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Silicon (Si) accumulation is known to alleviate various biotic and abiotic stressors in plants with potential ecological consequences. However, for dicotyledonous plants our understanding of Si variation remains limited. We conducted a comparative experimental study to investigate (1) interspecific variation of foliar Si concentrations across 37 dicotyledonous forbs of temperate grasslands, (2) intraspecific variation in foliar Si concentration in response to soil Si availability, the influence of (3) phylogenetic relatedness, and (4) habitat association to moisture. Foliar Si differed markedly (approx. 70-fold) across the investigated forbs, with some species exhibiting Si accumulation similar to grasses. Foliar Si increased with soil Si availability, but the response varied across species: species with higher Si accumulation capacity showed a stronger response, indicating that they did not actively upregulate Si uptake under low soil Si availability. Foliar Si showed a pronounced phylogenetic signal, i.e., closely related species exhibited more similar foliar Si concentrations than distantly related species. Significant differences in foliar Si concentration within closely related species pairs nevertheless support that active Si uptake and associated high Si concentrations has evolved multiple times in forbs. Foliar Si was not higher in species associated with drier habitats, implying that in dicotyledonous forbs of temperate grasslands high foliar Si is not an adaptive trait to withstand drought. Our results demonstrated considerable inter- and intraspecific variation in foliar Si concentration in temperate forbs. This variation should have pervasive, but so far understudied, ecological consequences for community composition and functioning of temperate grasslands under land-use and climate change. Supplementary Information The online version contains supplementary material available at 10.1007/s00442-021-04978-9.

Non-native plants may benefit, briefly or permanently, from natural enemy release in their invaded range, or may form novel interactions with native enemy species. Likewise, newly arrived herbivores may develop novel associations with native plants or, where their hosts have arrived ahead of them, re-establish interactions that existed previously in their ancestral ranges. Predicting outcomes from this diversity of novel and re-established interactions between plants and their herbivores presents a major challenge for invasion biology. We report on interactions between the recently arrived invasive planthopper Prokelisia marginata, and the multi-ploidy Spartina complex of four native and introduced species in Britain, each representing a different level of shared evolutionary history with the herbivore. As predicted, S. alterniflora, the ancestral host, was least impacted by planthopper herbivory, with the previously unexposed native S. maritima, a nationally threatened species, suffering the greatest impacts on leaf length gain, new leaf growth and relative water content. Contrary to expectations, glasshouse trials showed P. marginata to preferentially oviposit on the invasive allododecaploid S. anglica, on which it achieved earlier egg hatch, faster nymphal development, larger female body size and greatest final population size. We suggest P. marginata is in the process of rapid adaptation to maximise its performance on what is now the most abundant and widespread host in Britain. The diversity of novel and re-established interactions of the herbivore with this multi-ploidy complex makes this a highly valuable system for the study of the evolutionary ecology of plant–insect interactions and their influence on invasion dynamics.

Although microbial participation in litter decomposition is widely known within terrestrial soils, the role and significance of microorganisms during the aerial standing litter phase of decomposition remains poorly investigated. We examined the fungi inhabiting standing leaf litter of Schizachyrium scoparium and Schizachyrium tenerum in a Longleaf Pine savanna ecosystem and estimated their contribution to litter decomposition. We identified fungal phylotypes associated with leaf litter and quantified leaf C mass loss, fungal biomass production, and microbial respiration during decomposition. These data were used to construct budgets estimating C flow into and through fungi. Significant losses in S. scoparium (55%) and S. tenerum (67%) leaf C mass were observed during standing decomposition along with concomitant increases in fungal biomass, which reached a maximum of 36 and 33 mgC/g detrital C, respectively. Cumulative fungal production during decomposition totaled 99 ± 6 mgC/g initial detrital C in S. scoparium and 73 ± 5 mgC/g initial detrital C in S. tenerum, indicating that 18 and 11% of the litter C was converted into fungal biomass, respectively. Corresponding estimates of cumulative fungal respiration totaled 106 ± 7 and 174 ± 11 mgC/g initial detrital C in S. scoparium and S. tenerum, respectively. Next generation sequencing identified several fungal phylotypes, with the majority of sequences belonging to the Ascomycota (Dothideomycetes) and Basidiomycota (Agaricomycetes). Fungal phylotypes were similar between litter species and changed over time, showing a successional pattern. These findings extend our understanding of fungal processes to standing litter in terrestrial ecosystems, and highlight the quantitative importance of fungi in C cycling processes.

