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Anthropogenic changes in biodiversity and atmospheric temperature significantly influence ecosystem processes. However, little is known about potential interactive effects of plant diversity and warming on essential ecosystem properties, such as soil microbial functions and element cycling. We studied the effects of orthogonal manipulations of plant diversity (one, four, and 16 species) and warming (ambient, +1.5°C, and +3°C) on soil microbial biomass, respiration, growth after nutrient additions, and activities of extracellular enzymes in 2011 and 2012 in the BAC (biodiversity and climate) perennial grassland experiment site at Cedar Creek, Minnesota, USA. Focal enzymes are involved in essential biogeochemical processes of the carbon, nitrogen, and phosphorus cycles. Soil microbial biomass and some enzyme activities involved in the C and N cycle increased significantly with increasing plant diversity in both years. In addition, 16-species mixtures buffered warming induced reductions in topsoil water content. We found no interactive effects of plant diversity and warming on soil microbial biomass and growth rates. However, the activity of several enzymes (1,4-β-glucosidase, 1,4-β-N-acetylglucosaminidase, phosphatase, peroxidase) depended on interactions between plant diversity and warming with elevated activities of enzymes involved in the C, N, and P cycles at both high plant diversity and high warming levels. Increasing plant diversity consistently decreased microbial biomass-specific enzyme activities and altered soil microbial growth responses to nutrient additions, indicating that plant diversity changed nutrient limitations and/or microbial community composition. In contrast to our expectations, higher plant diversity only buffered temperature effects on soil water content, but not on microbial functions. Temperature effects on some soil enzymes were greatest at high plant diversity. In total, our results suggest that the fundamental temperature ranges of soil microbial communities may be sufficiently broad to buffer their functioning against changes in temperature and that plant diversity may be a dominant control of soil microbial processes in a changing world.

Microorganisms mobilize phosphorus (P) in soil by solubilizing bound inorganic P from soil minerals and by mineralizing organic P via phosphatase enzymes. Nitrogen (N) inputs are predicted to increase through human activities and shift plants to be more P limited, increasing the importance of P mobilization processes for plant nutrition. We studied how the relative abundance of P-solubilizing bacteria (PSB), PSB community composition, and phosphatase activity respond to N and P addition (+N, +P, +NP) in grassland soils spanning large biogeographic gradients. The studied soils are located in South Africa, USA, and UK and part of a globally coordinated nutrient addition experiment. We show that the abundance of PSB in the topsoil was reduced by 18 % in the N and by 41 % in the NP treatment compared to the control. In contrast, phosphatase activity was significantly higher in the N treatment than in the control across all soils. Soil C:P ratio, sand content, pH, and water-extractable P together explained 71 % of the variance of the abundance of PSB across all study sites and all treatments. Further, the community of PSB in the N and NP addition treatment differed significantly from the control. Taken together, this study shows that N addition reduced the relative abundance of PSB, altered the PSB community, and increased phosphatase activity, whereas P addition had no impact. Increasing atmospheric N deposition may therefore increase mineralization of organic P and decrease solubilization of bound inorganic P, possibly changing P mobilization processes in grassland soils. Consequently, in ecosystems in which plant P nutrition depends on bound inorganic P, increased N inputs might diminish P supply and thus aggravate P limitation and constrain plant productivity.

1. In grasslands and savannas, fire regime—frequently a major determinant of woody encroachment, herbaceous species composition and diversity, and nutrient cycling—is influenced by the quantity and characteristics of plant fuel. Laboratory studies reveal variation in flammability among herbaceous species, but field experiments are needed to assess whether herbaceous species composition meaningfully affects ecosystem-scale fire behaviour. 2. In our North American tallgrass prairie study system, grasses' thinner leaves and longer leaf retention appeared to create a finer, more aerated, more connected fuel bed than forbs. We tested the hypothesis that grasses promote fire spread area, fire intensity, and associated facets of fire behaviour more strongly than an equivalent mass of forbs. 3. We characterized spring fires over multiple years in 315 annually ignited plots spanning profound gradients of plant biomass, cover, and grass:forb ratio that resulted from species richness and composition treatments, in a 20-year grassland biodiversity experiment. 4. Grasses increased fire spread and associated facets of fire behaviour, compared with an equivalent biomass or cover of forbs. Grass dominance increased fire spread area—or equivalents increased fire frequency at any given point. For fire to spread through 50% of the 9 m × 9 m plot area required approximately twice as high an abundance of forbs as of grasses. Grass dominance also resulted in fires that advanced faster, were more intense (higher rates of heat release per unit fireline length), caused more damage to plants, and released heat to greater heights. Fire temperature at 50 cm above-ground was about twice as high in plots with only grasses as in plots with the same biomass of forbs. 5. Synthesis. Even within herbaceous ecosystems that may appear homogenously flammable compared with less flammable woody ecosystems, fuel quality—specifically, the proportional abundance of grasses—combines with fuel quantity and ignitions to determine effective fire regime at a given point. In spring burns, grassdominated plots burn more completely and generate higher temperatures, and thus better suppress woody plants and volatilize more nutrients, than forb-dominated plots (holding all else equal).

