s Dust Bowl drought in the central United States, species with the C3 photosynthetic pathway expanded throughout C4-dominated grasslands. This widespread increase in C3 grasses during a decade of low rainfall and high temperatures is inconsistent with well-known traits of C3 vs. C4 pathways. Indeed, water use efficiency is generally lower, and photosynthesis is more sensitive to high temperatures in C3 than C4 species, consistent with the predominant distribution of C3 grasslands in cooler environments and at higher latitudes globally. We experimentally imposed extreme drought for 4 y in mixed C3/C4 grasslands in Kansas and Wyoming and, similar to Dust Bowl observations, also documented three- to fivefold increases in C3/C4 biomass ratios. To explain these paradoxical responses, we first analyzed long-term climate records to show that under nominal conditions in the central United States, C4 grasses dominate where precipitation and air temperature are strongly related (warmest months are wettest months). In contrast, C3 grasses flourish where precipitation inputs are less strongly coupled to warm temperatures. We then show that during extreme drought years, precipitation-temperature relationships weaken, and the proportion of precipitation falling during cooler months increases. This shift in precipitation seasonality provides a mechanism for C3 grasses to respond positively to multiyear drought, resolving the Dust Bowl paradox. Grasslands are globally important biomes and increasingly vulnerable to direct effects of climate extremes. Our findings highlight how extreme drought can indirectly alter precipitation seasonality and shift ecosystem phenology, affecting function in ways not predictable from key traits of C3 and C4 species.

Linking changes in species composition and biomass in a globally distributed grassland experiment.

Global change drivers, such as anthropogenic nutrient inputs, are increasing globally. Nutrient deposition simultaneously alters plant biodiversity, species composition and ecosystem processes like aboveground biomass production. These changes are underpinned by species extinction, colonisation and shifting relative abundance. Here, we use the Price equation to quantify and link the contributions of species that are lost, gained or that persist to change in aboveground biomass in 59 experimental grassland sites. Under ambient (control) conditions, compositional and biomass turnover was high, and losses (i.e. local extinctions) were balanced by gains (i.e. colonisation). Under fertilisation, the decline in species richness resulted from increased species loss and decreases in species gained. Biomass increase under fertilisation resulted mostly from species that persist and to a lesser extent from species gained. Drivers of ecological change can interact relatively independently with diversity, composition and ecosystem processes and functions such as aboveground biomass due to the individual contributions of species lost, gained or persisting.

Soil carbon stocks in temperate grasslands differ strongly across sites but are insensitive to decade-long fertilization.

Enhancing soil carbon (C) storage has the potential to offset human-caused increases in atmospheric CO2 . Rising CO2 has occurred concurrently with increasing supply rates of biologically limiting nutrients such as nitrogen (N) and phosphorus (P). However, it is unclear how increased supplies of N and P will alter soil C sequestration, particularly in grasslands, which make up nearly a third of non-agricultural land worldwide. Here, we leverage a globally distributed nutrient addition experiment (the Nutrient Network) to examine how a decade of N and P fertilization (alone and in combination) influenced soil C and N stocks at nine grassland sites spanning the continental United States. We measured changes in bulk soil C and N stocks and in three soil C fractions (light and heavy particulate organic matter, and mineral-associated organic matter fractions). Nutrient amendment had variable effects on soil C and N pools that ranged from strongly positive to strongly negative, while soil C and N pool sizes varied by more than an order of magnitude across sites. Piecewise SEM clarified that small increases in plant C inputs with fertilization did not translate to greater soil C storage. Nevertheless, peak season aboveground plant biomass (but not root biomass or production) was strongly positively related to soil C storage at seven of the nine sites, and across all nine sites, soil C covaried with moisture index and soil mineralogy, regardless of fertilization. Overall, we show that site factors such as moisture index, plant productivity, soil texture, and mineralogy were key predictors of cross-site soil C, while nutrient amendment had weaker and site-specific effects on C sequestration. This suggests that prioritizing the protection of highly productive temperate grasslands is critical for reducing future greenhouse gas losses arising from land use change.

Richness, not evenness, varies across water availability gradients in grassy biomes on five continents.