Physiological integration of connected plants of the same clone, or ramets, often increases clonal fitness when ramets differ in resource supply. However, review of the literature found that no study has directly tested the hypothesis that integration can increase the ability of clones to compete against other species. To test this, we grew two-ramet clonal fragments of the stoloniferous, perennial herb Fragaria chiloensis in which none, one, or both of the ramets had neighbors of a naturally co-occurring, dominant grass, Bromus carinatus, and connections between ramets were either severed to prevent integration or left intact. We also grew four-ramet fragments in which all ramets had neighbors and connections were severed or intact. Severance decreased the final leaf mass and area of two-ramet fragments by 25% and their final total mass by 15% when just one ramet was grown with B. carinatus. Severance had no significant effect on the total mass of fragments when none or all of the ramets were grown with the grass. This provides the first direct evidence that physiological integration can increase the competitive ability of clonal plant species, though only when competition is spatially heterogeneous. Integration may thus enable plant clones to grow into plant communities and to compete within communities with fine-scale disturbance. However, integration may not increase the competitive ability of clonal plants within uniformly dense communities of taller species.

Ecosystems are faced with an onslaught of co-occurring global change drivers. While frequently studied independently, the effects of multiple global change drivers have the potential to be additive, antagonistic, or synergistic. Global warming, for example, may intensify the effects of more variable precipitation regimes with warmer temperatures increasing evapotranspiration and thereby amplifying the effect of already dry soils. Here, we present the long-term effects (11 years) of altered precipitation patterns (increased intra-annual variability in the growing season) and warming (1 °C year-round) on plant community composition and aboveground net primary productivity (ANPP), a key measure of ecosystem functioning in mesic tallgrass prairie. Based on past results, we expected that increased precipitation variability and warming would have additive effects on both community composition and ANPP. Increased precipitation variability altered plant community composition and increased richness, with no effect on ANPP. In contrast, warming decreased ANPP via reduction in grass stems and biomass but had no effect on the plant community. Contrary to expectations, across all measured variables, precipitation and warming treatments had no interactive effects. While treatment interactions did not occur, each treatment did individually impact a different component of the ecosystem (i.e., community vs. function). Thus, different aspects of the ecosystem may be sensitive to different global change drivers in mesic grassland ecosystems.

Alpine plants in Australia are increasingly exposed to more frequent drought and heatwaves, with significant consequences for physiological stress responses. Acclimation is a critical feature that allows plants to improve tolerance to environmental extremes by directly altering their physiology or morphology. Yet it is unclear how plant performance, tolerance, and recovery are affected when heat and water stress co-occur, and whether prior exposure affects responses to subsequent climate extremes. We grew a common alpine grass species under high or low watering treatments for three weeks before exposure to either none, one, or two heat stress events. We determined photosynthetic heat and freezing tolerance (LT

During environmental change, invasive species may be favored by increased resource input or reduced resource use of the resident community. Plasticity in certain plant traits of invasive species may be one possible mechanism behind their ability to quickly exploit unused resources. We tested whether rainfall manipulations (severe drought, moderate drought, watering) alter the growth and reproductive success of the invasive annual Conyza canadensis, and if it translates into a change in the abundance of the species in a semiarid perennial grassland in Central Hungary. Overall, C. canadensis exhibited greater individual performance and higher abundance in drought plots than in control and watered plots. At individual level, plants showed the strongest response to moderate drought: they grew 2.5-times taller than in control and watered plots, and produced twice and 2.5-times more seeds than in watered and control plots, respectively. Reproductive phenology was advanced in response to rain exclusions. Although severe drought caused 40% mortality, the cumulative performance of C. canadensis, expressed as plot-level aboveground biomass, was consistently greater in severe drought plots than in control and watered plots throughout the 3 years of the study. The higher performance of C. canadensis in drought plots is most likely due to the decreased abundance and, thus, decreased competitive effect of previously dominant perennial grasses caused by the rain exclusions. We conclude that drier summers that suppress perennial grasses will favor this alien annual forb, and phenotypic plasticity in growth and reproduction may contribute to its invasion success. Electronic supplementary material The online version of this article (10.1007/s00442-020-04711-y) contains supplementary material, which is available to authorized users.

Recent observational evidence suggests that nighttime temperatures are increasing faster than daytime temperatures, while in some regions precipitation events are becoming less frequent and more intense. The combined ecological impacts of these climatic changes on crassulacean acid metabolism (CAM) plants and their interactions with other functional groups (i.e., grass communities) remain poorly understood. Here we developed a growth chamber experiment to investigate how two CAM-grass communities in desert ecosystems of the southwestern United States and northern Mexico respond to asymmetric warming and increasing rainfall variability. Grasses generally showed competitive advantages over CAM plants with increasing rainfall variability under ambient temperature conditions. In contrast, asymmetric warming caused mortality of both grass species (Bouteloua eriopoda and Bouteloua curtipendula) in both rainfall treatments due to enhanced drought stress. Grass mortality indirectly favored CAM plants even though the biomass of both CAM species Cylindropuntia imbricata and Opuntia phaeacantha significantly decreased. The stem's volume-to-surface ratio of C. imbricata was significantly higher in mixture than in monoculture under ambient temperature (both P 0.0014); however, the difference became insignificant under asymmetric warming (both P 0.1625), suggesting that warming weakens the negative effects of interspecific competition on CAM plant growth. Our findings suggest that while the increase in intra-annual rainfall variability enhances grass productivity, asymmetric warming may lead to grass mortality, thereby indirectly favoring the expansion of co-existing CAM plants. This study provides novel experimental evidence showing how the ongoing changes in global warming and rainfall variability affect CAM-grass growth and interactions in dryland ecosystems.