1. Following the removal of invasive plant species, most land managers rely on natural succession to re-establish native plant communities. However, insufficient native propagule pressure combined with legacy effects of invasive plant species means that passive approaches to restoration are often inadequate to establish native communities and prevent reinvasion. 2. In this paper, we review literature evaluating the ability of active revegetation to suppress re-establishment of invaders in grasslands and woodlands. 3. We find that existing literature consistently demonstrates reduced performance of invasive plant species in revegetated grasslands, but that the magnitude of impact on invasive plants is highly variable. In contrast, the efficacy of revegetation in woodlands has rarely been reported, but the small number of such studies are consistent with results from grasslands. 4. Synthesis and applications. Our review highlights the mechanisms that lead revegetation suppressing invasive plants in grasslands and identifies knowledge gaps associated with revegetation using woody species or targeting woody invaders. We recommend concerted efforts be made to evaluate the viability of woody plant revegetation and the efficacy of revegetation in woodlands. Furthermore, we suggest that land managers may need to embrace novel species assemblages in order to prevent reinvasion.

The persistence of invasive plant species in soil seedbanks can pose a significant obstacle to effectively managing invasive plant populations. Rhamnus cathartica (common buckthorn, hereafter ‘buckthorn’) is a wide-spread invader of forest understories in North America that can quickly re-establish following removal, in part due to germination of buckthorn seedbanks. Although empirical evidence seems slight, influential organizations communicate that buckthorn seedbanks endure for at least six years. In order to assess the accuracy of such messaging, we characterize the duration of buckthorn in soil seedbanks by monitoring germination of both planted and naturally-occurring seeds. Across the 13,232 buckthorn seeds planted, germination occurred almost entirely in the first two years after planting (96.6% and 3.3% in the first and second year after planting, respectively). Our observations of naturally-occurring seedbanks displayed similar patterns, with 97.9% and 1.9% of all newly emerged seedlings found in the first and second years following removal of mature buckthorn stands, respectively. These findings indicate that re-occupation by buckthorn following removal results more from incomplete removal, an initial flush of germinants, and dispersal from outside the site than from long-lived seedbanks. Therefore, if buckthorn is removed comprehensively during two years of intensive initial management then subsequent buckthorn re-invasion is likely to be sporadic and to require less intensive, targeted follow-up management.

Humans dominate many important Earth system processes including the nitrogen (N) cycle. Atmospheric N deposition affects fundamental processes such as carbon cycling, climate regulation, and biodiversity, and could result in changes to fundamental Earth system processes such as primary production. Both modelling and experimentation have suggested a role for anthropogenically altered N deposition in increasing productivity, nevertheless, current understanding of the relative strength of N deposition with respect to other controls on production such as edaphic conditions and climate is limited. Here we use an international multiscale data set to show that atmospheric N deposition is positively correlated to aboveground net primary production (ANPP) observed at the 1-m 2 level across a wide range of herbaceous ecosystems. N deposition was a better predictor than climatic drivers and local soil conditions, explaining 16% of observed variation in ANPP globally with an increase of 1 kg N·ha −1 ·yr −1 increasing ANPP by 3%. Soil pH explained 8% of observed variation in ANPP while climatic drivers showed no significant relationship. Our results illustrate that the incorporation of global N deposition patterns in Earth system models are likely to substantially improve estimates of primary production in herbaceous systems. In herbaceous systems across the world, humans appear to be partially driving local ANPP through impacts on the N cycle.