We sought to understand the role that water availability (expressed as an aridity index) plays in determining regional and global patterns of richness and evenness, and in turn how these water availability-diversity relationships may result in different richness-evenness relationships at regional and global scales. We examined relationships between water availability, richness and evenness for eight grassy biomes spanning broad water availability gradients on five continents. Our study found that relationships between richness and water availability switched from positive for drier (South Africa, Tibet and USA) vs. negative for wetter (India) biomes, though were not significant for the remaining biomes. In contrast, only the India biome showed a significant relationship between water availability and evenness, which was negative. Globally, the richness-water availability relationship was hump-shaped, however, not significant for evenness. At the regional scale, a positive richness-evenness relationship was found for grassy biomes in India and Inner Mongolia, China. In contrast, this relationship was weakly concave-up globally. These results suggest that different, independent factors are determining patterns of species richness and evenness in grassy biomes, resulting in differing richness-evenness relationships at regional and global scales. As a consequence, richness and evenness may respond very differently across spatial gradients to anthropogenic changes, such as climate change.

Global change effects on plant communities are magnified by time and the number of global change factors imposed.

Global change drivers (GCDs) are expected to alter community structure and consequently, the services that ecosystems provide. Yet, few experimental investigations have examined effects of GCDs on plant community structure across multiple ecosystem types, and those that do exist present conflicting patterns. In an unprecedented global synthesis of over 100 experiments that manipulated factors linked to GCDs, we show that herbaceous plant community responses depend on experimental manipulation length and number of factors manipulated. We found that plant communities are fairly resistant to experimentally manipulated GCDs in the short term (<10 y). In contrast, long-term (≥10 y) experiments show increasing community divergence of treatments from control conditions. Surprisingly, these community responses occurred with similar frequency across the GCD types manipulated in our database. However, community responses were more common when 3 or more GCDs were simultaneously manipulated, suggesting the emergence of additive or synergistic effects of multiple drivers, particularly over long time periods. In half of the cases, GCD manipulations caused a difference in community composition without a corresponding species richness difference, indicating that species reordering or replacement is an important mechanism of community responses to GCDs and should be given greater consideration when examining consequences of GCDs for the biodiversity-ecosystem function relationship. Human activities are currently driving unparalleled global changes worldwide. Our analyses provide the most comprehensive evidence to date that these human activities may have widespread impacts on plant community composition globally, which will increase in frequency over time and be greater in areas where communities face multiple GCDs simultaneously.

Fire frequency, state change and hysteresis in tallgrass prairie.

Hysteresis is a fundamental characteristic of alternative stable state theory, yet evidence of hysteresis is rare. In mesic grasslands, fire frequency regulates transition from grass- to shrub-dominated system states. It is uncertain, however, if increasing fire frequency can reverse shrub expansion, or if grass-shrub dynamics exhibit hysteresis. We implemented annual burning in two infrequently burned grasslands and ceased burning in two grasslands burned annually. With annual fires, grassland composition converged on that of long-term annually burned vegetation due to rapid recovery of grass cover, although shrubs persisted. When annual burning ceased, shrub cover increased, but community composition did not converge with a long-term infrequently burned reference site because of stochastic and lagged dispersal by shrubs, reflecting hysteresis. Our results demonstrated that annual burning can slow, but not reverse, shrub encroachment. In addition, reversing fire frequencies resulted in hysteresis because vegetation trajectories from grassland to shrubland differed from those of shrubland to grassland.

Determinants of community compositional change are equally affected by global change.

Global change is impacting plant community composition, but the mechanisms underlying these changes are unclear. Using a dataset of 58 global change experiments, we tested the five fundamental mechanisms of community change: changes in evenness and richness, reordering, species gains and losses. We found 71% of communities were impacted by global change treatments, and 88% of communities that were exposed to two or more global change drivers were impacted. Further, all mechanisms of change were equally likely to be affected by global change treatments-species losses and changes in richness were just as common as species gains and reordering. We also found no evidence of a progression of community changes, for example, reordering and changes in evenness did not precede species gains and losses. We demonstrate that all processes underlying plant community composition changes are equally affected by treatments and often occur simultaneously, necessitating a wholistic approach to quantifying community changes.

Water and nitrogen shape winter annual plant diversity and community composition in near-urban Sonoran Desert preserves.