Climate warming and increased atmospheric nitrogen deposition are both predicted to alter the primary productivity of grass-dominated systems in the coming decades. In field experiments, while both factors can have substantial effects on productivity in the initial years, further changes can be delayed by lags in plant species composition responses. However, the effects of experiment age can be confounded by annual variability in environmental conditions, and a replicate experiment established at a later time is therefore needed to separate these effects. We added new warming and nitrogen plots to an ongoing (14 year-old) field experiment in a grass-dominated temperate old field to compare the short-term vs. long-term treatment effects on plant productivity and species composition, while controlling for interannual environmental variability. We predicted treatment effects on relative species abundances would be most pronounced in the old plots. Although treatment responses of productivity (specifically to N addition) were highest in the old plots in the first year, by the second year there were no interactions between treatment and plot age. Moreover, there were no plot age effects on plant species composition, which was associated with continued dominance of non-native grasses. Our results therefore suggest that despite initial increases in productivity in response to global change, dominance by non-native grasses can suppress the emergence of further long-term treatment effects on productivity by inhibiting the responses of other species.

Displacement of diverse native plant communities by low-diversity invasive communities is a global problem. In the western United States, the displacement of sagebrush-dominated communities by cheatgrass has increased since the 1920s. Restoration outcomes are poor, potentially due to soil alteration by cheatgrass. We explored the poorly understood role of plant-soil feedbacks in the dominance of cheatgrass in a greenhouse study where uninvaded sagebrush soils were conditioned with either cheatgrass, a native bunchgrass or sagebrush. Sagebrush seedlings were grown in the soils that remained following the removal of conditioning plants. We expected cheatgrass to strongly suppress sagebrush due to a change in belowground microbial communities, conspecifics to have neutral effects and the native bunchgrass to have intermediate effects as it coevolved with sagebrush but belongs to a different functional group. We assessed the effects of conditioning on sagebrush growth, tissue nutrients, and carbon allocation. We also characterized the abundance, diversity and community composition of root microbial associates. Cheatgrass strongly suppressed sagebrush growth at high and low conditioning densities, the native bunchgrass showed suppression at high conditioning densities only and conspecific effects were neutral. Tissue nutrients, amount of root colonization by soil fungi or root microbial community composition were not associated with these plant-soil feedbacks. Although we did not identify the precise mechanism, our results provide key evidence that rapid soil alteration by cheatgrass results in negative plant-soil feedbacks on sagebrush growth. These feedbacks likely contribute to cheatgrass dominance and the poor success of sagebrush restoration.

Increases in nitrogen (N) inputs to the biosphere can exacerbate the introduction and spread of invasive non-native plant species. Often, with elevated soil N levels, invasive plants establish and further enrich soil N pools, changing overall ecosystem function. This study examined the relationship between soil N cycling and an increasingly prevalent, invasive plant species, tall oatgrass (Arrhenatherum elatius subsp. elatius), in foothills ecosystems between the Colorado Rocky Mountains and the Denver-Boulder Metropolitan area-similar to many Western US grasslands and woodlands. It focused on investigating differences in soil N transformations, inorganic N pools, and vegetation characteristics across invaded and uninvaded plots at three sites in two seasons (summer and autumn). There was a statistically significant effect of invasion on rates of net N mineralization, but it was dependent on site and season (p = 0.046). Site had a statistically significant effect on soil moisture and aboveground biomass C:N (p 0.04). The interactions of invasion x site were statistically significant for ammonium pools (p 0.03). These findings suggest that A. elatius invasion can be associated with accelerated N cycling, but that the nature of the relationship differs by location and season in the foothills. More broadly, this study contributes to determining how the N cycle is shifting in grassland ecosystems subject to increasing pressures from anthropogenic change.

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.

Cover crops are increasingly being used in ecological restoration projects, and are hypothesized to facilitate establishment of sown species by reducing weed abundances without competing with the target mix. We tested these predictions and examined the role of cover crop species on later species composition and diversity using cover crop seed treatments. Treatments included a fall seeding of one annual (Raphanus sativus or Avena sativa), one biennial (Oenothera biennis), one perennial species (Elymus canadensis), two grass-forb species combinations, or nothing as a control. All plots received the same diverse tallgrass prairie seed mix in March of the following year. Plant communities were sampled through five growing seasons. We found that cover crop treatments influenced community assembly, and that cover crop species varied in their effectiveness at reducing weed abundances, with the perennial grass E. canadensis being especially effective at reducing weeds. After 5 years, treatments were neutral in their effects on species diversity. However, composition of establishing plants (not including the cover crop) differed significantly among cover crop treatments on all sampling dates, indicating that treatments had long-term effects. Plots containing the C