Modeling fire spread as an infection process is intuitive: An ignition lights a patch of fuel, which infects its neighbor, and so on. Infection models produce nonlinear thresholds, whereby fire spreads only when fuel connectivity and infection probability are sufficiently high. These thresholds are fundamental both to managing fire and to theoretical models of fire spread, whereas applied fire models more often apply quasiempirical approaches. Here, we resolve this tension by quantifying thresholds in fire spread locally, using field data from individual fires (n = 1,131) in grassy ecosystems across a precipitation gradient (496 to 1,442 mm mean annual precipitation) and evaluating how these scaled regionally (across 533 sites) and across time (1989 to 2012 and 2016 to 2018) using data from Kruger National Park in South Africa. An infection model captured observed patterns in individual fire spread better than competing models. The proportion of the landscape that burned was well described by measurements of grass biomass, fuel moisture, and vapor pressure deficit. Regionally, averaging across variability resulted in quasi-linear patterns. Altogether, results suggest that models aiming to capture fire responses to global change should incorporate nonlinear fire spread thresholds but that linear approximations may sufficiently capture medium-term trends under a stationary climate.

Transitions from wind pollination to insect pollination were pivotal to the radiation of land plants, yet only a handful are known and the trait shifts required are poorly understood. We tested the hypothesis that a transition to insect pollination took place in the ancestrally wind-pollinated sedges (Cyperaceae) and that floral traits modified during this transition have functional significance. We paired putatively insect-pollinated Cyperus obtusiflorus and Cyperus sphaerocephalus with related, co-flowering, co-occurring wind-pollinated species, and compared pairs in terms of pollination mode and functional roles of floral traits. Experimentally excluding insects reduced seed set by 56—89% in putatively insect-pollinated species but not in intermingled wind-pollinated species. The pollen of putatively insect-pollinated species was less motile in a wind tunnel than that of wind-pollinated species. Bees, beetles and flies preferred inflorescences, and color-matched white or yellow models, of putatively insect-pollinated species over inflorescences, or color-matched brown models, of wind-pollinated species. Floral scents of putatively insect-pollinated species were chemically consistent with those of other insect-pollinated plants, and attracted pollinators; wind-pollinated species were unscented. These results show that a transition from wind pollination to insect pollination occurred in sedges and shed new light on the function of traits involved in this important transition.

Plant stoichiometry, the relative concentration of elements, is a key regulator of ecosystem functioning and is also being altered by human activities. In this paper we sought to understand the global drivers of plant stoichiometry and compare the relative contribution of climatic vs. anthropogenic effects. We addressed this goal by measuring plant elemental (C, N, P and K) responses to eutrophication and vertebrate herbivore exclusion at eighteen sites on six continents. Across sites, climate and atmospheric N deposition emerged as strong predictors of plot-level tissue nutrients, mediated by biomass and plant chemistry. Within sites, fertilization increased total plant nutrient pools, but results were contingent on soil fertility and the proportion of grass biomass relative to other functional types. Total plant nutrient pools diverged strongly in response to herbivore exclusion when fertilized; responses were largest in ungrazed plots at low rainfall, whereas herbivore grazing dampened the plant community nutrient responses to fertilization. Our study highlights (1) the importance of climate in determining plant nutrient concentrations mediated through effects on plant biomass, (2) that eutrophication affects grassland nutrient pools via both soil and atmospheric pathways and (3) that interactions among soils, herbivores and eutrophication drive plant nutrient responses at small scales, especially at water-limited sites.

Fosamine ammonium (Krenite®) is a foliar herbicide that primarily targets woody plant species; however, formal evaluations of its efficacy and potential for non-target impacts are scarce in the literature. The few tests of fosamine ammonium that exist focus primarily on its use in open environments, and the value of fosamine ammonium in controlling invasive understory shrubs is unclear. Here, we test the impact of fosamine ammonium on invasive common buckthorn (Rhamnus cathartica L.) and co-occurring herbaceous plants across six forest sites in Minnesota, USA. Rhamnus cathartica treated with fosamine ammonium had a 95% mortality rate, indicating high efficacy of fosamine ammonium for use against R. cathartica. Non-target impacts varied between forbs and graminoids such that forb cover was reduced by up to 85%, depending on site, whereas graminoid cover was sparse and impacts of fosamine ammonium on graminoids were unclear. These results indicate that while fosamine ammonium can provide effective control of R. cathartica and other understory shrubs, there is potential for significant non-target impacts following its use. We therefore suggest that land managers carefully consider the timing, rate, and application method of fosamine ammonium to achieve desired target and non-target impacts.