Increased nitrogen (N) deposition threatens global biodiversity, but its effects in arid urban ecosystems are not well studied. In addition to altered N availability, urban environments also experience increases in other pollutants, decreased population connectivity, and altered biotic interactions, which can further impact biodiversity. In deserts, annual plant communities make up most of the plant diversity, support wildlife, and contribute to nutrient cycling and ecosystem processes. Functional tradeoffs allowing coexistence of a diversity of annual plant species are well established, but maintenance of diversity in urban conditions and with increased availability of limiting nutrients has not been explored. We conducted a 13-year N and phosphorus (P) addition experiment in Sonoran Desert preserves in and around Phoenix, AZ, to test how nutrient availability interacts with growing season precipitation, urban location, and microhabitat to affect winter annual plant diversity. Using structural equation modeling and generalized linear mixed modeling, we found that annual plant taxonomic diversity was significantly reduced in N-enriched and urban plots. Water availability in both current and previous growing seasons impacted annual plant diversity, with significant interaction effects showing increased diversity in wetter years and greater responsiveness of the community to water following a wet year. However, there were no significant interactions between N enrichment and water availability, urban location, or microhabitat. Lowered diversity in urban preserves may be partly attributable to increased urban N deposition. Changes in biodiversity of showy species like annual wildflowers in urban preserves can have important implications for connections between urban residents and nature, and reduced diversity and community restructuring with N enrichment represents a challenge for future preservation of aridland biodiversity.

Divergent responses of primary production to increasing precipitation variability in global drylands.

Interannual variability in precipitation has increased globally as climate warming intensifies. The increased variability impacts both terrestrial plant production and carbon (C) sequestration. However, mechanisms driving these changes are largely unknown. Here, we examined mechanisms underlying the response of aboveground net primary production (ANPP) to interannual precipitation variability in global drylands with mean annual precipitation (MAP) <500 mm year-1 , using a combined approach of data synthesis and process-based modeling. We found a hump-shaped response of ANPP to precipitation variability along the MAP gradient. The response was positive when MAP < ~300 mm year-1 and negative when MAP was higher than this threshold, with a positive peak at 140 mm year-1 . Transpiration and subsoil water content mirrored the response of ANPP to precipitation variability; evaporation responded negatively and water loss through runoff and drainage responded positively to precipitation variability. Mean annual temperature, soil type, and plant physiological traits all altered the magnitude but not the pattern of the response of ANPP to precipitation variability along the MAP gradient. By extrapolating to global drylands (<500 mm year-1  MAP), we estimated that ANPP would increase by 15.2 ± 6.0 Tg C year-1 in arid and hyper-arid lands and decrease by 2.1 ± 0.5 Tg C year-1 in dry sub-humid lands under future changes in interannual precipitation variability. Thus, increases in precipitation variability will enhance primary production in many drylands in the future.

Is a drought a drought in grasslands? Productivity responses to different types of drought.

Drought, defined as a marked deficiency of precipitation relative to normal, occurs as periods of below-average precipitation or complete failure of precipitation inputs, and can be limited to a single season or prolonged over multiple years. Grasslands are typically quite sensitive to drought, but there can be substantial variability in the magnitude of loss of ecosystem function. We hypothesized that differences in how drought occurs may contribute to this variability. In four native Great Plains grasslands (three C4- and one C3-dominated) spanning a ~ 500-mm precipitation gradient, we imposed drought for four consecutive years by (1) reducing each rainfall event by 66% during the growing season (chronic drought) or (2) completely excluding rainfall during a shorter portion of the growing season (intense drought). The drought treatments were similar in magnitude but differed in the following characteristics: event number, event size and length of dry periods. We observed consistent drought-induced reductions (28-37%) in aboveground net primary production (ANPP) only in the C4-dominated grasslands. In general, intense drought reduced ANPP more than chronic drought, with little evidence that drought duration altered this pattern. Conversely, belowground net primary production (BNPP) was reduced by drought in all grasslands (32-64%), with BNPP reductions greater in intense vs. chronic drought treatments in the most mesic grassland. We conclude that grassland productivity responses to drought did not strongly differ between these two types of drought, but when differences existed, intense drought consistently reduced function more than chronic drought.

Sensitivity of primary production to precipitation across the United States.

Primary production, a key regulator of the global carbon cycle, is highly responsive to variations in climate. Yet, a detailed, continental-scale risk assessment of climate-related impacts on primary production is lacking. We combined 16 years of MODIS NDVI data, a remotely sensed proxy for primary production, with observations from 1218 climate stations to derive values of ecosystem sensitivity to precipitation and aridity. For the first time, we produced an empirically-derived map of ecosystem sensitivity to climate across the conterminous United States. Over this 16-year period, annual primary production values were most sensitive to precipitation and aridity in dryland and grassland ecosystems. Century-long trends measured at the climate stations showed intensifying aridity and climatic variability in many of these sensitive regions. Dryland ecosystems in the western US may be particularly vulnerable to reductions in primary production and consequent degradation of ecosystem services as climate change and variability increase in the future.

Microbial processing of plant remains is co-limited by multiple nutrients in global grasslands.