Changing precipitation regimes can profoundly affect plant growth in terrestrial ecosystems, especially in arid and semi-arid regions. However, how changing precipitation, especially extreme precipitation events, alters plant diversity and community composition is still poorly understood. A 3-year field manipulative experiment with seven precipitation treatments, including - 60%, - 40%, - 20%, 0% (as a control), + 20%, + 40%, and + 60% of ambient growing-season precipitation, was conducted in a semi-arid steppe in the Mongolian Plateau. Results showed total plant community cover and forb cover were enhanced with increased precipitation and reduced under decreased precipitation, whereas grass cover was suppressed under the - 60% treatment only. Plant community and grass species richness were reduced by the - 60% treatment only. Moreover, our results demonstrated that total plant community cover was more sensitive to decreased than increased precipitation under normal and extreme precipitation change, and species richness was more sensitive to decreased than increased precipitation under extreme precipitation change. The community composition and low field water holding capacity may drive this asymmetric response. Accumulated changes in community cover may eventually lead to changes in species richness. However, compared to control, Shannon-Weiner index (H) did not respond to any precipitation treatment, and Pielou's evenness index (E) was reduced under the + 60% treatment across the 3 year, but not in each year. Thus, the findings suggest that plant biodiversity in the semi-arid steppe may have a strong resistance to precipitation pattern changes through adjusting its composition in a short term.

There is limited understanding of the combined effects of discrete climate extremes and chronic environmental changes on ecosystem processes and functioning. We assessed the interactions of extreme drought timing (45 days, in spring or summer) and nitrogen (N) addition in a full factorial field experiment in a Leymus chinensis-dominated meadow steppe in northeast China. We evaluated the resistance and recovery of the grassland (calculated in terms of aboveground biomass) to these two drought events. The spring drought reduced aboveground biomass by 28% in the unfertilized plots and by 33% in the fertilized plots, and the effects persisted during the subsequent post-drought period within the same growing season; however, the summer drought had no significant influence on aboveground biomass. Although there were no significant interactive effects between drought timing and N addition, we observed a potential trend of N addition increasing the proportion of aboveground biomass suppressed by spring drought but not summer drought. Moreover, the drought resistance of the aboveground biomass was positively correlated with the response of the belowground biomass to drought. One year after the extreme drought events, the spring drought effects on aboveground and belowground biomass were negligible. Our results indicate that the drought sensitivity of productivity likely depends on the phenological and morphological traits of the single highly dominant species (Leymus chinensis) in this meadow steppe.

Phylogenetic analyses are essential for disentangling how environmental filtering and competition determine species coexistence across spatial scales. Inner Mongolia steppe has strong environmental gradients, but how the phylogenetic relatedness of co-occurring species and phylogenetic signals of functional traits change across spatial scales remains unclear. We investigated the phylogenetic structure of grass assemblages along environmental gradients from regional to local scales, and measured functional traits within assemblages. We compared phylogenetic signals of plant traits between the same numbers of species randomly selected from the regional pool and species observed at the local scale, did phylogenetic principal component analysis to infer the main factors driving species coexistence, and examined the key plant trait-environment relationships across the phylogeny to reveal ecological adaptation mechanisms. Regionally, grass species were phylogenetically clustered with contrasting climate preferences. With decreasing spatial scales, species richness declined, changing from phylogenetically clustered to overdispersed, and phylogenetic signals of plant traits became weaker. At the local scale, grass assemblages were structured by soil water content and neighbor density, and the trait-environment relationships were less clear than those at the regional scale. This study demonstrated that at smaller scales, co-occurring grass species in the steppe tended to be more phylogenetically overdispersed, and that phylogenetic signals of plant functional traits became weaker with increasing abiotic and biotic interactions. Our findings contributed evidence for understanding species coexistence and maintenance at scales spanning regional to local communities in the East Asia steppe biome.

Exotic species are often predicted to successfully invade when their functional traits differ from species in recipient communities. Many studies have related trait differences among natives and invaders to competitive outcomes. Few studies, however, have tested whether functionally similar invaders have similar competitive impacts on natives. We investigated interactions in communities of a native annual forb Waitzia acuminata (Asteraceae) and two invasive annual grasses that are ecologically similar and co-occur in southwestern Australia. Using a combination of field and laboratory experiments and several performance measures, we assessed impacts of these grasses on W. acuminata. We also examined differences among species in their responses to intraspecific versus interspecific competition, including their frequency dependence. The two similar exotic grasses differed in interaction impacts, with one facilitating and the other suppressing the native. In general, intraspecific competition was stronger than interspecific competition for the native, while evidence of competition was weak for the exotics. These patterns may reflect that W. acuminata does well in these communities due to the combined impacts of stabilization and facilitation, whereas the exotics benefit from limited stabilization (mediated by their weak intraspecific competition) or weak interspecific competition with W. acuminata. We found divergent impacts of the exotic species despite their similar functional traits. We demonstrate that a native species may benefit from interactions with an exotic "benefactor" species, highlighting the potential importance of positive interactions in invaded communities. Our findings underscore the necessity of considering neutral and positive interactions in addition to competition in understanding invasion dynamics in real plant communities.

Priority effects provide an advantage to early establishing species and are thought to significantly affect the course of succession. We conducted a 20-year long experiment sowing high- and low-diversity mixtures in an ex-arable field. We ask how long the effect of sowing persists and which sown species affect the course of succession. The experiment was established in the Czech Republic in five replicate blocks, each containing three random 10 × 10 m plots with three treatments: natural colonisation, sowing low- and high-diversity seed mixtures. The species cover was annually estimated in 12 permanent 1 m2 quadrates within each plot. To identify the effects of sowing, we used an innovative method analysing the data separately for each year using Redundancy analysis (RDA) with identity of sown species as explanatory variables. In the first year, the effect of sowing was small; the peak of explained variability occurred between third and fifth year. The legacy of sowing was detectable in the natural colonisers for 18 years and in the sown species for the whole 20-year period. For some species, the difference between the plots where they were and were not sown remained significant for the whole 20-year period (e.g. Lathyrus pratensis) although the plots were adjacent and the area was mown with the same machine. Other ones (e.g. Trisetum flavescens) colonised all the plots evenly. The long-lasting effect of the initial sowing confirms contingency of successional pathway on the propagule pressure in the time of start of succession due to the priority effects.

Increasing availability of reactive nitrogen (N) threatens plant diversity in diverse ecosystems. While there is mounting evidence for the negative impacts of N deposition on one component of diversity, species richness, we know little about its effects on another one, species evenness. It is suspected that ecosystem management practice that removes nitrogen from the ecosystem, such as hay-harvesting by mowing in grasslands, would mitigate the negative impacts of N deposition on plant diversity. However, empirical evidence is scarce. Here, we reported the main and interactive effects of N deposition and mowing on plant diversity in a temperate meadow steppe with 4-year data from a field experiment within which multi-level N addition rates and multiple N compounds are considered. Across all the types of N compounds, species richness and evenness significantly decreased with the increases of N addition rate, which was mainly caused by the growth of a tall rhizomatous grass, Leymus chinensis. Such negative impacts of N addition were accumulating with time. Mowing significantly reduced the dominance of L. chinensis, and mitigated the negative impacts of N deposition on species evenness. We present robust evidence that N deposition threatened biodiversity by reducing both species richness and evenness, a process which could be alleviated by mowing. Our results highlight the changes of species evenness in driving the negative impacts of N deposition on plant diversity and the role of mowing in mediating such negative impacts of N deposition.

Forecasting the effects of climate change on the distribution of invasive species can be difficult, because invaders often thrive under novel physical conditions and biotic interactions that differ from those in their native range. In this study, we experimentally examined how rising temperatures and sand burial could alter the abundance and biotic interactions of two invasive beachgrasses, Ammophila arenaria and A. breviligulata, along the U.S. Pacific Northwest coast. We asked whether the current geographic ranges of the two congeners, and thus their effects on dune morphology and coastal ecosystem services, might shift as a consequence of climate driven changes in warming and sand supply. Our results show that A. breviligulata had lower biomass and tiller production when exposed to warming and high rates of sand burial, while A. arenaria showed neutral or positive responses to those treatments. Nevertheless, under all experimental combinations, A. breviligulata had strong negative effects on A. arenaria, while A. arenaria had weaker effects on A. breviligulata. Our models predict that although A. breviligulata mostly excludes A. arenaria, elevated temperatures and high rates of sand burial also increase the likelihood of species coexistence. We suggest that under climate change, the differences in physiological tolerance and the mediation of species interactions could expand the northern distributional limit of A. arenaria but restrict the southern limit of A. breviligulata. Moreover, because beachgrass abundance has direct effects on biophysical functions of dunes, reductions in vigor from warming could alter coastal protection, biodiversity, and carbon sequestration.

The presence of invasive species reduces the growth and performance of native species; however, the linear or non-linear relationships between invasive abundance and native population declines are less often studied. We examine how the amount and spatial distribution of experimental N deposition influences the relationship between non-native, invasive annual grass abundance (Bromus hordeaceus and Bromus diandrus) and a dominant, native perennial grass species (Stipa pulchra) in California. We hypothesized that native populations would decline as invasion increased, and that high nitrogen availability would cause native species to decline at lower invasion levels. We predicted that the rate of population decline would be slower in heterogeneous, compared to homogeneous, environments. We employed a field experiment that manipulated the amount and spatial heterogeneity of N addition across a range of invasive/native-dominated communities. There were strong negative and non-linear associations between level of invasion and S. pulchra proportional change (PC). Stipa pulchra PC was more negative and seedling survival was lower when N was added, and the negative effects of N addition on PC became larger in the final year of the study when S. pulchra had the largest declines. There was not strong evidence showing reduced competition in heterogeneous, compared to homogeneous, N treatments. Soil moisture was similar between S. pulchra and B. hordeaceus plots under ambient N, but B. hordeaceus under added N reduced soil moisture. Under N addition, Bromus spp. take up N earlier, reduce soil moisture, and create dry conditions in which S. pulchra declines.