Microbial processing of aggregate-unprotected organic matter inputs is key for soil fertility, long-term ecosystem carbon and nutrient sequestration and sustainable agriculture. We investigated the effects of adding multiple nutrients (nitrogen, phosphorus and potassium plus nine essential macro- and micro-nutrients) on decomposition and biochemical transformation of standard plant materials buried in 21 grasslands from four continents. Addition of multiple nutrients weakly but consistently increased decomposition and biochemical transformation of plant remains during the peak-season, concurrent with changes in microbial exoenzymatic activity. Higher mean annual precipitation and lower mean annual temperature were the main climatic drivers of higher decomposition rates, while biochemical transformation of plant remains was negatively related to temperature of the wettest quarter. Nutrients enhanced decomposition most at cool, high rainfall sites, indicating that in a warmer and drier future fertilized grassland soils will have an even more limited potential for microbial processing of plant remains.

Direct and indirect effects of temperature and precipitation on alpine seed banks in the Tibetan Plateau.

Plant community responses to global environmental change focus primarily on aboveground vegetation; however, the important role of the seed bank is frequently neglected. Specifically, the direct and indirect effects of changes in temperature and precipitation on seed banks remain poorly understood, yet seed banks provide a vital source of ecosystem resilience to global environmental change. We used a structural equation model to explore the direct and indirect effects of temperature, precipitation, and other biotic and abiotic factors on soil seed bank community composition using 1,026 soil seed bank samples from 57 sites along an elevation gradient that served as a space-for-time substitution for changing climate in the Tibetan Plateau. Seed bank richness was negatively correlated with both precipitation and temperature, but neither climate factor affected seed bank density. Temperature was also negatively correlated with vegetation species richness, which was positively correlated with seed bank richness and density. Both precipitation and temperature were positively correlated with soil total N, and total N was negatively correlated with vegetation richness. Both precipitation and temperature were negatively correlated with soil pH, and soil pH was negatively correlated with vegetation richness, but positively correlated with seed bank richness and density. Increasing precipitation and temperature would decrease seed bank diversity through direct effects as well as indirectly by decreasing vegetation diversity. Soil pH and total N emerged as the most important soil abiotic factors for seed bank diversity. Increasing precipitation and temperature under climate change may increase the extinction risk of some species in the seed bank by altering bet-hedging and risk-spreading strategies, which will degrade natural restoration ability and ultimately ecosystem resilience. This research is important because it identifies the potential underlying mechanistic basis of climate change impacts on seed banks through effects of aboveground vegetation and belowground biotic and abiotic factors.

Soil heterogeneity increases plant diversity after 20 years of manipulation during grassland restoration.

The "environmental heterogeneity hypothesis" predicts that variability in resources promotes species coexistence, but few experiments support this hypothesis in plant communities. A previous 15-yr test of this hypothesis in a prairie restoration experiment demonstrated a weak effect of manipulated soil resource heterogeneity on plant diversity. This response was attributed to a transient increase in richness following a post-restoration supplemental propagule addition, occasionally higher diversity under nutrient enrichment, and reduced cover of a dominant species in a subset of soil treatments. Here, we report community dynamics under continuous propagule addition in the same experiment, corresponding to 16-20 yr of restoration, in response to altered availability and heterogeneity of soil resources. We also quantified traits of newly added species to determine if heterogeneity increases the amount and variety of niches available for new species to exploit. The heterogeneous treatment contained a factorial combination of altered nutrient availability and soil depth; control plots had no manipulations. Total diversity and richness were higher in the heterogeneous treatment during this 5-yr study due to higher cover, diversity, and richness of previously established forbs, particularly in the N-enriched subplots. All new species added to the experiment exhibited unique trait spaces, but there was no evidence that heterogeneous plots contained a greater variety of new species representing a wider range of trait spaces relative to the control treatment. The richness and cover of new species was higher in N-enriched soil, but the magnitude of this response was small. Communities assembling under long-term N addition were dominated by different species among subplots receiving added N, leading to greater dispersion of communities among the heterogeneous relative to control plots. Contrary to the deterministic mechanism by which heterogeneity was expected to increase diversity (greater variability in resources for new species to exploit), higher diversity in the heterogeneous plots resulted from destabilization of formerly grass-dominated communities in N-enriched subplots. While we do not advocate increasing available soil N at large scales, we conclude that the positive effect of environmental heterogeneity on diversity can take decades to materialize and depend on development of stochastic processes in communities with strong establishment limitation.