Strictly vertically transmitted (hereditary) Epichloë spp. fungal endophytes are symbionts with cool-season pooid host grasses. Such endophytes may increase host invasiveness in the non-native, introduced ranges. However, because costs and benefits for the host can vary with the growing conditions, the endophyte may become locally or temporally extinct when costs outweigh benefits. Our long-term field experiment involved the introduction of seven Schedonorus pratensis (meadow fescue) cultivars hosting Epichloë uncinata endophyte, which represent host-grass populations differing in genetic backgrounds and Epichloë infection frequencies, to an unmanaged old field. In the first 6 years, the host grasses persisted but did not become invasive in the plant community, regardless of their endophyte infection frequency. Subsequently, we hypothesized that increasing nutrient availability would decrease endophyte costs and thus increase the host's success and abundance. We fertilized half of the plots for four additional years and re-examined S. pratensis invasiveness. We predicted that increased nutrient availability would increase S. pratensis abundance and E. uncinata frequency and concentration, as well as decrease plant community diversity, relative to unfertilized plots. Fertilization increased endophyte concentrations in three low-endophyte host populations. However, E. uncinata did not enable S. pratensis populations to achieve high abundance or to reduce plant community diversity in the old field, with or without fertilization. Thus, nutrient availabililty and host invasiveness appear to be decoupled in this study system.

The cool season rainfall at our study site should favour C

High propagule availability compared to native species is often critical to invasion success, but it is unclear if this has contributed to invasions by African grasses in Neotropical savannas. We compared patterns of occurrence in the vegetation, seed rain and seed bank among African and native grasses in Cerrado sites in southeastern Brazil. In grasslands and savannas, we obtained the abundance of grasses in the vegetation, in the seed rain (monthly for one year) and in the seed bank (rainy and dry season), and assessed seed limitation and relationships among compartments. Invasive grasses showed low abundance in all compartments and high seed limitation in grasslands, where the seed bank and seed rain were dominated by small-seeded native grasses, but were at least as abundant as the natives in the seed bank and seed rain in savannas, mostly due to high abundance of Melinis minutiflora at these compartments. Native grasses dispersal occurred in the rainy season, whereas invasive grass dispersal occurred from mid rainy to mid dry season (Urochloa decumbens) and in the dry season (M. minutiflora). Melinis minutiflora showed a more persistent seed bank than U. decumbens and natives in savannas. Abundance of invasive and most of the native grasses in the vegetation was positively related to their abundance in the seed rain. Differences in seed production, the timing of seed dispersal and seed bank persistence compared to native grasses seem to favour invasive African grasses in the Cerrado, but this role may differ between grasslands and savannas.

Soil respiration, the major pathway for ecosystem carbon (C) loss, has the potential to enter a positive feedback loop with the atmospheric CO

Understanding the processing of limiting nutrients among organisms is an important goal of community ecology. Less known is how human disturbances may alter the stoichiometric patterns among organisms from different trophic levels within communities. Here, we investigated how livestock grazing affects the C:N:P ecological stoichiometry of soils, plants (Leymus chinensis), and grasshoppers (Euchorthippus spp.) in a semi-arid grassland in northeastern China. We found that grazing significantly enhanced soil available N and leaf N content of the dominant L. chinensis grass by 15% and 20%, respectively. Grazing also reduced (soluble) C:N of L. chinensis leaves by 22%. However, grazing did not affect total C, N, or P contents nor their ratios in Euchorthippus grasshoppers. Our results reveal that the effects of grazing disturbances on elemental composition attenuated from lower to higher trophic levels. These findings support the theory that organisms from higher trophic levels have relatively stronger stoichiometric homeostasis compared to those from lower trophic levels. Moreover, grasshopper abundance dropped by 66% in the grazed areas, and they reduced the feeding time on their host L. chinensis grass by 43%, presumably to limit the intake of excess nitrogen from host plants. The energetic costs associated with the maintenance of elemental homeostasis likely reduced grasshopper individual performance and population abundance in the grazed areas. A comprehensive investigation of stoichiometric properties of organisms across trophic levels may enable a better understanding of the nature of species interactions, and facilitate predictions of the consequences of future environmental changes for a community organization.

Differentiation in physiological activity is a critical component of resource partitioning in resource-limited environments. For example, it is crucial to understand how plant physiological performance varies through time for different functional groups to forecast how terrestrial ecosystems will respond to change. Here, we tracked the seasonal progress of 13 plant species representing C3 shrub, perennial C3 and C4 grass, and annual forb functional groups of the Colorado Plateau, USA. We tested for differences in carbon assimilation strategies and how photosynthetic rates related to recent, seasonal, and annual precipitation and temperature variables. Despite seasonal shifts in species presence and activity, we found small differences in seasonally weighted annual photosynthetic rates among groups. However, differences in the timing of maximum assimilation (Anet) were strongly functional group-dependent. C3 shrubs employed a relatively consistent, low carbon capture strategy and maintained activity year-round but switched to a rapid growth strategy in response to recent climate conditions. In contrast, grasses maintained higher carbon capture during spring months when all perennials had maximum photosynthetic rates, but grasses were dormant during months when shrubs remained active. Perennial grass Anet rates were explained in part by precipitation accumulated during the preceding year and average maximum temperatures during the past 48 h, a result opposite to shrubs. These results lend insight into diverse physiological strategies and their connections to climate, and also point to the potential for shrubs to increase in abundance in response to increased climatic variability in drylands, given shrubs’ ability to respond rapidly to changing conditions.

Understanding how global change drivers (GCDs) affect aboveground net primary production (ANPP) through time is essential to predicting the reliability and maintenance of ecosystem function and services in the future. While GCDs, such as drought, warming and elevated nutrients, are known to affect mean ANPP, less is known about how they affect inter-annual variability in ANPP. We examined 27 global change experiments located in 11 different herbaceous ecosystems that varied in both abiotic and biotic conditions, to investigate changes in the mean and temporal variability of ANPP (measured as the coefficient of variation) in response to different GCD manipulations, including resource additions, warming, and irrigation. From this comprehensive data synthesis, we found that GCD treatments increased mean ANPP. However, GCD manipulations both increased and decreased temporal variability of ANPP (24% of comparisons), with no net effect overall. These inconsistent effects on temporal variation in ANPP can, in part, be attributed to site characteristics, such as mean annual precipitation and temperature as well as plant community evenness. For example, decreases in temporal variability in ANPP with the GCD treatments occurred in wetter and warmer sites with lower plant community evenness. Further, the addition of several nutrients simultaneously increased the sensitivity of ANPP to interannual variation in precipitation. Based on this analysis, we expect that GCDs will likely affect the magnitude more than the reliability over time of ecosystem production in the future.

Variation in color pattern between populations of cryptic animals is common and typically attributed to selection pressures from visual predators combined with variation in substrate composition. However, little is known about how cryptic color pattern relates to varied rates of predation, and few studies simultaneously analyze patterns of escape behavior and microhabitat use along with variation in color pattern, even though these traits evolve in tandem. Here, we use a combination of calibrated photographs and spectrometry to examine the influence of spatial heterogeneity in rates of predation on dorsal brightness in the Florida scrub lizard (Sceloporus woodi), a cryptic and sexually dimorphic species. Simultaneously, we analyze patterns of escape behavior and microhabitat use measured in the field. The results of this study indicate that populations inhabiting environments of increased predation have less color variation and more closely match the color of local substrate than populations sampled in environments of relaxed predation. Populations exposed to increased predation also show more pronounced escape behavior and are more selective in their use of microhabitat. Interestingly, geographic variation of dorsal brightness, escape behavior, and microhabitat use were greater for females than for males. Our results not only provide empirical evidence for theories of adaptive coloration, but suggest that sexual dichromatism can be maintained by selection pressures related to predation.

Ecosystems in the southwestern U.S. are predicted to experience continued warming and drying trends of the early twenty-first century. Climate change can shift the balance between grass and woody plant abundance in these water-limited systems, which has large implications for biodiversity and ecosystem processes. However, variability in topo-edaphic conditions, notably soil texture and depth, confound efforts to quantify specific climatic controls over grass vs. shrub dominance. Here, we utilized weather records and a mechanistic soil water model to identify the timing and depth at which soil moisture related most strongly to the balance between grass and shrub dominance in the southern Colorado Plateau. Shrubs dominate where there is high soil moisture availability during winter, and where temperature is more seasonally variable, while grasses are favored where moisture is available during summer. Climate change projections indicate consistent increases in mean temperature and seasonal temperature variability for all sites, but predictions for summer and winter soil moisture vary across sites. Together, these changes in temperature and soil moisture are expected to shift the balance towards increasing shrub dominance across the region. These patterns are strongly driven by changes in temperature, which either enhance or overwhelm effects of changes in soil moisture across sites. This approach, which incorporates local, edaphic factors at sites protected from disturbance, improves understanding of climate change impacts on grass vs. shrub abundance and may be useful in other dryland regions with high edaphic and climatic heterogeneity.

Climate change will alter global precipitation patterns, making it increasingly important that we understand how ecosystems will be impacted by more frequent and severe droughts. Yet most drought studies examine a single, within-season drought, and we know relatively little about the impacts of multiple droughts that occur within a single growing season. This distinction is important because many plant species are able to acclimate physiologically, such that the effects of multiple droughts on ecosystem function deviate significantly from the effects of cumulative, independent droughts. Unfortunately, we know relatively little about the ability of dominant species to acclimate to drought in drought-sensitive ecosystems like semi-arid grasslands. Here, we tested for physiological acclimation to multiple drought events in two dominant shortgrass steppe species: Bouteloua gracilis (C4) and Elymus elymoides (C3). Neither species exhibited physiological acclimation to drought; leaf water potential, stomatal conductance, and photosynthesis rates were all similarly affected by a single, late period drought and a second, late period drought. Biomass was lowest in plants exposed to two droughts, but this is likely due to the cumulative effects of both an early and late period drought. Our results suggest that late period droughts do exert weaker effects on biomass production of two dominant shortgrass species, but that the weaker effects are due to ontogenetic changes in plant physiology as opposed to physiological acclimation against multiple droughts. As a consequence, current ecosystem models that incorporate grass phenology and seasonal physiology should provide accurate predictions of primary production under future climates.

Climatic patterns are expected to become more extreme, with changes in precipitation characterized by heavier rainfall and prolonged dry periods. Yet, most studies focus on persistent moderate changes in precipitation, limiting our understanding of how ecosystems will function in the future. We examined the effects of extreme changes in precipitation on leaf-level and ecosystem CO2 and H2O exchange of three native C4 bunchgrasses (Andropogon gerardii, Panicum virgatum, and Sorghastrum nutans) over 3 years. Grasses were grown in three precipitation treatments: extreme dry, mean, and extreme wet based on historical rainfall records. After 3 years, plants were 45% smaller in the extreme dry treatment relative to the mean and extreme high treatment, which did not differ. We also found that an extreme decrease in precipitation caused reductions of 55, 40, and 40% in leaf-level photosynthesis (Anet), stomatal conductance (gs), and water use efficiency (WUE), respectively. Extreme increases in precipitation inhibited leaf-level WUE, with a 44% reduction relative to the mean treatment. At the ecosystem level, both an extreme increase and decrease in precipitation reduced net CO2 and water fluxes relative to plants grown with mean levels of precipitation. Net water fluxes (ET) were reduced by an average of 74% in the extreme dry and extreme wet treatment relative to mean treatment; net carbon fluxes followed a similar trend, with average reductions of 68% (NEE) and 100% (Re). Unlike moderate climate change, extreme increases in precipitation may be just as detrimental as extreme decreases in precipitation in shifting grassland physiology.

Interactions with resident species can affect the rate that expanding species invade novel areas. These interactions can be antagonistic (biotic resistance), where resident species hinder invasive establishment, or facilitative (biotic assistance), where residents promote invasive establishment. The predominance of resistance or assistance could vary with the abiotic context. We examined how the effects of a resident ecosystem engineer interact with abiotic stress to resist or assist the establishment of an expanding competitor. In Florida salt marshes, native cordgrass, Spartina alterniflora, is an influential ecosystem engineer that, when dead, exerts a legacy effect by forming persistent wrack patches. We examined how the legacy effect of Spartina wrack varies with spatial context and abiotic conditions to influence establishment of the northward-expanding black mangrove, Avicennia germinans. Field surveys documented that Spartina wrack and Avicennia propagules co-occur in the high intertidal zone, and we conducted two outdoor mesocosm experiments to investigate this association. Wrack positively affected propagule establishment when propagules were beneath wrack, but negatively affected establishment when propagules were above wrack. The abiotic tidal regime influences the magnitude of wrack effects by controlling ambient moisture, and the positive and negative effects of wrack were stronger in low moisture conditions that simulated desiccation stress during harsh neap tides. Thus, the same resident engineer can either resist or assist an expanding competitor and the magnitude of these effects depends on abiotic conditions. We propose that under harsh conditions, there is greater scope for an engineer's mediating influence to affect associated species, both positively and negatively.

Climate change and consumer outbreaks are driving ecosystem collapse worldwide. Although much research has demonstrated that these factors can interact, how heterogeneity in top-down control intensity and physical forcing modulates ecosystem resilience to climate stress remains poorly understood. Here, we explore whether the nocturnal herbivorous crab Sesarma reticulatum can control spatially dominant cordgrass (Spartina alterniflora) growth and how its top-down effects vary with crab density, drought stress, and large-scale disturbance in southeastern US salt marshes. In multiple field experiments and surveys, we show that Sesarma depresses cordgrass growth and that its effects increase in a saturating manner with increasing crab density, such that the highest naturally occurring densities of this consumer can trigger local cordgrass die-off. This top-down effect of Sesarma is similar in magnitude to what is thought to be the dominant grazer in the system, the marsh periwinkle snail Littoraria irrorata. In a drought stress by Sesarma density experiment, we further show that salinity stress and intensive crab herbivory additively suppress cordgrass drought resistance. After drought subsides, surveys and experiments reveal that Sesarma also stifles cordgrass re-growth into existing die-off areas. Together, these results show that multiple grazers powerfully regulate the productivity and drought resilience of these intertidal grasslands and that heterogeneity in physical stress and consumer density can dictate when and where top-down forcing is important. More generally, this work provides a rare, experimental demonstration of the critical role top-down control can play across the initiation and recovery stages of ecosystem die-off.